Orchids of Britain and Ireland PDF

Orchids
of Britain and Ireland
A Field and Site Guide
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For Rita

Orchids
of Britain and Ireland
A Field and Site Guide

Second Edition
Anne and Simon Harrap
A & C BLACK
LOndOn

ACKNOWLEDGEMENTS
We would especially like to thank Dr Tony Leech for much wisdom as well as practical help and advice (he read and re-read
most of the book), and Lenny Thomson for her equally good-humoured helping hand. Julie Dando very, very patiently per-
formed her design alchemy and has worked wonders, and at A&C Black Nigel Redman can claim the credit for initiating the
project and provided help and support throughout, while Jim Martin exercised his considerable editorial skills. For the pro-
vision of additional photographs we would like to thank Richard Bateman, Robin Chittenden www.harlequinpictures.co.uk,
Sean Cole, Bob Gibbons/Natural Image, Richard Gulliver, Michael Frost, Nigel Redman, Craig Robson, Paul Sterry/Nature
Photographers Ltd and Peter Wakely (English Nature). We also thank Richard Millington for his delightful illustration of Bee
Orchid varieties. We are very grateful to the Botanical Society of the British Isles and CEH Monks Wood for allowing us
reproduce the maps contained in the recent New Atlas of the British and Irish Flora (Preston et al. 2002), and for facilitating
their reproduction here; we would particularly like to thank Henry Arnold, the Database Manager, for his time and trouble.
Mark Van Beirs assisted with French translations, Keith and Karen Ashby with German, and Gareth Burnell with Latin. For
providing references we thank the Library of the Royal Botanic Gardens, Kew and also Jan Gebbie (Scottish Natural Heritage)
and Joan Chapman (Glasgow Natural History Society). The staff at Holt Public Library in Norfolk assisted cheerfully with
our many requests. Regarding matters botanical we especially thank Richard Bateman for answering many queries regarding
taxonomy and nomenclature and for providing advance copies of papers; similarly Pete Hollingsworth assisted with our
research on helleborines. Mavis and Richard Gulliver gave advice and unpublished material on Irish Lady’s Tresses and Richard
Hedley provided much information on Sword-leaved Helleborine (and suggested a translation for Cephalanthera), as did
David Ball (Hampshire County Council). Plantlife International kindly provided copies of their Back from the Brink dossier
on Sword-leaved Helleborine and Reg Land advised on Fen Orchid. Sean Cole supplied much information on Ghost Orchid
as well as other species. Jill Sutcliffe (English Nature) kindly read through all the proofs and made many useful suggestions.
We would also like to thank Richard Abbott, Steve Alton, Debbie Allan, Norman Baldock, Ian Bonner, Janet
Canning, David Carrington, Clive Chatteris, Robin Chittenden, Peter Clarke, Sean Cole, Les Colley, Richard Collingridge,
David Cottridge, Phil Davey, Charles David, Ian Denholm, Tim Dixon, Stuart Dunlop, Bob Ellis, Tom Ennis, Rachael
Fickweiler, Michael Foley, Maggie Gamble, Paul Hackman, Frank Hunt, James R. Hunter, Kevan Joynes, Brian Laney, Mandy
Leivers, Ian Livermore, Alan Lewis, Alex Lockton, Tim McGrath, Steve Madge, Stephen Martin, Nigel Milbourne, Dr E.
Charles Nelson, Malcolm Ogilvie, John Oxenford, Gavin Peplow, Michael Parsons, E. G. Philip, Chris Pogson, Rachel
Remnant, A. J. Richards, Craig Robson, Dr Francis Rose, Martin Sanford, Brendan Sayers, M.-A. Selosse, Alan Showler, Terry
Smith, Ron Stanbridge, Jon and Lucinda Starling, Malcolm Storey, Giles Strother, Ian Taylor, Peter Thompson, Chris Sydes,
Darrell Watts, Dave White, Steve Whitehouse, Martin Woodcock and Julian Woodman.
Of course, we take responsibility for any errors in the text and would be delighted to receive comments and
corrections. We can be contacted via our website: www.norfolknature.co.uk, or via the publishers. For assistance in the prepa-
ration of the second edition we would like to thank Richard Bosanquet, Tom Ennis, John Knowler, Geoff Musker and Andy
Scobie, but especially Richard Bateman and Sean Cole for their continuing support and for answering many questions, and
Roland de la Moussaye for facilitating a visit to North Uist.
Published in 2005 by A&C Black Publishers Ltd,

36 Soho Square, London W1D 3QY
www.acblack.com
Second edition published in 2009; electronic edition 2010
Copyright © 2005 and 2009 Anne and Simon Harrap
Photographs © 2005 and 2009 Simon Harrap except where indicated otherwise
ISBN 978-1-408-10571-9 eISBN 978-1-4081-3323-1
A CIP catalogue record for this book is available from the British Library
All rights reserved. No part of this publication may be reproduced or used in any form or by any means – photographic,
electronic or mechanical, including photocopying, recording, taping or information storage and retrieval systems –
without permission of the publishers. This book is produced using paper that is made from wood grown in managed, sus-
tainable forests. It is natural, renewable and recyclable. The logging and manufacturing processes conform to the environ-
mental regulations of the country of origin.
Printed in China by Lion Productions Ltd.

Designed by Fluke Art, Cornwall
10 9 8 7 6 5 4 3 2 1
Contents
INTRODUCTION 7
ORCHID BIOLOGY 8
ORCHID HABITATS 12
ORCHID CONSERVATION 18
FIELD GUIDE
Notes on the species accounts 20
Species Accounts
Lady’s-slipper Cypripedium calceolus 25
Red Helleborine Cephalanthera rubra 34
Sword-leaved Helleborine Cephalanthera longifolia 41
White Helleborine Cephalanthera damasonium 48
Lesser Twayblade Neottia cordata 56
Common Twayblade Neottia ovata 62
Bird’s-nest Orchid Neottia nidus-avis 67
Marsh Helleborine Epipactis palustris 78
Dark-red Helleborine Epipactis atrorubens 85
Broad-leaved Helleborine Epipactis helleborine 92
Violet Helleborine Epipactis purpurata 101
Narrow-lipped Helleborine Epipactis leptochila 107
Dune Helleborine Epipactis dunensis 113
Lindisfarne Helleborine Epipactis sancta 122
Green-flowered Helleborine Epipactis phyllanthes 126
Ghost Orchid Epipogium aphyllum 136
Fen Orchid Liparis loeselii 142
Bog Orchid Hammarbya paludosa 152
Coralroot Orchid Corallorhiza trifida 160
Creeping Lady’s-tresses Goodyera repens 166
Irish Lady’s-tresses Spiranthes romanzoffiana 174
Autumn Lady’s-tresses Spiranthes spiralis 181
Summer Lady’s-tresses Spiranthes aestivalis 188
Musk Orchid Herminium monorchis 193
Man Orchid Orchis anthropophora 202
Monkey Orchid Orchis simia 207
Military Orchid Orchis militaris 214
Lady Orchid Orchis purpurea 222
Early Purple Orchid Orchis mascula 228
Small White Orchid Pseudorchis albida 236
Lesser Butterfly Orchid Platanthera bifolia 243
Greater Butterfly Orchid Platanthera chlorantha 249
Common Fragrant Orchid Gymnadenia conopsea 256
Heath Fragrant Orchid Gymnadenia borealis 263
Marsh Fragrant Orchid Gymnadenia densiflora 267
Early Marsh Orchid Dactylorhiza incarnata 276

Frog Orchid Dactylorhiza viridis 288
Common Spotted Orchid Dactylorhiza fuchsii 294
Heath Spotted Orchid Dactylorhiza maculata 303
Southern Marsh Orchid Dactylorhiza praetermissa 309
Pugsley’s Marsh Orchid Dactylorhiza traunsteinerioides 316
Hebridean Marsh Orchid Dactylorhiza ebudensis 325
Northern Marsh Orchid Dactylorhiza purpurella 328
Irish Marsh Orchid Dactylorhiza occidentalis 334
Dense-flowered Orchid Neotinea maculata 341
Burnt Orchid Neotinea ustulata 345
Lizard Orchid Himantoglossum hircinum 354
Loose-flowered Orchid Anacamptis laxiflora 363
Pyramidal Orchid Anacamptis pyramidalis 366
Green-winged Orchid Anacamptis morio 372
Small-flowered Tongue Orchid Serapias parviflora 381
Greater Tongue Orchid Serapias lingua 384
Fly Orchid Ophrys insectifera 392
Bee Orchid Ophrys apifera 398
Late Spider Orchid Ophrys fuciflora 405
Early Spider Orchid Ophrys sphegodes 411
Other species 416
Orchid family tree 417
Map of vice-counties 418
Orchid flowering periods 420
SITE GUIDE
Introduction to the site accounts 421
Useful addresses 422
Botanical organisations 423
Abbreviations 423
Site Accounts
Southeast England 424
Southwest England and Channel Islands 432
East Anglia 438
Central England 442
Northern England and Isle of Man 447
Wales 452
Scotland 456
Ireland 460
GLOSSARY 465
SOURCES OF INFORmATION AND BIBLIOGRApHY 469
INDEx OF SpECIES 479

InTROdUCTIOn l 7
Introduction
why this book?
Orchid! The very word conjures up an image of the exotic, beautiful and, above all, rare. It is associated
with wealth, power and the eminently desirable. Few people realise that orchids are very much part of
the natural heritage of Britain and Ireland, and that there are no fewer than 56 species of native wild
orchid. They range from the tiny, green Bog Orchid to the flamboyant Marsh Helleborine and the
gorgeous Green-winged Orchid. They also include Lady’s-slipper and Ghost Orchid, probably the
two rarest native wild flowers in the British Isles, and Common Spotted and Early Purple Orchids,
species found throughout the land.
The aim of this book is both to introduce wild orchids to a wider audience and to show those who
think they know about orchids that there is always more to learn. For orchids need friends. Despite
the many schemes designed to alleviate its effects, modern ‘industrial’ agriculture, combined with
urbanisation, insensitive forestry practices and a rain of nutrients from car exhausts, has resulted in an
ever-increasing homogenisation of the countryside. The beautiful, delicately woven tapestry of fields,
pastures, woods and marshes created unwittingly by the hand of man over many generations is being
put into the equivalent of a food blender that is reducing everything to a monotonous and anonymous
wildlife desert. In the face of this assault, the majority of wild plants are in retreat, including all
orchids. Do not be fooled by cheeky television presenters and glossy wildlife magazines. Although
there is now more money for conservation in general and nature reserves are at last receiving some
desperately needed cash, orchids and much other wildlife besides have disappeared from most of the
landscape. If we can help more people to enjoy, appreciate and value orchids and the places where they
grow, our purpose will have been served.
upper sepal
what is an orchid? petal petal
Orchids (the family Orchidaceae) are among lateral lateral
the most diverse groups of plants, with over sepal sepal
1000 genera and at least 25,000 species. Indeed,
the Orchidaceae is the largest and most highly-
evolved family of flowering plants. column
The family derives its name from the lip
Greek orchis, meaning ‘testicle’, a reference to
the appearance of the underground tubers of
some species. The term orchis was first used by
Theophrastus (c. 370-285 BC) in his Natural
History of Plants; he was a student of Aristotle pollinia
and is considered to be the ‘father’ of botany. pollen
masses rostellum
Most people would recognise an orchid, even
bursicle
those without any particular interest in botany or caudicle stigma
gardening – at least the gaudy, hot-house hybrids mouth of
viscidium
and some of the more colourful wild orchids. the spur
Giving a precise definition of an orchid is more pollinium (detail) column (detail)
difficult, especially a non-technical definition, but
European orchids all share the following features: M A typical orchid flower.
l They are perennial herbs rather than trees or shrubs and lack any woody parts.
l The leaves are ‘simple’ and not divided into lobes or smaller leaflets.
l The leaves have no stalk and are arranged alternately along the stem.
l The flowers are carried in a single spike at the tip of the stem.

8 l ORCHIdS OF BRITAIn And IRELAnd
l The ovary is inferior, that is, placed below the sepals and petals.
l Male and female parts of the flower, the stamens and the stigma, are not separate but are fused
together into a single structure, the column, that lies in the centre of the flower.
l The flower is made up of three sepals and three petals but one of the petals differs from the others,
usually significantly so, and forms the lip (sometimes known as the labellum). This is often
brightly coloured and patterned, and intricately shaped. The lip is actually the uppermost petal
but usually lies at the bottom of the flower because either the ovary or its stalk is twisted (the
flower is therefore resupinate).
Orchid Biology
orchids And fungi
The relationship that orchids have with fungi impacts on all aspects of their biology and, more than
anything else, defines them.
Their seeds require fungi in order to germinate and grow. The seedling spends months or years
underground and during this period is completely dependent on the nutrients that it obtains from
fungi and is mycotrophic (a term deriving from the Greek mukes meaning ‘fungus’ and trephein ‘to
feed’). Seed must, however, be produced in large quantities to ensure that some, at least, will find the
correct conditions for successful germination and growth, including the presence of the correct fungi.
In turn, the need to produce large quantities of seed has powered the evolution of elaborate flowers
and complex pollination mechanisms.
Even when the orchid has appeared above ground as an adult or near-adult plant and is able
to photosynthesise and manufacture its own carbohydrates, in many species it still maintains a
relationship with fungi. In a few orchids the adult plant continues to be entirely dependent on fungi
(the so-called saprophytic orchids). In others, it is probably largely independent of fungi and gains
its nutrients almost entirely from photosynthesis (these are phototrophic). Most orchids, however,
fall somewhere between these two extremes, with both sources of nutrition being utilised, perhaps
in varying proportions depending on the season. The ability to utilise two sources of nutrition allows
orchids to thrive in marginal habitats; some grow in heavy shade and many are found on poor soils.
In the tropics, orchids have extensively colonised the soil-less trunks and branches of trees and are
epiphytes. Finally, the ability to fall back on fungi as a source of nutrition explains why many orchids
are able to become ‘dormant’ underground for a year or sometimes longer.
mycorrhizas
It is thought that around 90% of the world’s plants have a relationship with fungi. Such a relationship
is known as a mycorrhiza and the fungi that form these attachments are known as mycorrhizal.
Mycorrhizal fungi live in the soil and, unable to manufacture their own carbohydrates by photosynthesis
or obtain sufficient for their needs by the decomposition of organic matter, they invade the root
systems of green plants. But, rather than being parasitic, the fungus actually benefits the host plant
by functioning as an extended root system. The mycelium of the fungus extends far into the soil and
is able to provide the plant with minerals, especially phosphorus; it may also confer some degree of
drought, pest or disease resistance. The plant in turn provides the fungus with carbohydrate that it
has produced through the process of photosynthesis; most plants are able to divert up to 20% of their
carbohydrate to fungi without coming to harm. The relationship between the plant and the fungus
is therefore mutualistic (such mutually beneficial relationships were once termed symbiotic, but this
term is now used for a wider range of interactions).

ORCHId BIOLOGY l 9
Fungi form several kinds of mycorrhiza:

l Ectomycorrhizal fungi form a sheath or mantle over the plant’s roots (the Greek ectos means
‘outside’). Ectomycorrhiza are the dominant type of mycorrhiza formed by forest trees and
are critical to their growth. Most ectomycorrhizal fungi are macrofungi and many produce
recognisable mushrooms.
l Endomycorrhizal fungi (also known as VAM fungi) penetrate the cells of the plant’s roots (the
Greek endon means ‘within’). They are microfungi and do not produce distinctive fruiting bodies.
Poorly-known, they cannot be seen without a microscope and have proved impossible to cultivate
and study in vitro.
l Ericaceous mycorrhiza are formed with the roots of various ericaceous plants.
l Orchidaceous mycorrhiza are formed with orchids.
orchid fungi
Until very recently, orchid mycorrhizas were thought to differ fundamentally from other mycorrhizal
systems in that the orchid does not provide the fungus with carbohydrates: this must be true of orchid
seedlings which develop entirely underground and have no green leaves. Rather, it is the fungus that
provides the orchid with energy; there is good evidence that carbohydrate obtained by fungi from the
decomposition of organic matter (or from trees, see below) is transferred to the orchid. In short, it
seemed that orchids are parasitic on fungi (or, to use the terminology of recent scientific papers, they
‘cheat’ in their relationship with the fungus). There was no evidence for the transfer of nutrients from
the orchid to the fungus, even in mature plants which are able to photosynthesise, but very recently it
was shown that carbon does pass from Creeping Lady’s-tresses to their associated fungi, and this has
re-opened the debate regarding the orchid-fungus relationship. Throughout the text we have placed
‘infection’ and ‘partner’ in inverted commas to emphasise uncertain nature of the relationship between
orchids and fungi.
The physical relationship between orchids and fungi is very sophisticated. Fungal hyphae pass
through the outer layers of the orchid’s root, rhizome or other underground organs and penetrate
the cell walls to form loops and coils, called pelotons. At intervals, the orchid digests these pelotons
and receives water, mineral salts, carbohydrate and other organic compounds from the fungus. So
sophisticated is the orchid’s use of fungi that it is able to control its spread and confine it to specialist
cells; indeed, some orchids produce phytoalexins which act as a fungicide and prevent the fungi from
reaching tubers and other storage organs. The main fungal associates of orchids are Basidiomycetes
of the Rhizoctonia group (other members of the Rhizoctonia group are soil saprotrophs or
pathogens).
‘saprophytic’ orchids
Some species of orchid take the relationship with fungi to an extreme. Bird’s-nest, Coralroot and
Ghost Orchids have no green leaves (or have the green pigments very much reduced) and throughout
their lives depend entirely on their fungal ‘partner’ for nutrition (they are fully mycotrophic).
Furthermore, these species do not form associations with the usual orchid fungi of the Rhizoctonia
group. It has been shown that both Bird’s-nest and Coralroot Orchids form relationships instead with
ectomycorrhizal fungi which are simultaneously in partnership with nearby trees. Via these fungi the
orchids acquire carbohydrates from the trees and therefore they are, in effect, parasitic on the trees.
It has been suggested that forming associations with such ectomycorrhizal fungi may provide the
orchid with a stabler and more reliable source of nutrients, which is particularly important when it
has no other source of nutrition.
Bird’s-nest, Coralroot and Ghost Orchids are frequently but incorrectly described as saprophytic.
Saprophytes (nowadays more properly known as saprotrophs) derive their nutrition from dead
organic matter; these orchids acquire nutrients from a living fungus.

woodland orchids
It has also been shown recently that some other orchids, such as Red, White and Broad-leaved
Helleborines, also form relationships with ectomycorrhizal fungi and are therefore able to utilise
nutrients provided unwittingly by nearby trees. This may explain their ability to thrive in low light
levels and to become ‘dormant’ underground for long periods. Many are particularly associated with
Beech trees.
fungi and rare orchids

There is mounting evidence that some orchids are very specific about their fungal ‘partners’. Various
orchids in the genera Liparis, Goodyera and Spiranthes have been shown to associate with just one species
of fungus and to use the same species, both as germinating seeds and as adult plants. Other orchids
form a relationship with many different fungi as adult plants but are very specific as germinating seeds
(and may associate as seeds with a different species of fungus to any of those used by adult plants). Yet
other orchids seem to use a broad range of fungi for both germination and as adults.
Orchids that require a specific fungus in order to germinate and grow may be very limited in where
they can live, compared to the more generalist orchids, and this has clear conservation implications.
We would venture to guess that some of Britain and Ireland’s orchids are rare and localised because
their fungal partners are rare and localised. This may well explain why, for example, Burnt Orchid
seldom colonises new sites. Any conservation programme, especially if it involves a reintroduction,
may ultimately be unsuccessful if it fails to take this into account.
orchid seeds
Orchid seeds are rather small and, indeed, are often known as ‘dust seeds’. They typically weigh 2-8
micrograms and, in British species, are 0.35-1.4mm in length. They are made up of a relatively simple
embryo enclosed in a hardened carapace and surrounded by the much larger testa, a honeycomb of
dead cell walls that traps air.
Small size confers several advantages. Large
numbers of seeds can be produced at relatively testa
little cost; in British orchids counts of between
376 and 25,000 seeds per capsule have been
recorded (Lesser Twayblade and Greater Butter- embryo
fly Orchid being the extremes). With many 0.5mm
air spaces the seeds are ideally suited to wind
dispersal and can travel long distances. They can M A typical orchid seed.
also float on water, another effective means of
dispersal. However, small size also imposes limitations. The seeds are so tiny that they contain very
little in the way of food reserves (merely a few lipids and proteins) and they depend entirely upon
fungi to provide their nutrients when they germinate.
germinAtion And underground growth

The orchid seed germinates 2-10cm below the surface of the soil. The seedlings are very vulnerable to
desiccation and require the presence of fungi, so it is important for the seed to avoid germinating on
the soil’s surface. To ensure this, almost all orchid seeds will only germinate in darkness, once rain has
washed them down into the soil or they have been covered by fallen leaves. Unlike other plants, they
do not require light following germination because they do not produce green leaves.
It is thought that most European orchids germinate in the spring (although there is only limited
direct evidence for this) and therefore the seeds must have a mechanism to keep them dormant over
the winter. Germination depends upon the uptake of water by the seed and the hard carapace forms
a barrier against water, slowing or preventing germination for a while. In some orchids the carapace is
incomplete and these seeds germinate much more easily and rapidly.

ORCHId BIOLOGY l 11
With its very limited reserves of nutrients, the orchid seed is dependent on fungi from the outset.
In some species, such as Bird’s-nest Orchid, seeds start to germinate before ‘infection’ by a fungus,
although the breaking of the seed’s dormancy almost certainly requires the fungus to be present in the
immediate vicinity of the seed, which perhaps responds to a chemical signal from the fungus. In other
species, such as Coralroot Orchid, the seeds will only start to germinate after it has been ‘infected’ by
the specific fungal ‘partner’. From the outset the seedling is able to control the extent of ‘infection’ by
the fungus, confining it to certain areas.
Some species, including Bird’s-nest and Coralroot Orchids, form relationships with a very
restricted range of fungi, perhaps just one species. Their seeds will only germinate in the presence
of the appropriate fungus. (The implication is that the seeds can remain dormant for relatively long
periods, ‘waiting’ for the correct fungus to appear.) Other orchids form a relationship with a variety of
common soil fungi that probably occur in all suitable habitats. Examples include the marsh orchids;
their seeds will germinate and start to grow in the absence of the appropriate fungi, presumably
because there is a very high probability that a suitable fungus will be encountered very soon.
Upon germination the seed forms a protocorm. This is a small, often parsnip- or top-shaped
structure with a scatter of root-hairs on its surface; these are single-celled projections that facilitate
fungal ‘infection’. The protocorm usually goes on to develop roots and at this stage is often known as
a mycorhizome. The primary function of these first roots is to host fungal activity rather then the
supply of nutrients and water. In most orchids, as the seedling continues to develop, fungal activity is
increasingly confined to the roots, and the mycorhizome, now free of fungal ‘infection’, is known as a
rhizome. In some orchids the adult plant grows from a rhizome, but in many species the rhizome is
largely replaced by tubers. In adult orchids, whether they grow from a rhizome or from a tuber, fungal
activity is usually confined to the roots and, in some species, to the slender extremities of the tubers.
PollinAtion
Orchids are renowned for the beauty and complexity of their flowers. These flowers have not, however,
evolved to amaze and delight us, but to fulfil the primary function of the plant, which is to reproduce
itself.
cross-pollination versus self-pollination

Most flowering plants reproduce sexually and the production of the next generation depends upon
the successful unification of male and female gametes to produce seed; the male gamete is pollen
and the female gametes are contained within the ovary. In most plants, including orchids, each
flower produces both male and female gametes. It would seem a simple process therefore for pollen
to be transferred within the same flower to the ovary. Indeed, why bother with the resources and
energy needed to produce a flower at all when a small drab structure would be sufficient to bring
pollen and ovary together? The answer lies in the advantages that cross-pollination brings to the
species.
If pollen and ovary come from different individuals they will each carry a different set of genetic
material and the resulting offspring will not, therefore, be genetically identical to its parents. This
continual mixing of genetic material creates a large ‘gene pool’ which gives the species flexibility and
adaptability. It also means that individual mutations are disseminated through the population rather
than being passed, unchanged, to the offspring; cross-pollinated species therefore tend to be relatively
uniform in appearance.
If a flower is pollinated with its own pollen (i.e. it is self-pollinated) its offspring will have limited
genetic variation. In turn, their offspring will be similarly limited. In self-pollinated species the overall
genetic variation is rather low and the ‘gene pool’ is limited, giving the species very little flexibility.
Paradoxically, however, self-pollinated orchids can show more variation between colonies and are
more likely to produce distinct varieties as mutations; these might produce, for example, differently
coloured flowers and are passed down unchanged to the next generation. Bee Orchid is perhaps the
best example of this.

There are circumstances where self-pollination is an advantage. If, for example, a single wind-
blown seed produced a flowering Lizard Orchid many miles from other plants of the same species, it
would have no chance of reproducing if it could only be cross-pollinated. If it is able to self-pollinate,
it can produce seed and reproduce itself. Not surprisingly, self-pollination tends to be commoner in
species that have scattered populations, or are at the edge of their range. Another factor may be a
lack of pollinators, and species adapted to deep shade may well benefit from self-pollination; White
Helleborine, a species which is routinely self-pollinated, is successful in Britain, whereas Red and
Sword-leaved Helleborines, which are always cross-pollinated, have declined as increasing shade has
made their woodland homes too dark for their pollinators. Self-pollination may therefore be a useful
strategy in the short-term and it may only be over an evolutionary time scale, involving thousands of
generations, that the advantages of cross-pollination come to the fore.
Most orchids are cross-pollinated but, rather cunningly, a large proportion have adaptations that
prevent self-pollination but then allow the flowers to self-pollinate if, after a few days, a suitable
pollinator has not come along. Conversely, the small number of routinely self-pollinated orchids
are, at least occasionally, cross-pollinated by insects. In the world of orchids, nothing is straight-
forward.
insects as pollinators
Orchid flowers have evolved to use insects to carry pollen from one plant to another and employ
a variety of mechanisms to attract suitable pollinators. Bright colours and scents advertise their
presence and the lip of the flower acts as a convenient landing platform. Many offer their insect
visitors a reward of nectar but a large proportion do not. Their bright colours and scents are instead
a ‘deceit’ and they rely on the stupidity of insects, which are slow to learn that the flowers offer no
reward. Fly, Early Spider and Late Spider Orchids (genus Ophrys), have evolved even more elaborate
mechanisms that take advantage of the sexual behaviour of insects (see p.388).
Compared to most other flowering plants, orchids produce large quantities of seeds. Therefore
large quantities of pollen have to be moved between the flowers because each pollen grain can only
produce one seed. In most orchids the pollen grains are amalgamated together in large numbers to
form pollinia. Due to their size and weight, these pollinia must be carried to other flowers by an insect
and must be very securely attached if the process is to be completed successfully. The insect must be of
the right size and shape, both to pick up the pollinium in the first place and to be positioned correctly
in the next flower so that the pollinium makes contact with the stigma and effects fertilisation.
European orchids are not usually pollinated with a whole pollinium, rather just fragments, and thus
a single pollinium can, provided it remains on the insect, pollinate several flowers. But, if too small
a quantity of pollen is deposited the capsule will mature without all the ovules being fertilised, and
there will be many non-viable seeds.
Orchid Habitats
Britain and Ireland form a landscape that has been tamed, with virtually no truly wild places; even
the moors and mountains of Wales and Scotland are not natural, having been stripped of their trees
many generations ago. Orchids therefore have to find a niche in the habitats which mankind has
moulded; some of these habitats are very rich in orchids whereas others are devoid of their grace and
beauty.
It is possible to identify five factors which determine the suitability of any habitat.
Soil chemistry: Many orchids are fussy about the soil chemistry and, as any gardener knows, whether
the soil is acid or alkaline determines to a great extent which plants will thrive. A high proportion of
orchids favour soils that are rich in calcium; these are known as calcareous soils and they are alkaline
with a high pH. Calcium is a major component of chalk and limestone and therefore soils derived

ORCHId HABITATS l 13
from these are often excellent for orchids. A similar chemistry is produced by magnesium and some
limestones are rich in this mineral; although they are not ‘calcareous’, the terms ‘alkaline’ or ‘base-rich’
can also be used to cover these soils. In practice, however, ‘calcareous’, ‘alkaline’ and ‘base-rich’ are used
interchangeably. A small minority of orchids favour acid conditions, usually soils that are damp or
wet, and these are usually to be found where peat is formed from bog mosses (Sphagnum spp.), or in
pinewoods.
Soil fertility: Like most wild plants, orchids do best in a soil that is very low in nutrients. This is
largely because a soil that is high in nutrients, either naturally or through the application of fertilisers,
allows a limited range of larger and more aggressive species to thrive, and these will swamp any
orchids. There is also evidence that some nutrients are actually toxic to some orchids (for example,
high levels of phosphorus may be toxic to Green-winged Orchid).
Age: Orchids are great colonists and their dust-seeds enable them to spread long distances.
Nevertheless, there are some unknown factors which limit the ability of many species to move into
new habitats. Colonisation often takes place very slowly, probably over hundreds of years for some
species. It follows that the older a habitat is, the more likely it is to support orchids and the greater
the diversity of species.
Stability: Orchids are relatively long-lived plants and most have a period of immaturity lasting several
years which they spend underground. Both as underground seedlings and as adult plants they cannot
tolerate ground disturbance which would destroy their rhizomes or tubers. Therefore orchids can
only occupy a habitat if it has a degree of permanence; ground that is regularly ploughed or otherwise
disturbed cannot support orchids.
Disturbance: Despite the preference of many species for older, undisturbed habitats, our impression
is that orchids are often found on areas that have been disturbed in the past, albeit a long time ago
in some cases. Old quarries and sand and chalk pits are good examples of this. It is also increasingly
obvious that the continuous, low-level ground disturbance produced by grazing animals is critical to
the long-term survival of many species.
deciduous woodland
Deciduous woodland can be rich in orchids but is, all too often, a disappointment. Woodland in
Britain and Ireland falls into two broad categories; ancient woodland and secondary woodland.
Ancient woodland is found on sites which have probably had an uninterrupted cover of trees
for thousands of years. Throughout Europe, only fragments of this primeval ‘wildwood’ remain and
Britain and Ireland are no exception, with just 1-2% of the original cover left. Man has continuously
been at work in these woodland fragments, which have been cut for timber, coppiced and grazed,
but the continuity of tree cover has allowed a rich flora and fauna to survive and an ancient wood
will undoubtedly have the greatest diversity of orchids. Such woods can usually be recognised on
the ground by their substantial boundary banks and diverse flora, and on a map will often have a
somewhat irregular outline; historical research, looking at old maps and documents, is often needed
to confirm their provenance, but this is not necessary for the orchid hunter.
Secondary woodland grows where the continuity of tree cover has been broken. If a field that
has been cultivated for decades or even centuries is planted with trees or allowed to ‘tumble down’
to woodland it may look like a wood but it will seldom, if ever, acquire the richness and diversity of
ancient woodland (hence the planting of new ‘community forests’ and ‘millennium woodlands’ will do
little for orchids). Many woodland plants and animals are very poor colonisers and are unable cross
stretches of inhospitable territory. Importantly, a secondary wood may well lack the complex soil
structure and diversity of soil fungi that is found in an ancient wood and thus lack suitable fungal
‘partners’ for some orchids. Overall, with the exception of a few species, secondary woodland will be
poor in orchids (and will often have none).
Management is another factor that can determine how orchid-rich a wood is, whether ancient
or secondary. Until the early 20th century most woodland was intensively managed. Many woods

were coppiced – cut in small blocks on a rotation of around ten years to provide timber for hand-
crafts, charcoal and firewood. Coppice stools grow again after each round of cutting, providing an
endless supply of timber. In coppiced woodland periods of sunny, well-lit conditions are followed
by increasing shade as the stools grow again, ideal for some orchids. Abundant labour also allowed
rides to be cut and cleared for access and the limited technology of the day meant that felling and
replanting for large timber was carried out at a relatively slow pace and in a piecemeal manner. All
in all, the more extensive management of this period resulted in woods that were a patchwork of
habitats and much sunnier due to the larger number of open rides and glades. Modern management
techniques depend on the use of minimal labour supplemented by machinery. Coppicing has largely
been abandoned (although now increasingly reinstated by conservation bodies), ride management is
negligible and felling is undertaken in large blocks using heavy machinery. Under such regimes woods
are darker than before and often suffer heavy ground disturbance.
A final factor to affect the suitability of woods is biological. There are ever increasing numbers
of deer in Britain; indeed, it is said that there are more deer now than at any time since William the
Conqueror. Deer eat orchids and some scarce species, for example Narrow-lipped Helleborine, can
lose all their flowers to deer year after year. Another increasingly negative factor, at least in some parts
of the country, is pheasant rearing. In a commercial pheasant-shoot large numbers of young birds are
released into a wood and these can decimate the flora as they scratch and root en masse.
In spite of the many factors that can negatively effect a wood’s suitability, there are still many
wonderful woods in Britain and Ireland, and the best are often reserves. Characteristic species of
ancient woodland include Common Twayblade, Common Spotted and Fly Orchids, and, especially in
coppiced woodland, Early Purple and Greater Butterfly Orchids. In Kent, Lady Orchid is widespread
and some favoured woods, from southern England to northern Scotland, hold the exquisite, sun-
loving Sword-leaved Helleborine. Two of our rarest orchids, Red Helleborine and Ghost Orchid, are
confined to deciduous woodland. The overall increase in the level of shade in modern woodland is not
bad for all orchids: Bird’s-nest Orchid and White, Violet and Narrow-lipped Helleborines can thrive
in very shady woods.
Secondary woodland and plantations of deciduous trees are likely to be much poorer in species,
although two of the more mobile species, Common Spotted Orchid and, in beech plantations, White
Helleborine, can occur in large numbers. The strangely fickle Broad-leaved Helleborine and the
enigmatic Green-flowered Helleborine seem to occur at random.
coniferous woodland
The only coniferous trees that are native to Britain and Ireland are Juniper, Yew and Scots Pine and it
is the last of these that forms a special habitat for orchids. Native Scots Pine is confined to Scotland,
where the remnants of the so-called ancient ‘Forest of Caledon’ are concentrated on Speyside and
Deeside in the east and around Beinn Eighe in Ross & Cromarty. These are home to Creeping Lady’s-
tresses and, more locally, Lesser Twayblade and Coralroot Orchid, with Heath Spotted Orchid in
some of the more open areas.
Conifer plantations have been established all over Britain and Ireland but are usually planted with
non-native species such as Sitka Spruce and Douglas Fir. They were often established on areas that had
been open ground, such as heaths, moorland and sand dunes, but in the post-war period the Forestry
Commission also had a policy of ‘coniferisation’ and converted large areas of ancient woodland into
conifer plantations. Plants, including orchids, are great survivors and where an ancient woodland has
been ‘coniferised’ there may sometimes be orchids in rides and along edges, with Common and Heath
Spotted Orchids and Broad-leaved Helleborine the likeliest species. Otherwise, conifer plantations
south of the Scottish border hold little interest with one notable exception: in parts of Norfolk and
northern England, mature pine plantations hold some impressive colonies of Creeping Lady’s-tresses.
In Scotland, plantations of pines may acquire a great deal of interest as they mature, taking on some
of the characteristics of native pine woods.

carr woodland
Carr woodland is dominated by willows and Alder and develops in the wet, waterlogged conditions
found on and around mires, bogs, rivers and lakes. These fast-growing trees are pioneer species and
in a natural system carr woodland usually eventually dries out and other species of tree become
dominant. Carr may hold orchids such as Common and Heath Spotted Orchids, ‘inherited’ from
the open ground it colonised, but it also has two specialities. In Scotland, Coralroot Orchid is
most abundant in carr and in southern England carr may be the ‘natural’ habitat of Green-flowered
Helleborine.
grassland
Grassland is a prime orchid habitat, with a large range of species possible, including Common and
Heath Spotted Orchids, Greater and Lesser Butterfly Orchids, Common Fragrant, Green-winged,
Early Purple, Pyramidal, Bee and Frog Orchids and Common Twayblade. Calcareous grassland,
especially the chalk downs of southern England, is the richest; the specialities are Musk, Burnt,
Man, Monkey, Early Spider and Late Spider Orchids and Autumn Lady’s-tresses. Neutral grassland
can also be productive and even acid grassland can hold species such as Heath Spotted and Heath
Fragrant Orchids and, in northern Britain, Small White Orchid.
Grassland will always tend to develop into scrub and eventually woodland unless prevented by
either grazing or mowing. It is possible to distinguish between a ‘pasture’, grassland which is grazed
for some or all of the year but not cut, and a ‘meadow’, grassland from which stock are excluded so
that it can be cut for hay (and often then grazed later in the year). The timing and intensity of grazing
and mowing is very important in determining which orchids are able to thrive and even the animals
concerned can be important; cattle are heavy beasts and can damage the turf but they are selective
eaters and do not eat orchids (although they may stand on them); sheep, lighter on their feet, eat
everything. Some grazing regimes produce a very short turf beloved of Musk and Frog Orchids and
Autumn Lady’s-tresses, while longer grass can support Pyramidal, Man and even Lizard Orchids.
It is always worth investigating the various micro-habitats in any area of grassland, which may
be favoured by different orchids. Steeper slopes are often particularly interesting, as are any ancient
earthworks, which may have remained undisturbed for centuries.
limestone pavements
This very specialised habitat is found widely in the north and west of Britain and in Ireland. The
limestone bedrock was first scoured flat by the ice sheets and then weathered by the action of water
and frost to produce blocks of rock (clints) with deep and sometimes treacherous crevices between
them (grykes). Over time, soil may accumulate in the grykes and most pavements would, in a natural
state, be wooded, but many were cleared of trees long ago and some are now rather bare. The range
of orchids present on a limestone pavement depends in part on how much naked rock is exposed;
many pavements have a substantial covering of turf and this holds all the species typical of calcareous
grassland. Limestone pavements also have two specialities: the very local Dark-red Helleborine and,
in the Burren in western Ireland, Dense-flowered Orchid, which grows on short turf.
marshes and fens

‘Marsh’, ‘swamp’, ‘fen’ and ‘bog’ are all vague terms and are used interchangeably by the layman (and
most botanists too) to describe any sort of waterlogged or seasonally flooded ground. In terms of
their flora, however, there are many different varieties, only some of which are good for orchids. Wet
ground can be categorised in terms of the source of its water, the water chemistry and the management
that it receives. Broadly, the same factors which determine the orchids that can thrive in grassland
operate in marshes too.
All the water may come from rainfall, in which case a ‘bog’ is formed, either a blanket bog of the
sort which carpets the landscape in northern and western Britain and Ireland, or a raised bog, a much

rarer habitat. Bogs are acidic and usually dominated by bog-mosses Sphagnum spp., the dead remains
of which lead to an accumulation of peat. In general, bogs are poor for orchids, although Heath
Spotted Orchid may be frequent on the higher and drier hummocks.
Alternatively, the bulk of the water may come from springs and seepages in the ground, in
which case its chemistry is strongly influenced by the rocks it has passed through on the way to the
surface. This flushing water may be acid or very alkaline or anything in between, but is usually low in
nutrients. Such spring-fed marshes are often called ‘mires’ in the technical literature and can support
some scarce and local orchids. In southern England valley mires are commonest, in which water seeps
from the ground along the sides of a valley at the boundary of pervious and impervious rocks to form
a long, narrow mire with a central stream. Valley mires are often found in heathland but formerly
would have been widespread and almost every parish would have had its area of boggy ground; most
were reclaimed for agriculture long ago. In the north and west spring-fed mires can be found within
more extensive areas of acid bog and often form a focus for interesting plants.
Because there is a whole range of water chemistry each variety of mire grades into the next in a
complex and often poorly understood manner. Similarly the mix of orchids changes subtly as the
habitat becomes more or less favourable. And, even within a particular mire, there may be many
micro-habitats; in an otherwise very alkaline mire hummocks of Sphagnum may produce locally
acidic conditions.
Where mires are acidic they may hold the diminutive Bog Orchid, as well as the purple-flowered
form of Early Marsh Orchid (subspecies pulchella) and Heath Spotted Orchid. Slightly more neutral
conditions and drier ground favour Lesser Butterfly Orchid and the very local Heath Fragrant Orchid,
as well as the ubiquitous Common Spotted Orchid. Alkaline mires support Marsh Helleborine, and
Marsh Fragrant, Southern Marsh and Northern Marsh Orchids and, in some areas, the very local
Pugsley’s Marsh Orchid. Of the real rarities, Fen Orchid and the creamy-flowered subspecies of Early
Marsh Orchid cling on in East Anglia.
As a third alternative, the water in a marsh may come from rivers and streams, either overflowing
their banks from time to time or percolating through the soil. In this case the water may be acid or
alkaline but is usually nutrient-rich, either naturally or frequently due to the run-off of fertilsers from
agricultural land or the discharge of treated sewage. The nutrient-rich water leads to the growth of
lush, tall vegetation, often dominated by Common Reed or by Nettles, Meadowsweet and Hemp
Agrimony. This vegetation may be controlled, however, by mowing or grazing, in which case such
waterside meadows can hold Southern, Northern and Irish Marsh Orchids, Early Marsh Orchid of
the pink-flowered subspecies incarnata and Common Spotted Orchid.
heaths and moors

Heathland is dominated by heather, one or more species of gorse and sometimes various grasses, and
is generally found on sandy or gravely soils in the lowlands of southern England, whilst moorland is
dominated by heather, grasses and a variety of dwarf shrubs and is generally found in the uplands of
northern and western Britain and Ireland. Both heaths and moors are acidic and are likely to hold
few orchids. Heathland itself is usually very dry but is frequently interspersed by valley mires, which
can be rich in orchids (see above). Moorland tends to be wetter and may hold Heath Spotted Orchids
where it grades into bog. In the north and west, damp moorland, especially on north-facing slopes,
can hold Lesser Twayblade.
Pits, quarries and railway lines

When extraction has come to an end, quarries have large areas of bare rock or sparse soil and where
the bedrock is chalk and limestone these can, over time, form superb orchid habitats. Any of the
grassland orchids can occur and, if there is scrub, ‘woodland’ species such as Fly, Military and Lady
Orchids are also possible. The timescale involved can be lengthy, with medieval workings now forming
excellent sites. Railway lines go through cuttings which may also expose bare rock or produce thin,

skeletal soils, while embankments may have been built up from calcareous spoil; disused railway lines
are always worth investigating. Sandpits are unlikely to be calcareous but the bare ground can be
favourable to some species, such as Bee Orchid, which can form substantial colonies so long as scrub
does not take over.
Sadly, even these man-made habitats are under threat as any hole in the ground may be seen as a
suitable site for landfill and many important orchid sites have been buried by rubbish.
road verges, green lanes, churchyards and lawns

In the last few decades most hay meadows and permanent pastures have been lost, especially in
England. These unimproved grasslands were ideal for orchids and species such as Green-winged
Orchid, which were common and widespread not so long ago, are now highly localised. Road verges,
green lanes and churchyards are now often the only areas where permanent, unimproved grassland
can be found, although in many cases they are drenched with nutrients from farming operations
and car exhausts, and too overgrown with rank vegetation to support orchids or, in the case of
some churchyards, over-tidied and mown far too often. Nevertheless, Early Purple, Green-winged,
Pyramidal and Bee Orchids and even rarities such as Man and Lizard Orchids can be found in these
habitats. Some Wildlife Trusts have ‘conservation churchyard’ schemes that may help to identify
the best sites and most counties operate a Roadside Nature Reserve policy to protect and correctly
manage the best verges. Garden lawns are usually cut too frequently and managed with herbicides
and fertilisers and are therefore unlikely to support orchids. On suitable soils, however, unimproved
lawns, even relatively new ones, can be graced by Autumn Lady’s-tresses. Increasingly, Common
Spotted and Bee Orchids seem to be turning up on lawns and, once found, are often cherished by the
householders.
dunes and dune slacks

In coastal sand dunes the depressions are often damp and may be flooded in the winter. If the dune
systems are made up from sand that is rich in shell fragments they will be neutral or calcium-rich, and
such dune slacks and the surrounding slopes can then support a very rich variety of orchids. Specialities
include Dune Helleborine, a British endemic found in the dune systems of Anglesey and northwest
England; Lindisfarne Helleborine, endemic to the dunes of Holy Island in Northumberland, and Fen
Orchid, found on the coast of South Wales, as well as the red-flowered subspecies of Early Marsh
Orchid (subspecies coccinea) and, in northern England and Scotland, Coralroot Orchid. Other species
may occur, sometimes in large numbers; Marsh Helleborine, and Common Spotted, Southern Marsh
and Northern Marsh Orchids can be abundant and other species to look for include Broad-leaved
and Green-flowered Helleborines, Common Twayblade, the pale pink subspecies of Early Marsh
Orchid (subspecies incarnata), Frog, Pyramidal and Bee Orchids and even, in one or two areas, Man
Orchid. In many places dune systems have been planted with conifers and although this is bad news
for the specialised dune flora, Dune, Green-flowered and Broad-leaved Helleborines all take readily
to the plantations.
machair
Machair is species-rich grassland found on the wet and windy west-facing coasts of Scotland and
Ireland but nowhere else in the world. Sand that contains many fragments of seashells, and hence rich
in calcium, is blown onshore by Atlantic gales and settles on the low-lying coastal areas. Grassland
develops on these low dunes and periods of extensive grazing have been interspersed with cultivation
for crops. The uncultivated areas often support large numbers of orchids, including two specialities,
the ‘Hebridean Spotted Orchid’ (the hebridensis subspecies of Common Spotted Orchid) and the very
localised Hebridean Marsh Orchid, endemic to North Uist in the Outer Hebrides. Other orchids
which can be found include the red-flowered subspecies of Early Marsh Orchid, Northern Marsh,
Heath Fragrant, Lesser Butterfly and Frog Orchids, and Common Twayblade.

Orchid Conservation
Orchids face three major threats: habitat destruction, habitat change and human predation.
Habitat destruction has clearly taken the greatest toll. Farming, forestry and other developments
have destroyed innumerable orchid sites, especially in the period since World War Two. Most of the
destruction has been state-sponsored through the operations of the Common Agricultural Policy
(CAP) and the Forestry Commission. Between 1945 and 1980 the Forestry Commission attempted
to destroy and re-plant with conifers 200,000 hectares of ancient woodland, to say nothing of the
tens of thousands of hectares of heathland, moorland and sand dunes that were destroyed. The
CAP has been reformed in recent years and attitudes and policies at the Forestry Commission have
changed (although there is still a great reluctance to undo much of the damage done in the name
of near-worthless timber). But, despite much lip-service in recent years, few politicians have any
commitment whatsoever to conservation and when push comes to shove development almost always
takes precedence over wildlife. No wonder the government conservation agencies (English Nature,
the Countryside Council for Wales and Scottish Natural Heritage) have often been accused of being
lame ducks in the face of their political masters, despite the best efforts of their staff. In terms of
habitat destruction, the prospects for orchids remain bleak.
Habitat change has only recently been acknowledged as a major issue. It has come to be recognised
that orchids do not live in stable, ‘climax’ communities of plants, at least in the British Isles, rather
in habitats that were created and maintained, albeit inadvertently, by people. Grassland, marshes,
heathland and woodland are all the product of traditional land-use. Once these traditions died out,
habitats started to change, slowly at first but then rapidly, and many have become unsuitable for
orchids. Ironically, this applied especially to reserves, where a fence and a ‘keep out’ sign were often
the limit of any management. Now, conservationists try to replicate the traditional land-uses, often at
great expense, that created and maintained the habitats they manage. The reinstatement of grazing is
often the single most important measure that can be taken to help orchids.
Human predation has often been seen as a major threat to orchids, be it innocent ramblers picking
bunches of flowers or avaricious botanists determined to get another specimen for their collections or
gardens. More recently, photographers and even visitors keen to merely look at plants have joined the
list of ‘threats’. The answer has traditionally been secrecy, and details of the locations of the greatest
rarities were and still are jealously guarded; even the location of huge colonies of species such as Burnt
and Early Spider Orchids was veiled in secrecy.
Human predation certainly poses a threat to those species that occur in such small numbers that
a significant part of a population (or even the whole population) can be stolen. There are still cases
where plants are dug up illegally, from Bog Orchid to Lizard Orchid. Perhaps the most notorious in
recent years was the attack on the single Lady’s-slipper growing in Silverdale in Lancashire. English
Nature had made the bold decision to allow limited publicity and a large number of people had been
able to admire this beautiful orchid. The fact that this particular plant was probably originally of
garden origin does not in any way lessen the damage done.
Despite the odd incident, however, we are convinced that for most orchids human predation is, in
the final analysis, irrelevant to their fortunes, especially in the face of habitat destruction and habitat
change. Unnecessary secrecy has indeed probably led to the destruction or degradation of many sites,
as those responsible for the land remain in ignorance of its importance. It has also deprived many
people of the enjoyment of seeing the orchids and many potential friends for orchid conservation
have surely been lost in this way.
There have been a few special conservation initiatives involving orchids. In 1983 the Sainsbury
Orchid Conservation Project was established at the Royal Botanic Gardens, Kew. This has involved
research into the propagation of orchids with a view to reintroducing some of the rarer species. A
range of orchids has been involved, including Military and Fen Orchids, with a substantial effort
going into the reintroduction of Lady’s-slipper. Reintroductions are controversial, however, with

ORCHId COnSERVATIOn l 19
some conservationists arguing that the time and effort could be better spent on conserving existing
populations. A second initiative at the Royal Botanic Gardens was the establishment in 1997 of the
Millennium Seed Bank. This is intended to store viable seeds for as many of the world’s plants as
possible, including, of course, British wild orchids. Techniques have been developed which should allow
orchid seeds to be stored for long periods, although some species cannot yet be cultivated successfully.
what you can do
The first step in orchid conservation is accurate and up-to-date information on their distribution
and abundance. Amateur botanists provide the vast majority of information on plant distribution
in Britain and Ireland via the system of county recorders organised by the Botanical Society of the
British Isles (BSBI); they are always pleased to receive records, with details of the species involved,
numbers, date and location (for contact details see p.422). There is also always a need for volunteers
to undertake practical habitat management on reserves and other sites; the local wildlife trusts are the
first contact if you are keen to get involved.
The greatest contribution individuals can make to orchid conservation is, in our opinion, to
become a ‘local champion’. Getting to know an area intimately, finding and recording orchids and other
wildlife, and then badgering local councils, wildlife trusts, government agencies or church-wardens to
sit up and do what is necessary to safeguard the good areas. This may not make you popular in some
quarters but may, in the end, get things done.
orchids and the law
All orchids and, indeed, almost all wild plants, are protected in Britain by the Wildlife and Countryside
Act, 1981 and cannot be uprooted unless you are the owner or occupier of the land or have their
permission to do so (although it is legal to pick them). In addition, some of the rarer orchids enjoy
much greater protection under Schedule 8 of the Act and it is illegal for anyone, even the owner or
occupier of the land, to uproot, destroy or pick these orchids. The term ‘pick’ is defined to include
gathering or plucking any part of the plant, including collecting seeds. It is also illegal to posses any
live or dead wild plant in Schedule 8, or any part of or anything derived from such a plant, or to trade
in such items. Schedule 8 includes: Lady’s-slipper, Red Helleborine and Ghost, Fen, Monkey, Military,
Lizard, Late Spider and Early Spider Orchids (it also currently includes ‘Young’s Helleborine’, now
shown to be a poorly defined variant of Broad-leaved Helleborine, and ‘Lapland Marsh Orchid’,
now a subspecies of Pugsley’s Marsh Orchid). In Northern Ireland, Bird’s-nest, Bog, Small White,
Pugsley’s Marsh, Green-winged and Bee Orchids, Marsh and Green-flowered Helleborines and Irish
Lady’s-tresses are specially protected under Schedule 8 of the Wildlife (NI) Order, 1985. In the
Republic of Ireland, Sword-leaved Helleborine, Irish Lady’s-tresses, Bog, Small White and Green-
winged Orchids are specially protected under the Flora Protection Order, 1999.
M Derbyshire, June. Grazing is critical to the survival of many orchid populations and is increasingly the linchpin of
conservation management. Paradoxically, however, in the north and west, overgrazing has decimated species such as
Small White Orchid.

Field Guide
notes on the sPecies Accounts
conservation designations
The latest UK conservation designations are indicated in red. Several species are also BAP species,
that is included in the UK List of Priority Species and Habitats.
introductions to the genera
Each genus has an introductory section giving some general information about the genus, including
notes on identification for the more difficult species, such as the Epipactis helleborines and the marsh
and spotted orchids. It also includes notes on various aspects of biology, such as growth, reproduction
and pollination, that are common to all the members of the genus. For some genera that contain just
one species worldwide, or where only one species is represented in Britain and Ireland, much briefer
notes on distribution and the generic name are given.
names
The scientific names used in the text and the order of the species accounts follow the list produced by
Richard Bateman (Bateman 2005). Scientific names reflect the relationships between species and in
recent years new evidence, especially from genetic studies, has greatly improved our understanding of
these. This is reflected in new scientific names for some species.
Under the heading Formerly we give scientific names as used in publications that pre-date the
Bateman list. We have tried to include most older scientific names if they have been in general use
since the publication in 1968 of the second edition of Summerhayes’ Wild Orchids of Britain. We
have not, however, provided an exhaustive list of synonyms. In addition, some older English names
are also mentioned. Under the heading Other names we give some alternative names in current use,
especially in North America.
habitat
The likely habitats for each species are outlined and more detail on many orchid habitats can be
found above. The information applies to Britain and Ireland; in some cases orchids occupy a rather
broader range of habitats elsewhere. Most maximum recorded heights above sea level come from the
New Atlas (Preston et al. 2002).
flowering period
A guide is given to the period in which the species is likely to be in flower (see also chart of flowering
periods on p.420). Flowering times do vary, however, both predictably and unpredictably:
l Orchids growing further to the north and at higher altitudes tend to flower a little later (although
there is often surprisingly little difference between southern England and Scotland).
l Orchids growing in wetter habitats will flower later than the same species growing in drier habitats.
l Orchids growing on or very near to the coast will tend to come into flower a little earlier than
those inland.
l Orchids growing on sheltered south-facing slopes will flower earlier than those with an exposed,
westerly aspect or those facing north.
There can also be marked and unpredictable variations between colonies, even those close to each
other, and, in recent years, perhaps as a result of ‘global warming’, many orchids have been coming into
flower earlier; it is worth bearing this in mind if you are hoping to see a species at its best.
Orchids are notorious for the wide variations from year to year in the number of plants in flower.
It used to be thought that this was related to fluctuations in the size of the population and that some
species, such as Bee Orchid, were monocarpic and therefore flowered just once before dying. It is now

SpECIES ACCOUnTS l 21
known that most orchids are relatively long-lived and the total population, including non-flowering
plants and those ‘dormant’ underground, is often fairly stable. Fluctuations in the numbers flowering
are related to growing conditions both in the current year and in the previous growing season (which
may be either the previous summer or the previous winter, depending on the species). Growing
conditions are, in turn, usually related to rainfall. Wet weather is conducive to growth but prolonged
dry spells can be very bad for orchids and in some cases can severely restrict flowering.
range
Details of the range is given for Britain and Ireland. A problem faced by all biologists (and many other
people as well) is how to sort and classify records. County boundaries have changed several times over
the years and it becomes very difficult to keep track; was a plant recorded in the ‘Lincolnshire’ of the
19th century in the same region as one recorded in ‘Humberside’ or ‘North Lincolnshire’ today? This
problem is particularly acute in Wales and Scotland where there have been radical changes, and some
of the modern administrative units are very large.
Wisely, most botanical recording uses the system of Watsonian vice-counties. This was devised
in 1852 by Hewett Cottrell Watson, who divided Britain into 112 similarly-sized areas. He followed
traditional county boundaries where he could and divided larger counties such as Yorkshire into
smaller units. The value of the vice-county system is its stability, allowing past and present to
be compared. In the majority of cases we have given records by vice-county, although we have
amalgamated some (e.g. we often refer to Norfolk rather than West Norfolk and East Norfolk) and
we have amended some vice-county names to become more recognisable (for example, West and
East Ross becomes Ross & Cromarty; Westerness and Easterness becomes Inverness-shire). See
p.418 for full details.
We have used two main sources for distributional information: The New Atlas of the British and
Irish Flora (Preston et al. 2002) and the Vice-county Census Catalogue (Stace et al. 2003). We have
also consulted a variety of county floras. These sources do not always agree, especially regarding the
validity of older records and the origin of out-of-range species.
Orchid seeds have evolved for wind or water dispersal and can, in some circumstances, travel
long distances. Nevertheless, a great deal of scepticism attaches to records of orchids away from
their normal range. In some quarters, these are inevitably attributed to deliberate introductions, even
where there is no evidence whatsoever for this (for example, the two tongue orchids). On the other
hand, a non-natural origin has either been proven or is very strongly suggested for some controversial
records (see Other species p. 416). It seems best to keep an open mind but our inclination is to give
such records the ‘benefit of the doubt’.
Under the subheading World range, details of the orchids’ distribution outside Britain and
Ireland is given. We have used several sources for this, including Brown (2003), Davies et al. (1983),
Delforge (1995, 2001) and Hultén & Fries (1986). Again, these sources do not always agree and we
have tried to present the best and most likely compromise.
Accompanying the text is a range map, where the distribution is given by 10km squares in three
data classes: 1987-1999, 1970-1986 and pre 1970. These maps have been supplied by the Biological
Records Centre.
Key to range maps:
Native Distribution
1987-99
1970-86
pre 1970
description
A detailed description of the orchid is given, taken in many cases from the living plant. We have
depended on the literature for some details, especially measurements. It is worth remembering that
many orchids can be taller than the range quoted; even in our limited experience we have found plants
that are bigger than the books suggest.

subspecies
Some orchids show definite patterns of variation which may be either geographical or ecological;
plants in a particular region or a particular habitat may differ consistently in appearance from other
areas. In these cases we have recognised different subspecies. There is much disagreement about
which subspecies are worthy of recognition and we have not followed any one authority in this.
Variation and varieties

We give brief details of the normal range of variation which can be encountered. We also give details
of some of the ‘varieties’. Like all organisms, orchids are subject to random mutations of their DNA
which may produce a variety of aberrant and deformed plants, and, like stamp collectors (and the
butterfly collectors of old), orchidologists have traditionally been fascinated by these abnormalities.
We do not share that obsession and consider that the odd plant that produces flowers that are deformed
or unusually coloured in some way to be of little special interest. However, some deformities shed
light on more primitive stages in the orchid’s evolution. In addition, species such as Bee Orchid throw
up the same mutation again and again in widely separated localities, and in such cases these ‘varieties’
(var. for short) are often given names. Unlike subspecies, varieties do not show definite geographical
or ecological patterns and can pop up anywhere. In most cases we have followed Ettlinger (1997) in
the use of varietal names.
hybrids
Hybrids have, like varieties, long fascinated orchidologists and in some cases are either very attractive
or throw light on the relationships between orchids. Hybrids between species in the same genus tend
to occur much more frequently than hybrids between species in different genera and, for example,
the frequency with which Frog Orchid hybridised with the spotted and marsh orchids was a long-
standing clue to its true relationships. On the other hand, many hybrids are undistinguished and
their true parentage, and even their status as a hybrid, may be the subject of guesswork, sometimes
highly ambitious guesswork. All too often, plants are diagnosed as hybrids when they are merely
aberrant individuals (or even within the range of normal variation). We have usually only included
hybrids that are listed in Stace (2004) and have given the names used in that work.
Name and classification

The origin of the scientific name is discussed under this heading. In some cases the derivation is
clear but in others there is disagreement (or the stated origin seems highly unlikely). We have used
Delforge (1995), Gledhill (2002) and Société Française d’Orchidophilie (1998) as our main sources.
Notes are also given on the classification of the species, including any recent changes, and on its
relationships (see also the orchid family tree on p.417).
history and conservation

The date and, where known, the location of the first record for Britain and Ireland is given (largely
from Clarke 1900). For the rarer species we have also given extensive details of their past and current
distribution.
The conservation status of each species is mentioned. ‘Nationally Scarce’ denotes that the species
is included in Scarce Plants in Britain (Stewart et al. 1994) which indicated that it has been recorded
from 16 to 100 10km squares in Britain from 1970 onwards (the project did not include Ireland).
Rarer species were treated in the British Red Data Book (Wiggington 1999), which includes all plants
recorded from 15 or fewer 10km squares in the period from 1987 onwards. Each species is assigned
to a threat category (Endangered, Vulnerable etc.).
We have extracted data from the New Atlas (Preston et al. 2002) to draw up a table for each species
indicating the extent of its historic and current range and the amount by which it has declined. This
information is put in context by a discussion of the possible reasons for decline (all orchids have
declined), and current threats and conservation measures.

GenUs CYPRIPEDIUM
Lady’s-sLippers
001 systematic.indd 23 29/1/09 11:59:25

24 l GENUS CYPRIPEDIUM
Lady’s-slippers are the most primitive orchids in Europe, differing markedly in their
floral structures from the other species, and have a cunning method of trapping insect
pollinators.
Distribution produces two buds at its tip: the larger will

The genus Cypripedium contains 45 species produce the next aerial shoot, the smaller may
distributed in Europe, temperate Asia and form a new branch of the rhizome but usually
North and Central America. Around 30 species eventually dies. The aerial shoot will also die
are found in China and 12 in North America off at the end of each season, but the rhizome
but just three extend to Europe and only one continues to grow from a bud that forms on
occurs in Britain. its base (sympodial growth, as opposed to
Classification monopodial growth, in which the main stem
The lady’s-slippers belong to the subfamily continues to grow indefinitely behind the
Cypripedioideae and are the only representatives same growing point). In the lady’s-slipper,
of that subfamily in Britain and Europe. the rhizome grows 5-10mm each year in a
characteristic zig-zag pattern as the position of
Floral structures the larger bud alternates from left to right. The
Lady’s-slippers are distinguished from all other rhizome may die off slowly and rot at the rear
British and European species by having two while growing at the front, but up to 20 living
fertile stamens, one on either side of the column, segments may be present at any one time
plus a large, modified, third, sterile stamen (and by counting the number of segments
known as the staminode. This is flattened, its age may be estimated). The rhizome also
tongue-like, often boldly marked and projects branches, roughly every five years, and each
into the large opening of the ‘slipper’. The pollen branch may eventually put up an aerial stem.
is granular, sticky and glutinous. There are Thus as a plant ages, the number of flowering
three receptive stigmas and the lip is shaped shoots increases.
into a slipper or moccasin-like pouch. The two
lateral sepals are joined for almost their entire Fungal partners
length with just a notch at the tip to suggest Poorly known, and this lack of information is
their origin (this structure is called a synsepal). an obstacle to the conservation of the species.
Members of the other orchid subfamilies have Vegetative reproduction
just one fertile stamen, pollen that is aggregated The rhizome branches as it grows and
together into two pollinia and two stigmas
eventually the basal portion dies, leaving the
(although these are often joined together).
terminal branches as independent plants, clones
Pollination of the ‘mother plant’. In at least some species of
There is no nectar, and pollinating insects are Cypripedium, buds can also form on the tips of
tricked into entering the flower through the the roots as a means of propagation.
large opening on the top of the lip. They can
Name
only leave via the small rear openings, picking
The generic name Cypripedium derives from the
up pollen on the way.
Greek Kypris, a name for Aphrodite (goddess
Growth pattern of love), and Latin pes, meaning ‘foot’ and thus
All lady’s-slippers grow from a rhizome that ‘Aphrodite’s slipper’.
has slender, fleshy roots. Each year the rhizome
O 31 May, Khabarovsk, eastern Sibera. Lady’s-slipper

occurs right across Europe and Asia but is threatened
almost everywhere.
001 systematic.indd 24 29/1/09 11:59:25

LADY’S-SLIPPER l 25
Lady’s-sLipper Critically Endangered, BAP

Cypripedium calceolus
This spectacular species is Britain’s rarest orchid with just one group of plants of native
origin surviving at a closely guarded site in Yorkshire. Originally fairly widespread in the
limestone districts of northern England, its large, showy flowers have been its downfall.
Plants were pillaged from the wild for hundreds of years, either to be dried, pressed and
stored away in dusty herbaria or to be transplanted to gardens.
Identification north-facing slope. They nestle in a sheltered

Unmistakable. limestone valley that has some trees and lies on
Similar species the northern edge of a large block of woodland.
In Europe, Lady’s-slipper favours a variety of
The leaves of non-flowering plants could be
woodland, usually on calcareous soils, but in
overlooked and resemble the leaves of Lily-of-
the northern parts of the range is also found in
the-valley.
the open, in fens and marshy grassland. Overall,
Habitat it appears to require relatively well-lit areas
The surviving native plants are found in species- but likes to have its roots in cool, moist soil. In
rich grassland on a fairly steep, well-drained, England suitable areas would have been found
in open woodland on north or northwest facing
slopes or where subsurface ‘flushing’ kept the
ground damp but not waterlogged; the former
English, sites were in ash, hazel and oak woods
on steep, rocky slopes, always on limestone, at
150-260m above sea level. The correct balance
of sun and shade seems to be critical. Too little
light and the plant fails to flower and may even
become ‘dormant’ underground, too much
light and rank herbs and shrubs will swamp it,
although in a more natural environment grazing
animals may have suppressed much of the
competing vegetation.
Flowering period
Late May to early or mid-June. Plants are in
flower for two or three weeks and each flower
lasts 11-17 days, withering on the sixth day
after pollination.
Range
Confined to a single site in Yorkshire,
although formerly more widespread.
Artificially propagated plants have recently
M 30 May, Lancashire (Sean Cole). This plant, probably been planted out at several localities within
of garden origin, had been growing in this spot for perhaps
a century but was vandalised a few weeks after this
its former range. World range: Europe and
picture was taken in 2004. southern Siberia, extending eastwards to
001 systematic.indd 25 29/1/09 11:59:26

DESCRIPTION
1987-99
1970-86 Height: 15-70cm but usually around 30cm and
pre 1970
rarely more than 60cm.
Stem: Glandular-hairy with three to four green
or brown sheaths at the base. The species forms
clumps of shoots, sometimes quite large, which
may belong to one or more plants.
Leaves: Green and oval, elongated into a
pointed tip; the three to four (sometimes five)
leaves are arranged alternately up the stem.
They are sparsely hairy (especially on the
underside), ciliate along the margins, wavy
edged and very prominently veined.
Spike: Each stem usually produces one or two
flowers, very rarely three.
Bract: Leaf-like, longer than the flower and
held erect behind it.
northern Mongolia, northeast China, Korea,
Ovary: Long, slender, six-ribbed, curved (but not
the Russian Far East and Sakhalin. It is
twisted), with glandular hairs and a short stalk.
essentially a northern or boreal species, and
Flower: Large and conspicuous. The sepals
in Europe the Lady’s-slipper ranges north to
are purplish-brown or claret with wavy edges,
northernmost Scandinavia. It is absent from
downy on their inner surface and hairy at
the Mediterranean lowlands but is found in
the base. The upper sepal is lanceolate with
the mountains as far south as northern Spain,
a pointed tip and is held erect, whereas the
central Italy, northern Greece and Bulgaria.
two lateral sepals are fused and hang vertically
There are also outposts in the Crimea and
below the lip (their tips forming two small teeth
Caucasus. It is also absent from the Atlantic
at the tip of the combined synsepal). The petals
fringes of Europe, for example central and
are purplish-brown, mottled with olive-yellow
western France, and the English populations
towards the base and with downy midribs and
were therefore always out on a limb.
long hairs at the base. They are strap-shaped
How to find it with pointed tips, variably twisted (through up
Potential visitors are asked to keep away from to 360°) and hang at ‘four’ and ‘eight o-clock’
the native site in Yorkshire due to the fragility on either side of the lip. The lip is yellow and
of the habitat. Visitors were welcome at a site at looks like a bag or clog (the ‘slipper’). There is
Silverdale in Lancashire where one can admire a large entrance on the upper side towards the
a plant, albeit probably of European origin, in rear and two small openings on either side of
a natural setting (it has hopefully survived the the column at the base. The edges of the large
attack by vandals of 2004). Otherwise, we must upper opening are rolled down and under, and
hope that the reintroduction programme is a the interior of the slipper is covered in sticky
success and that self-sustaining populations of hairs with lines of reddish dots along its floor.
flowering plants will once more grace the dales The column projects forwards into the slipper
of northern England. At present, just one of and is divided into two parts: the staminode,
the reintroduction sites has public access which is yellowish-white variably marked with
– Ingleton Glen in North Yorkshire – but red spots and very conspicuous, and the large
the young Lady’s-slipper plants have not yet
flowered there. P Yorkshire (Peter Wakely, English Nature).
001 systematic.indd 26 29/1/09 11:59:27

001 systematic.indd 27 29/1/09 11:59:36
fleshy stigma, which lies on the lower part of picking up a load of pollen in the process. The
the column hidden inside the slipper. The two bee goes on to visit another flower; when it
remaining stamens lie on either side of the base eventually leaves this its back rubs against the
of the staminode, adjacent to the two small stigma, which projects down into the slipper,
rear openings into the slipper. The flowers are and pollen from the first flower is deposited
delicately scented, and the scent is said to be there; the surface of the stigma has minute, stiff,
sweet, recalling oranges. pointed papillae that act as a ‘brush’ to remove
pollen from the bee’s back. As it escapes, more
Subspecies
pollen is carried away, ready to be deposited on
None in Britain.
the next flower and continue the process.
Variation and varieties The mechanism is precise and in order to
None in Britain. effect pollination the bee has to be a specific
species that is the right size; bees that are
BIOLOGY too large or too small can escape without
pollinating the flower. A wide variety of other
Pollination and reproduction insects also enters the slipper but these too
It is thought that Lady’s-slipper is pollinated
are the wrong size and shape and either leave
by small bees, especially bees of the genus
unharmed or may be trapped and die. Self-
Andrena. The flowers do not produce nectar,
pollination is unlikely; the bee would have to
however, and there is debate as to what it is
reverse back into the flower just as it was on the
that attracts the insects. A strong contender is
point of escape. In addition, it seems that the
the flower’s scent, which comprises a complex
flowers are, to a great extent, self-sterile.
mixture of chemicals that may mimic the
The pollination strategy is not efficient
bees’ pheromones, chemical signals that are
and seed set is rather poor with few fertile
associated with feeding and mating behaviours.
capsules being produced. Bees are attracted
Other possible attractions, which could
to large groups of flowers, especially those
indeed combine with the scent lure, include
in sunlight, but even in large populations in
the red spots on the staminode and floor of
Europe an average of just 10% of flowers set
the slipper that may act as ‘false’ nectar guides.
seed. Nevertheless, each capsule contains 6,000-
Alternatively, the bees may actually receive a
17,000 seeds which may be dispersed by rain
reward for their visit, perhaps the oil secreted
as the seedpods seem to close up when dry and
by the small hairs inside the slipper. Bees may
open when wet.
also find shelter in the slipper in cold weather
Lady’s-slipper also reproduces vegetatively
or overnight.
through division of the branching rhizome,
Whatever the attraction, bees land on the
and in many populations in Europe this is
edge of the ‘large opening’ or try to land on
thought to be more important than seed in the
the staminode and fall into the slipper. After
recruitment of new plants to the population.
a few minutes the bee tries to leave. However,
the sides of the slipper are very smooth and Development and growth
slippery and the rim of the large opening curls The Lady’s-slipper grows from a slender,
over and inwards, making escape via this route creeping, branched rhizome. Each branch of
impossible. The bee can only leave through the the rhizome may eventually put up an aerial
small openings on either side of the column stem, and so as a plant ages the number of
where there are small stiff hairs to give it a flowering shoots increases. Alternatively,
foothold. The openings are only just big enough a clump of shoots may arise from several
for the bee, which is forced to make contact separate clones, each produced by vegetative
with one of the stamens as it makes its escape, reproduction, or by the development of
001 systematic.indd 28 29/1/09 11:59:37

M 6 June, Lancashire.The ‘large opening’ of the flower, with the tongue-like staminode to the rear, is obvious.
001 systematic.indd 29 29/1/09 11:59:39

seedlings at the base of the ‘mother’ plant. Past and present occurrence of Lady’s-slipper in Britain
and Ireland (based on presence or absence in 10km
The first green leaves are reported to appear squares of the National Grid; data from the New Atlas).
in the fourth year after germination by some
authors and in the first year by others. The Britain Ireland
immature plants have a slender stem with one
total historical range, 22 0
or two small leaves and may remain in this state 1500-1999
for several years. In England a seedling has been current range 1 (0.04%*) 0
noted to flower nine years after it first appeared
above ground, and in Europe the young plant % lost, 1500-1969 95%
takes six to ten years to produce flowers. Plants % lost, 1970-1986 0%
are long-lived and many are over 30 years old, % lost, total 95%
with some over 100 years. Indeed, a lifespan * current range as a % of the total number of 10km squares
of 192 years has been determined from the
examination of the growth of a single rhizome of northern England. There was a single site
in Estonia. in Derbyshire at the Heights of Abraham
overlooking Matlock, and it was recorded in
Hybrids southern Cumbria around Whitbarrow and
None in Britain. Scout Scar (northwest of Levens). The bulk of
Name and classification the records came, however, from three areas:
The specific name calceolus means ‘little shoe’ first, West Yorkshire around Ingleborough
or ‘little slipper’ and like the English name (including Helkes Wood) and the Upper
refers to the slipper-like appearance of the Wharfedale region around Litton, Kettlewell
lip. An old name for the species was ‘Calceolus and Grassington; second, North Yorkshire
Mariae’ or ‘Mary’s Shoe’. Yellow Lady’s-slipper on the southern flanks of the Cleveland Hills,
C. parviflorum of North America is very closely especially the valley of the River Rye and its
related and is sometimes amalgamated with tributaries north and west of Helmsley; third,
this species. Castle Eden Dene in Co. Durham.
Such a large, conspicuous and attractive
HISTORY AND flower as the Lady’s-slipper was an obvious
CONSERVATION subject for curiosity and avarice, and plants
A Red Data Book species that is classed as were picked or dug up from at least the 16th
Critically Endangered and fully protected century onwards. In the late 18th and early
under Schedule 8 of the Wildlife and 19th centuries, they were ruthlessly stripped
Countryside Act 1981. The species is also rare from the wild, both for horticulture and as
and threatened throughout much of its range herbarium specimens. By the mid-19th century
in Europe and Asia. the species was rare and in 1917 the Lady’s-
The first British record dates from 1629 slipper was declared extinct in Britain.
when John Parkinson, a London apothecary, In 1930, Lady’s-slipper was resurrected
recorded the species in his Paradisi in Sole; from the dead when a single plant was found
Paradisus Terrestris (‘Park-in-sun’s Earthly in a remote Yorkshire dale (this is the plant
Paradise’): ‘In a wood called the Helkes in that has survived to the present day). However,
Lancashire neere the border of Yorkeshire.’ the last note of its rediscovery in print was
This was the same wood that supplied the first in 1937, and the species quietly slipped from
British record of Sword-leaved Helleborine in the botanical world’s attention. As well as the
1666. Both species are now gone from there. site being a closely-guarded secret, the Lady’s-
The Lady’s-slipper was subsequently found slipper itself was being very coy. From 14 stems
widely but locally in the limestone districts and just one flower in 1930 it dwindled to two
001 systematic.indd 30 29/1/09 11:59:40

to five stems by the late 1940s and 1950s and aids the Lady’s-slipper seeds in germination and
hardly ever bothered to flower; a single bloom growth could not be identified, however, and
was produced in 1934 and 1943 but not again this initially thwarted efforts to cultivate plants
until 1959. from seed. But, after much trial and error, a
Despite the secrecy, in the 1960s word method of germinating seed in the absence of
started to get out and the Lady’s-slipper fungi was developed. This involves supplying
again faced the old threat from collectors the nutrients directly to the seedling in a sterile,
and a new threat from visiting botanists with asymbiotic medium. Although only about 10%
big feet and heavy cameras. Indeed, the site of seeds germinate, large numbers of seedlings
was raided and half the plant was removed. can now be produced from the hand-pollinated
Various individuals and groups attempted to capsules of the wild plant.
protect the orchid and the ensuing conflict The major problem has become the
and confusion prompted the late Edgar Milne- development of techniques to introduce these
Redhead, President of the Botanical Society laboratory-grown seedlings into a natural
of the British Isles, to set up the ‘Cypripedium environment. The first six seedlings were
Committee’ in 1970. The committee included planted out at the native site in Yorkshire in
representatives of various conservation and autumn 1989, and by 2003 approximately
botanical interests with the aim of coordinating 2,000 seedlings had been introduced into
the orchid’s conservation. The first priority 23 locations (although some of the sites are
was to safeguard the sole remaining wild plant, very close to each other). Survival rates are
and this has been guarded every year since not high, and slugs and snails are a particular
then. Visitors are asked to keep away, and problem. Up to 2004, just 105 plants survived
even the committee has only rarely visited the at ten sites. Two have flowered, the first in the
site, whilst ‘management visits’ have also been summer of 2000, 11 years after being planted
curtailed for fear of damaging seedlings. out. Early reintroductions used two-year old
With careful protection and habitat seedlings (one year ‘in flask’ and another in
management the Lady’s-slipper has slowly compost), and survival rates were particularly
increased in vigour at the wild site, with a poor as the tiny seedlings fell foul of slugs or
steady increase in the number of shoots and bacterial and fungal infections. Fewer older
flowers. In 1996, the peak year to date, there seedlings are now used (three or even more
were 65 shoots and 23 flowers on the main years old). These are being planted in drier and
clump (which may be just one plant, several more open micro-habitats and appear to have
clones or even include seedlings). Few or much better prospects for survival.
no flowers were being pollinated naturally, A single plant has been present at Silverdale
however, and hand-pollination began in 1970. in Lancashire for many years, although it is
This has resulted in good seed set and the thought that it was planted there in the late
production of many capsules, some of which 19th or early 20th century; its DNA suggests
are left to mature on the plant while others are that it is from either Austria or possibly the
sent to Kew Gardens. Pyrenees. It did not flower for many years
As part of the ‘Species Recovery Programme’, but slowly increased in vigour and by 2004
organised by English Nature, and with the goal produced nine flowers. Sadly, later in the 2004
of establishing self-sustaining populations of season this plant was vandalised and probably
Lady’s-slipper in the wild, ex-situ propagation partially removed although fortunately it has
began in 1983, when a donation from Sir survived this.
Robert and Lady Sainsbury established the
Sainsbury Orchid Conservation Project at the
Royal Botanic Gardens, Kew. The fungus that
001 systematic.indd 31 29/1/09 11:59:40

GENUS
CEPHALANTHERA
HELLEBORINES (I)
001 systematic.indd 32 12/2/09 16:51:29

GENUS CEPHALANTHERA l 33
A genus of relatively primitive woodland orchids that are probably heavily dependent on
fungi throughout their lives. Their flowers look very simple but are surprisingly similar to
those of the apparently more complex and highly-evolved Epipactis helleborines.
Distribution pollinia. As in the Epipactis helleborines, the lip

Around 15 species are found in Europe, is divided by a narrow waist into the hypochile
North Africa and temperate Asia with a single at the base and epichile at the tip.
representative in western North America. Eight
Pollination
species are found in Europe, with three in Britain.
White Helleborine is largely self-pollinated but
Classification Red and Sword-leaved Helleborines are cross-
Considered a relatively primitive genus due pollinated. Their flowers do not produce nectar,
to the rather unspecialised reproductive however, and depend upon deceit, displaying
structures. Closely related to the helleborines of bright yellow pseudopollen on the lip to entice
the genus Epipactis, but the flowers are stemless potential pollinators to visit. The mechanism is
(sessile), the ovary is often twisted and the rather simple: visiting insects brush against the
column is simpler in structure. stigma and pick up some of the sticky substance
that it exudes, then, as the insects back out of
Floral structures
the flower, the pollinia stick to this sticky mess
upper and are carried away.
sepal
Growth pattern
anther The aerial stems grow from rhizomes that put
petal
column out slender, fleshy roots.
pseudopollen Fungal partners
They are probably heavily dependent on fungi,
lateral sepal
and recent studies have shown that adult plants
of both Red and White Helleborines do acquire
nutrients via a fungal route and also have an
ovary lateral
upper sepal association with ectomycorrhizal fungi; they
sepal
may gain nutrients from the roots of nearby
trees via these fungi. One member of the genus,
petals the Phantom Orchid C. austiniae of western
ovary lip North America, produces no green leaves and is
fully dependent on fungi.
Development from seed
bract lateral sepals Poorly known, and seedlings are rarely found.
Vegetative reproduction
The column has a simple, sticky stigma and no
White Helleborine may produce new shoots
rostellum or viscidium. The pollen is aggregated
from buds on the roots, but Sword-leaved
together into pollinia but as single grains
Helleborine is thought to lack any method of
that form a powdery mass; in more advanced
vegetative propagation.
orchids the pollen is grouped into tetrads that
are held together by elastic threads, and it is Name
these tetrads that are bound together in the The name Cephalanthera means ‘flowery-
headed’ and derives from the Greek kephale
O 26 May, Cambridgeshire.White Helleborine ‘head’ and antheros ‘flowery’ or ‘blooming’.
001 systematic.indd 33 29/1/09 11:59:42

34 l GENUS CEPHALANTHERA
red HeLLeborine Critically Endangered, BAP

Cephalanthera rubra
With flowers that are closer to pink than red, this is one of the most striking and attractive
of all British and Irish orchids. Sadly, it is also one of the rarest, with just a handful of plants
known at three sites in southern England. Red Helleborine has always been rare in Britain
but even in its traditional stronghold in the Cotswolds it has now declined to just a single
site. On the edge of its range in Britain, it is capricious and seems to require exactly the right
habitat with exactly the right mix of light and shade, but in Europe it is much less picky.
Identification
This species is so rare that identification is likely
to be academic. If you are fortunate enough to
be looking at a Red Helleborine you will have
travelled specifically to see it. It is, however,
possible that new sites may still be found in
or around suitable woodland in southern
England. Flowering plants are unmistakable,
but non-flowering plants are very hard to find
and difficult to distinguish with certainty from
White Helleborine or even indeed from the
Epipactis helleborines.
Habitat
Found in beechwoods growing either on
chalk, as in the Chilterns and Hampshire, or
on limestone, as in the Cotswolds, usually on
free-draining slopes. At all three current sites it
grows at the boundary of the chalk or limestone
and the overlying acidic clay drift. All three sites
are probably ancient woodland.
Although a woodland orchid, too much
shade will prevent it from flowering regularly
or successfully. It has long been known that
the finest specimens were to be found in rather
open spots within the woods, sometimes
amongst tall grass, brambles and other
undergrowth alongside paths or in scrubby
places and on open banks. In such open
situations, however, the growth of scrub can
eventually overwhelm the helleborines. It may
well be adapted to flower in the gaps caused
by tree-falls, where a sudden increase in light
allows it to flower for a year or two before the
M 24 June, Buckinghamshire. The plant appears spindly
canopy closes again, light levels fall and the and fragile, almost top-heavy, and seemingly struggles to
plant retreats to a ‘dormant’ state underground support such a large, elegant flower spike.
001 systematic.indd 34 29/1/09 11:59:43

RED HELLEBORINE l 35
until conditions become favourable again.

Alternatively, grazing animals in the primeval
forests may have kept the development of rank
vegetation and scrub at bay for long periods
of time. In France and Germany, where Red
Helleborine is locally common, it thrives best in
situations where it receives a few hours of direct
sunlight each day, in glades or along forest roads
and tracks.
Flowering period
Mid-June to mid-July, sometimes late July, but
usually at its best in the last week of June and
first week of July.
Range
Currently known from single sites in north
Hampshire, Buckinghamshire and east
Gloucestershire. World range: Essentially a
European species, ranging north to c. 60°N in
southern Scandinavia, southernmost Finland
and the Baltic States, east to the Urals and
south to the Mediterranean, including Corsica, M 28 June, Buckinghamshire. Confined to beechwoods,
Red Helleborine acquires nutrients from its fungal
Sardinia, the Balearic Islands, Crete and ‘partner’ throughout its life.
Cyprus, and also to the Crimea and Caucasus.
Found in North Africa in Morocco, Algeria How to find it
and Tunisia and in Turkey, Israel, Syria and Red Helleborine is a shy flowerer, and even
northern Iran. It appears to prefer a continental when it flowers, there may be many more
climate and is largely absent from the Low non-flowering plants in the vicinity, and it can
Countries, the milder western seaboard of remain in a vegetative state for many years. It
France and, of course, Britain. is nevertheless worth looking for, especially
in beechwoods on chalk or limestone, where
recent felling or tree-falls have let in more
1987-99
1970-86 light. But realistically, the only chance to see
pre 1970
this species is at one of the known sites. The
populations are very small and vulnerable and
although visitors are not encouraged, low-key
open days are organised at the Hampshire
site during the flowering season (contact the
Hampshire Wildlife Trust for details).
DESCRIPTION
Height: 15-60cm.
Stem: Slender, often wavy and with abundant
short glandular hairs on the upper portion. The
stem is dusky green, variably washed brownish-
purple towards the tip and sometimes with
violet sheaths at the base.
001 systematic.indd 35 29/1/09 11:59:44

001 systematic.indd 36 29/1/09 11:59:48
Leaves: Dusky grey-green, held in two ranks and petals form a rather loose hood. The lip
alternately up the stem at about 45°. Most of is divided into hypochile and epichile by a
the five to eight leaves are long, narrow and constriction around the mid-point. The basal
lanceolate, but the lower ones are shorter, portion (hypochile) is gutter-shaped and the
blunter and duller green. sides curl up on either side of the purplish
Spike: Rather open, with two to nine flowers column and anther; it is white with fine yellow
(occasionally as many as 17). The ovaries are veins and pale-pink sides. The epichile is flatter
held more-or-less erect, and the flowers face (although still concave), arrow-shaped and
upwards and outwards. The unopened tapers to a pointed tip that is bent downwards.
buds are purplish-pink, becoming creamy at It is whitish with variably pinker edges, a deep
the base. lilac-pink tip and seven to nine longitudinal
Bract: Dusky green, with numerous short yellow ridges. The column is violet-rose.
glandular hairs, especially towards the base.
The bracts are very narrow and pointed, the
Subspecies
None.
lower about one-and-a-half times the length
of the ovary, the upper roughly equal in length. Variation and varieties
Ovary: Dusky green, becoming brownish- None.
purple at the base and with the ribs variably
washed purplish. Slim, cylindrical, ribbed
BIOLOGY
and twisted, the ovary has numerous
glandular hairs. Pollination and reproduction
Flower: A beautiful shade of pink, the flowers Red Helleborine is pollinated by small solitary
open widely. The sepals are elongated ovals, bees, with members of the genus Chelostoma
narrowing both towards the pointed tips and being important, at least in Europe. The flowers
towards the base. They are lilac-pink, becoming do not produce nectar, but studies in Sweden
whiter around the base, with numerous short suggest that the bees are attracted to the
glandular hairs on their outer surface. The helleborines because to the bees’ eyes (which are
petals are similar, although shorter, broader not sensitive to the red end of the spectrum)
and less tapering at the base. When fully they resemble certain blue bellflowers
open the lateral sepals are held horizontally Campanula sp. that do produce nectar. And,
like outstretched arms, while the upper sepal not only do bees visit the bellflowers to collect
nectar, but also male bees search for females
around the flowers. The Red Helleborine may
therefore be able to exploit the bee’s sexual
urges as well as its foraging behaviour. The
pollination mechanism is, however, not efficient
in England, perhaps due to an absence of bees
of the right size and shape, and few flowers set
seed. Many flowers remain unfertilised and fall
off the plant complete with their ovaries.
Vegetative reproduction may be more
important than reproduction by seed. If the
central rhizome dies off, the short side roots,
densely ‘infected’ with fungi, can remain alive
M 28 June, Buckinghamshire.The lip has a series of par-
allel raised ridges, probably related to the attraction of and produce a bud at the tip that will grow into
pollinating insects. a new rhizome and eventually produce a new
O 24 June, Buckinghamshire. leafy shoot.
001 systematic.indd 37 29/1/09 11:59:50

O 28 June, Buckingham-
shire. The lateral sepals
are spread wide.
Development and growth Hampshire site the plant first discovered in

The aerial stems grow singly from a slender, bloom in 1986 survived for at least another 18
horizontal rhizome. There is a heavy presence years and flowered in all but four seasons.
of fungi in the roots, especially the short side There is no information on germination or
roots. Recent isotope studies have confirmed early development. It has been stated that the
that mature plants acquire nitrogen and carbon first leaves are not produced until about six
via their fungal ‘partner’, gaining around 60% years after germination, and flowering does not
of their nitrogen and about 25% of their occur until the plant is ten years of age. But, as
carbon in this way. Unexpectedly, the Red with all such reports, the actual time-scale may
Helleborine has a particular association with be much shorter.
ectomycorrhizal fungi and thus, like Coralroot
Hybrids
and Bird’s-nest Orchids, may gain nutrients
None.
via these fungi from the roots of nearby trees,
especially Beech. But unlike the tree, it ‘cheats’ Name and classification
the fungus by giving nothing in return. The specific name rubra means ‘red’.
Red Helleborine may become ‘dormant’
underground, and gaps of up to four years HISTORY AND
between appearances have been recorded CONSERVATION
from Hampshire (although it is possible that A Red Data Book species that is classed as
shoots did emerge but were quickly grazed off ). Critically Endangered and fully protected under
Indeed, it may be able to persist underground Schedule 8 of the Wildlife and Countryside
for much longer periods. After some trees were Act 1981. The population is tiny, and in 2004
felled at the Hampshire site, a plant appeared there were just over 30 shoots at three sites,
in 2003 that had not been recorded in any producing ten flower spikes.
of the previous 17 years. With 14 flowers, it The first British record was published in
was one of the most robust plants ever seen 1797 in English Botany: ‘Gathered last June
at this site, suggesting that it was not a newly on Hampton Common, Gloucestershire,
emerged seedling. In addition, Red Helleborine by Mrs. Smith, of Barnham House in that
may be relatively long-lived, and again at the neighbourhood.’ The Cotswolds were for a long
001 systematic.indd 38 29/1/09 11:59:51

time the stronghold of the species with records was located. It is thought that this plant may
from ‘a great many of the Beech woods from have been stimulated into flowering by the
Nailsworth to Birdlip, but it has seldom been felling of a few trees some years before; the
seen in quantity and flowering is extremely wood where it grows had been heavily shaded
erratic’ (Lousley 1969). It was common enough by Beech and Yew. Subsequently the whole
at times, however: ‘I once saw some fifty or sixty area was thinned and then clear-felled, mown
plants together, but only about ten bore spikes and hand-weeded. Wire cages have been used
of flowers, and somebody cut those before the to protect the plants, but despite this the
next morning’ (Riddelsdell et al. 1948). There flower spikes have been damaged by careless
are also old records from south Somerset, west photographers on three occasions. And, despite
Gloucestershire, West Sussex and Kent, but the careful management, there has been just
no specimens have survived from the latter a modest increase with one to eight plants
two counties to confirm the identifications appearing and producing one or two flower
and although this is a very distinctive plant the spikes almost every year.
reliability of these reports is uncertain. Red Helleborine is clearly sensitive to light
Following its discovery, the number of levels. Another factor that affects the plants is
sites in the Cotswolds gradually dwindled grazing: deer, rabbits and slugs undoubtedly
and there is now just one (although it is quite damage the helleborines and prevent flowering.
possible that the plant is lurking unseen at Slug pellets are used in the Cotswolds for this
others). At this beechwood reserve there were reason, although their effect on the fungal
up to 40 shoots in the 1970s, but by the late ‘partner’ is unknown. Conversely, grazing
1990s, probably due to increased shade, it had also keeps down the surrounding vegetation
declined to only three shoots with just one of and reduces competition, and the scratching
these flowering. Happily, there has since been and scraping of rabbits may provide suitable
a partial recovery to around ten shoots with at conditions for the establishment of new plants.
least one flowering each year. This may have Thus in the Cotswolds the management regime
been encouraged by some limited felling to let now includes some limited raking and also
in more light. fencing the area against deer over the summer
Red Helleborine was found in the Chilterns and then removing the fences in an attempt to
in Buckinghamshire in 1955 when three ‘get the best of both worlds’.
flowering spikes were discovered in a clearing British populations of Red Helleborine
in a beechwood. There were ten the following produce few young plants and seed production
year, together with 64 non-flowering plants is low. An individual plant in Hampshire
within about 25m. After this spectacular produced 96 flowers over 14 seasons but just
discovery, however, there was a sharp decline, nine capsules were recorded, despite some
and no flowers were recorded from 1960. flowers being hand-pollinated in nine of those
Indeed, no plants were seen at all for several years. In the Chilterns natural pollination levels
years in the 1970s. But from 1980 onwards odd are also low, and seed is only occasionally set;
non-flowering plants reappeared and flowering hand-pollination was also undertaken for a
resumed in 1983. There was a distinct increase while but analyses of the seeds showed that
from 1989 (probably encouraged by the felling only around 20% were viable.
of a handful of trees in 1987), and since then The capricious flowering and poor seed-
about 10-14 plants have appeared each year, a set of Red Helleborine in England recalls
little over half of them producing flowers. Sword-leaved Helleborine, a species that is
Red Helleborine was recorded in also pollinated by small solitary bees. Both
Hampshire in 1926 and then refound in north helleborines will flower once a certain level
Hampshire in 1986 when a single flower spike of light is achieved, but much brighter, sunlit
001 systematic.indd 39 29/1/09 11:59:52

conditions with a variety of genuinely nectar- Past and present occurrence of Red Helleborine in Brit-
ain and Ireland (based on presence or absence in 10km
producing flowers in the vicinity are probably squares of the National Grid; data from the New Atlas).
required if they are to be visited by the correct
pollinators and set seed. The spectacle of Britain Ireland
large numbers of flowering plants may also
boost pollination rates and the tiny British 1500-1999
populations of Red Helleborine may be current range 3 (0.1%*) 0
especially handicapped if this is true.
With so few plants, management of the three % lost, 1500-1969 50%
populations is necessarily tentative. The most % lost, 1970-1986 20%
effective techniques may be to mimic ‘natural % lost, total 70%
systems’: limited felling to produce a mosaic of
* current range as a % of the total number of 10km squares
woodland and glades; winter grazing to suppress
rank vegetation; and protection from deer and
rabbits when flowering and fruiting.
O 28 June, Buckingham-
shire. As in all helleborines,
the lip is divided into two
about half way along its
length.
001 systematic.indd 40 29/1/09 11:59:54

SWORD-LEAVED HELLEBORINE l 41
sword-Leaved HeLLeborine
Cephalanthera longifolia Vulnerable, BAP
Other names: Narrow-leaved Helleborine

This is among the most striking and attractive orchids, but sadly also one of the most
threatened. Sword-leaved Helleborine occurs very widely in Britain and Ireland but has
suffered a dramatic decline over the last 200 years, and sites for the species are now few
and far between. It has only been recorded recently from around a quarter of its historical
range, and many populations are so small that they are in imminent danger of extinction.
Fortunately, our understanding of the biology of this species has improved, and with
appropriate management some sites have made a spectacular recovery.
Identification
A distinctive orchid, with graceful, gently
arching foliage, long leaves alternating all the
way up the stem and spires of pure white
flowers that open just enough to display a large
golden-yellow patch on the lip.
Similar species
White Helleborine is rather similar and
sometimes found growing together with
Sword-leaved Helleborine in southern England.
It has duller, creamier flowers, blunter sepals,
and its leaves, especially the lower ones, are
on average shorter and broader. The best
distinction is the length of the bracts, which are
longer than the ovary in all White Helleborine
flowers, even at the top of the spike. On Sword-
leaved Helleborine the bracts are shorter than
the ovary, at least in the upper part of the spike
(they may be very long and leaf-like on the
lowest two or three flowers), and therefore the
flower spike is well-demarcated from the leafy
part of the stem.
Habitat
Sword-leaved Helleborine is a ‘woodland’
orchid but its optimum habitat is the interface
of woodland and grassland. It does best in
glades, clearings, rides and on the margins of
roads and tracks, in areas where the ground
vegetation is not too dense. It depends on small
solitary bees for pollination and these are found
in species-rich grassland, visiting the sunnier M 27 May, Hampshire. The stately, leafy stems, clearly
parts of adjacent woodland to forage. Suitable distinct from the leafless flower spike, are characteristic.
001 systematic.indd 41 29/1/09 11:59:55

sunlit, open conditions are naturally transient scrub and occasionally on chalk grassland, but
in woods as open areas are invaded by scrub the plants are very small on grassland with few
or the canopy closes overhead, increasing the flowers. On the coast of Co. Mayo in western
level of shade. Sword-leaved Helleborine will Ireland, it once grew on wind-blown shell-sand
persist and can flower well under the shade overlying peat, both in stunted hazel and oak
of a high, closed canopy but in the absence scrub and on an exposed summit (where the
of successful pollination and reproduction, plants were even smaller, just 5-7.5cm high
will eventually disappear from such sites. It with only one or two flowers). It has also been
can also persist in dense shade under scrub, found in pine plantations at Newborough
but flowering will be very much reduced, the Warren on Anglesey.
plants remaining in a vegetative state or even Many sites for Sword-leaved Helleborine
becoming ‘dormant’ underground. However, it are on calcareous soils overlying chalk and
can flower spectacularly if the scrub is cleared limestone, but in the north and west it is found
and the light intensity rises above a certain level. on a variety of other soils, although probably
Notably, in Hampshire, rather than being in still with an alkaline influence. It is tolerant of
ancient woodland many of its sites are along both wet and dry conditions, from damp woods
ancient trackways or in secondary woods that or wet scrub in the north and west to dry chalky
were arable fields in the 19th century but were slopes in southern England. Sword-leaved
allowed to ‘tumble down’ to woodland. Helleborine is confined to the lowlands.
In southern England, Sword-leaved
Helleborine favours beechwoods on chalk, but Flowering period
elsewhere it grows under a variety of deciduous Mid-May to mid-June, being at its best in
trees. In the Wyre Forest in Worcestershire, the last two weeks of May, even in Scotland,
it once thrived in coppice cut on a relatively although sometimes still in flower in early July
long, 18-year cycle. It is sometimes found in at exposed sites in the north.
M 27 May, Hampshire. Although sadly much reduced, Sword-leaved Helleborine responds very favourably to appropriate
management, especially good levels of light combined with grazing.
001 systematic.indd 42 29/1/09 11:59:56

Range
Sword-leaved Helleborine has always had
an unusually scattered distribution in the
British Isles but is now very much reduced.
The strongholds are in Hampshire and Argyll,
each with around a dozen sites. Other minor
concentrations are found in Worcestershire (the
Wyre Forest holds a few small populations),
Merionethshire and Cumbria. There are
one or two sites each in West Sussex,
Surrey, west Gloucestershire, Warwickshire,
Montgomeryshire, Caernarvonshire, Anglesey
and, in Scotland, Perthshire, Inverness-shire,
Ross & Cromarty and west Sutherland, and
also Arran, Islay, Jura and Skye in the Inner
Hebrides. In Ireland it is very scattered, with
a handful of sites in Co. Kerry, Co. Clare, Co.
Wexford, Co. Galway, Westmeath and Co.
Donegal. World range: Essentially a European
species but with scattered records to the east,
in northern Iran, the mountains of Central
Asia and through the Himalayas to southwest
China. In Europe it occurs north to 63°N in
Scandinavia, southern Finland and the Baltic
States, east to western Russia (around Moscow)
and south to the Mediterranean, including the
Balearic Islands, Corsica, Sardinia and Sicily,
and also to the Crimea and Caucasus. It is
found in North Africa in Morocco, Algeria M 27 May, Hampshire. The lower flowers can have very
long bracts, but they are always very short on the upper
and Tunisia, in a few places in Turkey and in flowers, a clear distinction from White Helleborine.
Lebanon and Israel.
How to find it
1987-99
1970-86
The Hampshire Wildlife Trust reserve at
pre 1970 Chappett’s Copse holds by far the largest British
population (2,185 flowering plants in 2003)
and is undoubtedly the best place in southern
England to look for this stunning orchid. In
Scotland it can be found at Knapdale and at
Ballachuan Hazelwood (both Argyll & Bute).
DESCRIPTION
Height: 15-65cm but can be as short as 5cm in
very exposed situations.
Stem: Green, with some short hairs on the
upper part and two to four whitish, often green-
tipped sheaths at the base. Stems grow singly.
Leaves: Clear ‘grass’ green with fairly prominent
001 systematic.indd 43 29/1/09 11:59:57

001 systematic.indd 44 29/1/09 12:00:00

veins, strap-shaped, keeled, long and rather Subspecies

narrow, the upper ones with a pointed tip, the None.
lowermost shorter, broader and blunter. The
seven to 20 leaves are spaced alternately up
None.
the stem and arranged in two variably defined
opposite rows that may twist around the stem in
a spiral pattern. They are held at 30°-60° above
BIOLOGY
the horizontal and arch gracefully outwards. Pollination and reproduction
Spike: Rather loose, with three to 15 flowers, The flowers are pollinated by solitary bees.
sometimes 25 or very exceptionally even 40, the Sword-leaved Helleborine produces no nectar
lower are held at 30°-45° above the horizontal, and relies on deceit to attract its pollinators;
the unopened upper buds tending to be nearer the golden-yellow pseudopollen on the lip
the vertical. may be especially important in luring insects.
Bract: Green, narrow and pointed. On the lowest However, for the deceit to function effectively
few flowers the bract can be either very long and there must be a good supply of genuinely
leaf-like or much shorter, only half to two-thirds nectar-producing flowers in the vicinity.
the length of the ovary, but on all plants the bracts Furthermore, bees will only visit flowers in
are very short at the tip of the spike. bright sunlight and forage mostly between
Ovary: Green, long and slender, prominently 10am and 3pm, and both the helleborines and
twisted and boldly six-ribbed, with short hairs; the surrounding nectar-producing plants have
the very short stalk is minutely hairy. to be in sunlight for some of that period; sites
Flower: Large and white with a golden-yellow that catch the morning sun may be especially
patch on the lip. The sepals and petals are a favoured. Pollination rates in Britain are very
delicate white, the sepals lanceolate and pointed variable, and a study in Hampshire found that
and the petals slightly shorter and broader. The an average of 35% of flowering plants in sunny
vast majority of flowers do not open widely, glades produced at least one seed capsule;
the sepals and petals cupping the lip with the under a high canopy this fell to 16% and in
tips of the sepals flared outwards, but in some scrub to just 7%. The maximum recorded in
plants the lateral sepals are held spreading. The any one season was 56% of flowering plants in a
lip is white, short and broad and divided into
inner (hypochile) and outer (epichile) halves
by a distinct narrowing or ‘waist’. The hypochile
and base of the epichile are held parallel to the
column with their sides curved upwards and
inwards to form a deep gutter. The epichile is
spade-shaped with a projecting central lobe or
‘tooth’. The base of the hypochile is washed a
rich golden-yellow, and towards the tip of the
epichile there is an extensive patch of dense
papillae with a golden-yellow frosting, although
the fringe of the lip remains white. Towards the
base of this golden-yellow patch there are five
to six (rarely seven) parallel longitudinal ridges
that are also washed golden-yellow. The column
is whitish, long and projects forward (like a M 27 May, Hampshire. The flowers sometimes open
more widely, showing the yellow pseudopollen on the lip.
boat’s figurehead) and the anther is tinged pale-
yellowish. There is no scent. O 27 May, Hampshire.
001 systematic.indd 45 29/1/09 12:00:01

There is no information from wild plants on

the length of the period between germination
and the first appearance above ground but it
may be four or more years. In the laboratory the
interval is two and a half years. Plants need to
have reached a certain height and minimum leaf
area before they can flower but do sometimes
bloom in their first year above ground.
Sword-leaved Helleborine is thought to lack
any method of vegetative propagation.
Hybrids
C. x schulzei, the hybrid with White
Helleborine, has been recorded very rarely from
Hampshire and West Sussex.
The specific name longifolia means simply
‘long-leaved’.
HISTORY AND
CONSERVATION
The first British record was published by
Christopher Merrett in 1666 in his Pinax
Rerum Naturalium Britannicarum (‘A picture
M 27 May, Hampshire. Some plants are very robust. of British natural history’): ‘In Helk wood in
Yorkshire, not far from Ingleborough’.
sunny glade, although many of these may have Sword-leaved Helleborine is classified as
produced just one seed capsule. As well as sunlit Vulnerable in Britain and is specially protected
sites, higher rates of pollination are associated in Eire under the Flora (Protection) Order. It
with larger congregations of flowers, and it may is the subject of a Plantlife International ‘Back
be that the spectacle of many helleborines in from the Brink’ project.
flower is more attractive to bees. Conversely, at
many sites, especially where there are very small Past and present occurrence of Sword-leaved Helleborine in
populations, virtually no seed is produced. Britain and Ireland (based on presence or absence in 10km
squares of the National Grid; data from the New Atlas).
Self-pollination has not been recorded and is
Britain Ireland
unlikely due to the structure of the flower.
Development and growth 1500-1999
Poorly known. The aerial stems grow from
current range 34 (1.2%*) 7 (0.7%*)
a rhizome that produces long and sparsely
branched roots, with fungi concentrated in the % lost, 1500-1969 60.5% 68%
tips of the roots and in the few short side roots. % lost, 1970-1986 13.5% 9.5%
Plants are able to remain ‘dormant’ underground % lost, total 74% 77.5%
for about one year. Nothing specific is known * current range as a % of the total number of 10km squares
about its fungal associates but it seems
probable that like Red and White Helleborines Sword-leaved Helleborine has been in decline
it acquires a substantial proportion of its throughout the 19th and 20th centuries, initially
nutrients via a fungal route. due to woodland losses and the replanting of
001 systematic.indd 46 29/1/09 12:00:02

woodlands with conifers. At a few sites, small

populations were given the final death-knell ExtINCt?
by collectors. The decline has continued in (the date of the last record is given where known)
the last 30 years, probably due to the lack Devon (before 1939)
of woodland management. This has led to Dorset (1876)
Wiltshire (1958)
the disappearance of glades and rides and
East Sussex (1982)
the development of a denser canopy, and the Kent (1980s)
management that does take place often involves Oxfordshire (1970)
clear-felling large areas at once. The loss of Norfolk (1883)
nearby hedgerows and permanent pastures may Herefordshire (1986)
Shropshire (last date uncertain)
have indirectly affected the species too, reducing
Staffordshire (before 1901)
the populations of suitable pollinating bees. Derbyshire (before 1903)
Other threats include heavy browsing by deer, Nottinghamshire (before 1839)
now commoner in Britain than since the time Lancashire (1898)
of William the Conqueror. Yorkshire (1941)
Sword-leaved Helleborine can now be found Co. Durham (1882)
Northumberland (1841)
at less than 50 sites, with around 20 each in Monmouthshire (1987)
England and Scotland and five in Wales. Cardiganshire (1985)
Research in the 1990s showed that 31% of sites Montgomeryshire (1978)
supported just one plant and 61% held less than Dumfries-shire (1854)
ten. Many are likely to be long-lived individuals Ayrshire (1972)
Midlothian (1892)
clinging on at a site no longer suitable for Fife (1847)
reproduction, and the prospects of such tiny Co. Cork
populations surviving are bleak. By 2003 Co. Carlow
only nine sites in Britain held more than 50 Co. Wicklow
plants, and in Ireland the species was formerly Co. Dublin
Co. Tipperary
recorded from 15 counties but is now extinct in
Co. Roscommon
ten of them. Co. Mayo
It is difficult to fashion effective Co Down
conservation measures for this orchid. Its Co. Antrim (prior to c. 1840)
long-term survival depends on adequate levels
of flowering and fruiting and the subsequent to suppress rank vegetation and protection
recruitment of enough new plants to the from deer and rabbits when flowering and
population. It will flower well once the light fruiting. Recent successes give hope for this
levels reach a critical threshold (which is rather very special orchid: at the Little Shoulder of
lower than the levels produced, for example, Mutton in Hampshire, 31 plants were found in
by coppicing). Its pollinating bees require 1987 after Beech trees and scrub were cleared
higher levels of sunlight, however, and at these to increase the size of an existing small patch of
higher light levels there is a tendency for rank chalk grassland. A programme of further scrub
vegetation and scrub to take over and this will control, mowing and protecting individual
overwhelm the helleborines. ‘Nutrient creep’ plants with wire netting resulted in a negligible
from surrounding agricultural land can also increase, but dramatic results came when winter
promote the growth of rank vegetation that can grazing was introduced, with a jump to 240
smother the orchids. plants by 2004. It now seems to be the ‘perfect’
The most effective management regime site with a sunny, south-facing scrubby edge
probably comprises limited felling to produce a adjoining chalk grassland (which can provide
mosaic of woodland and glades, winter grazing nectar and nest sites for the pollinating bees).
001 systematic.indd 47 29/1/09 12:00:02

wHite HeLLeborine Vulnerable, BAP

Cephalanthera damasonium
This subtly attractive orchid is relatively common in beechwoods in southeast England and
is sometimes almost the only flowering plant to be found beneath the dense canopy of the
trees. It has recently been found to have a special relationship with nearby trees, extracting
nutrients from them via a mutual fungal ‘partner’.
Identification
The loose spike of rather egg-shaped, upward-
pointing, creamy-white flowers, most of which
do not open widely, is very distinctive. Most
flowers set seed, and the stout, elongated cap-
sules, held upright, can identify the species late in
the season, well after the flowers have withered.
Similar species
Sword-leaved Helleborine is much rarer but is
sometimes found together with White
Helleborine. The flowers of Sword-leaved
Helleborine are always pure white and tend to
open more widely. In addition, the sepals have
more pointed tips, and the leaves, especially the
lower ones, are on average rather longer and
narrower. The best distinction is the length of the
bracts: on Sword-leaved Helleborine these are
shorter than the ovary, at least in the upper part
of the spike (on the lowest two or three flowers
they may be very long and leaf-like); on White
Helleborine the bracts are all longer than the ovary.
Habitat
Strictly confined to well-drained calcareous
soils on chalk and limestone and found in
woodland and shelter belts, sometimes in scrub
and occasionally on nearby grassland (especially
on north-facing slopes). It tolerates quite deep
shade, but the most robust plants are found
where the shade is not too intense. It is strongly
associated with Beech, even solitary trees, and
can happily grow in the dense shade cast by
this species. Its truly classic habitat is a beech
hanger on a steep slope with a sparse or
non-existent ground cover, the helleborines
M 26 May, Cambridgeshire. The flowers are usually
creamy and egg-shaped, with the sepals and petals hardly growing through a carpet of dead leaves or on
opening at all. bare stony or mossy ground.
001 systematic.indd 48 29/1/09 12:00:03

WHITE HELLEBORINE l 49
Flowering period
1987-99
Mid-May to late June, exceptionally from late 1970-86
April or to mid-July, with flowers in the open or pre 1970
in very dry woods being the earliest.

Range
Southern England, especially the North and
South Downs, the chalk of Hampshire and
Wiltshire, and the Chilterns and Cotswolds.
Ranges west to Dorset, Somerset and
Gloucestershire, south to the Isle of Wight
and north to Herefordshire, Worcestershire,
Northamptonshire and Cambridgeshire. World
range: Almost confined to central and southern
Europe, extending north to Denmark, Gotland
in southern Sweden and the Baltic States, east
to Belarus and the western Ukraine and south
to the Mediterranean, including the Balearic the southeast include Beechwood in Cambridge-
Islands, Corsica, Sardinia, Sicily and Crete, and shire and Brown’s Folly in ‘Avon’, and at Friston
to the Crimea and Caucasus. Also found in Forest in East Sussex it is rapidly colonising a
Turkey, Syria, Israel and northern Iran. large area of beech plantations.
How to find it
Usually an easy species to find in suitable habitat, DESCRIPTION
although often growing well away from paths Height: 8-67.5cm but typically 15-40cm, and
and tracks in heavy shade in the depths of the many are just 13-18cm tall with only one or
forest. Notable sites away from its heartland in two flowers.
M 26 May, Cambridgeshire. White Helleborine has a special association with Beech trees, extracting nutrients via a
mutual fungal link; it may occur in large numbers where little else can grow.
001 systematic.indd 49 29/1/09 12:00:04

001 systematic.indd 50 29/1/09 12:00:08

Stem: Green, the upper part is ridged and

either hairless or slightly hairy. There are one
to three brownish, membranous sheaths at the
base, the upper of which is sometimes tipped
green. Stems grow singly.
Leaves: Oval to broadly lanceolate, tapering to
a moderate point and becoming narrower and
more bract-like towards the spike; the lowest
leaf is rather short and cowl-like. The three to
five leaves are placed alternately up the stem,
more or less in two opposite rows, and curve
gracefully upwards to lie horizontally. They
are greyish-green, sometimes with a bluish
tinge, and have prominent veins (especially
on the underside).
Spike: Loose, with one to12 relatively large
flowers, sometimes as many as 16, most
pointing vertically upwards.
Bract: Greenish, narrowly lanceolate and
relatively long, often much longer than the
flower, but becoming shorter towards the tip of
the spike, although still longer than the ovary.
Ovary: Pale green, either not twisted or only
moderately so, slender, cylindrical and boldly
six-ribbed.
Flower: Relatively large and creamy-white.
The sepals are white to creamy-white with a
hint of green. They are tear-shaped to oval-
lanceolate with the broader end at the base and
a blunt tip. The petals are slightly shorter and
more oval in shape. The lip is short and broad
and divided into inner (hypochile) and outer
(epichile) halves by a distinct narrowing or
‘waist’. The hypochile and base of the epichile
are held parallel to the column with their sides
curved upwards and inwards to form a gutter,
the base of which is washed golden-yellow. The
epichile is heart-shaped, rather broader than M 31 May, Cambridgeshire. Although often forming a tall
spire, the shape of the spike is variable.
long, with the tip curving gently downwards.
Towards the tip of the epichile there is an O 27 May, Hampshire.
extensive patch of golden-yellow frosting that
column, but in a few plants the lateral sepals
has three to five longitudinal ridges and furrows
spread apart like outstretched arms and the
towards its base; these ridges are also washed
upper sepal and petals then form a loose hood.
golden-yellow. The column is long, slender and
There is no scent.
whitish, with the anther tinged pale yellow. The
flowers do not normally open widely, the sepals, Subspecies
petals and lip forming an egg-shape around the None.
001 systematic.indd 51 29/1/09 12:00:09

flower is fully fertilised, the lip folds up again to

shut the ‘door’. The mechanism is very efficient,
unlike Red and Sword-leaved Helleborines,
and almost all flowers produce seed. Vegetative
propagation also occurs. The roots produce
buds, and these develop into new aerial shoots.
Development and growth
The aerial stem grows from a deeply buried,
short, woody rhizome. This sends up to 95
roots vertically down into the soil; these are
thick, corky and 30-60cm long. The mature
plant was thought to be independent of fungi,
but recent isotope studies have shown that
it acquires the majority of its nitrogen and
roughly half its carbon from a fungal ‘partner’.
This undoubtedly explains the ability of
M 31 May, Cambridgeshire.The spout-like tip of the lip is
revealed. White Helleborine to thrive in densely shaded
sites where it is sometimes the only green
Variation and varieties plant present. Albino, chlorophyll-less White
Var. chlorotica is deficient in chlorophyll, and Helleborines are sometimes found and these
its stem and leaves are very pale green or even must be fully mycotrophic and completely
yellowish-white. It has rarely been recorded. dependent on their fungal ‘partner’.
Although it appears to form
BIOLOGY relationships with a wide range of fungi,
Pollination and reproduction

Probably largely self-pollinated in a process
that was described by Charles Darwin. The
anther opens while the flower is still in bud
and releases the pollinia, which sit on top
of the column (the flower is held pointing
upwards) with their front edge resting against
the stigmatic zone. The pollinia are very friable
and at least some of the pollen adheres to the
stigma and effects pollination. Once the flower
is mature, however, the outer part of the lip
folds down to form a landing platform for
insects, especially bees. These are attracted by
the golden-yellow mass of pseudopollen on
the tip of the lip (the ridges are said to taste of
vanilla and are often gnawed by some creature;
Darwin found minute, bitten-off fragments but
could not be sure of the culprit). It seems that
visiting insects may not only spread more pollen
onto the stigma within the flower but also M 31 May, Cambridgeshire. Rather rarely, the flowers
open widely, showing the yellow pseudopollen and raised
sometimes transfer pollen from flower to flower, ridges on the lip and the long column projecting into the
thus causing cross-pollination. But, once the flower.
001 systematic.indd 52 29/1/09 12:00:14

White Helleborine preferentially forms Nottinghamshire, Derbyshire, Warwickshire,

associations with ectomycorrhizal fungi, Essex and Devon. This species is still, however,
especially Basidiomycetes; these in turn have relatively common within its core range and
a relationship with nearby trees. Thus, like can occur in large numbers. Furthermore, it
Coralroot and Bird’s-nest Orchids, it gets can colonise newly available habitats, such as
nutrients from the roots of these trees via the maturing beech plantations.
fungi but unlike the trees, it ‘cheats’ the fungus
by giving nothing in return. In view of this, the Past and present occurrence of White Helleborine in Brit-
well-known association of White Helleborine squares of the National Grid; data from the New Atlas).
with Beech takes on a more sinister significance.
Britain Ireland
After germination the seedling is reported
to spend eight years underground before total historical range, 233 0
1500-1999
sending up aerial shoots, and flowers are not
produced for another two or three years. There current range 136 (4.8%*) 0
is, however, probably little direct evidence for % lost, 1500-1969 33%
these timings. % lost, 1970-1986 8.5%
Hybrids % lost, total 41.5%
C. x schulzei, the hybrid with Sword-leaved * current range as a % of the total number of 10km squares
Helleborine, has been recorded rarely from
Hampshire and West Sussex.
The origin of the specific name damasonium is
obscure. The usual derivation has no connection
whatsoever with orchids: ‘damasonium’ was used
by Pliny (AD 23-79, author of the encyclopedic
Natural History) for ‘Alisma’, which in turn was
a name given by Dioscorides in around AD 64
to a ‘plantain-leaved water plant’. An alternative
derivation is from damaso meaning to ‘subdue’,
i.e. subdue evil, as the plant was considered an
antidote to the venom of toads.
HISTORY AND
CONSERVATION
The first British record was published in
1670 when John Ray noted this species in
his Catalogus Plantarum Angliae et Insularum
adjecentium (‘A catalogue of plants found
around Cambridge’), ‘In the woods near
Stokenchurch, Oxfordshire...’
White Helleborine has been lost from
over 40% of its historical range, largely due to
woodland clearance and coniferisation, and
is classified as Vulnerable. Many of the losses
have been on the edges of the range, and it is M 31 May, Cambridgeshire. Plants may be tiny, with just
now extinct in southern Yorkshire, Shropshire, one flower (this was 11cm high).
001 systematic.indd 53 29/1/09 12:00:14

GenUs NEOTTIA
bird’s-nest orCHids
and twaybLades
001 systematic.indd 54 29/1/09 12:00:18

GENUS NEOTTIA l 55
This genus contains two apparently very different groups, the bird’s-nest orchids and the
twayblades, but they are united by a very similar flower structure and almost identical
pollination mechanism.
Distribution Pollination
Europe, temperate Asia and North America, All three British species produce nectar and
just creeping into North Africa. There are have a ‘hair-trigger’ mechanism. This fires a
c. 33 species in this genus, but only three occur drop of glue onto the head of a visiting insect,
in Europe, all of which are found in Britain and at the same time the pollinia are released.
and Ireland. Pollination is efficient and seed-set is high.
Classification Growth pattern
The genus can be conveniently divided The aerial stem grows from a rhizome that
into two groups. There are nine species of puts out numerous roots. The growth pattern
bird’s-nest orchids from Europe and Asia, is sympodial, and each year the rhizome grows
all of which lack green leaves and are fully upwards to form a new aerial stem that may
dependent on fungi throughout their life, flower and always dies off. Underground the
and 25 species of twayblades, all of which rhizome continues to grow from a lateral bud at
have a pair of green leaves held opposite each the base of the aerial stem.
other roughly midway up the stem. Until Fungal partners
very recently the twayblades were placed in In adult plants the roots have a heavy fungal
a separate genus, Listera, although they had ‘infection’, and Bird’s-nest Orchid is dependent
long been considered close to the bird’s-nest
on fungi throughout its life and is fully
orchids (the original Neottia), due to their
mycotrophic.
very similar flower structure and pollinating
mechanism. Recent genetic research has Vegetative reproduction
shown that the two groups are so closely The roots are heavily ‘infected’ with fungi and
related that they should all be in the same are nutritionally independent. They are also
genus, which takes the name Neottia, the older able to form buds at their tips (a facility that is
and more senior of the two names available. unique to orchids) and these can develop into
new rhizomes. Thus, if the central rhizome dies
Floral structures
after flowering (as in Bird’s-nest Orchid) or is
The column has a wide flat rostellum and
otherwise fragmented, each piece can grow into
there are two stalkless pollinia. There is,
a new plant.
however, no detachable viscidium, rather the
rostellum expels its contents in a sticky drop Name
when triggered. There is no spur, and nectar is The generic name Neottia means ‘nest-of-
produced in a central groove or slight hollow at fledglings’, a reference to the appearance of the
the base of the lip. The pollen is grouped into roots of Bird’s-nest Orchid. (The genus Listera
tetrads and these are loosely connected by a few was named after Martin Lister (1638-1711),
weak threads. The pollen is rather powdery. the English doctor and botanist.)
O 20 May, Norfolk. Bird’s-nest Orchid has a complex

relationship with fungi and forest trees, in particular
Beech. It can be hard to spot among the leaf-litter of the
woodland floor.
001 systematic.indd 55 29/1/09 12:00:19

56 l GENUS NEOTTIA
Lesser twaybLade
Neottia cordata
Formerly: Listera cordata: Other names: Heart-leaved Twayblade (North America)

This species occurs widely on wet moorland and woodland in Scotland, as well as in northern
England, Wales, Ireland and on Exmoor. It is very small and often hard to find, but it is well
worth the effort. On close examination one can see that the flower is a perfect miniature,
with every part reproduced exactly.
Identification
This little orchid is very distinctive. It has two
heart-shaped leaves set opposite each other
rather high on the stem and tiny, more-or-less
reddish flowers, each sitting on a large, globular
ovary. On close inspection with a hand-lens the
flowers resemble a tiny elfin figure. The deeply
forked lip forms the ‘legs’, the two hornlike
projections at its base the ‘arms’, while the sepals
and petals spread star-like around the column
to form a ‘hat’ around the ‘head’.
Similar species
None, but there are usually a significant number
of non-flowering plants in any population, with
the paired leaves lying at the tip of the stem.
These are very like young Bilberry plants.
Habitat
Lesser Twayblade is found in two, apparently
distinct, habitats, but both offer the same
combination of cool, humid shade and acid
soils. The first and most frequent habitat is wet
moorland or peat bog, where it grows on the
cushions of moss, usually Sphagnum, found
under and between mature, leggy bushes of
Heather, Bell Heather and Bilberry. The best
conditions are usually found on north-facing
slopes. In the oceanic climate of Shetland it is
sometimes also found on short, heathy pastures.
The second habitat is damp woodland, where
the orchid can be found growing among a
variety of mosses, sometimes in open areas
and sometimes again among an understorey of
Heather, Bilberry and scattered Bracken fronds.
Willow, birch and alder woods are favoured
M 7 June, Merioneth. A tiny plant, sometimes with just
but Lesser Twayblade is also found in ancient a few flowers in a reddish spike; no wonder they are
‘Caledonian’ pinewoods and mature pine hard to spot.
001 systematic.indd 56 29/1/09 12:00:20

LESSER TWAYBLADE l 57
plantations. It occurs up to 1,065m above sea

level (Stob Coire Easain, Inverness-shire), and
most sites are now in the hills, as those in the
lowlands have largely been destroyed.
Flowering period
Mid-May to mid-July, exceptionally from
late April, but generally peaking from late
May. Once the flower has been pollinated the
column quickly withers and blackens but the
remainder of the flower sometimes persists
until September.
1987-99
1970-86
pre 1970
Range
Lesser Twayblade occurs throughout Scotland,
including the Outer Hebrides, Orkney and
Shetland, although it is absent from the central
lowlands. In England it is found from north
Lancashire and Yorkshire northwards, with an
isolated population in southwest England on
Exmoor (Somerset/Devon). In Wales it occurs
very locally from northern parts of Breconshire
and Cardiganshire northwards, including
Anglesey. In Ireland Lesser Twayblade is fairly
widespread from Co. Sligo, Co. Cavan and Co.
Down northwards but is very local in the south:
in Co. Dublin and Co. Wicklow in the east, Co.
Galway in the west, and from Co. Limerick and
Co. Tipperary southwards.
M 7 June, Midlothian. Lesser Twayblade needs cool, moist
There are a few records of Lesser Twayblade conditions and is found on north-facing slopes on damp
from elsewhere in southern England. In moorland or in woodland.
001 systematic.indd 57 29/1/09 12:00:22

58 l GENUS NEOTTIA
001 systematic.indd 58 29/1/09 12:00:24

Hampshire it was recorded in 1853 and 1895 even harder to spot. This species is very local in
near Bournemouth and then it turned up in Wales but one well-known site is Roman Steps
the New Forest near Brockenhurst in 1927-30 (Gwynedd).
and was reported again there in the 1970s (a
record from Bratley in about 1980 is suspected DESCRIPTION
to have been a deliberate introduction). The Height: 3-25cm but usually 5-10cm; tends to
species has also been recorded from Baldwin’s be tallest in sheltered woodland.
Wood in Hertfordshire in 1980 and from Stem: Green or reddish-purple, ridged
Gravetye Woods in East Sussex in about 1975 towards the tip and with fine glandular hairs
and then again in 1989. Some of these records for a short distance above the leaves. There are
are assumed to involve plants introduced one or two membranous, brownish sheaths at
accidentally when pines or rhododendrons the base of the stem. Usually grows singly but
were planted but wind-blown seed is a possible occasionally two or three stems grow from the
source and in Hampshire there could have been same rhizome.
relict populations on the New Forest heaths. Leaves: Two, lying opposite each other,
World range: Found throughout the boreal one third to halfway up the stem and held
regions of the Northern Hemisphere in Europe, either horizontally or at up to 45° above the
Asia and North America, ranging furthest horizontal. They are dark, shiny green and
south in the mountains. In Europe it extends roughly heart-shaped, with a prominent midrib
south to the Pyrenees, northern Italy, northern that terminates in a tiny projecting point
Greece, the Crimea and Caucasus and adjacent (mucro). The leaves are also faintly net-veined
northern Turkey, and north to Greenland, (reticulate) and often have undulating margins.
Iceland and northernmost Scandinavia. In Spike: Relatively open, with three to 20 flowers.
Asia it is found in Siberia eastwards to Lake Bract: Tiny, triangular and greenish.
Baikal and in the Russian Far East, Sakhalin Ovary: Green, spherical, with six reddish ribs,
and Japan. In North America it occurs south held on a reddish or greenish stalk that is a little
to California and New Mexico in the west and longer than the ovary, ribbed and twisted.
along the Appalachians to North Carolina in Flower: Very small and variable in overall
the east. coloration; it usually has a pronounced reddish
tone but can be much plainer and greener. The
How to find it
Locally common in Scotland, it is worth
looking for Lesser Twayblade wherever there
is rank Heather on damp, north-facing slopes,
as well as in suitable woodland, although the
number of spikes can fluctuate markedly from
year to year. It is often to be found under the
Heather, or at least just under its eaves, and
it may be necessary to move the vegetation
aside to see the orchids. Scattered plants can
be exceedingly difficult to find but, fortunately,
the species often occurs in loose colonies and at
least one or two plants may be more obvious.
The key to success is patience and perseverance.
Once a Lesser Twayblade is spotted, a careful
M 7 June, Midlothian.The flowers form tiny, elfin figures.
search of the area will usually produce more,
although a majority may be non-flowering and O 7 June, Merioneth.
001 systematic.indd 59 29/1/09 12:00:25

60 l GENUS NEOTTIA
sepals are greenish, variably washed reddish in in a disc-shaped nectary at the base of the
the centre and around the edges, and are oval lip just below the column; there are two very
with blunt tips. The petals are narrower and short horn-shaped lobes on either side of this
more strap-shaped and tend to be redder. Both nectary and a longitudinal nectar-filled groove
sepals and petals are widely spread and form running from it to the base of the fork. The lip
a star-like pattern around the column. The lip is held pointing downwards, more or less at
is coppery or pale green, washed red, relatively right angles to the column. The column is short,
large and triangular and divided more than stubby and whitish, with a large, thin, leaf-like
halfway to the base into two sharply pointed rostellum that extends forward over the base
lobes. Tiny amounts of nectar are produced of the lip, above which lie the yellow anther cap
and yellow pollinia. The pollinia are shed by
the anther when the flower is still in bud and lie
loose on top of the rostellum, held in position
by its incurved margins. The flowers have a
faint but unpleasant foetid odour, probably
originating from the nectar.
Subspecies
None.
Var. trifoliata has a third leaf above the
usual two. It is rare, but has been recorded in
Scotland.
BIOLOGY
The flowers are pollinated by a variety of small
insects, including flies and gnats, attracted by
the nectar. Three pressure-sensitive hairs project
from the tip of the rostellum and these act as a
trigger. The slightest touch by an insect causes
a droplet of ‘glue’ to be squirted explosively
onto the insect’s head, and the pollinia are
simultaneously released and fall onto this ‘glue’.
The glue dries in just a few seconds and the
pollinia are carried off by the startled insect.
When the flower first opens the flap-
like rostellum physically blocks access to the
stigma and any insect visitor will trigger the
mechanism the moment it touches the hairs on
the rostellum. The rostellum remains in place
once the pollinia have been removed but is now
spread flat, having released the pollinia. This
prevents self-pollination should the pollen-
carrying insect return to the flower immediately
M 7 June, Midlothian. The ovary is roughly spherical and after it has left. About 24 hours later the
almost as big as the flower. rostellum slowly moves upwards, allowing
001 systematic.indd 60 29/1/09 12:00:26

insects to deposit pollen from other flowers Name and classification

onto the stigma, which has now become very The specific name cordata means ‘heart-shaped’,
sticky. Insects are reported to work upwards a reference to the shape of the leaves.
from the bottom of the spike towards the Together with Common Twayblade, this
top. They therefore start with the lowest and species was formerly placed in the genus Listera,
most mature flowers (i.e. those likely to have but genetic studies have indicated that Listera
receptive stigmas), before moving on to younger should be united with Bird’s-nest Orchid in the
flowers that still have pollinia waiting to be genus Neottia, as it was in the past.
removed. If this is so, an insect cannot pollinate
flowers on the same plant. If by any chance HISTORY AND
the pollinia are not removed by an insect, the CONSERVATION
rostellum lifts upwards anyway after a few days, The first British record was published in 1666
allowing the flower to be self-fertilised. by Christopher Merrett in his Pinax Rerum
The individual flowers of Lesser Twayblade Naturalium Britannicarum (‘A picture of British
are self-compatible and artificial self-pollination natural history’): ‘Neer the Beacon on Pendle
will produce viable seed, but studies in Hill in Lancashire’.
California suggested that it is not usually self-
pollinating. However, it is possible that, as in Past and present occurrence of Lesser Twayblade in Brit-
Common Twayblade, small quantities of pollen squares of the National Grid; data from the New Atlas).
may occasionally fall from the pollinia on to the
Britain Ireland
stigma or be carried there by tiny insects.
Seed-set is very efficient and the capsules 1500-1999
mature and split open within five weeks. The
current range 454 (16%*) 48 (4.8%*)
lowest, oldest capsules on a spike may be
ripe and shedding seed before the uppermost % lost, 1500-1969 25% 41.5%
flowers are even pollinated. Indeed, the capsules % lost, 1970-1986 19.5% 8.5%
swell so quickly and seed is produced so % lost, total 44.5% 50%
efficiently that some authors suggest that self- * current range as a % of the total number of 10km squares
pollination must occur routinely.
Lesser Twayblade also reproduces Since 1666 the species has disappeared
vegetatively, from buds on the roots. from many sites, especially in the lowlands
and on the periphery of the range in northern
Development and growth England, where the drainage and reclamation
The aerial stem grows from a slender, creeping of bogs and heaths caused many losses in the
rhizome that lies near the surface of the soil. 19th century. Lesser Twayblade has gone from
This puts out a few long, slender, hairy roots. 44.5% of its historical range in Britain and
Usually only one stem is produced, but buds 50% in Ireland, with a rather large proportion
can form at the tip of the roots and develop of the losses in Britain being comparatively
into additional aerial shoots that flower in recent. Given that it has been better recorded
their third year. Lesser Twayblade is apparently in recent years because more people are actively
short-lived and there is little information on the looking for it, the actual decline has surely been
interval between germination and flowering, substantially greater. It is extinct in Shropshire,
although the first green leaf is reported to Derbyshire, Cheshire, Flintshire, southern parts
appear after two or three years of underground of Yorkshire and the Isle of Man, and probably
development. also Montgomeryshire.
Hybrids
None.
001 systematic.indd 61 29/1/09 12:00:26

62 l GENUS NEOTTIA
Common twaybLade
Neottia ovata
Formerly: Listera ovata

This rather unassuming orchid is one of the commonest and most widely distributed species
in Britain and Ireland. Its small, green, inconspicuous flowers belie the fact that it is very
long-lived and has an intricate and efficient means of using insects as pollinators.
Identification
Straightforward. It is green or greenish-yellow
overall with two large, egg-shaped leaves held
opposite each other at the base of the stem
and a tall spike of small flowers, each of which
resembles a tiny green figure. Non-flowering
plants, with just two leaves opposite each other
at the tip of the stem, are fairly frequent.
Similar species
Lesser Twayblade is rather similar but tiny, with
heart-shaped leaves, and flowers that are usually
reddish and have sharply pointed tips to the
lobes of the lip.
Man, Frog, Fen and Bog Orchids have
greenish flowers but all of these differ markedly
in the structure of the lip. Man and Frog
Orchids also differ in having a basal rosette of
leaves and Bog Orchid has merely tiny clasping
leaves at the base of the stem.
Habitat
Possibly more varied than any other British
orchid. It is found on short chalk grassland,
machair, dune slacks, limestone pavements,
permanent pastures, road verges and fens, and
also in scrub, hedgerows and moist deciduous
woodland, sometimes in deep shade. It has a
preference for calcareous soils but will grow in
mildly acidic conditions, occasionally amongst
Bracken and Heather. It can sometimes be
found in relatively new habitats, such as disused
railway lines, quarries and sand-pits or in
plantations, even of pine. It occurs up to 670m
above sea level (Ben Lawers, Perthshire).
M 29 June, Hampshire. Twayblades have two leaves and
Flowering period the English name originates from ‘tway’, an archaic and
obsolete word for ‘two’.
Late April to early August, latest in the north,
exceptionally even to September. P 23 May, Co. Clare.
001 systematic.indd 62 29/1/09 12:00:27

COMMON TWAYBLADE l 63
001 systematic.indd 63 29/1/09 12:00:29

64 l GENUS NEOTTIA
Stem: Green to mid-brown, thicker and whiter

1987-99
1970-86
at the base with two or three membranous,
pre 1970 scale-like basal sheaths and numerous short,
white, glandular hairs. The stems usually
grow singly, although not infrequently there
are clusters.
Leaves: The two large, egg-shaped or elliptical
sheathing leaves are held opposite each other
towards the base of the stem; sometimes they
lie flat and sometimes they are angled at up to
45° above the horizontal. The leaves are green
in the shade and more yellowish-green in sunny
places, with three to five prominent veins. There
are also one to three tiny, triangular, bract-like
leaves higher on the stem.
Spike: Loose to fairly compact, with 15-30
small green flowers, sometimes as many as 100.
Range Bract: Lanceolate and short; usually shorter than
Occurs almost throughout Britain and Ireland, the flower stalk and sometimes much shorter.
including the Outer Hebrides and Orkney, but
absent from Shetland, much of the Scottish
Highlands and many coastal regions of southern
Ireland. World range: Essentially a European
species but the range extends into Asia. In
Europe it occurs north to Iceland and c. 70°
N in northern Scandinavia and south to the
Mediterranean, including Corsica, Sardinia,
Sicily and Crete, and to the Crimea and
Caucasus. In Siberia it ranges to c. 95° E,
and in southern Asia it is found in Turkey,
northern Iran, the Altai Mountains and western
Himalayas. Introduced in southern Ontario in
Canada.
How to find it
One of the commonest and most widespread
orchids and usually fairly easy to find, although
in regions away from chalk and limestone
soils it is often rather local. The best places to
look will be in ancient woodlands, marshes
or in grasslands that have not been ‘improved’,
ploughed or cut too closely. Being green,
including the flowers, odd plants can be
inconspicuous but Common Twayblades are
often found in large numbers.
DESCRIPTION M 15 June, North Uist. Twayblades are found in a wide

variety of habitats, from woodland to chalk downland or
Height: 10-75cm but usually 20-60cm. (as here) coastal machair.
001 systematic.indd 64 29/1/09 12:00:31

COMMON TWAYBLADE l 65
Ovary: Green, short and rounded, with six

prominent reddish-brown ribs. The reddish-
brown flower stalk is twisted and a little longer
than the ovary; both are variably hairy.
Flower: Small, green and vaguely man-like.
The sepals are bluntly oval and dull green,
sometimes tinged or fringed dull reddish
or brown. The petals are dull green and
narrower and more strap-shaped. Both sepals
and petals curve inwards to form a very
loose hood around the short, thick column.
The lip is yellowish-green, long and strap-
shaped, sharply folded down and backwards
below the flower and divided for around half
its length into two blunt-tipped lobes (the
‘legs’); a shallow nectar-bearing groove runs
down the centre of the lip to the base of the
fork. The column is greenish. The flowers are
variously described as scented (light, musky or
repellent) or as odourless.
Subspecies
None.
Var. trifoliata has three leaves and is not
uncommon. The third leaf is smaller, more
pointed and placed above or, less frequently,
below the main leaves.
Var. platyglossa has a short lip that broadens M 5 July, Norfolk. The tiny green flowers look very much
into blunt diverging lobes, sometimes with like a human figure.
a small tooth between them. It was formerly extremely sensitive and the slightest touch will
found in dunes in South Wales. trigger it. The startled insect flies away with
the pollinia attached, often to another plant
BIOLOGY altogether. Meanwhile, once the pollinia have
Pollination and reproduction been removed, the rostellum bends forwards
Pollination was intensively studied by C. K. and downwards, thereby hindering access to
Sprengel (published in 1793) and later by the stigma and preventing self-pollination. It
Charles Darwin. The flowers are pollinated then slowly shrinks away upwards again to
by small insects, especially ichneumon wasps expose the stigma to visiting insects. If they
but also by sawflies and beetles, attracted are carrying pollinia these can then pollinate
by the flowers’ scent. Once it has landed the the flower as they try to get at the nectar at the
insect follows the nectar-filled groove up the base of the lip. The mechanism is efficient and
lip and makes contact with the projecting many flowers set seed. Self-pollination may also
rostellum. The pollinia are then stuck to its occur occasionally; if the pollinia dry out small
head by the sudden secretion or ‘explosion’ fragments of pollen may fall on to the stigma
of a drop of sticky liquid that dries in just and effect pollination. The seed capsules each
two or three seconds. The mechanism is contain an average of 1,240 seeds.
001 systematic.indd 65 29/1/09 12:00:32

66 l GENUS NEOTTIA
study in Sweden 20 out of 29 plants were still

alive after 40 years. Mature plants have been
recorded spending one or two years ‘dormant’
underground and then reappearing.
Hybrids
None.
The specific name ovata meaning ‘egg-shaped’ is
a reference to the shape of the leaves. Together
with Lesser Twayblade, this species was
formerly placed in the genus Listera, but recent
genetic studies have indicated that this should
be united with the genus Neottia.
HISTORY AND
CONSERVATION
Together with Autumn Lady’s-tresses, this
was the first orchid to be recorded in Britain.
In 1548 William Turner noted in his Names of
Herbes: ‘Martagon...in many places of Englande
in watery middowes and in woddes’.
Common, widespread and with a very
catholic choice of habitats, this species would
seem to be well-placed to survive changes
to the countryside. Nevertheless, Common
M 27 June, Norfolk.Vegetative multiplication can produce
clumps of spikes.
Twayblade has vanished from almost 30%
of its historical range in Britain and Ireland,
with a relatively large proportion of the British
Vegetative propagation also occurs, with
losses being recent.
buds on the roots producing new rhizomes, and
group of clones can be formed, sometimes a Past and present occurrence of Common Twayblade in Brit-
dense circular cluster of dozens of plants ain and Ireland (based on presence or absence in 10km
Britain Ireland
The aerial stem grows from a short, thick
rhizome which has large numbers of long, fairly total historical range, 1,869 512
1500-1999
thick roots. Seeds are thought to germinate
current range 1,354 (47.5%*) 362 (36%*)
in spring, and early estimates of the interval
between germination and flowering were seven % lost, 1500-1969 16% 22.5%
to 20 years or more, with the seedling spending % lost, 1970-1986 11.5% 6.5%
the first three or four years underground. These % lost, total 27.5% 29%
estimates may be rather inaccurate, however, * current range as a % of the total number of 10km squares
and in the laboratory plants can produce leaves
less than a year after germination.
Common Twayblade can be extremely long-
lived. The remains of 24 old flower spikes have
been counted on a single rhizome, and in a
001 systematic.indd 66 29/1/09 12:00:33

BIRD’S-NEST ORCHID l 67
bird’s-nest orCHid Near Threatened

Neottia nidus-avis
Spotting a Bird’s-nest Orchid in the woodland gloom is somehow very reassuring; it is a

sign that you are in a special place. The species is locally common in mature woodland in
parts of southern England, but in much of the remainder of the British Isles it is scarce and
very local. One of the three British orchids that have no green leaves and are completely
dependent throughout their life on nutrients provided by fungi, Bird’s-nest Orchid only
appears above ground in order to flower and set seed. Both the English and the scientific
name relate to this subterranean existence, as the roots of this orchid form an untidy mass
that vaguely resembles the nest of a Wood Pigeon or Rook.
Identification
The honey-brown spikes are unique among
British orchids. On careful examination the
flowers are typical of an orchid, with the petals
and sepals forming an open, fan-shaped hood,
and the lip vaguely resembling a human torso.
The dried stem and open seed capsules of the
previous season’s blooms may be found nearby
and these can remain intact for almost two years.
Similar species
Confusion is possible with Yellow Bird’s-nest,
a more-or-less similarly coloured but totally
unrelated plant that is often found in the
same habitats, although the spike of Yellow
Bird’s-nest is bent over and ‘nodding’ until it is
very mature. The various broomrapes (family
Orobanchaceae) also superficially resemble
Bird’s-nest Orchid, especially Knapweed
Broomrape, which is closest in colour. These
chlorophyll-less parasitic herbs are found in
open, grassy habitats and could occur on the
edge of woods or in woodland rides. A quick
look at the structure of the flower will settle the
matter, however, as neither Yellow Bird’s-nest
nor the broomrapes have a flower with a hood
and a two-lobed lip.
Habitat
The classic habitat for Bird’s-nest Orchid is the
heavy shade of a mature beechwood, the orchids
emerging from the leaf-litter and deep humus of
a woodland floor otherwise devoid of vegetation.
It also grows in mixed deciduous woodland and M 2 May, Oxfordshire.
001 systematic.indd 67 29/1/09 12:00:34

68 l GENUS NEOTTIA
bloom in the latter half of May. Bird’s-nest

Orchids have been recorded flowering and
setting seed underground if the route to the
surface has become blocked by a large stone
or other obstacle. As no systematic searches
have been made, it is not known whether this
is exceptional or a regular event (Australian
orchids of the genus Rhizanthella routinely
flower underground).
Range
Found almost throughout Britain and Ireland
but absent from the Isle of Man, Shetland,
Orkney and the Inner and Outer Hebrides
(with the exception of Skye and Mull); there
are, however, only a few sites in northern
Scotland, much of Ireland and in a belt from
the northern parts of East Anglia through
Lincolnshire and the north Midlands to
north and west Wales. Its strongholds are
undoubtedly southern England (especially
central-southern England) and southeast
Wales. World range: Occurs in Europe
and western Siberia, reappearing in the Far
East. In Europe found north to c. 65°N in
M Yellow Bird’s-nest, 16 June, Norfolk. Another fully Scandinavia and Finland and south to the
mycotrophic (fungus-dependent) species of the wood-
land floor, the similarity to Bird’s-nest Orchid ends with
Mediterranean, including Corsica, Sardinia,
the honey colour. Sicily and the Balearic Islands, and to the
Crimea and Caucasus; it is largely confined to
overgrown hazel coppice or sometimes under the mountains in the south of the range and
shady old hedges, shelter-belts or planted absent from the Mediterranean lowlands. In
conifers, especially if there are still deciduous
trees present (and in Europe it regularly grows
1987-99
in conifer woods). Rarely, it has been recorded 1970-86
pre 1970
from grassland just outside woods, but this is
exceptional, as exposure to direct sunlight tends
to dry out the ground too much. Conversely,
it is not found in areas where the soil becomes
waterlogged. Bird’s-nest Orchid is commonest
on chalk and limestone soils but also grows on
clays and sands that have a chalky or limestone
component, such as boulder clay. Generally it is
a lowland species, but it has been recorded up to
250m above sea level in Cumbria.
Flowering period
Early May to late June, exceptionally from
late April or to early July, but most are in
001 systematic.indd 68 29/1/09 12:00:35

Siberia it extends to c. 85°E and it is also found DESCRIPTION

in Algeria in North Africa and at scattered sites Height: 15-52cm but usually 20-40cm.
in Turkey and northwest Iran. In eastern Asia it Stem: Yellowish-brown, slightly glandular-
occurs in the Russian Far East, Sakhalin, Japan hairy towards the tip. Grows singly, although
and Korea. two spikes occasionally develop from the same
How to find it rhizome.
In most of Britain and Ireland the species is Leaves: No green leaves are present. The lower
rather local and only found in small numbers. part of the stem is enclosed by three to five long,
It often emerges in the darker and shadier roughly oblong, scale-like, yellowish-brown,
parts of a wood and this, combined with loosely sheathing leaves (the upper ones longer
its pale-brown colour, can make it hard to and blunter).
find. Bird’s-nest Orchid is totally dependent Spike: Cylindrical and crowded, with up to 100
on the activity of its fungal ‘partners’ and flowers in large plants, but the lower flowers
not surprisingly does best in the warm, wet in the spike are usually more widely spaced
conditions beloved of fungi in general. Warm, and there are odd single flowers further down
wet springs can encourage larger numbers the stem.
of plants to flower, and conversely very dry Bract: Papery, lanceolate and roughly as long as
periods can result in a reduction in the above- the ovary and stalk together, but inconspicuous.
ground population. Ovary: Oval, subtly six-ribbed, glandular-hairy
M 3 June, Norfolk.Whether through seed or vegetative reproduction via division of the underground rhizome, Bird’s-nest
Orchid sometimes occurs in groups.
001 systematic.indd 69 29/1/09 12:00:38

70 l GENUS NEOTTIA
BIOLOGY
Pollination is by insects, including flies,
attracted by the nectar. The mechanism is
very similar to that of Common Twayblade.
The visiting insect makes contact with the
projecting rostellum where there are six minute,
rough, touch-sensitive points and the pollinia
are stuck to its head by the sudden secretion
or ‘explosion’ of a drop of sticky liquid. After a
while the rostellum, which has hitherto blocked
access to the stigma, rises to allow visiting
insects, complete with pollinia attached to their
heads, to make contact with the stigma. If the
mechanism is not triggered, after a few days
the pollinia fragment and pollen can then fall
onto the stigma below, effecting self-pollination
(autogamy); pollen may also be carried to
the stigma by small insects such as thrips.
M 26 May, Norfolk. Comparison of the flower structure Ants have been noted carrying pollen from
with Common Twayblade shows how close they are. one flower to another on the same spike and
this may also effect self-pollination (this time
and held on a twisted stalk that is about half
geitonogamy, as it is pollen from a different
the length of the ovary.
flower on the same plant). Occasionally, self-
Flower: Entirely yellowish-brown. The sepals
pollination may take place in the bud before the
and petals are roughly oval-spatulate in shape
flowers have even opened. Pollination is very
and form a loose fan-shaped hood over the
efficient and almost all flowers will set seed.
column. The lip is slightly darker brown and Bird’s-nest Orchid is thought to be
has a nectar-producing bowl-shaped depression monocarpic, that is the plant dies after
at the base (representing a rudimentary spur). flowering once. But, although the rhizome dies,
The lip is divided towards the tip into two the numerous roots can remain alive and go on
broad, rounded lobes that spread widely, to produce new plants from buds at their tip.
especially on the lower flowers, to form a lyre-
shape; there may also be a subtle point or tooth Development and growth
on either side of the lip half way towards the The aerial stem develops from a short rhizome
base. The lip is held pointing outwards and that lies horizontally in the soil and is almost
downwards at c. 90° to the column, which is entirely surrounded by an untidy mass of short,
pale brown, long and slender. The pollinia are thick, fleshy roots that stick out more or less at
yellow and project conspicuously from beneath right-angles.
the anther cap. The flowers have a pleasant but Throughout its life Bird’s-nest Orchid is
sickly, honey-like scent. entirely dependent on fungi for nutrition;
in adult plants fungi are found exclusively in
Subspecies the roots of the orchid (in the three cortical
None.
cell layers, just below the epidermis). Recent
Variation and varieties studies have shown that the orchid is very
Var. pallida has a yellowish-white stem and specific about its fungal ‘partner’ and only
flowers, and white pollinia. It is rare. forms an association with a species of Sebacina.
001 systematic.indd 70 29/1/09 12:00:39

This obtains its carbohydrates by forming a

symbiotic ectomycorrhizal association with
the roots of trees, particularly Beech: the tree
produces carbohydrates through the process of
photosynthesis and passes these to the fungus,
which in turn contributes mineral nutrients to
the tree. The orchid invades this relationship
and, via the fungus, gets nutrients from the tree.
Bird’s-nest Orchid ‘cheats’ in its partnership
with the fungus because, unlike the tree, it
does not contribute anything to the fungus
(see also p.9).
It is often stated that Bird’s-nest Orchid
lacks chlorophyll, but a small amount of
chlorophyll is present, although no effective
photosynthesis can occur. It is also often said
that Bird’s-nest Orchid is saprophytic and
obtains its nutrients from decaying organic
material. This is not correct. The orchid gets its
nutrients by digesting the living fungus.
Seeds germinate in the spring and require
the presence of the same fungus that supports
the growth of the adult plant. The seedling
initially takes the form of a torpedo-shaped
protocorm a few millimetres long. It then
begins to develop short, fat rootlets that stick
out at about 90° and start to take on the
appearance of a ‘bird’s nest’. At an early stage
M 13 May, Norfolk. When shoots emerge in the spring
the bud that will produce the flower spike the flower buds are already fully formed.
appears in the axil of a scale leaf at the tip of
the rhizome. As the rhizome grows the number exact spot that previously held a flowering plant
of roots progressively increases and the period may be the product of this, or alternatively, of
from germination to flowering is probably three the break-up of a single rhizome to produce
to five years. several new plants.
Seed can only germinate where the Sebacina
Hybrids
fungus is present and this may have a localised
None.
and restricted distribution, even within a
single wood, at least partly because the fungus Name and classification
is entirely dependent on its host tree species The specific name nidus-avis means ‘bird’s nest’
and cannot grow without it. The distribution (as the generic named Neottia means ‘nest-of-
of the orchid is therefore also tied to that of fledglings’ the scientific name is a tautology).
suitable host trees. Adult Bird’s-nest Orchids With a similar flower structure and
always harbour the correct fungus and form pollination mechanism, the Bird’s-nest
a convenient source of ‘infection’ for the seed, Orchid is clearly related to the twayblades.
and germination is far more prolific in the DNA studies have recently shown that the
immediate vicinity of adult plants. The clusters relationship is a close one, and they are all now
of Bird’s-nest Orchids that often grow on the placed in the same genus, Neottia.
001 systematic.indd 71 29/1/09 12:00:41

72 l GENUS NEOTTIA
HISTORY AND
CONSERVATION
First recorded from Britain in 1597 by John
Gerard in his Herball: ‘I found it growing in
the middle of a wood in Kent two miles from
Gravesend.’
With its requirement for heavy shade and a
stable, generally moist environment, the species
has undergone a significant decline, especially
in the period after 1945. This is due to the
grubbing out of woodland and the conversion
of deciduous woodland to conifer plantations.
More subtle changes, such as the use of heavy
machinery in forestry operations, could also
have been detrimental. The decline has been
most marked in southeast England, especially
Kent and the eastern Chilterns, and Bird’s-nest
Orchid has gone from 54% of its total historical
range in Britain and 45% in Ireland. As well as
being more widespread, populations may have
been larger in the past. For example, ‘thousands’
were recorded on the south-facing slopes of Box
Hill in Surrey in 1947 at a site which nowadays
only supports about 200 spikes. Bird’s-nest
Orchid is classified as Near Threatened in
Britain and is specially protected in Northern
Ireland under Schedule 8 of the 1985 Wildlife
Order (NI).
Past and present occurrence of Bird’s-nest Orchid in
Britain and Ireland (based on presence or absence in
10km squares of the National Grid; data from the New
Atlas).
Britain Ireland

1500-1999
current range 340 (12%*) 54 (5.4%*)
% lost, 1500-1969 41% 29.5%

% lost, 1970-1986 13% 16%
% lost, total 54% 45.5%
O 6 June, Norfolk.The dead spike may persist for at least

a year.
P 8 July, Norfolk. Marsh Helleborines.
001 systematic.indd 72 29/1/09 12:00:42

GENUS EPIPACTIS
HELLEBORINES (II)
002 systematic.indd 73 12/2/09 16:53:23
74 l GENUS EPIPACTIS
A genus of essentially woodland orchids which has diversified to occupy a variety of

other situations. Recent research has shown that fungi contribute very significantly to
their nutritional budgets, even as adults, with several having connections via their fungal
‘partners’ to neighbouring trees. As a genus, Epipactis is distinctive, with flowers that are
rather different to those of ‘typical’ orchids, but it is necessary to have a good grasp of its
flower structure in order to identify some of the species with confidence.
Distribution
Largely confined to Europe and Asia with single
representatives in Africa and North America.
The number of species is uncertain, with the
most conservative authors listing around 11
species in Europe, whereas Delforge (2005)
details 59. There are eight species in the British
Isles, including two endemics.
Classification
Epipactis is closely related to the genus
Cephalanthera, and, although the flowers look
quite different, both have the lip divided into
outer and inner portions by a narrow ‘waist’.
Epipactis differs in having more complex
reproductive structures and an ovary that is not
twisted; rather it has a distinct stalk which is
twisted to bring the lip around to the bottom
of the flower.
The genus is sometimes divided into two
sections: Arthrochilium, which contains Marsh M Violet Helleborine, 29 June, Buckinghamshire. The
Helleborine, and Euepipactis, which contains the flower’s spike is fully formed when the stem breaks
remainder of our species. In Marsh Helleborine throughthesoilsurfacebutitmaybeseveralweeksbefore
itelongates,becomesuprightandtheflowersopen.
the inner and outer halves of the lip are joined
by a flexible hinge, whereas in all the other very numerous. All the helleborines bloom
species the joint is rigid and the inner portion of from mid- to late summer, later than most
the lip does not form such a distinct bowl. other orchids.
Identification Floral structures
Epipactis helleborines are relatively easy to All the Epipactis helleborines have a similar
recognise as such. They have upright stems 10- flower structure. The ovary forms the
120cm tall and oval leaves with obvious parallel apparent ‘stalk’ of the flower. It actually
veins. In some species, several spikes can arise narrows into the real stalk, the pedicel,
from the same rootstock. When it emerges just before it reaches the main stem. Once
from the soil the stem is bent double and as it fertilisation has occurred, the ovary swells
grows it continues to ‘weep’. Eventually, however, conspicuously to form a capsule while the
the stem becomes fully upright and the flowers, flower shrivels to a few brown wisps. In all
which remain in bud for a frustratingly long species the ovary is usually obviously ribbed.
time, begin to open. In many species the flowers The three sepals and two petals are rather
are relatively small and drab but they may be similar in size, shape and coloration; in
002 systematic.indd 74 29/1/09 12:11:01

GENUS EPIPACTIS l 75
some species they form a five-pointed star stalk

(petiole) ovary
around the column and lip, in others the column anther cap
flowers do not open widely and form a cup
pollinia
or bell shape. In Epipactis the lip is divided
viscidium
into two parts. The inner section is known rostellum
as the hypochile; it is cup-shaped and often bract
boss
contains nectar. The outer part, or epichile,
is flatter, more or less triangular in shape and
hypochile epichile
often reflexed (bent under) at the tip. At the
base of the epichile there are raised areas lip
known as bosses and in some species these MBroad-leavedHelleborineflowerinprofilewithsepals
are contrastingly coloured. The hypochile and andpetalsremoved.
epichile are connected by a narrow strip that
is rigid in most species. is an obvious structure, in Epipactis and other
orchids it is just a region on the front of the
Reproductive structures
column that becomes sticky when it is receptive
The column is a robust structure which projects
to pollen.
from the ovary into the centre of the flower.
In Epipactis it is often conspicuous and can be Pollination
important for identification. It is topped by the In cross-pollinated helleborines, the clinandria
anther or ‘anther cap’, which is attached to the or grooves on top of the column are relatively
remainder of the column by a short stalk or deep and the pollinia lie there securely. The
narrow flexible hinge. The two pollinia develop rostellum functions as an additional barrier,
side-by-side within the anther. Shaped like a preventing the pollinia from falling onto the
fat sausage, each pollinium is in turn divided stigma below. In these species the pollinia
into two segments by a fine groove, and the two remain intact and lie with their tips in contact
pollinia are joined together at one end in the with the viscidium. A wasp visits the flower to
form of a wishbone. drink nectar from the cup-shaped hypochile
Below the anther cap, the top of the column but to reach this it must rub its head or the top
has a system of grooves, the clinandria. These of its thorax against the rostellum. In doing
form a platform on top of the main part of so it ruptures the viscidium and the sticky
the column. As the pollinia mature, the anther contents glue the pollinia to the insect’s head.
splits open and the pollinia drop down to rest Thus, when the wasp flies off, it carries one
in the clinandria, which are moulded to or both of the pollinia away with it. The wasp
their shape. The pollinia remain protected goes on to visit another flower, deposits parts
from above by the anther cap and are often of the pollinia on the wet surface of the stigma
largely hidden. and pollination takes place; once pollinated, the
At the front of the column, separating stigma changes from whitish to brownish.
the clinandria from the stigma below, lies In self-pollinated helleborines, the
the rostellum. (Literally the ‘little beak’, clinandria are shallow and do not hold the
the rostellum is actually a greatly modified pollinia securely, the rostellum is usually
sterile third stigma.) The rostellum secretes reduced in size and the viscidium either never
a detachable white cap, the viscidium. This is develops or is present in the bud but disappears
a thin-skinned sack containing a sticky liquid when the flowers open; there is therefore little
and the slightest touch will rupture the skin to prevent the pollinia, or parts of them, from
and release the ‘glue’, which sets hard on contact falling onto the stigma below. The pollinia also
with the air. Below the rostellum lies the fertile tend to be more friable and easily fall to pieces.
stigma. Unlike other flowers, where the stigma The pollinia may either disintegrate where they
002 systematic.indd 75 29/1/09 12:11:01

lie (the pollen germinating in the clinandrium

and the pollen tubes growing down to the
stigma and ovary), or the pollen may fall on to
the stigma below and pollinate it. Indeed, in
r some Green-flowered Helleborines the flowers
r
never open and pollination occurs within the
unopened bud. Contrastingly, it is probable that
the Dune Helleborine, although self-pollinating,
occasionally has a functional viscidium and is
therefore sometimes cross-pollinated.
In the self-pollinated species, cross-
pollination can occur from time to time, even
without a viscidium. Their flowers, although
MBroad-leavedHelleborine, 28July, Norfolk.Theviscid-
typically small, dull and not opening widely,
iumandpolliniaareconspicuous. are visited by insects in order to drink nectar.
The pollinia may stick to their bodies despite
the lack of the viscidium as ‘glue’ and the insects
may carry the pollinia away with them.
Separation of cross-pollinated and
self-pollinated helleborines
r Four of the Epipactis helleborines are cross-
pollinated: Marsh, Dark-red, Broad-leaved
and Violet. When the flowers first open they
have neat, undamaged, creamy-yellow pollinia
and a well-formed whitish viscidium. Although
the pollinia are nearly concealed by the anther
cap, they can usually be seen poking out
slightly, like fat yellow sausages. After a short
while it is likely that a visiting wasp will remove
MNarrow-lippedHelleborine, 12July, Buckinghamshire. the viscidium and the pollinia, although the
Thisflowerisfreshlyopened,butneverthelesslacksavis- yellow anther cap remains. The flower will
cidium,althoughthepolliniaarepresent.
probably have been fertilised by this stage
and the whole column will start to look tatty
and brown.
Narrow-lipped, Dune, Green-flowered and
Lindisfarne Helleborines are self-pollinated.
They have no viscidium (if it is present, it is
r
almost always small and vanishes before the
flower opens). The pollinia are released by the
anther and sit on top of the column. By the
time the flower opens the pollinia have begun
to swell and crumble, fragments falling over the
edge onto the stigma below. The pollinia are
largely hidden by the anther cap but appear to
‘foam-out’ from under it. The flower, having
M Narrow-lipped Helleborine, 22 July, Oxfordshire.The
polliniaarevisiblydisintegratingin situ, andthereisno been fertilised, starts to go-over and the column
viscidium. turns brown.
002 systematic.indd 76 29/1/09 12:11:05

GENUS EPIPACTIS l 77
Growth pattern developed outer shell or carapace and this slows

The aerial stem grows from a rhizome which the uptake of water and delays germination,
puts out numerous roots, each of which can suggesting that they have a period of dormancy.
live for around three years. The growth pattern Seed probably germinates in spring, forming
of the rhizome is sympodial; the tip of the first a protocorm and then a mycorhizome (the
rhizome grows upwards to form a flower spike earliest and most heavily infected stage in the
that withers and dies off once seed has been set, development of the rhizome). The first roots
while the rhizome continues to grow from one develop in the autumn and, as the rhizome
or more buds that are formed at the base of the grows, fungal activity is transferred to the roots
aerial stem. These buds are formed at least a so the rhizome itself becomes ‘infection-free’.
year before the next stem appears above ground
but may remain dormant for one or more years,
This may take place via several mechanisms.
in which case the plant remains underground
1. The roots develop buds, which go on to
for a year or more. The probability of a plant
form a secondary rhizome (recorded for Dark-
appearing above ground and blooming is
red Helleborine). 2. The rhizome produces
determined by growing conditions over the 12
two flower spikes that each produce buds and
months previously.
adventitious roots below ground, and these
Fungal partners buds go on to produce new rhizomes (recorded
There is disagreement about the extent to for Dune Helleborine). 3. The rhizome
which adult helleborines obtain nutrients from branches (as in Marsh Helleborine). In all these
fungi. Examination of rhizomes and roots led examples, if the central ‘mother’ plant dies off
to a consensus that some species were ‘free’ or the rhizome is broken up in some other way,
from fungi, including Violet Helleborine, two or more new plants may result. However,
which thrives in deep shade. However, several vegetative reproduction is not recorded for
species occasionally throw up variants that several of the helleborines and appears to be
lack chlorophyll (including var. rosea of Violet relatively unimportant for most species.
Helleborine). These cannot photosynthesise
Name
but prosper nevertheless, so at least some
Rather obscure. The name Epipactis was
helleborines are able to gain substantial
first given to a plant used to curdle milk by
quantities of nutrients from fungi. Recently,
Theophrastus (c. 370-285 BC; often regarded as
sophisticated techniques have been developed
the ‘father of botany’). It is not certain what he
to identify fungi genetically and to assess their
was referring to, but it may have been a Veratrum,
contribution to the helleborine’s nutrition. The
a genus now known as ‘false helleborines’ but
evidence produced so far suggests that fungi
formerly called ‘hellebores’. When the German
do play a very significant role. Several Epipactis,
botanist Johann Zinn described the genus
including Broad-leaved and Dune Helleborines,
Epipactis in 1757 he may have been struck by
display consistent associations with one
the resemblance of Broad-leaved Helleborine to
group of fungi, the Ascomycetes. Some
a Veratrum, especially the broad, prominently
Epipactis, such as Broad-leaved and Dark-red
veined leaves, and therefore chose to use
Helleborines, may also have an association
Theophrastus’s name for his new genus.
with ectomycorrhizal fungi and therefore gain
The word ‘helleborine’ has been used in
nutrients from nearby trees via their fungal
English since the 16th century and means
‘partners’, which they ‘cheat’.
a plant resembling a ‘hellebore’; presumably
Development from seed again, a reference to its similarity to a Veratrum
Poorly known, as the subterranean seedlings (rather than the modern ‘hellebores’, genus
are difficult to find. The seeds have a well- Helleborus).
002 systematic.indd 77 29/1/09 12:11:05

Marsh helleborine
Epipactis palustris
This is one of the most attractive orchids, and when examined closely the individual flowers
are simply stunning, being miniature versions of the gaudy hothouse hybrids. Marsh Helle-
borine occurs widely across England, Wales and Ireland, but due to its specialised marshy
habitats it is very local. It has declined significantly but can still occur in large numbers,
especially in coastal dune slacks.
Identification striking purple veins at the base, and the tip

Distinctive. Marsh Helleborine is easily has a distinctive frilled edge.
identified by its colourful, purple and white Similar species
flowers, habitat and mid-summer flowering
None.
period. The flowers resemble those of the
other Epipactis helleborines in shape but Habitat
are relatively large and bright. The lip shows Marsh Helleborine is found in a wide variety
of wet, marshy habitats but requires neutral
to alkaline ground water and relatively short,
open, vegetation to thrive. The two most typical
habitats are dune slacks and spring-fed fens. In
dune slacks the ground water is calcium-rich
due to the presence of shell fragments in the
sand. In spring-fed fens the ground water is
both nutrient-poor and calcareous and such
fens can be found nestled in heathland valleys
or within more extensive acid bogs, as well
as on more obviously chalk or limestone-rich
soils. Marsh Helleborine may also be found in
meadows which are seasonally flooded with
chalky water, but it cannot compete with tall
vegetation and such habitats must be regularly
mown or grazed for it to survive. Sometimes,
it may be found growing among Common
Reeds, but these, too, are likely to take over and
eventually crowd it out. Marsh Helleborine
occasionally grows in other habitats. These
include wet, slumped, clay cliffs (as in Dorset
and the Isle of Wight), gravel pits and fly-
ash pools. Very occasionally it is found in
small numbers on ‘dry’ chalk grassland (as in
Bedfordshire, Kent, Surrey and Wiltshire),
especially where quarrying and excavations have
left a compacted surface prone to becoming
O 8July,Norfolk.Growingonclose-croppedturf,itispos-
sibletoseethestructureofthewholeplant.
002 systematic.indd 78 29/1/09 12:11:06

MARSH HELLEBORINE l 79
waterlogged; it also grows in large numbers on

1987-99
the floor of an old chalk pit in Norfolk where 1970-86
there is standing water in the winter months. pre 1970
Flowering period
Late June to early August, very exceptionally to
early September but mostly in July.
Range
Widespread in England, Wales and Ireland
but often very local and absent from large
areas (much of Kent, Sussex, Surrey,
Wiltshire, Devon, Cornwall, the Midlands,
Co. Durham, Cumbria, mid-Wales, Co. Kerry,
Co. Cork, Co. Waterford and Co. Wexford in
southern Ireland and Co. Antrim in Northern
Ireland). The strongholds are in Norfolk and
Hampshire, but even in these areas it has Colonsay. World range: Europe and Asia. In
declined significantly. Very rare in Scotland Europe it is found north to Denmark, southern
and recently only recorded from a handful of Scandinavia and the Baltic States and south to
sites: Perthshire in the Central Highlands, Portugal, northern Spain, southern Italy, central
Argyll and the Inner Hebrides on Islay and Greece, Bulgaria, the Crimea and Caucasus,
M8July,Norfolk.Summerrainhascausedshallowfloodinginthisabandonedquarry.
002 systematic.indd 79 29/1/09 12:11:08

002 systematic.indd 80 29/1/09 12:11:11
and also Corsica and Sicily, but absent from sepal

ovary
the Mediterranean lowlands. Also found in
Turkey, northern Iran, Central Asia and extends
petals
eastwards to Lake Baikal in eastern Siberia. stem
sepal
bract
How to find it
This is a showy, conspicuous orchid that often sepal
grows in large numbers, especially in coastal hypochile epichile
dune slacks. Notable sites include Lindisfarne
in Northumberland, Sandscale Haws in lip
Cumbria, Ainsdale in Lancashire, Sandwich the lowest one or two are short, egg-shaped
Bay in Kent and Wells-Holkham, Beeston and formed into a cowl around the stem; the
Common and Buxton Heath in Norfolk. remainder are large, pointed, broadly strap-
shaped with a distinct keel, ‘papery’ in texture
DESCRIPTION but rather stiff; these are held erect, c. 45-60°
Height: Usually 20-45cm, occasionally up to above the horizontal, arranged in a spiral
82cm. Dwarf forms, only 10cm high but with around the stem and grade into narrower and
normal-sized flowers, have been recorded, more bract-like non-sheathing leaves towards
especially in dry habitats. the spike.
Stem: Green, flushed with brownish-purple, Spike: Up to 25 flowers, although usually
especially towards the tip, prominently hairy, rather fewer, form a lax spike; all face more
ridged towards the tip and with one or two or less to one side and are initially held
purplish sheaths at the base. Several stems may horizontally but slowly droop.
grow from the same rhizome. Bract: Green, lanceolate and pointed, the
Leaves: Mid-green, sometimes washed purple lowest a little longer than the ovary, the upper
around the base or sheaths, with three to five much shorter and also blunter.
prominent veins. Of the four to eight leaves, Ovary: Brownish-purple, prominently hairy,
O 21July, Norfolk.Apar-
ticularly richly coloured
plant.
P 8 July, Norfolk. The

large boss at the base of
the outer half of the lip
(epichile)andthedelicate
purpleveiningontheinte-
riorofthecup(hypochile)
areclearlyvisible.
002 systematic.indd 81 29/1/09 12:11:13

cylindrical, with six fine ribs, narrowing at each

end and tapering to a short, purple stalk that is
twisted through 180°.
Flower: Purplish with a white, frilled lip
and prominent purple veins; the flowers
open widely and the lip is held projecting
horizontally outwards. The sepals are oval-
lanceolate, the lateral sepals asymmetrical.
On the outer surfaces they are sparsely hairy
and dull greenish-yellow, flushed, veined and
mottled purple, especially towards the base;
on the inner face they are more solidly washed
purple, especially towards the tip, with a neat
whitish border. The petals are slightly shorter
and blunter, white, washed pinkish-purple
towards the base on the outer surfaces (shining
through to the inner surface) and also delicately
veined with purple towards the base on the
inner face. The hypochile is formed into a
dish, the base of which is longer than wide and
yellow, variably blotched with reddish nectar-
producing swellings. The sides, including the
erect, triangular side-lobes, are white with
prominent purple veins on the inner surface.
The epichile is white with a slight purple wash
towards the edges at the base; it is more or less
circular but with the sides turned upwards
and strongly frilled or crimped. At the base of
the epichile there is an irregular boss which is
whitish with yellow edges and bisected by a
deep, narrow groove; this also produces small M 9 July, Norfolk.Var. albiflora.This rare variety lacks
anthocyaninpigmentsandhasagreenstem, ovaryand
quantities of nectar. The hypochile and epichile sepalsandunmarkedwhitepetalsandlip, lackingeven
are connected by a narrow, flexible hinge. The purplebanding.
column is very pale yellowish-white with a
well-developed white viscidium, dull yellow Var. albiflora is similar but lacks the purple
anther cap and primrose yellow pollinia. veins. It is rare.
Subspecies BIOLOGY
None.
Variation and varieties Marsh Helleborine is cross-pollinated but, despite
Var. ochroleuca lacks brown and purple several investigations, there is disagreement as to
pigments and is very pallid. The stem and ovary which insects are the most effective pollinators
are green, the sepals yellowish-white to pale and what role the unusually flexible outer part of
green, and the petals and lip are white, although the lip plays in the mechanism.
the interior of the hypochile still has purple The flowers are visited by a wide variety
veins. It is uncommon, but where it is found it of insects, including flies and beetles. Some of
occurs in large numbers. them, such as hover flies and honeybees, groom
002 systematic.indd 82 29/1/09 12:11:14

themselves extensively and even if they pick up sufficient to bring their heads into contact with
pollinia are not effective in delivering them to the viscidium.
other flowers. Honeybees also feed their young Vegetative reproduction may occur if the
on pollen. Solitary bees and wasps are probably rhizome breaks up into several sections.
the most efficient pollinators, the nectar on the
large boss on the lip acting as a guide to entice
The aerial stems grow from a relatively slender,
the insect into assuming the correct position for
well-branched rhizome that creeps horizontally
pollination. Ants also visit the flowers and may
near the surface of the soil. A single plant may
cause self-pollination, or may carry fragments
have an extensive rhizome and produce several
of pollinia away and pollinate other flowers on
aerial stems. Indeed, it has been claimed that
the same spike (geitonogamy). Self-pollination
over 100 flower spikes may grow from the
may also occur spontaneously as pieces of
same plant (Davies et al. 1983). Roots are
pollinia fall onto the stigma below. An average produced at many points along the rhizome,
of over 80% flowers set seed. both horizontal roots that penetrate the more
Charles Darwin suggested that the outer organic surface layers and vertical roots that
part of the lip hinged downwards due to the often grow deep into the mineral soil. The
weight of the visiting insect, allowing it to roots are reported to have little or no fungal
enter the flower without removing the pollinia. ‘infection’, except in soils deficient in nitrogen.
Once the insect was within the hypochile, And indeed, recent isotope studies have shown
however, the epichile hinged back up to its that Marsh Helleborine may acquire around
original position. The insect, as it backed out 30% of its nitrogen from its fungal ‘partner’ but
of the flower, was therefore forced upwards, it does not appear to receive any carbohydrates
allowing the pollinia to become attached to its via that route.
head. Later authors suggested other functions The subterranean seedling stage has never
for the hinged lip, such as causing the insects been observed in the wild and there is no infor-
to struggle to keep their balance and this being mation on the early development of this species.
Hybrids
None.
The specific name palustris means ‘of swampy
ground’.
HISTORY aND
CONSERVaTION
The first published record was in the 1633
edition of Gerard’s Herball, where John Goodyer
recorded it: ‘within a mile of…Peters-field, in a
moist meadow named Wood-mead, neere the
path leading from Peters-field towards Beryton’
(a record in Mathias de Lobel’s Stirpium
Illustrationes (‘Illustrations of plants’) dated from
1601, but this was not published until 1655).
Marsh Helleborine has declined
substantially and is now gone from 60% of its
M 8July, Norfolk.Theflowerinprofile, showingthetwo
historical range in Britain and 39% in Ireland.
halvesofthelip,connectedbyaflexiblehinge. The decline has affected all areas but perhaps
002 systematic.indd 83 29/1/09 12:11:15

especially those away from the coast. It is they quickly become too overgrown for the
extinct in the Channel Islands, Bedfordshire, helleborine to survive.
Huntingdonshire, Northamptonshire, Marsh Helleborine is specially protected in
Worcestershire, Radnorshire, Dumfries and Northern Ireland under Schedule 8 of the 1985
Galloway, Roxburghshire, Berwickshire, the Wildlife Order (NI).
Lothians and Fife.
PastandpresentoccurrenceofMarshHelleborineinBrit-
Many of the losses occurred in the 19th ainandIreland(basedonpresenceorabsencein10km
century due to the drainage and destruction squaresoftheNationalGrid;datafromthe New Atlas).
of marshes and fens. Drainage and the subtler
Britain Ireland
effects of water abstraction continued to cause
losses in the 20th century. More recently, total historical range, 450 160
eutrophication, that is the enrichment of 1500-1999
ground water by fertiliser run-off or even the current range 180 (6.3%*) 98 (9.7%*)
discharge of sewage, has caused suitable fens % lost, 1500-1969 53% 29.5%
to become overgrown with more vigorous % lost, 1970-1986 7% 9.5%
vegetation. The abandonment of grazing or
% lost, total 60% 39%
mowing compounds this effect and has led to
the invasion of fens by scrub, in which case * current range as a % of the total number of 10km squares
O 8July,Norfolk.Arather
dusty-pinkplant.
002 systematic.indd 84 29/1/09 12:11:17

DARK-RED HELLEBORINE l 85
Dark-reD helleborine
Epipactis atrorubens
This orchid is found very locally in open, rocky places in the north and west of Britain and
Ireland and is strongly associated with outcrops of limestone. Whether they are emerging
from the grykes in a limestone pavement or set against a grassy slope, the spikes of reddish-
purple flowers and rather dusky-green foliage are very striking. In Europe it is often a
woodland plant, and the absence of trees in our denuded landscape may be a reason for its
very scattered and local distribution in the British Isles. Dark-red Helleborine is the county
flower of Banffshire.
Identification
With its attractive reddish-purple flowers,
Dark-red Helleborine is easy to identify. It has
very obvious yellow anthers and pollinia, and is
cross-pollinated.
Similar species
Broad-leaved Helleborine is the only other
helleborine to occur in the same rocky habitats,
albeit only occasionally. It may sometimes have
rather dark-reddish flowers and, conversely,
Dark-red Helleborine may rarely have paler
pinkish or greenish-red flowers, similar to
those of some Broad-leaved Helleborines.
However, Dark-red Helleborine can always be
distinguished by its leaves, which are darker,
more markedly folded and held in two opposite
rows. It also has larger and rougher bosses on
the lip and a very hairy ovary.
Habitat
Dark-red Helleborine is very strongly associated
with limestone, growing on cliff ledges, scree
slopes, rocky hillsides, in old quarries and in the
shelter of the grykes of limestone pavements.
It is usually found in the immediate vicinity
of bare rock but sometimes also on well-
drained grassy slopes with scattered scrub
or even in meadows or on road verges. And,
although most sites are open and sunny, it is
also found in moderate shade on well-wooded
limestone pavements, in open ash woodland
or in pine plantations. Indeed, light woodland
P 17July, Co. Durham.Theleavesareclearlyarranged
intotwooppositeranks.
002 systematic.indd 85 29/1/09 12:11:18

002 systematic.indd 86 29/1/09 12:11:20

and woodland edges may be the more typical

habitat, as in Europe; most British sites are
deforested and heavily grazed and the species
has a very fragmented, ‘relict’ distribution. Most
sites are between sea level and 270m, but it
is present at 400m in eastern Cumbria, over
500m on Cronkley Fell in upper Teesdale and
at 610m in Gleann Beag in east Perthshire.
Flowering period
Early June to early August, but mostly late June
to late July.
Range
Very local and scattered. In North Wales
recorded from Caernarvonshire and Flintshire
(including the Great Orme), and in England
found in the Peak District of Derbyshire,
the Yorkshire Dales, in west Lancashire in
the Morecambe Bay area, in Cumbria in the
southern Lake District (around the Kent
estuary and at Hodbarrow), the north Pennines
(upper Eden Valley) and in Co. Durham. In
Scotland found in the eastern Highlands in east
Perthshire, in Banffshire and in the northwest
on the north coast of Sutherland, in west Ross
& Cromarty and on Skye. In Ireland confined
to the Burren region of Co. Clare and Co.
Galway. World range: Primarily a European
species, extending a little way into western
Asia. Occurs north to northernmost Norway
1987-99
1970-86
pre 1970
M 5 July, Lancashire. Dark-red Helleborine will grow in

quitewoodedsituations.
O 5July,Lancashire.Aclassicplantintheclassichabitat,
limestonepavement.
002 systematic.indd 87 29/1/09 12:11:22

and in Russia to the Arctic Circle, and south There are one to three funnel-shaped basal
to southern Spain, southern Italy, southern sheaths, the uppermost often green towards the
Greece, Romania and patchily to the Caucasus tip. Stems usually grow singly but sometimes
but absent from the Mediterranean lowlands. two or three may arise from the same rhizome.
Also found in western and northern Turkey, Leaves: Around five to ten, arranged more or
northern Iran, and southern Siberia to less in two opposite rows towards the base of
about 85°E. the stem. The leaves are dark green, variably
washed reddish-purple on the underside
How to find it and are sometimes purple at the base. They
Dark-red Helleborine is usually a relatively easy
are always distinctly longer than wide (the
species to locate when it is growing in the open.
lower elongated-oval, the upper tending to be
Undoubtedly the best site to see the species
narrower and more lanceolate), strongly folded
is Bishop Middleton Quarry in Co. Durham
and keeled, and held stiffly at about 30° above
where it is abundant, creating a great spectacle.
the horizontal.
Spike: Rather lax, with six to 45 flowers
DESCRIPTION set loosely to one side of the stem. Plants
Height: 11.5-60cm, sometimes to 100cm. in sheltered localities tend to have the most
Stem: Dull green, variably washed purple, flowers but there are often fewer than ten in
especially towards the base (sometimes entirely more exposed situations. There is usually a
purple), with a dense covering of whitish hairs, distinct gap between the uppermost leaf and
particularly on the upper part of the stem. the lowermost flower.
M 17July,Co.Durham.Thecontrastingyellowanther-capisverynoticeable;thelarge,roughbossatthebaseofthelip
less so.
002 systematic.indd 88 29/1/09 12:11:24

upper sepal sunlight, but normal plants always have some

petal anther traces of dark red. Two aberrant varieties have
been named:
petal Var. albiflora has white or creamy flowers and
pollinia
has been recorded in the Kishorn area of west
ovary viscidium Ross and Cromarty.
stigma Var. lutescens has yellowish or buff flowers and
has been found in Cumbria and The Burren in
Ireland.
sepal
hypochile boss sepal
(cup)
epichile BIOLOGY
Bract: Lanceolate, green, sometimes washed
This species is cross-pollinated. Both the colour
purple at the base. The lower bracts may be a
and the scent of the flowers may be important
little longer than the flowers but they become
in attracting suitable pollinators and wasps,
shorter towards the tip of the spike.
bees and hover flies visit to feed on the nectar,
Ovary: Pear-shaped, six-ribbed, green, variably
removing the pollinia in the process.
washed purple (sometimes deep blackish-purple)
Vegetative reproduction may take place, new
with abundant short, pale hairs. The flower stalk
plants developing from buds on the roots.
is short and greenish to purplish-black.
Flower: Usually a distinctive, rich, wine red. The Development and growth
flowers often do not open very widely and may The aerial stem grows from a short, thick,
assume a slightly drooping, bell-like shape. The hard rhizome that puts out 40-50 long,
sepals are oval with slightly pointed tips and slender, widely spreading roots. These
a downy outer surface. The petals are shorter sometimes form irregular swellings from
and broader, more oval-triangular in shape. which fresh rootlets grow.
Both sepals and petals are deep purple, slightly The roots of this species are reported to
paler on the inner surfaces and towards the have a rather limited fungal ‘infection’, and it
base of the petals. The hypochile is dull green was thought to be largely phototrophic as a
on the outer surface of the cup, becoming rich mature plant, depending on photosynthesis to
purple towards the front, with the interior pale supply its nutrients rather than fungi. Recent
greenish-white and mottled purple. The epichile isotope studies have shown, however, that Dark
is heart-shaped, broader than long and variably Red Helleborines acquire about 65% of their
turned under at the tip. It is rich purple with nitrogen and 15% of their carbon from fungi.
two elaborate wrinkled bosses at the base which And, unexpectedly, they have an association
sometimes merge into a V or heart shape. The with ectomycorrhizal fungi and thus, like
column is pale greenish-white, flushed purple, Coralroot and Bird’s-nest Orchids, may gain
with the anther cap contrastingly yellow with a nutrients from the roots of nearby trees via
narrow brown stripe at the sides; the pollinia are these fungi (see p.8). These studies took place
also yellow. There is a functional viscidium, and in Europe, where Dark Red Helleborines are
the flowers have a vanilla-like scent. often found in wooded environments. British
and Irish plants may grow well away from any
Subspecies trees, and the absence of suitable host trees may
None.
be a limiting factor in its distribution in the
Variation and varieties British Isles.
Occasionally the flowers may be paler red or There is no information on the length of the
even greenish, especially in sites exposed to full period between germination and flowering.
002 systematic.indd 89 29/1/09 12:11:24

002 systematic.indd 90 29/1/09 12:11:28
Hybrids Denbighshire, and old records for west

E. x schmalhausenii, the hybrid with Broad- Gloucestershire and Herefordshire are not
leaved Helleborine, has been reported from always accepted as valid.
several parts of the range, notably Cumbria. It
is fertile and therefore very difficult to confirm
because infertility cannot be used to distinguish
potential hybrids from plants that are merely
aberrant.
The specific name atrorubens means simply ‘very
dark red’.
HISTORY aND
CONSERVaTION
The first British record was published in
1677 in Ray’s Catalogus Plantarum Angliae
et Insularum adjecentium: ‘On the sides of the
mountains near Malham 4 miles from Settle in
great plenty’.
PastandpresentoccurrenceofDark-redHelleborinein
10kmsquaresoftheNationalGrid; datafromtheNew
Atlas).
Britain Ireland
1500-1999
current range 42 (1.5%*) 8 (0.8%*)
% lost, 1500-1969 18% 0%

% lost, 1970-1986 12% 38%
Dark-red Helleborine is very local and

absent from many apparently suitable sites
within its restricted range. In Britain it is
Nationally Scarce. Most populations are small,
with many non-flowering plants, and many
plants that would flower are prevented from
doing so by sheep, deer or rabbits. Bishop
Middleton quarry in Co. Durham holds 2,000
or more plants, probably more than all the
other British sites put together.
M 17 July, Co. Durham.The leaves have been largely
Some sites have been lost due to quarrying, grazed off, but nevertheless it is able to flower well,
and overgrazing threatens others; both perhapsaidedbyitsfungal‘partner’.
the British and Irish populations may still O 17July, Co. Durham. Usuallyarisessingly, buttwoor
be in decline. Extinct in Breconshire and threespikesgrowingtogetherisnotuncommon.
002 systematic.indd 91 29/1/09 12:11:30

broaD-leaveD helleborine
Epipactis helleborine
The commonest and most widespread of the helleborines, this species is found in and
around woodland and, in the north and west, sometimes also in more open habitats. But,
although at heart a forest orchid, it is a species that you stumble upon in unexpected places
rather than set out to find; a shady lane, a road verge, an old railway cutting – nowhere is too
humble for this adaptable helleborine. It is said to be commoner in the city of Glasgow than
anywhere else in Britain and has spread from coast to coast in North America since it was
introduced in 1879. ‘Young’s Helleborine’, described in 1982 and thought to be endemic to
Britain, is now known to be a minor variant of Broad-leaved Helleborine.
Identification
A very variable species. Broad-leaved
Helleborine can be a tall, robust, leafy plant or
a small, weedy specimen with just one or two
flowers. The flowers themselves can be almost
entirely green or almost completely purple
but are usually a mixture of pale green, pink
and purple. It is said that plants in deep shade
tend to be taller and greener whereas those in
sunnier locations are shorter with more red or
purple coloration in the flower, but the variation
often seems to be random.
Typically, Broad-leaved Helleborine has
several relatively large leaves which are a dull,
mid to dark green, sometimes washed purple
but lacking yellow tones. They are usually
obviously veined or ‘pleated’ and placed all
around the stem, in three rows or in a spiral
pattern, but are sometimes arranged into two
opposite ranks. As the name suggests, the leaves
are broad, especially the lowest, which may be
roughly as wide as they are long. The upper part
of the stem and the ovaries are hairy, although
hairs can be sparse on the ovaries. The outer
part of the lip is heart-shaped, broader than
long and usually turned under at the tip. There
are two bosses at the base of the lip which are
usually brownish and rough or wrinkled but
can be smooth and pink. Importantly, the base
of the flower stalk is washed purple. Broad-
leaved Helleborine is cross-pollinated; when
O 28July, Norfolk.Atypicalplant, withanopen, rather
sparsely-floweredspike.
002 systematic.indd 92 29/1/09 12:11:31

BROAD-LEAVED HELLEBORINE l 93
freshly opened, the flowers have an obvious held rigidly at about 45° above the horizontal.
and functional white viscidium, an important Its flowers are smaller, duller and do not
distinction from Narrow-lipped, Dune and open as widely. The petals and base of the lip
Green-flowered Helleborines (see p.76 for a are variably washed pink, but it never shows
discussion of the separation of cross-pollinated pink or purple tones to the sepals or a strong
and self-pollinated helleborines). purple wash on the lip. At inland localities the
Similar species variant of Dune Helleborine, known as Tyne
Helleborine, is found. This is very like Narrow-
Violet Helleborine may occur in the same
woods but is typically found in densely shaded lipped Helleborine and can be distinguished
areas. It is usually distinctive. Its leaves are more from Broad-leaved by the forward-pointing
greyish-green with a distinctive purple wash tip to its lip and a yellowish-green base to the
to the undersides, and also narrower, with the flower stalk. Dune Helleborine is normally
lowest leaf longer than wide. Its flowers are self-pollinated.
larger, brighter and cleaner; the sepals and petals Green-flowered Helleborine typically has
are pale greenish-white, lacking pink or purple green flowers which are held drooping and do
tones, and the lip is whitish with two smoothly not fully open. Sometimes, however, its flowers
pleated, pink bosses. Like Broad-leaved may be held more horizontally and may open
Helleborine, it is cross-pollinated. widely although they are still predominantly
Narrow-lipped Helleborine is a scarce green, with any pink tones restricted to a
inhabitant of southern beechwoods. It has delicate wash on the lip. Whatever, its upper
green flowers, often with a delicate pink wash stem and ovaries are hairless or there are just
to the petals and lip (but not the sepals). As a few, sparse hairs on the stem, the base of the
its name suggests, the tip of the lip is long, flower stalk is green and it is self-pollinating.
narrow and pointed, and held projecting Dark-red Helleborine is usually rather
prominently outwards. Its leaves are a paler and distinctive but Broad-leaved is occasionally
more yellowish-green and usually held in two found in the rocky habitats beloved of Dark-
opposite ranks. In combination these features red Helleborine and sometimes has rather
should be distinctive but the tip of the lip of dark-reddish flowers, too, while Dark-red
Broad-leaved Helleborine may not always be Helleborine rarely has pinkish or greenish-red
reflexed, especially when the flower has just flowers. If there is doubt, Dark-red Helleborine
opened. In Broad-leaved Helleborines with can be distinguished by its leaves, which are
largely green flowers, this can cause confusion darker, more markedly folded and held in two
but the outer part of the lip (epichile) in Broad-
leaved Helleborine is always broader than long.
In case of any doubt, the base of the flower
stalk is greenish-yellow in Narrow-lipped
Helleborine and the flowers are self-pollinated
and lack an effective viscidium.
Dune Helleborine is only found on
Anglesey, in northern England and in southern
Scotland, usually on dunes but also at inland
sites; conversely Broad-leaved Helleborine is
occasionally found on open dunes or under the
pines that are often planted on coastal sand
hills. In its typical form, Dune Helleborine can
be separated from Broad-leaved by its more M 26July, Norfolk.Arichlycolouredflower.Thepollinia
yellowish-green, two-ranked leaves that are havealreadybeenremoved.
002 systematic.indd 93 29/1/09 12:11:33

O 28 July, Norfolk. Two

plantssidebyside, onein
fullflower,theotherstillin
bud.
opposite rows. The bosses on its lip are also stream-sides. Broad-leaved Helleborine will
larger and rougher and its ovary is densely hairy. also grow in the open, on limestone pavements,
cliffs, scree and grassland, but only in the
Habitat cooler and damper conditions of the north and
Broad-leaved Helleborine is essentially a plant west. In Ireland and south Wales it is found
of deciduous woodland. It favours the better-lit in dune slacks. One of the most adaptable
areas along paths, rides and roadsides, in glades of the helleborines, it is an opportunist
and on the woodland fringe, but can grow in and can colonise newly available habitats,
deep shade. Like many of the helleborines it has such as mature birch scrub on spoil heaps,
an affinity for Beech trees. It can also be found willow and alder carr and conifer plantations
in suitable shady conditions in scrub, along (especially where conifers have replaced
well-grown hedges, banks, disused railways and ancient woodland). In Glasgow and a few
002 systematic.indd 94 29/1/09 12:11:35

Armagh, Co. Derry and Co. Antrim. World

1987-99
1970-86
range: Very widespread in Europe and
pre 1970 Asia and introduced to North America. In
Europe it occurs north to c. 71°N in Norway
and to southern Finland, and south to the
Mediterranean, Crimea and Caucasus. It is
absent from the Mediterranean lowlands but
found on the Balearic Islands, Corsica, Sardinia
and Sicily. Also scattered in Turkey, Lebanon
and Israel. In Asia it also ranges through
southern Siberia east to Lake Baikal and south
through the mountains of Central Asia to the
Himalayas. Populations from North Africa and
the Far East are often now treated as different
species. Introduced to North America, perhaps
deliberately as a remedy for gout. It was first
recorded in 1879 near Syracuse, New York,
other cities in southern Scotland and northeast and is now widespread; it reached California by
England, Broad-leaved Helleborine has 1950 and is still spreading.
moved into mature gardens, as well as parks,
cemeteries, golf courses, playing fields, rubbish How to find it
tips, roadsides and railway embankments. It The commonest of the helleborines but never-
is comparatively tolerant of soil pH and will theless often rather local, it is best looked for
grow in slightly acid conditions but is usually in and around undisturbed ancient woodland,
commonest on calcareous soils. Recorded up to especially along shady roads and tracks passing
350m above sea level (Ystradfellte, Breconshire). through suitable habitats. It is frequently robust
but despite this the relatively small, dull flowers
Flowering period can be surprisingly hard to see when it is
Early July to early September but mostly
growing among brambles or Bracken.
from mid-July to mid-August. Like all the
helleborines, it flowers earlier and more sparsely DESCRIPTION
in dry summers. upper sepal
Range petal anther
Widespread in England and Wales, although
petal
rather local in some areas, such as Norfolk,
Suffolk, Cambridgeshire and Lincolnshire, and pollinia
absent from west Cornwall and some upland ovary viscidium
areas in northern England. In Scotland it is stigma
well distributed in the Central Lowlands and
scattered in the Borders but to the north is
sepal
extremely local, with a few sites in southern hypochile boss sepal
Perthshire, Angus, Banffshire, Sutherland, west (cup)
epichile
Ross & Cromarty, Argyll and Kintyre, and on
the Inner Hebrides on Skye. Widespread in Height: 10-120cm but usually around 25-80cm.
Ireland but mostly found in the west and the Stem: Pale green, often washed purplish
north and rather local away from The Burren towards the base, with short, whitish hairs on
(Co. Clare and Co. Galway), Co. Fermanagh the upper portion and two or three leafless
and the region around Lough Neagh in Co. sheaths at the extreme base. Usually grows
002 systematic.indd 95 29/1/09 12:11:36

singly but clumps of two or three are quite

frequent and five or more may occasionally
grow from the same rhizome. Non-flowering
stems are frequently produced.
Leaves: Four to ten, more-or-less spirally
arranged around the stem. They are oval to
oval-lanceolate and less than twice as long
as wide; the lowest leaf is sometimes almost
rounded, but they become narrower and more
bract-like towards the flower spike. The leaves
are dull, mid to dark green, sometimes tinged
purple, with several prominent veins.
Spike: Up to 60 flowers, rarely as many as 100,
are arranged into a roughly one-sided spike that
is sometimes very dense but on other plants
rather lax.
Bract: Dull green, lanceolate, with a pointed
tip. The lowest are significantly longer than the
flower but they become shorter, roughly the
length of the ovary, towards the tip of the spike.
Ovary: Green, hairless or with a few short
hairs, boldly six-ribbed and rather pear-shaped,
tapering into a long, twisted stalk which is
washed purple at the base.
Flower: Very variable in colour but usually
with some purple tones and opening widely.
The sepals are oval, tapering to a point, with
their outer surface green mottled with variable
amounts of dull purple or pink (sometimes
none) and a prominent green midrib; their
inner face varies from pale green to dull, dusty
purple. The petals are similar but slightly
shorter and less tapering and tend to be paler,
‘cleaner’ and often pinker. They vary from pale,
dusty pink to purple and are often whiter
towards the centre; they also have a green
midrib on the outer face. The hypochile is pale
greenish-white, variably washed pink or purple,
with the interior of the cup purple to mid-
brown, glistening with nectar. The epichile is
heart-shaped, broader than long, with the major
part strongly curled down and under. There are
two bosses at the base of the epichile, usually
purplish-brown and wrinkled, but they may be
green or pink, and are sometimes smooth. The
M 1August, Northumberland. A spike of rather purple
flowers.Broad-leavedHelleborines,wheninspectedclose- epichile varies in colour from dull greenish-
ly,alltoooftenappearratherdishevelled. white, washed pink, to pale pink or dull purple.
002 systematic.indd 96 29/1/09 12:11:38

The column is greenish-white, the anther cap

dull pale yellow with brown stripes at the side,
and the pollinia are creamy-yellow. There is an
obvious and functional white viscidium. The
flowers usually have no scent.
Subspecies
None recognised.
Var. youngiana ‘Young’s Helleborine’
Young’s Helleborine was described as a new
species, ‘E. youngiana’, in 1982 from plants
found in Northumberland. It was subsequently
identified in Scotland and, more controversially,
in Yorkshire and south Wales. It was reported
to differ from Broad-leaved Helleborine in a
number of ways. Its wavy-edged, more-or-less
two-ranked leaves are on average paler and more
yellowish-green. The lowest leaf is usually longer
than wide, rather than wider than long. Its ovary
is sparsely hairy to hairless (only rarely hairless in
typical Broad-leaved Helleborines). The flowers
are similar to some Broad-leaved Helleborines
but relatively large, clean and bright. The critical
difference, however, was to be found in the
reproductive structures; in Young’s Helleborine
the viscidium is very small and disappears
rapidly. The pollinia remain in the flower
and disintegrate onto the stigma causing self-
pollination. The column has a long rostellum (as
long as the anther cap) which, together with two
pointed bosses at the base of the stigma, forms a
distinctive ‘three-horned’ shape.
Some botanists were, however, always
sceptical about the distinctiveness of ‘Young’s
Helleborine’ and recent genetic studies have
shown that it does not exist as a distinct
entity. At each site investigated, the ‘Young’s
Helleborines’ were genetically closer to the
local Broad-leaved Helleborines than they
were to ‘Young’s Helleborines’ at other sites.
Also, rather than being self-pollinated, it has a
high level of genetic diversity and a population
genetic structure that indicates that it is cross-
pollinated. Plants matching the description of
Young’s Helleborine in terms of their leaves, M 1August,Northumberland.A‘Young’s’type,withrather
ovaries and flowers are found at its classic sites yellowish-greenleavesandbrightflowers.
002 systematic.indd 97 29/1/09 12:11:39

002 systematic.indd 98 29/1/09 12:11:42
(for example, Settlingstones in Northumberland) mouthparts but can nevertheless easily reach
but almost all appear to have a large and fully the nectar produced in the hypochile; in the
functional viscidium and are presumably cross- process they rupture the viscidium and the
pollinated, supporting the genetic studies. With pollinia are stuck to their heads. Other insects,
the claimed differences in reproductive biology including short-headed wasps, bees, hover
gone, the remaining distinctions between flies and beetles, may visit the flowers but
‘Young’s’ and Broad-leaved Helleborine are very are the wrong size or shape to act as efficient
subtle and it is hard to justify even the status pollinators. Fermentation of the nectar in the
of variety for Young’s, especially given the wide helleborine’s flowers may produce ethanol and
variation in Broad-leaved. this can have a narcotic effect on visiting wasps,
Var. neerlandica ‘Dutch Helleborine’ Overall which become slow and sluggish and may even
deep green and rather short (15-40cm), with fall to the ground ‘drunk’.
short, stiff, rounded leaves that are held more- The flowers are self-compatible and are
or-less erect and grouped at the base of the frequently pollinated by wasps carrying
stem, which they closely sheathe. The leaves pollinia from flowers of the same spike (i.e.
have a border of tiny teeth that are irregular geitonogamy). It is sometimes stated that
and fused at the base (use a 20x hand-lens; Broad-leaved Helleborine may be self-pollinated
typical Broad-leaved Helleborines have more (i.e. fertilised by pollen from the same flower).
regular teeth). The spike is dense and the The evidence for this is contradictory but
flowers are dull purplish-pink, bell-shaped and experiments have shown that in normal
do not open widely. Found along the coast of circumstances self-pollination either does not
the North Sea from Pas-de-Calais in northeast occur or only takes place rarely when small
France to Denmark and on the Baltic coast insects carry pieces of pollinia onto the stigma
of northern Germany; similar plants growing below. It has also been reported that in drought
amongst Creeping Willow in the dune slacks conditions the flowers shrivel without opening
of south Wales have been identified as this and may be cleistogamous and self-pollinate in
form. Although it is treated as a distinct species, the bud (Ettlinger 1997). Seed-set is usually
E. neerlandica, by some Continental authors, good and almost all the flowers may produce
genetic studies reveal little difference between it ripe capsules, each containing up to 3,000 seeds.
and typical Broad-leaved Helleborines.
Var. monotropoides (also known as var. albifolia) Development and growth
lacks chlorophyll and is pale pink or straw- The aerial stem grows from a small, woody
coloured with white or rosy flowers. It is very rare. rhizome which sends numerous cordlike
Var. viridiflora lacks anthocyanins and has pale roots deep into the soil. The degree of fungal
green flowers with a greenish-white lip and ‘infection’ of the roots is reported to vary, being
shows no trace of red or purple. It is rare. high in plants growing in humus-rich soils
Var. purpurea has especially dark purple or and negligible in mineral soils, but at least
reddish-violet flowers. It is rare. some Broad-leaved Helleborines are heavily
dependent on their fungal ‘partner’. Recent
BIOLOGY isotope studies have shown that they acquire
Pollination and reproduction around 60% of their nitrogen and about 14%
Broad-leaved Helleborine is pollinated by of their carbon via fungi. Unexpectedly, Broad-
wasps, especially long-headed species of leaved Helleborine has an association with
the genus Dolichovespula. Wasps have short ectomycorrhizal fungi and thus, like Coralroot
and Bird’s-nest Orchids, may gain nutrients
O 28July,Norfolk.Abeautiful,richly-colouredplantwith
wine red petals and lip.The pollinia have already been from the roots of nearby trees via these fungi
removedfromalltheflowers. (see p.8). It is probably not very fussy about its
002 systematic.indd 99 29/1/09 12:11:42

Scotland and confirmed by genetic analysis.

This hybrid has no scientific name.
The specific name helleborine means ‘like
a hellebore’. The leaves of Broad-leaved
Helleborine do indeed look like those of
Veratrum album, a plant that was once called
‘White Hellebore’ but is now, rather ironically,
known as ‘White False Helleborine’. (Broad-
leaved Helleborine bears no resemblance to the
plants now called ‘hellebores’, quite unrelated
flowers belonging to the buttercup family.)
M 28 July, Norfolk.A very pallid plant, with just a hint HISTORY aND
of purplish-pink in the sepals and lip.The pollinia and CONSERVaTION
viscidiumhavealreadybeenremoved.
The first British record was in 1562 when
fungal ‘partners’ but might have a preference William Turner stated in his Herball: ‘I have
for Ascomycetes, including ectomycorrhizal seen it…in England in Soffock’.
fungi of the genus Tuber, better known by their
English name ‘truffle’. PastandpresentoccurrenceofBroad-leavedHelleborinein
BritainandIreland(basedonpresenceorabsencein10km
Broad-leaved Helleborine, presumably squaresoftheNationalGrid;datafromtheNew Atlas).
supplied by its fungal ‘partner’, may spend a
Britain Ireland
significant proportion of the time underground.
Plants may flower and then spend one or, rather total historical range, 1,218 161
1500-1999
less often, two or even three years ‘dormant’
current range 840 (29.5%*) 107 (9.7%*)
before appearing again. In a study in America,
25%-50% of the population appeared above % lost, 1500-1969 19% 24%
ground each year and around a third of these % lost, 1970-1986 12% 9.5%
flowered. Very few plants flowered every year, % lost, total 31% 33.5%
however, although annual flowering may be more
frequent where the soil is reasonably moist.
The interval between germination and The New Atlas states that the ‘overall
flowering can be as little as 18 months, although distribution is stable’ but although the
periods of eight or nine years, including several boundaries of the range may be largely
years above ground as a non-flowering plant, are unchanged there has been a significant decline,
also quoted. with the loss of around a third of the historical
Hybrids range. Losses have been concentrated in the
E. x schmalhausenii, the hybrid with Dark-red Midlands and northern England, where the
Helleborine, has been reported from several distribution is now rather fragmented, and
parts of the range, notably Cumbria. It is fertile more recently in the Home Counties. The
and very difficult to confirm. clearance or ‘coniferisation’ of woodland,
E. x schulzei, the hybrid with Violet Helleborine, increase in dense shade due to a lack of
has been recorded quite frequently but, like woodland management, ground disturbance by
E. x schmalhausenii, is fertile and hard to machinery and horses, and grazing by deer may
confirm. all have contributed to the decline, and losses
x Dune Helleborine has been found in seem to be ongoing.
002 systematic.indd 100 29/1/09 12:11:44

VIOLET HELLEBORINE l 101
violet helleborine
Epipactis purpurata
Thought to be a long-lived orchid, older and more mature plants may produce several flower-
ing spikes from the same rootstock, and a group of Violet Helleborines blooming in the
cathedral-like gloom of a late summer beechwood certainly makes a dramatic sight. Among
the last orchids to come into flower, it is endemic to western and central Europe and in Britain
is confined to the southern half of England, but it is always rather local and uncommon.
Identification
Relatively distinctive. The flowers are pale green
or whitish with the lip faintly washed pink
and bearing two pink bosses; the petals and
sepals spread widely and the large, pale flowers
contrast strongly with the dark purplish stem
and the rather small, dark leaves. It is cross-
pollinated, and the flowers have an obvious
and functional viscidium, a useful distinction
from Narrow-lipped and Green-flowered
Helleborines (see p.76 for a discussion of
the separation of cross-pollinated and self-
pollinated helleborines).
Similar species
Broad-leaved Helleborine is much commoner
than Violet Helleborine. It has broader leaves,
with the lowest more-or-less wider than long
(longer than wide in Violet Helleborine).
Its leaves are also a cleaner and brighter
green (duller, more greyish-green in Violet
Helleborine, with a distinctive purplish wash
to the underside). Its flowers are smaller and
often duller and darker with a purplish wash,
with the bosses on the lip usually rougher and
browner. In Violet Helleborine the flowers are
cleaner, brighter and paler, with two smoothly
pleated, pink bosses. Like Violet Helleborine, it
is cross-pollinated.
Narrow-lipped Helleborine is also found
in densely shaded beechwoods in southern
England but is easily separated by its long,
pointed lip and much paler stem, flower stalks
and leaves.
P 11August, Suffolk. Summer 2003 was hot and dry,

andthehelleborinesweresmallandfewinnumber.
002 systematic.indd 101 29/1/09 12:11:45

woodland relicts and in wild gardens. Although

it can tolerate acid soils, it is strongly associated
with areas of calcareous bedrock, especially
chalk, and its deep root system requires a
substantial thickness of soil. It can grow on
sands and gravels but is particularly associated
with clays, especially the ‘clay-with-flints’ found
on plateaus and hilltops overlying chalk. Within
woods, it is often found in areas of deep shade
where little else can grow; it can flourish in
much darker situations than Broad-leaved
Helleborine but may share its dimly-lit haunts
with Narrow-lipped Helleborine.
Flowering period
Mid-July to early September, exceptionally late
June to late September, but typically peaking in
early August. It tends to flower earliest in more
open situations and will be especially early in a
dry season.
Range
Found in southeast England (including the Isle
of Wight) and the Midlands, occurring west
to Dorset, north Somerset, Gloucestershire,
Herefordshire and Shropshire, and north
to south Staffordshire, Leicestershire, south
Lincolnshire, Huntingdonshire, Hertfordshire
and southern Suffolk. Also recorded from
Denbighshire. The species is commonest in
Kent, Surrey and the Chilterns. World range:
1987-99
1970-86
M 11August,Suffolk.Growingonarelativelybarewood- pre 1970
landfloorinoldhazelcoppice,atypicallyshadyspot.
Green-flowered Helleborine may be in

bloom late in the summer, like Violet Helle-
borine, but is easily separated by its more-or-less
hairless, green stem and ovaries, green flower
stalks and rather smaller flowers which are
usually held drooping and seldom open widely.
Habitat
Violet Helleborine is very much a woodland
orchid, favouring beech, hornbeam and oak
woods as well as overgrown hazel coppice. It
is occasionally found in hedgerows that are
002 systematic.indd 102 29/1/09 12:11:47

Confined to western and central Europe where

it occurs north to Denmark, west to France (to
a line between Mont St Michel and Grenoble),
south to the Alps, Balkans and Transylvanian
Alps in Romania, and east to the Baltic States.
How to find it
Despite its size and showy flowers, this can be
a tricky species to find. Like other helleborines,
it is loosely colonial, and although the colonies
sometimes grow alongside paths and roads or
on the edges of glades and clearings, they are
equally at home in deep shade and can easily
be missed. It is best looked for in wet seasons,
as the species is affected by drought and in dry
summers the size and number of spikes may be
greatly reduced.
DESCRIPTION
upper sepal
petal anther
petal
pollinia
ovary viscidium
stigma
sepal
hypochile boss M 10August, Suffolk. Of all the woodland helleborines,
sepal
(cup) Violethasthebrightestandcleanestflowers.
epichile
Height: 20-90cm but usually less than 70cm. like. The leaf posture is variable; they may be
Stem: Greyish-green, variably but often heavily held horizontally with the tips slightly drooping
washed purple, with dense, short, grey hairs or at about 30° above the horizontal. They are
on the upper part of the stem and one to three a rather dull, ‘cold’, greyish-green and may be
small, purplish-brown sheathing scales at the washed purple towards the tips. The undersides
base (the uppermost scale often tipped green). have a diagnostic purple wash and the leaf
Stems usually grow singly but multiple stems sheaths are also frequently tinged purple.
are fairly common, groups of six to eight not Spike: Slightly to moderately one-sided, there
unusual, and a cluster of 38 has been recorded. are usually seven to 40 flowers but some well-
Leaves: Well-spaced up the stem and arranged grown plants can carry over 100 blooms.
spirally (or sometimes in two opposite rows), Bract: Green, variably washed purple. The
the four to 14 leaves are relatively small, more- bracts are narrow, lanceolate, held roughly
or-less oval in shape and taper to a point. They horizontally and are longer than the flower in
are usually rather more than twice as long as the lower part of the spike, becoming shorter in
wide, with the upper leaves narrower and more the upper part.
bract-like and the lowest leaf short and cowl- Ovary: Green, with six prominent ribs, which
002 systematic.indd 103 29/1/09 12:11:48

002 systematic.indd 104 29/1/09 12:11:50

may be washed purple, and sparsely hairy. The Development and growth
flower stalk is purplish and variably twisted. Violet Helleborine grows from a rhizome
Flower: Rather large, opening widely, and which lies more-or-less vertically in the soil
overall greenish-white with a pinker lip. The and has up to 50 fleshy roots, each up to 70cm
sepals are triangular-oval, rather large and long (exceptionally 120cm), growing vertically
pale green, becoming paler towards the edges downwards. It is reported that the roots are
and with a prominent green midrib on the fungus-free and therefore the mature plant is
outer surface. The petals are smaller, whitish, phototrophic, depending on photosynthesis
becoming slightly greener towards the centre, rather than fungi for nutrition. However, given
with a fine green midrib. The hypochile is the dense shade in which it grows, it seems
translucent-whitish, slightly greener towards much more likely that fungi contribute a large
the base of the cup, with the interior variably part of its nutritional budget; the rare var. rosea
washed pale purplish-rose to pale brown or lacks chlorophyll but nevertheless is able to
pale greenish. This colour shines through to flower and fruit successfully and must depend
the outside. The epichile is short, triangular entirely on fungi.
or heart-shaped with the tip strongly folded Violet Helleborine is long-lived and only
downwards. It is whitish with two prominent, appears above ground when mature enough to
smoothly pleated, pink bosses at the base. The flower. Immature, non-flowering plants are very
column is whitish and the large, conspicuous rarely seen. A single-stemmed plant may be 30
anther cap is very pale yellowish-white years old, and it has been suggested that large,
with narrow brown stripes at the sides. The multi-stemmed plants are probably hundreds of
viscidium is whitish and the pollinia are pale years old.
yellow. The flowers are faintly scented. There is no information on the duration of
the period between germination and flowering.
Subspecies
None. Hybrids
E. x schulzei, the hybrid with Broad-leaved
Helleborine, has been recorded widely but this
Plants may sometimes have variegated leaves,
hybrid is fertile and therefore very difficult to
and some may have the leaves very extensively
confirm. (Infertility and the failure to produce
streaked violet. One variety has been named:
viable pollen and seed are a standard means of
Var. rosea is rare but stunning. It lacks
confirming a hybrid.)
chlorophyll and the entire plant is a rosy pink
with whitish flowers. Name and classification
The specific name purpurata means purplish.
BIOLOGY
HISTORY aND
Routinely cross-pollinated, often by wasps
CONSERVaTION
which are attracted to the nectar. As with This species was discovered by Rev. Dr. Abbot
Broad-leaved Helleborine, the nectar is ‘parasitical on the stump of a maple or hazel
reported to have a narcotic effect, with ‘drunken’ in a wood near the Noris farm at Leigh,
wasps falling to the ground. Pollination is Worcestershire, in 1807’ and was described
efficient and most or all of the flowers on a scientifically from English specimens in 1828
spike will set seed. There are no reports of by Sir J.E. Smith in The English Flora. Notably,
vegetative reproduction. the type specimen was probably the rare
variant rosea.
O 11August,Suffolk.Amulti-stemmedplant(therecord Violet Helleborine has declined steadily over
is38spikesgrowingtogether). the last 150 years as ancient woodlands have been
002 systematic.indd 105 29/1/09 12:11:51

destroyed or replanted with conifers, although be grazed off in some woods. The species can
most sites should now be safe from this particular be very persistent, however, and has even been
threat. It has vanished from the edge of the range recorded pushing its way through newly
and is extinct in Devon, Cambridgeshire, Norfolk laid tarmac.
and southwest Yorkshire, and is reduced to one PastandpresentoccurrenceofVioletHelleborineinBrit-
site in south Lincolnshire. ainandIreland(basedonpresenceorabsencein10km
squaresoftheNationalGrid;datafromtheNew Atlas).
Favouring the dense shade of closed-canopy
woodland, this is perhaps the only species of Britain Ireland
orchid to have benefited from the abandonment total historical 235 0
of coppicing in many British woods during range, 1500-1999
the 20th century. Conversely, the great current range 145 (5%*) 0
storms of 1987 and 1990 devastated many
% lost, 1500-1969 28.5%
woods, opening up the canopy and leading
% lost, 1970-1986 10%
to a great reduction in the numbers of Violet
% lost, total 38.5%
Helleborines in affected areas. Another threat
is deer, and whole populations of orchids can * current range as a % of the total number of 10km squares
O 10 August, Suffolk.

Violet Helleborine is
cross-pollinated,mainlyby
wasps.
002 systematic.indd 106 29/1/09 12:11:53

NARROW-LIPPED HELLEBORINE l 107
narrow-lippeD helleborine
Epipactis leptochila Data Deficient
Other names: E. muelleri

This enigmatic orchid is confined to southern England with an outpost in south Wales. It
is a characteristic species of the beechwoods of the Cotswolds and Chilterns but is scarce
and local and has declined markedly in recent years. For a while, a case of mistaken identity
allayed fears about this decline, but it is now clear that this surprisingly attractive helleborine
is in need of friends.
Identification
Narrow-lipped Helleborine looks almost
uniformly green and has relatively large, clean,
pale-green flowers with purplish-pink confined
to a delicate wash around the base of the lip
and a variably obvious tinge on the petals. The
flowers are held drooping and the tip of the
lip projects forward rather than being turned
under as in most other helleborines; the lip
therefore appears long and pointed. The upper
stem and ovaries are hairy (use a hand-lens to
check this), and the base of the flower stalk is
greenish-yellow. The leaves are usually carried
in two opposite ranks and are fresh green,
sometimes tinged with yellow. The flowers
are self-pollinated and lack a viscidium, the
pollinia crumbling apart where they lie and
falling piecemeal onto the stigma. This is a
useful distinction from Broad-leaved and Violet
Helleborines (see p.76 for a discussion of the
separation of cross-pollinated and self-pollinated
helleborines). Rarely, when freshly open, the
flowers of Narrow-lipped Helleborine can be
temporarily cross-pollinating (‘facultatively
allogamous’), with functional viscidia and intact,
cohesive pollinia; such plants must be identified
as Narrow-lipped Helleborine with caution.
Similar species
Broad-leaved, Violet and Green-flowered
Helleborines may be found in the same woods
although only Violet Helleborine is frequent
P 22 July, Oxfordshire. This can be a very stately

helleborinebutalltoooftenitsflowerspikesaregrazed
offbydeer.
002 systematic.indd 107 29/1/09 12:11:54

in the deep shade beloved of Narrow-lipped

1987-99
Helleborine. Broad-leaved Helleborine can 1970-86
have largely green flowers, but they are usually pre 1970
extensively washed with dull pink or purple,

and the tip of the lip is almost always turned
down and backwards to give the lip a very
short, blunt-ended appearance. Even if the
lip is not reflexed, the heart-shaped epichile
is always broader than long. The base of the
flower stalk is tinged with purple, and, in most
Broad-leaved Helleborines, the leaves are a
darker and duller green and are usually carried
spirally around the stem. Violet Helleborine
has flowers that are closer in coloration to
Narrow-lipped, but they are held more erect
and face outwards, giving the spike a different
character. The flower stalk is purple and the
leaves are a far duller greyish-green and often However, it has a hairless, or almost hairless,
have a faint purple wash on the underside. upper stem and ovaries, and the interior of
Green-flowered Helleborine is self- the cup at the base of the lip is pale greenish.
pollinated, like Narrow-lipped Helleborine, In many cases, the flowers of Green-flowered
and can be rather similar to it (especially var. Helleborine are very distinctive as they often
vectensis of Green-flowered), with drooping, hang down almost vertically and in some
bell-like green flowers and a green flower stalk. populations hardly open at all.
M 22July,Oxfordshire.GrowingonatypicallybarewoodlandfloorbelowBeeches.
002 systematic.indd 108 29/1/09 12:11:56

Habitat
Narrow-lipped Helleborine is always found on
calcareous soils derived from chalk or limestone,
especially on the steeper slopes where the soil is
very thin and ‘skeletal’. It is strictly a woodland
orchid and usually to be found in ancient
woodland. The classic habitat is a beechwood
on chalk, sometimes with a mixture of Yew, but
occasionally it is found under a variety of other
deciduous trees, including overgrown
ash-hazel coppice. Whatever the type of
woodland, Narrow-lipped Helleborines will be
found in areas of deep shade where the ground
cover is sparse or absent. It is intolerant of
direct sunlight.
Flowering period
Rather short, from the second week in July to
mid-August (exceptionally from late June) but
mostly in the last half of July.
M 12 July, Buckinghamshire. This species has largely
Range green flowers but with a pinkish-purple tinge to the lip
Highly localised. The strongholds are the andsometimesalsothepetals.
Cotswolds in Gloucestershire and the
Chilterns in Berkshire, Oxfordshire and the species at its best is the Warburg Reserve
Buckinghamshire. Away from these areas at Bix Bottom in Oxfordshire where a strong
it is rare with just a few widely scattered population is protected from deer.
populations in Surrey, Hampshire, Dorset,
south Wiltshire, north Somerset, Shropshire DESCRIPTION
and Glamorganshire. The species is incon-
upper sepal
spicuous, however, and could still be found at
new localities. Delforge (1995) reported the petal anther
presence of an unidentified helleborine of the petal
‘leptochila group’ from the Burren in Co. Clare.
World range: Although once thought to be pollinia
ovary
endemic to Britain, the species has now been stigma
found in Europe, north to Denmark, east to
Slovakia, Hungary and Yugoslavia, and south to
Italy and the Pyrenees. sepal
hypochile boss
How to find it (cup) sepal
epichile
With a restricted distribution, specialised
habitat and short flowering period, this can Height: 15-75cm but usually 30-60cm.
be a hard species to find. It is also superficially Stem: Green, with the upper part hairy. Usually
rather anonymous, although well-grown plants grows singly but sometimes there are two
are very attractive. The apparent size of colonies together and rarely up to five or six stems may
can vary dramatically from year to year but the arise from a single rootstock.
underground population may be much more Leaves: Three to seven, fresh green and arranged
stable and rather larger. One of the places to see in two opposite rows up the stem, although
002 systematic.indd 109 29/1/09 12:11:57

Bract: Pale green and lanceolate; in the lower

part of the spike the bracts are very long,
project well beyond the flower and often hang
downwards, but towards its tip they are shorter.
Ovary: Pale green, variably hairy, prominently
six-ribbed but not twisted; the flower stalk is
greenish-yellow.
Flower: Overall greenish, relatively large
and usually opening fairly widely, although
occasionally some flowers may not open fully,
especially towards the tip of the spike. The
sepals are oval-triangular and elongated into
pointed tips, pale green on the outer surface
and slightly paler and more whitish-green
on the inner. The petals are similar in shape
but slightly smaller. They are also paler, more
greenish-white, becoming even paler towards
the edges but greener towards the midrib
(which is prominent on the outer face) and
variably flushed pale pink with faint pink veins.
The hypochile is whitish-green on the outside
of the cup, variably flushed purplish-pink at
the sides, and similarly pale on the inside but
with a wine-red or chocolate-brown rear wall;
it contains nectar. The epichile is whitish-green,
sometimes delicately flushed pink, shaped like
an arrowhead, longer than wide, and its pointed
tip projects outwards. It has two relatively
small, smooth bosses at its base and these are
washed purple or purplish-pink; they flank
a longitudinal central groove connecting the
hypochile and epichile. The column is greenish-
white, the pollinia creamy-yellow and the anther
cap pale greenish-yellow with a narrow cream
and broad chocolate-brown stripe on either
M 22July,Oxfordshire.Aparticularlycrowdedspikeona
robustplant. side; it is attached to the column at the rear by
a projecting spur or stalk (and in profile looks a
sometimes not obviously so. The leaves are little like the ‘leaping jaguar’ motif of the famous
elliptical and mostly more than twice as long sports car, a distinction from Broad-leaved
as wide, with the uppermost long, narrow and Helleborine where the anther is unstalked). The
grading into the lowest bracts. They are rather rostellum is reduced in size (less than half as
‘floppy’ and held more-or-less horizontally but long as the anther) and the viscidium, although
the uppermost, bract-like leaves are pendant. present in the bud, has almost always withered
Spike: Usually rather lax, with four to 35 by the time the flower opens.
flowers, often facing to one side. Initially held
horizontally, they droop to a variable extent as Subspecies
they age. None.
002 systematic.indd 110 29/1/09 12:11:58

lacking chlorophyll, with a lemon-yellow stem

and flowers, and pale greenish-yellow leaves,
has been recorded (Young 1962b). It apparently
thrived and flowered. This observation, together
with the dense shade of its normal habitats,
suggests that Narrow-lipped Helleborine must
be able to acquire a large proportion of its
nutrients from fungi.
Hybrids
None. (E. x stephensonii, the hybrid with Broad-
leaved Helleborine, has been reported, but not
confirmed, in southern England.)
The specific name leptochila means ‘with a
slender lip’.
M 22July, Buckinghamshire. Showingthepointedtipto Initially confused with variants of Broad-
thelip,whichisnotbentbackunderneaththeflower,and
therelativelysmall,smoothbossesatitbase.
leaved Helleborine, it was not until 1921 that
Narrow-leaved Helleborine was described as
Variation and varieties a distinct species. It was then confused with
Var. cleistogama was described from plants Green-flowered Helleborine until the British
found in Gloucestershire on the steep western botanist Donald Young resolved the identity
escarpment of the Cotswolds near Wotton- and characteristics of the two species in the
under-Edge, but this population is apparently 1950s and early 1960s. Confusion then arose
now extinct. It had pendulous bracts, flowers again in the mid-1970s when plants resembling
that did not open and, within the flower, a Narrow-lipped Helleborine were found inland
greener lip. It has been treated as a distinct in northern England (the Tyne Helleborine, see
species but the upper flowers of some normal p.97). In some places these Tyne Helleborines
plants may never open, and it seems much more were found together with Dune Helleborines
likely to have been a minor variant. and intermediates were reported, leading
Var. cordata is poorly-known. It was described in to the conclusion that Narrow-lipped and
1950 as having smaller, more bell-shaped flowers Dune Helleborines must be one and the same
with a broader lip and less spreading sepals. species. However, more recent genetic studies
have shown that all the plants in northern
BIOLOGY England are Dune Helleborines, and that they
are genetically distinct from the true Narrow-
lipped Helleborine, which is once again
Narrow-lipped Helleborine is routinely self-
restricted to its classic habitat, the southern
pollinated but occasionally the flowers may
beechwoods.
have a functional viscidium. Visiting insects will
Narrow-lipped Helleborine is one of a
then carry away the pollinia, and some cross-
group of self-pollinating helleborines found in
pollination is therefore possible. Seed-set is
Europe, all of which have evolved independently
good and each capsule forms 1,000-2,000 seeds.
from within the cross-pollinated Broad-leaved
Development and growth Helleborine complex. Being self-pollinated,
The rootstock lies rather deeply in the soil they have a restricted genetic diversity but
and numerous roots grow from it. Otherwise, some, such as Narrow-lipped Helleborine, are
there is no information on this species. A plant successful species which have spread widely.
002 systematic.indd 111 29/1/09 12:12:00

These self-pollinating species have long caused

problems for both taxonomists and botanists,
and their identification and correct classification
is only now being resolved by genetic and
biochemical techniques. For example, it has
been suggested that Narrow-lipped and Dune
Helleborines are the same as Epipactis muelleri
of Continental Europe (most recently by Stace
2004). Recent genetic research shows that this
is not the case.
HISTORY aND
CONSERVaTION
The first British record comes from ‘Woods
at Bosmere pool, Salop’ in 1841. This was
published in Leighton’s Flora of Salop and was
later identified as Narrow-lipped Helleborine.
It is Nationally Scarce, generally rather
localised and declining. Indeed, following the
clarification of the status of Narrow-lipped
and Dune Helleborines, it is now known to be
much scarcer than was reported in Scarce Plants
in 1994.
PastandpresentoccurrenceofNarrow-lippedHelleborine
inBritainandIreland(basedonpresenceorabsencein
10kmsquaresoftheNationalGrid; datafromtheNew
Atlas).
Britain Ireland
total historical range, 58 0 M 22July,Oxfordshire.Twostemsgrowingtogether.
1500-1999
current range 29 (1%*) 0 its woodland habitat and the conversion of
suitable woods to conifer plantations, although
% lost, 1500-1969 26% these practices have largely ceased. The loss of
% lost, 1970-1986 24% suitable shaded woodland due to the opening
% lost, total 50% up of the canopy by severe gales, such as the
* current range as a % of the total number of 10km squares great storms of 1987 and 1990, is a subtler
threat. The compaction of woodland soils
The New Atlas records Narrow-lipped by the use of heavy machinery in forestry
Helleborine from just 29 10km squares from operations, horse riding, mountain biking and
1987 onwards, and it has vanished from 50% the spread of wild boar are other potential
of its historical range, with many of the losses hazards. However, the most obvious is the
being comparatively recent. It is now extinct in widespread damage to flowering plants by the
Devon, West Sussex and Monmouthshire, has ever-increasing population of deer. Unlike
not been seen in Kent and Hertfordshire for Violet Helleborine, it does not seem to have
some years and was recorded for the first time in benefited from the abandonment of coppicing
Herefordshire in 1970 only to disappear again. during the 20th century, perhaps because it is
Direct threats include the destruction of slow to move into overgrown coppice.
002 systematic.indd 112 29/1/09 12:12:01

DUNE HELLEBORINE l 113
Dune helleborine Data Deficient

Epipactis dunensis
Other names: E. muelleri

Often considered to be one of the less attractive helleborines due to its relatively small, dull
flowers, Dune Helleborine has the distinction of being endemic to the British Isles and has
a very curious distribution and ecology. Typical plants are found in damp dune slacks on the
coasts of north Wales and northwest England and at a few inland sites in northern England
and southern Scotland. A distinct subspecies, the Tyne Helleborine, is found at other inland
sites in northern England, usually on old spoil heaps and in areas contaminated with heavy
metals. It differs genetically, albeit only slightly, from typical dune slack populations. Dune
Helleborine is the county flower of Lanarkshire.
Identification
Dune Helleborine has yellowish-green, two-
ranked leaves which are held rather stiffly at
about 45° above the horizontal. The upper part
of the stem is distinctly downy, and the flowers
are relatively small, do not open very widely and
are yellowish-green with the petals and base of
the lip washed with pink. The outer part of the
lip (epichile) is heart-shaped, usually broader
than long, and its tip folds downwards to a
variable extent as the flower ages. The base of
the flower stalk has a violet tinge.
Tyne Helleborine is found at inland sites
and has greener flowers with an epichile that is
longer than broad and not folded downwards.
The base of its flower stalk is yellowish-green.
Both forms are usually self-pollinated (see
p.76 for a discussion of the separation of cross-
pollinated and self-pollinated helleborines).
Similar species
Broad-leaved Helleborine typically has broader,
darker and greener leaves, arranged all around
the stem and not held rigidly erect. It has
larger and more widely opening flowers, often
with pink or purple tones on the sepals and
a distinctly purple tinge to the lip. Unlike
Tyne Helleborine, the outer part of its lip is
almost always strongly bent down and under;
P 6July, Lancashire.AtypicalDuneHelleborineamong
CreepingWillow,withsmall,slightly‘sickly’,yellowish-green
leavesthatareheldstifflyerect.
002 systematic.indd 113 29/1/09 12:12:02

usually self-pollinated, only very occasionally

showing a functional viscidium.
Green-flowered Helleborine is, like Dune
Helleborine, self-pollinating and potentially
more confusing. However, its upper stem and
ovaries are either hairless or have a few sparse
hairs. Its leaves are a clean apple green and often
short and rounded, and its flowers hang more-
or-less downwards, both in bud and when
open. A feature to check in difficult cases is the
fringe on the edge of the leaves (use a 20x hand-
lens). In Green-flowered Helleborine the tiny,
transparent, whitish teeth (cilia) are arranged
into irregular groups separated by gaps, but in
Dune Helleborine there is an even fringe of
minute teeth.
Narrow-lipped Helleborine is very similar
to Tyne Helleborine, and indeed, for almost
30 years after its discovery, Tyne Helleborine
was thought to be Narrow-lipped Helleborine.
Fortunately, separation of the two is academic,
as Narrow-lipped Helleborine is confined to
southern England, with a few colonies on the
Welsh Marches and in south Wales. The sepals
and petals of Narrow-lipped Helleborine are
on average larger and slightly broader, its lip is a
little more attenuated into a point, the bosses at
the base of the lip are washed pink and the slot
between them (the ‘keyhole’) is rather broader.
Habitat
Dune Helleborine is typically found in damp
dune slacks where it grows on the slightly
M 17 July, Northumberland. Tyne Helleborine typically higher and drier areas among and through a
appearsneatandclean-cut. carpet of low, scrubby Creeping Willow. It has
also spread into nearby pine plantations. Tyne
Tyne Helleborine can also be separated by
Helleborine, on the other hand, is found at
the yellowish-green base to its flower stalk
inland sites, in light, regenerating woodland,
(washed violet in Broad-leaved). However,
especially birchwoods, where the ground is
Broad-leaved Helleborine is very variable and
kept relatively open by the presence of heavy
in northern England and southern Scotland
some have more yellowish-green, two-ranked metals, mine waste or clinker. For example, in
leaves, although they still have large, widely Northumberland most of the populations grow
opening flowers (these are the so-called ‘Young’s along the River South Tyne on well-drained
Helleborine’, see p.97). In tricky cases it is gravel soils that are heavily contaminated by
worth checking the reproductive structures
P 17July,Northumberland.AgroupofTyneHelleborines
within the flower as Broad-leaved Helleborine growing under birches and willows by the River South
is cross-pollinated and Dune Helleborine Tyne.
002 systematic.indd 114 29/1/09 12:12:03

002 systematic.indd 115 29/1/09 12:12:06

habitats as all the inland sites are of recent

origin and are man-made. And, although
seemingly very different, adaptations which
allow the species to grow in dune slacks prone
to the stresses of summer drought and salt
spray may be similar to those needed in habitats
stressed by the presence of toxic metals.
Flowering period
Late June to mid-August, usually peaking in the
second week of July in the dune populations
but a little later in adjacent conifer plantations
and at inland sites. The flowers are short-lived
and can be badly affected by drought.
Range
Dune Helleborine is found on Anglesey, the
coasts of Merseyside and Lancashire and
by the Duddon Estuary at Sandscale Haws
in Cumbria. It has also been reported from
the coast of Co. Dublin. Tyne Helleborine
occurs in Northumberland, Cumbria, Co.
Durham and northwest Yorkshire. Scattered
populations of typical dune-type plants
have also been recorded at inland sites in
north Lincolnshire, southeast Yorkshire, Co.
Durham, Cumbria, and in southern Scotland
in Lanarkshire, Midlothian and West Lothian.
However, the features that separate these from
Tyne Helleborine (especially the colour of the
base of the flower stalk) have only recently
been clarified and all the records of ‘Dune
1987-99
1970-86
pre 1970
M 6July,Lancashire.AtypicalDuneHelleborinegrowing
upandthroughthepinkflowersofRestharrow.
zinc and lead tailings, but it is also found on

wooded spoil-heaps contaminated with lead.
In Scotland Dune Helleborine is found on the
wooded slopes of old ‘pit bings’ (the shale-rich
spoil heaps produced by coal mining), and at
Almond Bing (Falkirk) and Carlisle (Cumbria)
it is also found on and around old railway lines,
both in deep shade and in more open areas.
It seems likely that Dune Helleborine has
spread inland from its semi-natural coastal
002 systematic.indd 116 29/1/09 12:12:07

Helleborine’ at inland sites merit

re-examination. World range: Endemic to
Great Britain.
How to find it
Usually easy to find in dune slacks, although
often hidden among Creeping Willow. The
dune populations are all protected on reserves,
including Newborough Warren on Anglesey,
Ainsdale in Lancashire and Sandscale Haws
in Cumbria. Inland, the most accessible sites
for Tyne Helleborine are the tiny Williamston
Reserve and Beltingham River Gravels (both
Northumberland).
DESCRIPTION
upper sepal
petal anther
petal
pollinia
ovary
stigma
sepal
boss sepal
hypochile
(cup) epichile
Due to the significant differences between the M 6 July, Lancashire. A typical Dune Helleborine. The
outerpartofthelip(epichile)isbroaderthanlongand,in
two subspecies, they are described separately. thiscase,stronglywashedpink.
1. ‘Typical’ Dune Helleborine
Height: 20-40cm, sometimes to 50cm, with the teeth (cilia) 0.03-0.06mm wide. The leaves
tallest plants occurring in sheltered sites. are often damaged by wind or drought by
Stem: Pale green, tinged violet towards the base flowering time.
and downy towards the tip, with fine, pale hairs. Spike: Rather lax, with six to 20 flowers,
Usually arises singly, although sometimes there sometimes as many as 30, set to one side of the
may be two or three together. stem. The flowers are initially held horizontally
Leaves: Three to ten, arranged in two (patent) but often droop and become pendant
opposite rows up the stem and held stiffly as they go over. At the Scottish sites the flowers
at about 45°. The leaves are oval-lanceolate, are characteristically held in a drooping position,
mostly more than twice as long as wide and but this could be evidence of hybridisation.
become relatively narrower towards the flower Bract: Strap-shaped with a pointed tip; the
spike, but the lowest is very short, broad and lower bracts are slightly longer than the flowers,
rounded and forms a cowl-shaped sheath. but they become shorter towards the tip of
They are yellowish-green, deeply veined and the spike.
their margins have fine, regular, whitish Ovary: Green, hairy, six-ribbed (but not
002 systematic.indd 117 29/1/09 12:12:08

distinct notch running between the epichile and

hypochile. The anther is yellowish-green and is
variably stalked. The pollinia are crumbling and
whitish. The viscidium, although present in the
bud, usually disappears as the flowers open.
2. Tyne Helleborine
Height: 17-52cm.
Stem: Green, with fine, whitish hairs on the
upper part. Usually grows singly but fairly
frequently two stems arise together.
Leaves: The three to seven fresh green leaves
are arranged in two opposite rows up the stem,
although sometimes not obviously so. They
are rather floppy and not held as rigidly erect
as in typical dune slack plants; indeed, the
uppermost leaves and lower bracts may be held
horizontally. The leaves are deeply veined but
not sharply folded, the lower elliptical, mostly
more than twice as long as wide, the uppermost
long, narrow and grading into the lower
bracts. The leaf margins have very fine, regular
teeth (cilia) 0.01-0.05mm wide, sometimes
imperceptible.
Spike: Usually rather lax, with five to 35 flowers
M 6July,Lancashire.AtypicalDuneHelleborinewiththe
tipofthelipbentdownandunder. (mostly between ten and 25), often facing to
one side. Initially held horizontally, they droop
twisted) and pear-shaped, tapering into a short as they age.
stalk that is washed violet at the base. Bract: Fresh green, lanceolate and several times
Flower: Relatively small, dull and not opening the length of the flower in the lower part of the
widely, the flower is cup-shaped. The sepals spike but shorter at its tip.
are yellowish-green, oval-triangular, relatively Ovary: Green, hairy, ribbed (but not twisted)
short and blunt with a prominent midrib on and tapering into a yellowish-green stalk.
the outer surface. The petals are similar in Flower: Greenish and relatively small but
shape but slightly smaller and very pale green, opening quite widely. The sepals are narrowly
often washed pink. At the base of the lip, the oval in shape, elongated towards the bluntly
interior of the hypochile is reddish-brown to pointed tip. They are pale green on the outer
dark brown and contains nectar; the exterior of surfaces, paler and more whitish-green on the
the hypochile is whitish, variably flushed pink inner surfaces with a diffuse whitish margin.
(the colour shining through from the inside). The petals are similar but slightly smaller
The epichile is whitish, sometimes with a pink and proportionally shorter, paler and whiter.
flush in the centre, with a greener tip. Heart- The hypochile is transparent-whitish with its
shaped, it is usually broader than long and the interior washed dirty chocolate-brown at the
tip is sometimes folded under, especially as the base and rear, this colour shinning through to
flower matures; often the entire epichile is bent the exterior of the cup. The epichile is heart-
downwards near the join with the hypochile. At shaped, variably longer than broad, with a
the base of the epichile there are two smooth, pointed tip that projects forward. At its base
pink or greenish bosses, and these frame a there are two small, smooth bosses that frame
002 systematic.indd 118 29/1/09 12:12:09

a narrow, longitudinal groove. The epichile,

including the basal bosses, is whitish with a
faint cream or greenish wash. The column is
greenish-white, the pollinia cream-coloured,
and the anther cap is dull pale ochre with a
brown stripe on either side. The rostellum is
short, around half as long as the anther. The
clinandrium (depression on the top of the
column) is much smaller than ‘typical’ Dune
Helleborine, and the viscidium is always absent.
Subspecies
E. d. dunensis is found in coastal dune systems
on Anglesey and in northern England, and
at a handful of inland sites in Lincolnshire,
Yorkshire, Cumbria and southern Scotland.
E. d. tynensis occurs at inland sites in
northern England (Yorkshire, Cumbria and
Northumberland, with genetic analysis of three
populations in Northumberland confirming
subtle differences from dune-slack populations).
There is, however, a near-continuous
gradation between inland sites that hold
classic Tyne Helleborines (e.g. Williamston in
Northumberland) to those with plants with
intermediate characters and those holding
plants much like dune-slack populations listed
above under E. d. dunensis.
Among dune populations, those growing
in the open have a distinctive, almost ‘sickly’
yellow cast and often look rather wind-blasted,
whereas those growing nearby in the shelter of
plantations are taller, have more flowers and
both the leaves and flowers are greener and
‘healthier’. Dune-slack type plants growing at
inland sites are also reported to be greener and
more robust.
M 17 July, Northumberland;Tyne Helleborine.The pale,
nearlywhitish,tipofthelipcontrastswiththesooty-brown
BIOLOGY interiorofthecupatitsbase(thehypochile).
Dune Helleborine is self-pollinated and even be removed by insects and cross-pollination to
in bud, fragments of the pollinia may fall onto take place. Even when no viscidium is present,
the stigma and effect pollination. However, in plants may be cross-pollinated, wasps or other
some plants the viscidium may persist until insects carrying away the pollinia that stick
the flower has opened, allowing the pollinia to to their bodies regardless. In one case, around
002 systematic.indd 119 29/1/09 12:12:11

P 17 July, Northumber-

land;TyneHelleborine.The
tip of the lip is pointed
and does not bend down
andunder;itisalsorather
narrowandusuallylonger
thanbroad.
two-thirds of the flowers in a population of no scientific name. At Bardykes Bing near

Tyne Helleborine had their pollinia removed Glasgow there is a hybrid swarm of Dune and
(Richards 1986). At a site near Glasgow, Broad-leaved Helleborines and the individual
genetic studies have shown that Dune plants are very difficult to identify.
Helleborine is interbreeding with Broad-leaved Name and classification
Helleborine to form a hybrid swarm. The specific name dunensis means ‘of dunes’.
Vegetative reproduction may also take place, The taxonomy of Dune Helleborine is
the rhizome producing two flower spikes and complex and confusing, and is best understood
these in turn producing buds and roots. in seven stages:
Development and growth Stage 1: In 1921 Dune Helleborine was first
The rhizome is short, slender and woody. It lies recognised as something different and described
deeply buried in the soil and bears up to ten as a subspecies of Narrow-lipped Helleborine,
thin, wiry roots. There is no information on the using the name E. leptochila dunensis.
Stage 2: In 1926 Dune Helleborine was
period between germination and flowering.
elevated to the status of a full species, E.
Hybrids dunensis, known only from coastal dune systems
Broad-leaved Helleborine has been found at on Anglesey and in northwest England.
most (or all?) of the sites in Scotland. Although Stage 3: Colonies of plants identified as
confirmed by genetic analysis, this hybrid has Narrow-lipped Helleborine, a species hitherto
002 systematic.indd 120 29/1/09 12:12:13

known only from mature woodland on chalk HISTORY aND

and limestone in southern England, were CONSERVaTION
found at inland sites in Yorkshire in 1967, Dune Helleborine is Nationally Scarce and
Northumberland in the early 1970s and recorded from just 26 10km squares in the
subsequently more widely in northern England. New Atlas.
(These would eventually become known as the Dune slack populations are confined to a
Tyne Helleborine.) small number of sites but at these localities
Stage 4: From the late 1970s onwards a few Dune Helleborine is often common, for
colonies of Dune Helleborines were discovered example, at Sandscale Haws in Cumbria over
at inland sites, from Lincolnshire northwards. 1,000 flowering plants have been counted. It
Stage 5: The discovery at a few sites in northern is, however, sensitive to the level of grazing,
England of populations apparently intermediate especially by rabbits. If this is too intense the
between Dune Helleborine and Narrow-lipped spikes are all nipped off and the plants fail to
Helleborine led to the conclusion that they set seed. Conversely, if grazing is too light,
must be one and the same species, and that scrub invades the habitat and may eventually
Dune Helleborine should therefore be treated shade it out.
as a variety of Narrow-lipped Helleborine, The inland populations in northern England
named E. leptochila var. dunensis. were found in the 1960s and 1970s. Some were
Stage 6: More recently, research, based in part large, with colonies of over 1,000 flowering
on DNA studies, has led to the conclusion plants. All these sites are in young, ‘secondary’
that all the populations in northern England, woodland, however, and as this matures the
both ‘Dune’ and ‘Narrow-lipped’, are closely accumulation of humus seems to buffer the
related, but not the same species as Narrow- effects of the toxic metals and the ground
lipped Helleborine in southern England. These cover increases; such changes may lead to the
northern helleborines take the name disappearance of the helleborines.
E. dunensis, a species endemic to Great Britain.
Stage 7: Tyne Helleborine is formally named PastandpresentoccurrenceofDuneHelleborineinBrit-
ainandIreland(basedonpresenceorabsencein10km
as a distinct subspecies, E. d. tynensis. There squaresoftheNationalGrid;datafromtheNew Atlas).
is support for this from genetic studies, which
do show some differences between Tyne Britain Ireland
Helleborine and typical dune slack plants. total historical range, 26 1?
However, they also indicate that they are 1500-1999
probably each other’s closest relatives, have the current range 24 (0.8%*) 0
same ancestor and have only recently separated
% lost, 1500-1969 0% 0
into two slightly different entities.
% lost, 1970-1986 8% 0
Note that the helleborines on Holy Island
off Northumberland, once thought to be % lost, total 8% 0
Dune Helleborine, have now been separated * current range as a % of the total number of 10km squares
as a distinct, endemic species, Lindisfarne
Helleborine (see p.122). Delforge (1995)
suggested that Tyne Helleborine, together
with Lindisfarne Helleborine, were the British
representatives of a European species, Muller’s
Helleborine E. muelleri. The genetic evidence
shows that this is not the case, but this name
has nevertheless been used in some recent
publications.
002 systematic.indd 121 29/1/09 12:12:13

linDisfarne helleborine BaP

Epipactis sancta
For almost 50 years after their discovery, the helleborines on Holy Island in Northumberland
were considered to be Dune Helleborines. However, studies in the 1980s highlighted some
apparently minor differences, and recent genetic work has confirmed that they are not the
same as the helleborines growing in the dune slacks along the coasts of northwest England
and Anglesey. Rather they are a distinct species, endemic to this one small island.
Identification
Straightforward, due to its extremely limited
distribution, although this species is very
similar to Dune Helleborine, with yellowish-
green leaves held in two opposite ranks and
rather dull, greenish flowers which are normally
self-pollinated.
Similar species
Dune Helleborine is very close in appearance,
but typical dune slack plants have slightly
larger flowers and the base of the flower stalk
is washed violet (greenish in Lindisfarne
Helleborine); the details of the column also
differ. Lindisfarne Helleborine is even closer to
Tyne Helleborine, which also has a greenish-
yellow flower stalk and a similar column
structure, but the lip of Tyne Helleborine tends
to be slightly longer and narrower, and does not
turn under at the tip.
Habitat
Dunes and dune slacks, especially the slightly
raised and more steeply sloping zone around
the perimeter of the slacks. It grows among
Creeping Willow and various grasses or, just as
frequently, on bare sand among Marram grass.
It may be associated with the disturbed ground
around rabbit burrows.
Flowering period
Late June and the first three weeks of July but
usually at its best early in July. Self-pollinated;
the flowers go over quickly.
P 16July,Northumberland.Towardstheendoftheflow-
eringperiod, withthelowerflowerswithered, butinthis
robustplanttheleavesareunusuallyundamaged.
002 systematic.indd 122 29/1/09 12:12:15

LINDISFARNE HELLEBORINE l 123
002 systematic.indd 123 29/1/09 12:12:19

Range
Only found on the island of Lindisfarne off
the coast of Northumberland, growing at the
Snook at the western end of the island. World
Range: Endemic to England.
How to find it
Found singly and in small groups scattered
through the dunes of the Snook, although
scarce and rather scattered, and absent from
much apparently suitable habitat. The species is
prone to drought and may not flower if it is too
dry, or the buds will shrivel before opening.
DESCRIPTION
upper sepal
petal anther
petal
pollinia
ovary
stigma
sepal M 16 July, Northumberland.The stem and ovary are

boss distinctly hairy, and the pollinia have apparently been
hypochile sepal removed.
(cup) epichile
Spike: Fairly loose, with most flowers facing
Height: 6.5-42cm.
to one side and held roughly horizontal or just
Stem: Yellowish-green, finely hairy towards
below the horizontal, although the flowers
the tip.
droop as they age. The spike consists of up to
Leaves: Yellowish-green, arranged in two
ten flowers, although large plants may have 27
ranks on either side of the stem and held at
or more.
around 45°. The lowest leaf is rather short,
Bract: Yellowish-green, lanceolate, longer than
broad and rounded and forms a cowl-shaped
the flowers in the lower part of the spike but
funnel near the base of the stem. The two to
rather shorter than the uppermost leaves and
six higher leaves are elongated-oval in shape,
becoming shorter towards the tip.
becoming narrower and more lanceolate
Ovary: Green, ribbed, hairy and quickly
towards the spike. They are clearly veined
inflating as the flower self-pollinates. The flower
and longitudinally folded but not sharply
stalk is green or yellowish-green.
keeled. The leaf margins have tiny, very fine,
Flower: Rather small and drab but opening
regular teeth (cilia), 0.01-0.05mm wide
fairly widely. The sepals are greenish and
(sometimes imperceptible). By flowering
roughly triangular with the petals smaller
time many leaves are wind-burnt, grazed or
and paler. The hypochile is whitish with a
otherwise damaged.
chocolate-brown inner rear wall to the cup and
sometimes a pink wash to the exterior. The
O 7 July, Northumberland. Two flower spikes growing
amongst CreepingWillow; note the blooms of the intro- epichile is heart-shaped with the tip variably
ducedPirri-pirri-burr,somethingofapestonLindisfarne. deflexed and whitish, washed green in the
002 systematic.indd 124 29/1/09 12:12:22

LINDISFARNE HELLEBORINE l 125
England and Wales. Delforge & Gévaudan

(2002) went on to propose the name Epipactis
sancta for the species. Genetic studies show that
not only are Holy Island plants distinct from
both classic dune slack populations and the
Tyne Helleborine but also that they probably
evolved independently of both of these.
HISTORY aND
CONSERVaTION
These helleborines were first found on Holy
Island in 1958. In recent years there has been
an increase in the number of flowering plants
due to a reduction in the number of rabbits
(which nip off the stems) and around 150-300
M 7July,Northumberland.Thelipistypicallywhitish,be-
cominggreentowardsthetip.
spikes appear annually. The presence of rabbits
may be important, however, despite their
centre towards the tip. The column is whitish, impact on flowering numbers; they prevent
the anther cap yellow with a narrow brown scrub from invading the dunes and the bare
stripe at the side, and the pollinia are ochre- ground created by their scrapings may help in
yellow. The clinandria (depressions on the the establishment of seedlings.
top of the column) are very reduced and the
rostellum is short, around half the length of
the anther.
Subspecies
None.
None.
BIOLOGY
Self-pollinated.
No information.
Hybrids
None.
The specific name sancta means ‘holy’ or ‘sacred’,
a reference to Holy Island.
Holy Island plants were initially identified
as Dune Helleborine, but the colour of their
flower stalks and details of the flower structure
led to suggestions from the 1980s onwards
that they were not the same as the Dune M 16 July, Northumberland.The yellow pollinia are still
Helleborines growing on the coast of northwest intactonthisplant.
002 systematic.indd 125 29/1/09 12:12:24

Green-flowereD helleborine
Epipactis phyllanthes
Formerly: E. pendula, E. vectensis. Other names: Pendulous-flowered Helleborine

This mysterious orchid has a habit of coming and going at its known sites and of popping
up unexpectedly in new places, so is always worth looking out for in almost any type of
woodland in England and Wales. It is also one of the most variable orchids, and this has
caused a great deal of confusion over the years.
Identification
Green-flowered Helleborines come in a
wide variety of sizes and shapes. Some are
diminutive plants with large, swollen, pear-
shaped ovaries and small flowers which hang
vertically downwards and never really open.
Others are rather robust, with wide-open,
saucer-shaped flowers that can be held facing
more outwards than downwards (with such
well-developed plants being commonest
in northern England). However, despite
this diversity, this species often has a fairly
distinctive ‘feel’.
Green-flowered Helleborine is typically
relatively slender with short leaves which are
a fresh apple-green colour. Leaf shape and
posture are variable, but some plants have
characteristically well-spaced leaves that are
very rounded and held stiffly horizontal. The
flowers are green with a whitish or sometimes
pinkish lip and often do not open widely or,
indeed, may not open at all; such plants appear
to be permanently in bud, although the large
ovaries swell conspicuously. The base of the
flower stalk is greenish, and in most plants
the flowers hang vertically downwards. The
lip shape is very variable. In some populations
it is almost identical to the petals (a feature
shown by no other British helleborine) and
in others the lips are fully formed and divided
O 1 August, Northumberland, var. pendula. A typically

small, few-flowered plant, with very rounded leaves held
horizontallyandflowersthatdonotopenwidely.
P 12July,Northumberland,var.pendula.Arobustplant,
withmanyflowers, butalmostallofthem, asusual, are
drooping.
002 systematic.indd 126 29/1/09 12:12:25

GREEN-FLOWERED HELLEBORINE l 127
002 systematic.indd 127 29/1/09 12:12:28

as normal into a cup-shaped hypochile and

heart-shaped epichile. The flowers are generally
self-pollinated, the pollinia crumbling within
the flower onto the stigma.
Whatever the appearance of the flowers,
three features will confirm an identification.
First, the upper part of the stem and the
ovaries are either hairless or, rather less
frequently, the upper stem may be sparsely
hairy (use a hand-lens). Second, the hypochile
is greenish-white, lacking an obvious dark
brown or purple lining. Third, in difficult
cases look at the edge of the leaves (a 20x
hand-lens is best for this). In all the other
helleborines there is an even fringe of tiny,
whitish, hair-like projections, but in Green-
flowered Helleborine these cilia are unevenly
distributed in groups.
irregularly bunched cilia along leaf margin
Similar species
Narrow-lipped Helleborine is found in
beechwoods in southern England and south
Wales and is also greenish overall. At all times
it can be distinguished by its rather hairy upper
stem and ovaries.
Dune Helleborine occurs on coastal dunes
in Anglesey and northwest England, and as the
distinct variety known as Tyne Helleborine at
inland sites in northern England and southern
Scotland. It always has a hairy upper stem and
ovaries and a dark lining to the hypochile at
the base of the lip. In addition, typical dune
slack plants have a violet wash to the base of
the flower stalk (yellowish-green in Tyne and
Green-flowered Helleborines).
Broad-leaved Helleborine sometimes has
greenish flowers but always has a densely hairy
upper stem and slightly hairy ovary. It is also M 26August,Norfolk,var.degenera.Verytypically,grow-
ing through a carpet of Ivy. In this season the flowers
cross-pollinated (see p.76 for a discussion of hardlyopenedatall.
002 systematic.indd 128 29/1/09 12:12:29

the separation of cross-pollinating and self- Flowering period

pollinating helleborines). Late June to early or even mid-September but
mostly from mid-July to mid-August, with
Habitat dune populations typically earliest.
Very varied, and although the species favours
alkaline soils, it is not confined to areas of Range
chalk or limestone and will grow on calcareous In England rather scattered and local but
to mildly acidic sands and clays and in silty found west to Dorset, Gloucestershire and
river valleys. It is mostly found in light to Herefordshire, east to Kent and Norfolk and
moderate shade but can occur in densely north to south Cumbria and Northumberland.
shaded situations, although it is often rather The bulk of the population is found in central
small and slight in such places. Green-flowered and northern Hampshire, Wiltshire, West
Helleborine frequents a wide variety of Sussex and Surrey, where Green-flowered
woodlands but has a definite preference for Helleborine can be locally frequent. In Wales
beechwoods in southern England and overall there are scattered sites in coastal dunes
favours smaller woods, copses, belts of trees in Glamorganshire (including Kenfig and
or tall hedges next to woodland, or the better- Whitford Burrows) and Merionethshire
lit edges of larger woods; it is often found on (Morfa Dyffryn), but it is otherwise only found
road verges. It prefer areas where the ground in Flintshire. There are also a handful of sites
cover is rather sparse or low, no taller than 15- in Ireland, in Co. Dublin, Co. Leitrim and
20cm, and is characteristically found growing Co. Ferrmanagh. World range: Restricted to
through a carpet of Ivy. western Europe, from northern Spain through
Rather than having an association with France and Belgium to Denmark. May occur
ancient woodland, many of its sites are more widely but the taxonomic confusion
relatively recent in origin, such as beech surrounding the species and its close relatives
and pine plantations and shelterbelts and, in Europe has clouded an understanding of its
in Northumberland, the maturing birch and true range.
hawthorn scrub found on old waste tips
contaminated with zinc and lead. Another
1987-99
favoured habitat is thickets of willows and 1970-86
pre 1970
other trees alongside rivers and streams
that are subject to occasional flooding, the
helleborines growing on the better-drained
ridges and banks. Indeed, this may be the
‘natural’ habitat of the species and it has
even been found in Hampshire growing
among reeds and willows in the tidal part of
the River Itchen. Conversely, Green-flowered
Helleborines also occur in very dry woods.
At a few sites in Wales, northwest England
and Co. Dublin, it grows in the open on
sand dunes, coming up through a blanket of
Creeping Willow on the drier hummocks.
However, it tends to look yellow and ‘sickly’
in this habitat, appearing rather healthier How to find it
where it has spread into adjacent conifer Its overall green coloration, relatively small size,
plantations. habitat and flowering period can make this a
002 systematic.indd 129 29/1/09 12:12:30

hard species to find, although the apple-green, frequently funnel-like, partially sheathing the
rounded leaves, which are held horizontally, and stem. The leaves may be arranged in two ranks
the mass of large, drooping buds often catch the on either side of the stem and on some plants
eye. Colonies are often small, and numbers can are held flat in a very characteristic horizontal
fluctuate, with fewer plants in dry years. Most plane. On others they may be held lightly folded
of the sites for the species in southern England and positioned nearer to 45°. The margins
are anonymous and have little or no other have tiny, whitish teeth (cilia) arranged into
orchid interest, and the most accessible sites to irregular groups.
see the species at its best are the dunes on the Spike: The buds may be held upright,
Lancashire and Cumbrian coasts. horizontally or drooping, but, once open, the
flowers usually hang down near to the vertical.
DESCRIPTION The spike contains between two and 25 flowers,
exceptionally as many as 35, and is often fairly
upper sepal crowded. The flowers often face in the same
petal anther
direction, and the uppermost buds in the spike
may fail to open.
petal Bract: Green and lanceolate; the lowest
bracts are much longer than the flowers but
pollinia
ovary they become progressively shorter, with the
stigma uppermost a little shorter than the flowers.
sepal
boss sepal
hypochile
(cup) epichile
Height: 5-75cm but mostly 15-50cm; taller

plants are commonest in northern England.
Stem: Robust, purple-brown at the base
(where there are one to three similarly coloured
sheaths) but becoming apple-green for most of
its length. The stem, including the upper part, is
hairless or has sparse, short hairs. Usually grows
singly but occasionally two or three stems may
arise together. We have seen a photograph of
a group of nine (Killick et al. 1998), although
such a large cluster may have resulted from the
rhizomes of several plants growing together.
Leaves: There are three to seven, exceptionally
as many as 16, apple-green leaves which are well
spaced but often rather high up on the stem.
They are very variable in shape and posture
but usually relatively small. Many plants have
leaves that are broadly oval or egg-shaped but
on others they are longer, narrower and more
pointed. On all plants, the upper leaves are
M 12 July, Northumberland, var. pendula.The ovary is
usually narrower and more bract-like, and boldly-ribbedbuthairless,andthebaseoftheflowerstalk
the lowest green leaf is often rather small and is green.
002 systematic.indd 130 29/1/09 12:12:33

The bracts are usually horizontally, especially on

less densely crowded spikes.
Ovary: Swells rapidly and becomes large and
pear-shaped. It is shiny, virtually hairless but
with six prominent ribs, tapering into the short,
curved and twisted green stalk.
Flower: Var. pendula (see below for other
flower shapes). Overall greenish. The sepals are
elongated-ovals, variably tapering to a pointed
tip, and pale green with a prominent green
midrib on the outer surface. The petals are a
little smaller, paler and more greenish-white.
The hypochile is whitish or dull olive-white,
almost translucent, with the interior of the
cup pale greenish (sometimes lightly washed
brown). The epichile is heart-shaped with a
pointed tip which is strongly turned under, and
there are two rough bosses at the base with a
central groove between them. It is whitish and
tinged with green or sometimes pale pink in the
centre or towards the tip, especially the bosses.
The column is whitish and the anther cap dull
yellowish-white with narrow brown stripes
at the side. The pollinia are cream or whitish,
quickly crumbling and inconspicuous in the
anther. A small and not very sticky viscidium
may be present in the bud, but this withers by
the time the flower opens.
Subspecies
None.
Green-flowered Helleborine is very variable,
both in terms of the structure of the flowers
and in the overall size and shape of the plant.
The flowers range from those where the lip is
well developed, with a clear distinction between
the basal cup (hypochile) and the heart-shaped
tip (epichile), to those where the lip is simple
and petal-like. There are also parallel differences
in the shape of the anther. Between the two
extremes the range of variation is almost
continuous. There are four named varieties,
although due to the variation it can be difficult
to identify plants. Notably, there is no real
M 1 August, Northumberland, var. pendula. The lip is
dividing line between var. pendula and var. well-formedinthisvarietyand, onthisplant, theflowers
vectensis, and flowers with mixed characters can haveopenedextraordinarilywidely.
002 systematic.indd 131 29/1/09 12:12:35

be found. It would probably be better to unite

the two varieties as var. vectensis.
There are broad geographical trends in
lip shape, with populations in the north and
west having the most robust plants, the best
var. pendula
developed lip structure and widely opening
flowers (var. pendula), whereas those in the
south tend to be smaller, with incompletely
developed flowers that open only partially. var. vectensis
Even in the same region, however, there can
be a good deal of variation, and at a few sites
a mixture of lip shapes can be found in the
same colony.
Var. pendula The lip is large and fully var. degenera
var. degenera
var. phyllanthes
var. phyllanthes
(After Young 1952b)
developed. The hypochile is well formed and

around 4mm long. The epichile is heart-
shaped, as long as the hypochile or only
slightly longer, with a pointed tip that is
normally strongly reflexed. The epichile is
wrinkled at the base or has two bosses. The
flowers open widely and cleistogamic flowers
are rare. This is the commonest variety in
north Wales and northern England but it has
been found further south and intermediates
with var. vectensis are not infrequent.
Var. vectensis As var. pendula but the
hypochile is smaller, 2.5-3.5mm long, and
more hemispherical so that it embraces the
column closely. The epichile is distinctly longer
than the hypochile, sometimes markedly so,
M 17 July, Norfolk, var. vectensis. In this variety the lip is and the tip is generally not reflexed so that the
well formed and it can, as here, have a pink tinge. whole lip points forwards. The flowers open to
002 systematic.indd 132 29/1/09 12:22:02

a variable extent and are often cleistogamous.

Mostly found in southern England and the
Midlands but has been recorded north to
Yorkshire. ‘E. cambrensis’ was described from a
few reportedly distinctive plants, delicate and
yellowish-green, with yellowish-white flowers,
found in the 1940s at Kenfig and Margam
Burrows in south Wales. Initially described
as a distinct species, it has more recently been
demoted to ‘E. p. var. cambrensis’, but was not
recorded again until similar plants were found
at Kenfig NNR in 2004. The most distinctive
feature of ‘cambrensis’ appears to be the leaves,
which are narrow and sharply folded along the
midrib, but whether it is appropriate to treat
it as a named variety has still to be confirmed.
Var. phyllanthes The lip is not divided into
hypochile and epichile and is oval or lanceolate
with a central midrib, resembling a petal in
size and shape. The flowers rarely open widely
and are usually cleistogamous. This form is
commonest in southern England.
Var. degenera This is intermediate between
var. vectensis and var. phyllanthes. The
hypochile is reduced to a shallow depression
at the base of the lip and although there may
be small bosses at the sides of the base of the
‘epichile’, the lip has either no waist at all or
only a rudimentary one and always lacks
the central groove between the two bosses.
The flowers rarely open widely and are
usually cleistogamous. It is mainly found
in southern England.
The size and shape of the plants and their
overall colour also varies, especially the size
and shape of the leaves. This variation seems
to be determined by the local environment
(for example, on exposed dunes the plants are
dwarfed and yellowish). In addition, individual
plants and whole populations vary in the
degree to which the flowers open and how
pendulous they are, although this can change
from season to season; plants in which the
flower buds never open are commonest
in dry seasons, whereas in very wet years
M 14August, Norfolk, var. degenera.The flowers have
openedasmuchastheyaregoingto,andtheovariesare more plants than normal may show widely
allswollen(theywerepresumablyself-pollinatedinbud). opening flowers.
002 systematic.indd 133 29/1/09 12:12:37

BIOLOGY in the number of records in recent years but

this is offset by the loss of known sites. It is
Pollination and reproduction extinct in the Isle of Wight, Somerset, Suffolk,
Green-flowered Helleborine is always self- Warwickshire, Derbyshire, Co. Antrim, Co.
pollinated. In some plants this occurs when Derry and Co. Wicklow.
the flowers are still in bud (the flowers are
cleistogamous); the buds may or may not open Past and present occurrence of Green-flowered Helle-
subsequently. Following pollination the whole borineinBritainandIreland(basedonpresenceorab-
sencein10kmsquaresoftheNationalGrid; datafrom
column, together with the lip, withers rapidly the New Atlas).
but the sepals and petals can remain intact for
Britain Ireland
a long time.
Development and growth 1500-1999
The aerial stem grows from a rhizome which current range 86 (3%*) 4 (0.4%*)
varies from nearly horizontal to nearly vertical, % lost, 1500-1969 21% 44.5%
with numerous thick, fleshy roots, both long % lost, 1970-1986 15% 11%
and short. There is no information on the
% lost, total 36% 55.5%
period between germination and flowering.
Hybrids
None. Losses may be due to the grubbing-out
Name and classification or coniferisation of woodland, but some
The specific name ‘phyllanthes’ may celebrate of its habitats are ephemeral in nature and
Phillis Wood near Treyford in West Sussex, become unsuitable as the woodland matures
the site from which the first specimens were and becomes more shaded. In general, there
collected in the 1830s, but is more probably has been a decrease in numbers in southern
from a compound of Greek words meaning England but some spread on the edges of the
‘leaf-like flower’. range. For example, it was first found in Norfolk
This species was misunderstood for a in 1969 at Santon and subsequently a number
long time and widely confused with Narrow- of colonies were discovered in the 1990s
lipped Helleborine. It was not until 1952 along the banks of the Rivers Yare and
that E. vectensis (described as a variety of Wensum in Norwich.
Narrow-lipped Helleborine in 1918 and then
as a distinct species in 1940) and E. pendula
(described in 1942) were shown by the British
botanist Donald Young to be the same as E.
phyllanthes, a species described in 1852 in the
Gardeners’ Chronicle but ignored for a hundred
years. ‘Epipactis cambrensis’ (described in 1950)
has since been added to the pot-pourri.
HISTORY aND
CONSERVaTION
Nationally Scarce in Britain and specially
protected in Northern Ireland under Schedule
8 of the 1985 Wildlife Order (NI). A better
awareness of the species has led to an increase
002 systematic.indd 134 29/1/09 12:12:38

Distribution
There are around five species in this genus, distributed in
Europe, Asia, Africa and through the Pacific Islands to
Australia. All are fully mycotrophic and depend on fungi
throughout their life.
Name
The generic name Epipogium derives from the Greek and
means ‘overbeard’, a reference to the position of the lip,
uppermost in the flower.
GENUS EPIPOGIUM
GHOST ORCHIDS
003 systematic.indd 135 12/2/09 16:55:27
136 l GENUS EPIPOGIUM
Ghost orchid Extinct (?)

Epipogium aphyllum
This may be the rarest wild plant in Britain. It has only been recorded in two widely separated
regions: the Chilterns and around Herefordshire and Shropshire. Fully dependent on fungi
throughout its life, it only appears above ground to flower and fruit. There have, however, been
no documented sightings in England for almost 20 years and it has become the Holy Grail for
many orchidophiles. Once known as ‘Spurred Coralroot’, the English name Ghost Orchid much
more accurately reflects its elusiveness. Its small size and pallid, ethereal appearance make it
extremely hard to find in its gloomy woodland haunts – truly ‘ghostly’ qualities.
Identification
Very distinctive, it is pale and ‘waxy’ with
relatively large, pinkish flowers and no green
leaves.
Similar species
None.
Habitat
In Herefordshire and Shropshire it has been
found in oak woodland on clay soils. The
Chiltern sites are in beechwoods on chalk or
clay-with-flints. Ghost Orchid often grows
where the soil is slightly deeper, in hollows
or on the top and sides of ditches (but not,
apparently, in the bottom of a ditch). The
margins of roads and tracks appear to be
favoured but this may reflect the fact that it is
easiest to search for the plants in such places. It
often grows in a deep mat of decaying leaves but
may also be found in areas with almost no leaf
mould, the rhizome lying in the mineral soil
itself. Several times it has been found growing
out of old tree stumps, and one has even been
recorded growing through a rotten mattress in a
roadside ditch.
Ghost Orchid is often said to grow in
heavily shaded woodland where the ground
is otherwise bare. In 1953, however, having
discovered the largest ever colony in Britain,
Rex Graham stated, ‘The canopy, locally heavy,
is on the whole rather more open than what one M 23 July, Scwartzwald, Germany (Sean Cole). The spur
might have expected to be ideal for this orchid, of the flower lies at the top and the lip faces downwards.
and a few plants grew in comparatively light O 14 August, Buckinghamshire (Nigel Redman). A
conditions and amongst the type of ground ghostly apparition on the beechwood floor.
003 systematic.indd 136 29/1/09 12:25:04

GHOST ORCHID l 137
vegetation that invades these woods whenever to the Kamchatka Peninsula, Russian Far East,
light gives a chance’. Sakhalin, Japan, northeast China and Korea
and is also found to the south scattered
Flowering period
through the Himalayas and southern China,
April to mid-October but mostly mid-July to
including Taiwan.
mid-September. Plants in Buckinghamshire
tend to flower from mid-July to the third week How to find it
of August and those in Oxfordshire a little later, Ghost Orchid is probably the hardest plant
from mid-August to mid-September. to find growing wild in Britain. Its small size
Flowering is said to follow a wet spring. makes it extremely inconspicuous and very
This allows the plant to store up water prior to difficult to see in the deep leaf-litter of the
developing an aerial stem. The flower bud is, woodland floor. Plants may even be hidden
however, initiated in the year prior to flowering, under wind-blown leaves. Some ingenious
so the conditions in two consecutive years may techniques have been used to try to locate it,
be critical in determining whether the plant including night forays where a powerful torch
is able to bloom. The Ghost Orchid may even is shone parallel to the ground in the hope of
flower underground, buried in the leaf-litter. highlighting the tiny spikes. The prolonged
period during which it may bloom does not
help, nor does the fact that slugs so often
1987-99
1970-86 destroy the flower spikes; even if they survive,
pre 1970
the flowers do not last more than a few days.
Knowledge of the precise location of previous
records is of only limited use, as they seldom
reappear in exactly the same spot. Finally,
even in the right place at the right time few
flower spikes are produced, with one to four,
exceptionally seven, at any one site in any one
year. The total of 25 near Marlow in 1953 was
quite unique.
DESCRIPTION
Height: Usually 5-10cm but sometimes to
12.5cm and exceptionally to 24cm.
Stem: Swollen at the base, thick but fragile and
Range translucent-white, washed dull rose-pink or
Confined to England, where it has been pinkish-brown and variably streaked pinkish.
found in Oxfordshire, Buckinghamshire, Above the uppermost flower the stem continues
Herefordshire and Shropshire. World Range: as a short, narrow projection (like an aborted
Central and northern Europe and through flower stalk). Stems arise singly but occasionally
northern Asia to the Far East, with a few two spikes grow from the same rhizome.
scattered sites south to the Himalayas and Leaves: There are no green leaves, merely two
China. Occurs north to northern Scandinavia or three brown, sheathing scales at the base of
and south to the Alps, central Italy, northern the stem and one or two longer, often dark-
Greece, Crimea and Caucasus; also found edged, tightly clasping, scale-like leaves higher
on Corsica. Largely absent west of the Alps on the stem.
and Massif Central, but there are odd sites, Spike: Most plants carry one or two flowers,
including the Pyrenees. In Siberia it ranges east but the more robust bear three or four blooms,
003 systematic.indd 137 29/1/09 12:25:04

which are held pointing outwards or nodding

downwards on short stalks. Neither the stalk
nor the ovary is twisted, hence the spur points
more-or-less upward and the lip similarly lies at
the top of the flower.
Bract: Roughly equal in length to the flower
stalk and ovary together, papery and translucent
with a yellowish-straw wash.
Ovary: Bag-like, pale straw, veined or spotted
with pink or violet; the flower stalk is noticeably
slender and curves to hold the ovary hanging
downwards.
Flower: The spur and lip lie at the top of the
flower, with the lip facing downwards and
outwards. The lip is whitish or very pale pink
and divided into two sections: at the base the
hypochile is short with two broad, spreading,
triangular side-lobes flanking the entrance to
the spur; the epichile is longer, broadly tongue-
or heart-shaped and deeply concave with
crimped margins and several rows of raised
magenta ridges. The lip is strongly curved to
lie parallel with the spur, which it may touch.
The spur is relatively large, sack-shaped, slightly M 23 July, Scwartzwald, Germany (Sean Cole). At some
curved, pale pink and filled with nectar. The sites in Europe this enigmatic orchid is rather predictable
in its appearances and almost common, although never
sepals and petals are very pale yellowish-straw commonplace.
with fine reddish dots or streaks; they are
narrow and strap-shaped with their edges cap aside, and the slender stalks of the pollinia
curled inwards, making them appear even are glued to its back or head. However, it has
narrower. The ‘upper’ sepal and two petals hang been suggested that bees, although they do
down below the rear of the flower, and the two visit the flowers, are unlikely pollinators. They
lateral sepals project downwards and variably are the wrong shape and size, especially given
outwards at the sides. The column is pale yellow that the flowers are ‘upside-down’ (and all
and short with a well-developed rostellum; the references to bees as pollinators may originate
anther is pale yellow and the two pollinia are with the observations of a Dr Rohrbach in
connected by a stalk at their base to the two the 19th century); if this is so, the mechanism
distinct viscidia. The nectar is said to smell of pollination is a mystery. Self-pollination
of fermenting bananas or vanilla but it is also in flowers in the normal pendant position is
described as foetid. prevented by the position of the stigma, above
the anther, because pollen cannot fall upwards.
However, self-pollination may be possible while
BIOLOGY the flowers are upright when still in bud.
Pollination and reproduction Whatever the mechanism, few flowers
Possibly pollinated by bumblebees and wasps. produce capsules, perhaps because of the
The insect lands on the lip and moves towards lack of suitable pollinators in the dark woods
the spur to access the nectar. As it backs out, where Ghost Orchids are found, or because
it ruptures the rostellum, pushes the anther of a lack of other simultaneously flowering
003 systematic.indd 138 29/1/09 12:25:05

GHOST ORCHID l 139
Ghost Orchids to cross-pollinate the flower. Subspecies

The species certainly reproduces None.
vegetatively and this may be the major means
of propagation, although the literature is
None.
contradictory about the mechanism. Some
sources state that new plants grow from buds Hybrids
at the tip of the roots. Others state that some None.
of the branches of the rhizome become slender,
attenuated and thread-like, and function as Name and classification
underground runners. Bulbils are formed at The specific name aphyllum derives from the
intervals along these runners and these may Greek and means ‘without leaves’.
become detached and grow into independent
new plants. After flowering, the major part of HISTORY aND
the rhizome usually dies, leaving the runners to CONSERVaTION
form new plants. A Red Data Book species that is classified as
‘Extinct’, it is fully protected under Schedule 8
Development and growth of the Wildlife and Countryside Act 1981.
Throughout its life Ghost Orchid is entirely The first British record dates from 1854,
dependent on fungi for its nutrition (that is, when Ghost Orchid was found by the Sapey
fully mycotrophic). It is often stated that Ghost Brook on the border of Herefordshire and
Orchid is ‘saprophytic’ and gets its nutrients Worcestershire, as detailed in the Journal of
from decaying organic material, but this is Botany: ‘The discovery was communicated …
not correct, as it feeds by consuming a living on the 9th of this month [September], by the
fungus. It seems likely that, like Bird’s-nest and Rev. W. Anderton Smith, of Tedstone, from
Coralroot Orchids, its fungal ‘partner’ will prove the Rectory, Delamere, Bromyard: - “…a few
to be ectomycorrhizal and have a symbiotic weeks since, Mrs. Anderton Smith found a
association with the roots of trees. This would specimen… For some time we looked in vain
allow the Ghost Orchid to acquire nutrients, via for other specimens; but, on the 23rd ult., I
its fungal ‘partner’, from the trees. was fortunate in detecting a considerable mass
The aerial stem grows from a whitish of it. All were found at the foot of a very steep
rhizome which has many branches. In turn, woody bank, close to a brook; the soil very wet
these are forked or tri-lobed. The lobes are and stiff. As the banks are very much trampled
rounded at the tip and often spread fanwise. on at present (timber and faggots being drawn
The much-branched rhizome is said to look along), I decided on digging it up, and planting
like certain types of coral, hence the old name, it in a similar spot in our own grounds.”’
‘Spurred Coralroot’. There are no roots, rather The next records of Ghost Orchid came
the rhizome has a sparse covering of fine, long from Bringewood Chase near Ludlow
hairs. Once the plant is mature enough to flower, (then in Shropshire, the area is now in the
a bud appears in the autumn and swells as administrative county of Herefordshire). It was
the food and water reserves from the rhizome found by a Miss Lloyd in 1876 and again in
are transferred to it. If the following season is the same wood by a Miss Peel in 1878, with a
sufficiently wet, a flower spike may be produced. third record from a different part of the wood
Seed probably germinates in the autumn, in 1892. A Ghost Orchid was also recorded
and up to ten years, possibly even twenty, may from the Wye Valley near Ross-on-Wye,
elapse before flowering, although no one has Herefordshire, in July 1910.
performed a long-term study of marked plants There was then a gap of 13 years before
to confirm this. the species was seen again in Britain, this time
003 systematic.indd 139 29/1/09 12:25:06

in Oxfordshire. In June 1923 two plants were there were reliable reports from near Marlow
found just north of Henley-on-Thames. After until 1989 and several reports since from both
a prolonged search, Dr. G. C. Druce, perhaps Buckinghamshire and Oxfordshire, covering
the most eminent botanist of the day, succeeded almost every year up to 2003. The evidence for
in tracking down a third plant in July. In late some of these is certainly limited, however, and
May of the following year two more plants the finders have displayed a great reluctance
were found there. Then, on 30 June 1931, in coming forward and allowing the record to
Ghost Orchid was found a few miles to the be confirmed, probably to the detriment of
west by Mrs Vera Paul (then a schoolgirl), in the plant. The relative lack of records in recent
beechwoods towards Stoke Row. There was a years may merely be part of the natural cycle
single large spike, 24cm high with three flowers, of a small population or more likely reflects a
growing from the middle of an old tree stump. genuine decline. It has been suggested that the
A single spike was also seen nearby in 1933. opening of the canopy following tree falls after
The trail went cold again for 20 years and it the great storms of 1987 and 1990 may have
was not until 1953 that Ghost Orchid appeared resulted in the exposure of the woodland floor
again, this time in Buckinghamshire (around to too much sun, conditions becoming too dry
ten miles from the Oxfordshire sites). Rex for the species (but see Habitat).
Graham had been searching for Ghost Orchids Ghost Orchid is on the very edge of its
for 20 years and on 18 July 1953 was in woods range in England and is naturally rare. Its
west of Marlow. He was lighting his pipe when, position here has not been helped by the
over the bowl of it, he saw a Ghost Orchid unsympathetic management of ancient
growing among the beech leaves. There proved woodland, which has been cleared or replanted
to be a scattered colony of 25 spikes belonging with conifers (for example, Bringewood Chase
to 22 plants. Three specimens were collected in Shropshire has largely been ‘coniferised’,
and at least three more spikes were felled by with little ancient woodland remaining). Other
slugs. It was found there again the following more immediate threats include horse riders
year but not in such great numbers. These and cyclists who create paths through areas
woods went on to become the most productive where the plants have flowered in the past, and
area for the species, although the number of forestry operations which involve the use of
plants never matched the 1953 total. Spurred heavy machinery and the dumping of materials.
on by the events in Buckinghamshire, Vera Some of these problems could be avoided if the
Paul searched the woods near Stoke Row in responsible authorities knew the location of the
1953 and two spikes were found. Plants were plants, but there is such secrecy surrounding
seen there in 1954 and 1956, too, while on 17 Ghost Orchid that many sightings are ‘hushed
September 1963 five spikes were found close up’. Humans can be a direct threat, too, with
to the stump where the species had been first photographers trampling the area around
found in 1931. Sightings in this complex of flower spikes, and in 1978 and 1979 collectors
woods have continued, with reports up to and apparently dug up several plants. Finally, there
including recent years. are natural hazards, with deer and slugs taking
On 19 September 1982, after a gap of a toll. Well-meaning naturalists have advocated
62 years, the Ghost Orchid reappeared in surrounding plants with slug pellets, but even a
Herefordshire, when a single spike was found moribund slug may cause damage and there is
in a cart track under oak and pine in a wood far no information on the effect of molluscicides on
removed from its old localities. By 1 October the the plant or its fungal ‘partner’.
stem had rotted, probably due to slug damage.
The Red Data Book states flatly that Ghost
Orchid has not been seen since 1986, but P 16 June, Norfolk. Fen Orchid, a relatively robust plant.
003 systematic.indd 140 29/1/09 12:25:07

Genus LIPARIS
Fen orchid
Distribution
This small, green-flowered orchid
grows from a pseudobulb, a structure
commonly found in tropical epiphytes
and, although Liparis has an almost
worldwide distribution, it is essentially
tropical. There are 300 or so species but
just one is found in Britain and Europe.
Name
The generic name Liparis is from the
Greek liparos and means ‘fatty’, ‘greasy’ or
‘shiny’, a reference to the appearance of
the leaves.
003 systematic.indd 141 29/1/09 12:25:10

142 l GENUS LIPARIS
Fen orchid Endangered, BaP

Liparis loeselii
Other names: Loesel’s Twayblade (North America)

This small and rather drab orchid has always been very localised in Britain and is now confined
to just a handful of sites in Norfolk and South Wales. Indeed, it is threatened throughout its
range in Europe and has declined in much of North America, too. Surprisingly, studies of its
life history suggest that it is a ‘weedy’ species, well-adapted to colonising and rapidly filling
newly available habitats, and that it thrives on disturbance.
Identification
Very rare and localised, this is an orchid highly
unlikely to be stumbled upon by chance. When
in flower, Fen Orchid is distinctive. The small,
pale green flowers appear to be a jumble of thin,
spidery projections and are unique. Importantly,
they are usually held facing upwards, with the
column vertical, and the three strap-shaped
sepals, two thread-like petals and the lip all
held more-or-less horizontally in a cross; two of
the sepals lie parallel, close together under the
tongue-like tip, to form one ‘arm’ of the cross.
Similar species
Non-flowering plants may form the bulk
of the population. They could be confused
with Common Twayblade or Lesser Butterfly
Orchid but are much smaller and their leaves
sheathe the swollen base of the stem to form a
pseudobulb.
Habitat
Fen Orchid is found in two distinct habitats;
marshes in the Norfolk Broads and damp dune
slacks on the coast of South Wales. Although
superficially very different, both offer the same
combination of bare ground which is kept
damp or wet by neutral or calcium-rich
ground water.
In the Norfolk Broads, Fen Orchid
occurs as a member of a particularly species-
rich community in areas of wet, peaty fen
dominated by Common Reed or Great Fen
Sedge. It grows on the mossy carpet at the
M 18 June, Norfolk. The aerial stem grows from a pseudo
bulb, a swelling in the stem that is concealed by the base base of the reeds, on the sides and tops of
of the leaves. sedge tussocks, or on the bare peat itself. It is
003 systematic.indd 142 29/1/09 12:25:12

FEN ORCHID l 143
flowers capsules
old new new

pseudobulb pseudobulb pseudobulb
early summer late early

spring summer winter
roots
M Annual replacement of pseudobulbs in Fen Orchid.
extremely fussy, however, and an analysis of its with Creeping Willow, Marsh Pennywort and a
current and former sites in East Anglia reveals variety of mosses. As in the Norfolk Broads, the
that they have several features in common: species has very specific habitat requirements:
1. All are wet, with a water table which is 1. The slacks must be calcareous; calcium is
typically just below the surface and relatively provided by fragments of seashells in the sand.
stable. Fen Orchid does not necessarily require 2. The slacks must be moist (although they may
waterlogged conditions and, indeed, may be flooded in the winter, sometimes for as long
grow on tussocks up to 15cm above the water as five months, the water table can fall to more
level, but the presence of surface water could than 50cm below ground level in August and
be important in protecting it from slugs and September). 3. The slacks must be young.
snails. 2. The ground water is neutral to mildly Fen Orchid can colonise newly formed
alkaline and very low in nutrients. 3. Most slacks very quickly, within a few years of the
sites were cut for ‘sedge’ (Great Fen Sedge was slack developing from bare sand. Indeed,
used for thatching) or ‘litter’ (hay). The cutting establishment from seed seems to be most
and removal of reed and sedge, combined with successful in these young slacks, perhaps
infertile ground water, slow the development of because some bare ground remains, and
scrub and keep the vegetation relatively sparse completely new populations of Fen Orchid
and open. 4. Most of the sites in the Broads are are only found in such new slacks. As the
on abandoned turbaries where peat was cut as vegetation becomes more established, however,
fuel in past times. These cut-over areas then it is less able to compete; fewer new plants
flooded, and gradually the resultant ‘turf ponds’ appear and eventually there are no new recruits
were colonised by vegetation to form semi- to the population. Fen orchid will usually die
floating mats or ‘hovers’. These mats are able to out in a slack about 50 years after its formation.
rise and fall with changing water levels, and this
may be important in keeping the Fen Orchids Flowering period
at roughly the same level relative to the water In south Wales, early June to late July,
table. (As more peat accumulates, however, exceptionally as late as mid-September. In
and despite continued cutting or grazing, the Norfolk, early or mid-June to early July, with a
vegetation inevitably changes, and Fen Orchid few appearing later, to late July. In dry seasons
will eventually disappear.) few plants flower, whereas flowering tends to
In South Wales (and formerly north Devon) be retarded in wet years if surface water lingers.
Fen Orchid is found in dune slacks, growing The individual flowers are rather short-lived.
003 systematic.indd 143 29/1/09 12:25:12

144 l GENUS LIPARIS
003 systematic.indd 144 29/1/09 12:25:15

FEN ORCHID l 145
Non-flowering plants, which may be much

1987-99
1970-86
more numerous, are exceedingly hard to
pre 1970 spot. Visitors are asked to keep way from the
Norfolk sites but are welcome at Kenfig in
Glamorganshire. Detailed and up-to-date
directions are required to find Fen Orchid at
this site, and it is best to contact the visitor
centre in advance.
DESCRIPTION
petal lip
anther lateral
sepals
column
‘upper’ ovary
Range
sepal
Much reduced in Britain and now found at flower
only three sites in the Norfolk Broads and at stalk petal
three sites in South Wales, in Glamorganshire
and Carmarthenshire, but only in significant Height: 3-18cm, exceptionally to 30cm.
numbers at Kenfig. World range: Temperate Stem: Green and three-angled, becoming
Europe, Asia and North America. Found almost winged towards the tip. The base of
north to c. 61°N in southern Scandinavia, the stem is enlarged and this elliptical, green,
Finland and the Baltic States and south shiny swelling is enveloped by the bases of the
to northernmost Spain, southern France leaves to form a pseudobulb, the top of which is
(including Corsica), northern Italy, Bosnia and sometimes visible.
Romania (and formerly also Bulgaria). In Asia Leaves: Two or very occasionally three, strap-
the range extends eastwards to c. 80°E in central shaped, tapering both towards the base and
Siberia and also recorded in far eastern Siberia the pointed tip. They are held erect and nearly
and southeast Kazakhstan. In North America opposite each other. The leaves are pale green,
widespread in the northeast, from Nova shiny and prominently keeled. Immature,
Scotia and New England to Saskatchewan and non-flowering plants, with just one leaf, are
Minnesota, ranging south to North Carolina, common.
Illinois and Iowa. There are also other scattered Spike: Up to 12 flowers form a loose spike, but
populations west to the Northwest Territories, there are usually rather fewer, sometimes just
British Columbia and Washington, and south one, although as many as 17 have been recorded.
to Alabama. Bract: Green, minute, lanceolate.
Ovary: Green, narrow and tubular, six-ribbed,
How to find it straight or slightly twisted at the base; the
This small, green orchid is hard to find. It flower stalk is around half the length of the
often grows in fairly dense vegetation and even ovary, three-angled and twisted.
flowering plants can be very inconspicuous. Flower: The flowers are entirely greenish-
yellow and face upwards so that all the petals
O 26 June, Norfolk. Flower spikes (or their remains) are
visible from three season’s growth; all presumably arise
and sepals lie more-or-less horizontally and the
from the same psudobulb. ovary and column point upwards. The sepals
003 systematic.indd 145 29/1/09 12:25:16

146 l GENUS LIPARIS
are long and narrow (c. 5.5mm long x 1mm a horizontal plane. The column is pale green,
wide) with their edges rolled under, making relatively large and prominent, slightly curved
them appear even narrower; the lateral sepals towards the lip (like an erect cobra), with the
lie parallel below the lip and are often twisted. anther cap sitting on top and pointing forwards.
The petals are even finer (c. 4.5mm x 0.5mm) There is a minute rostellum and two waxy,
and are usually curved, both downwards in a yellow pollinia, each of which is divided into
gentle bow and forwards in the direction of the two flat plates and attached to one of the two
lip. The lip is tongue-shaped, rather broader viscidia.
than the sepals (c. 5mm x 2.5mm), with a wide
Subspecies
longitudinal groove that is deepest towards the
L. l. loeselii is found in Norfolk and has up to
base, and has slightly wavy or frilly margins. It
12 flowers and pointed, relatively narrow leaves
points upwards, parallel with the column, and
which are at least four times as long as broad. It
then bends sharply over at right angles to lie in
is widespread but scarce and declining in fens in
continental Europe.
L. l. ovata occurs in South Wales and formerly
also in north Devon. On average it is shorter
and has fewer flowers (usually up to six,
rarely as many as ten). The leaves are rather
more broadly elliptical or egg-shaped, blunter
and more hooded at the tip and held more
consistently erect. It also occurs in dune slacks
in northern France and northern Brittany and
in the early 1900s was present in the Dutch and
German Waddenzee.
There has occasionally been speculation that
the two subspecies should be treated as distinct
species, but recent genetic analysis has shown
virtually no difference between them.
None. (Subspecies ovata is sometimes treated as
var. ovata.)
BIOLOGY
The Fen Orchid is probably routinely self-
pollinating, the process being assisted by rain.
Raindrops hit the anther cap, which lies on top
of the column like a tiny ‘lid’. This in turn knocks
the pollinia towards the stigma. The upturned
lip may function to deflect raindrops towards
the anther. The prolific numbers of seedlings
recorded suggest that seed is the major means of
reproduction; in South Wales up to 128 shoots
have appeared in a 0.25m2 plot.
M 21 June, Norfolk.The wispy petals and upwardpointig
column and lip result in flowers that do not resemble most Vegetative reproduction also occurs but
other orchids and are hard to make sense of. its importance relative to seed is unknown.
003 systematic.indd 146 29/1/09 12:25:17

FEN ORCHID l 147
M 16 June, Norfolk.The flowers are self-pollinated and the lip is thought to deflect raindrops onto the column to knock-
out the pollinia.
‘Buds’ (detachable propagules) are formed on sheaths. Each summer, the rhizome continues
the swollen stem of the pseudobulb, and these its growth from a bud at the base of the existing
are dispersed in the autumn, already carrying a pseudobulb, and this new section swells in turn
fungal ‘infection’. as it stores nutrients. Normally, only the two
Development and growth youngest sections of the rhizome, representing
The aerial stem grows from a pseudobulb, the last two year’s growth, are alive at any one
the swollen tip of the rhizome which acts as time, and so there are two pseudobulbs lying
a storage organ and is surrounded by the leaf side-by-side.
003 systematic.indd 147 29/1/09 12:38:18

148 l GENUS LIPARIS
P 16 June, Norfolk. Flow

ers from above, showing
the threadlike petals,
which, together with the
sepals and lip, form a cross
shape.
Fen Orchid remains partially dependent on Hybrids

fungi as an adult plant, fungal ‘infection’ being None.
concentrated in the rhizome; the pseudobulb
is always fungus-free. A few short, thick, hairy
The specific name loeselii celebrates Johann
roots, largely free of fungi, develop in the spring
Loesel (1607-1657), a Prussian professor of
from the base of the pseudobulb.
medicine and a botanist.
Seed germinates in the autumn and the
protocorm produces the first green leaf by the
following August. The first small pseudobulb HISTORY aND
develops in the autumn of that year and the CONSERVaTION
protocorm starts to wither away. No roots are The first British record was published in
developed in this first year and all the plant’s 1660, when John Ray found it ‘in the watery
requirements for water must be met by its places of Hinton & Teversham Moors’ in
minute root hairs or by its fungal ‘partner’. In Cambridgeshire (Catalogus Plantarum circa
the summer of the second year both roots and Cantabrigiam nascentium; ‘Catalogue of plants
small leaves develop and a new pseudobulb is found around Cambridge’).
formed. Fen Orchid can flower in the fourth year A Red Data Book species, it is considered
after germination (perhaps as early as the second Endangered and included in Schedule 8 of the
year), and plants reach their maximum size from Wildlife & Countryside Act 1981. It has always
the seventh year onwards. However, there is a been scarce and localised in Britain but was
high mortality among immature plants, and the formerly known from 34-36 sites in eastern
vast majority disappear before they can flower. England, one in Devon and a further nine in
But, once mature, most flowering individuals South Wales. Fen Orchid is now reduced to
reappear the next season, and they may flower three sites in the Norfolk Broads where the
for several years in succession; the average overall population is stable: two in the Ant
lifespan of mature Fen Orchids is ten years. Valley, one holding a few hundred plants, the
Occasionally, plants may remain underground other about a thousand, and a much smaller
for one year, very rarely two, although it is hard colony in the Bure Valley, typically with about
to be sure in such cases that the leaves have not 10 plants but up to 40 can appear in a very
appeared and been rapidly grazed off. good year. There are also three sites in South
003 systematic.indd 148 29/1/09 12:25:21

FEN ORCHID l 149
O 16 June, Norfolk. The

flower in profile, show
ing the erect column and
horizontal lip.
Wales where the population totalled over (indeed, probably especially in ‘protected’
10,000 plants until recently but has declined to areas such as Wicken Fen) this resulted in the
an unknown extent. development of rank vegetation, scrub and in
In Norfolk, always the stronghold of the some cases the eventual transformation of the
species, in addition to the three surviving fen into wet carr woodland. Thus, although
localities, there were formerly eight more sites quite a few of the old sites still exist as ‘fens’,
in the Broads, with a further eight colonies changes in the vegetation have rendered
scattered elsewhere in the county, especially the them totally unsuitable for Fen Orchid.
Waveney-Ouse Valley on the Norfolk-Suffolk Collecting also took its toll, as demonstrated
border. In Suffolk there were four or five sites by this account of a club outing to Burwell
but the species was last recorded in 1974 at Fen, Cambridgeshire, in 1835: ‘We had very
Thelnetham. In Cambridgeshire there were good sport both in plants and insects. Ophrys
eight sites, six of which were drained long ago, loeselii was found in great plenty. Between four
but Chippenham Fen and Wicken Fen remain hundred and five hundred specimens were
intact, although Fen Orchid was last recorded brought home. It was growing in the grass and
at these sites in 1928 and 1945 respectively. moss among the pits where they cut turf. There
In Huntingdonshire there were two sites, the were two bulbs to each plant, and the bulbs
last record coming from Whittlesea Mere in were scarcely in the ground at all, so that we
1849, but this too was subsequently drained. picked them out easily with our fingers’ (quoted
Elsewhere in eastern England, Fen Orchid was in Marren 1999). As with most orchids,
recorded from Ham Fen in Kent in 1802 and however, such wanton destruction was a minor
was reported in 1884 near Lincoln. factor in its decline compared with changes in
Early losses in East Anglia were largely due its habitat.
to the drainage of the fens and their conversion In South Wales, Fen Orchid was first
to agriculture. Although drainage and water recorded in 1897 from dunes at Pembrey in
abstraction continued to cause extinctions, Carmarthenshire and was eventually found
most of the more recent losses have been caused at a further eight ‘Burrows’ (Kenfig, Margam,
by the decline of peat and turf-cutting, reed Baglan, Crymlyn, Oxwich, Whiteford, Tywn
and sedge harvesting and grazing, all traditional and Pendine/Laugharne). It was still present
land uses. Even in areas protected as reserves in seven dune systems in the early 1970s but
003 systematic.indd 149 29/1/09 12:25:23

150 l GENUS LIPARIS
by 2001 was reduced to just three: Kenfig, A variety of conservation measures are being
which still holds a substantial population employed, such as reed and sedge harvesting
and, although there has been some decline, in the Broads and scrub removal, grazing, close
concerted management efforts have stabilised mowing and turf stripping in South Wales.
numbers; Whiteford, where it is on the point These should provide the plants, both adults
of extinction, with just two plants in 2004; and seedlings, with the appropriate niche.
and Pendine/Laugherne, where 76 plants were Many of these techniques are only possible on
counted in 1997 but just one was found in a relatively small scale, however, and are interim
2000. The other sites have either been buried measures which will slow but not stop the
by sand, dried out, become overgrown or have inevitable succession to unsuitable habitats. The
been reclaimed for heavy industry. only real hope for the Fen Orchid’s long-term
The dune slack subspecies ovata was also survival is more radical action: digging peat
found at Braunton Burrows in north Devon in from the Norfolk Broads to create fresh turf
1966, when two flowering spikes were found in ponds; and the destabilisation of the dunes in
a slack that had been virtually bare of South Wales to create a succession of young
vegetation in 1953. Fen Orchid was last slacks. If the right conditions were available,
seen there in 1987. reintroduction also becomes an option. There
Fen Orchid has declined in South Wales have been small-scale experimental trials at
as the habitat has become unsuitable. Over- sites in Norfolk and Suffolk but so far no full
stabilisation of the dunes has meant that few programme has been attempted.
new slacks have been formed and a lack of
grazing, exacerbated by a decline in the rabbit Past and present occurrence of Fen Orchid in Britain and
Ireland (based on presence or absence in 10km squares
population due to myxomatosis, has hastened of the National Grid; data from the New Atlas).
the ageing process in the existing slacks. At
Britain Ireland
Kenfig there was a large amount of mobile sand
after World War Two and many new slacks total historical range, 26 0
were formed. It is this era of slack formation 1500-1999
which has provided the current supply of current range 7 (0.25%*) 0
suitable habitat: in 1946 40% of the area was % lost, 1500-1969 61.5%
bare sand but this had declined to 2% by 1992
% lost, 1970-1986 1.5%
and inevitably there will soon be a sharp decline
% lost, total 63%
in suitable habitat as the existing slacks mature
and are not replaced. * current range as a % of the total number of 10km squares
In both Norfolk and South Wales the
problems of Fen Orchid conservation are
similar and so are the solutions. Fen Orchid is
a ‘weedy’ species adapted to grow in dynamic,
changing environments. It is able to colonise
wet, calcareous habitats with plenty of bare
ground and multiply quickly but is eventually
crowded out as the vegetation matures. There is
a high rate of turnover of individual plants, with
considerable ups and downs in overall numbers.
Many plants are short-lived but large numbers
of seedlings can be produced, and to maintain
P 13 July, New Forest. Bog Orchids often grow from tiny
a population conditions need to be right for bulbils which develop on the edge of the leaf and can thus
seedlings to become established. form small groups.
003 systematic.indd 150 29/1/09 12:25:24

Genus
HAMMARBYA
BoG orchid
Distribution
Bog Orchid is the only member of this
genus, which has a circumpolar distribution
in Europe, Asia and North America.
Name
The generic name Hammarbya has an
auspicious history. In the mid-18th century
the great Swedish naturalist Linnaeus gave
Bog Orchid the name ‘Ophrys paludosa’.
As the relationships between the various
orchids came to be better understood, Bog
Orchid was moved from Ophrys to its own
genus by Otto Kuntze in 1891. He named
the new genus ‘Hammarbya’ in honour of
Hammarby, Linnaeus’s summer residence
near Stockholm.
003 systematic.indd 151 29/1/09 12:25:28

152 l GENUS hAMMARbyA
BoG orchid
Hammarbya paludosa
Formerly: Malaxis paludosa. Other names: Bog Adder’s-mouth (North America)

This tiny green orchid is the exact opposite of the classic big, bright and blousy hothouse
hybrid. Indeed, it is the smallest orchid in the British Isles and one of the hardest to find.
But, despite its rather dull flowers and diminutive stature (or perhaps because of these),
it holds a particular fascination for botanists and is always a delight to see. It is still fairly
common, at least locally, in northwest Scotland but has declined markedly in the lowlands
and in England is now only frequent in the New Forest.
Identification
The small size, tiny green flowers and habitat
make this orchid distinctive – when and if you
can find it.
Similar species
Of the plants found in its boggy habitat, Marsh
Arrowgrass is closest in general appearance but
has very different flowers.
Habitat
As its name suggests, this species is found
in bogs. It is associated with a good cover
of Sphagnum (bog-mosses) and grows
alongside sundews, butterworts, cottongrasses,
White Beak-sedge and Cross-leaved Heath.
Importantly, it requires a bog which has a flow
of water through the peat, and it does not like
stagnant conditions; as well as streams and
runnels, the slow flow of water may even be
evident on the surface of the peat. Bog Orchid
is often found in the vicinity of such moving
water and also close to the shores of lakes and
lochs. The ground water is usually moderately
acidic but in parts of Wales the species is also
recorded from areas flushed with alkaline water.
Another essential requirement is that the bog
must not dry out, even in a hot summer. Bog
Orchids often grow on carpets of Sphagnum
but can also be found on bare peaty mud or in
denser vegetation amidst sedges, grasses and
O 13 July, New Forest. The flower spike continues to

grow as the flowers open and becomes significantly taller.
This plant was 12cm high.
003 systematic.indd 152 29/1/09 12:25:29

BOG ORCHID l 153
small shrubs. Generally found in the lowlands

1987-99
but it has been recorded up to 500m above sea 1970-86
level (Llyn Anafon, Caernarvonshire). pre 1970
Flowering period
Late June to mid or even late September but
perhaps most reliably from early July to mid-
August. It tends to flower early in a hot summer
and later in a cool, wet season. Numbers are
erratic, with wide variations between years.
Flowering at any one site is not necessarily
synchronised, however, and the flowers
themselves are long-lasting, and thus larger
colonies may have at least some plants in bloom
over a lengthy period.
Range
Bog Orchid is widespread in Scotland. There known from relatively few, scattered localities;
are a few sites in the southwest, in Dumfries- in the west in Co. Clare and from Co. Mayo
shire and Kircudbrightshire, and in the eastern to Co. Donegal; in the east in Co. Carlow,
Highlands in Angus and south Aberdeenshire. Co. Kilkenny, Co. Wicklow and Co. Dublin,
The majority are, however, in the northwest and in Northern Ireland in Co. Armagh, Co.
where it is found north to Sutherland and Down and Co. Antrim. In Wales it is also
extends to the Inner and Outer Hebrides and recorded at just a few, scattered sites, mostly in
to Yell in Shetland. In Ireland, Bog Orchid is the west, from Breconshire, Carmarthenshire
M 29 June, New Forest. Although often hard to find, Bog Orchids can be conspicuous at times, contrasting as here with
a carpet of russet Sphagnum.
003 systematic.indd 153 29/1/09 12:25:30

003 systematic.indd 154 29/1/09 12:25:33

BOG ORCHID l 155
and Pembrokeshire north and west to Sphagnum are not favoured but the orchids can
Caernarvonshire, but often there are just one or be found around their edges, especially if they
two sites in each county. In England the species are close to water.
is now very local, occurring in Cumbria, at a Once a Bog Orchid has been seen, before
handful of sites in northwest Yorkshire and diving in for a closer look, it is wise to make
Northumberland and at one site in Norfolk. a very careful inspection of its immediate
In southern England it is known from a surroundings. This species often grows in
very few localities in Cornwall and Devon scattered groups, and one or two relatively
(including Dartmoor, although it is very rare obvious plants may be accompanied by several
there) but occurs much more commonly in the others hidden in the vegetation at one’s feet.
Purbeck region of Dorset and especially in the
New Forest in Hampshire. World range: In DESCRIPTION
Europe it occurs north to the Faeroe Islands
and to c. 69°N in Scandinavia, and south in lateral
the mountains to France, the Italian Alps, the sepals
former Yugoslavia and Romania. In Asia there lip
ovary
are scattered records from across southern lip column
Siberia to Sakhalin and Japan. In North petals
America it occurs from Alaska eastwards to
column flower
western Ontario and south to Minnesota. ‘upper’
stalk
How to find it ‘upper’ sepal
sepal
This tiny green plant is one of the hardest
British orchids to find. It is so inconspicuous bract
that it is easy to tread on it unawares, and the
pseudobulbs, often only half-buried in the Height: 2-15cm but usually 4-8cm.
moss, can easily be dislodged. All in all, it is best Stem: Yellowish-green and three to five angled.
not to look for it until you are an experienced The stem grows from a pea-sized pseudobulb
orchid hunter. which is covered by the leaves and often only
Faced with a large area of apparently half-buried in the moss.
suitable habitat, the task of searching for Bog Leaves: There are two or three (occasionally four)
Orchid may be daunting. It is best, however, to oval to oblong, fleshy, pale-green or yellowish-
concentrate on the most suitable areas. These green basal leaves. They are prominently veined,
are likely to be along the edges of streams sheathe the pseudobulb at their base, and their
and runnels or where there is some obvious margins and tips are strongly curved inwards
movement of surface water and away from giving them a hooded appearance. One to three
dense stands of shrubs such as Cross-leaved minute, triangular, scale-like leaves are scattered
Heath and Bog Myrtle. Like all our smaller higher along the stem.
orchids, Bog Orchid is difficult to see from Spike: Up to 25 flowers are carried in a spike
a walking height and becomes much more which is dense at first but elongates and
obvious when viewed from a low vantage point. becomes much more open as it matures.
On open, flat Sphagnum carpets or ‘lawns’ it Bract: Green, narrow and pointed, about as
can then be fairly conspicuous, but plants may long as the ovary.
grow among grasses and sedges and these are Ovary: Green, ovoid and just a little fatter than
extremely difficult to see; dense hummocks of the flower stalk, which is about three times its
length and twisted through 360° (therefore the
O 29 June, New Forest. Growing as usual on a carpet of
Sphagnum, minute bulbils are starting to appear on the
lip is held uppermost and the flower is hyper-
margins of the leaves. resupinate).
003 systematic.indd 155 29/1/09 12:25:34

BIOLOGY
The flowers are probably pollinated by gnats
and tiny flies, attracted to the nectar at the base
of the lip, and the pollinia are usually removed.
Seed-set is good. Perhaps more importantly
in terms of reproduction, the leaves are often
fringed with numerous, minute, protocorm-like
buds, called bulbils. These drop off and can
develop into new plants if they are ‘infected’
from the soil by fungi of the right species
(unlike the propagules of Fen Orchid, they do
not carry fungi from the mother plant). This
may account for the frequency with which Bog
Orchid is found in small groups.
M 7 July, Norfolk.The lip is uppermost (the flower stalk is Development and growth
twisted through 360°), and the lateral sepals wrap around The rhizome lies almost vertically in the peat
the back of the flower. or Sphagnum, and a swelling is formed each
Flower: Greenish and tiny, about 2mm wide year at its tip. In the spring leaves develop
by 4mm tall. The sepals are yellowish-green around the base of this swelling and the
and tongue-shaped; the dorsal sepal points flower spike grows from a bud on the top;
downwards and is slightly longer than the lateral the swollen stem and leaf sheaths combine
sepals, which point upwards. The petals are to form a pseudobulb 4-8mm in diameter.
rather smaller and narrower, strap-shaped and After it has flowered the aerial stem dies off,
green. They are held spreading horizontally but but the rhizome continues to grow from a
curve sharply back around the sepals to clasp bud at the base of the pseudobulb and at the
the flower. The lip is dark green with paler green end of the growing season again terminates
longitudinal stripes and is rather shorter than in a swollen internode. Although in theory
the petals, triangular and curled upwards at a whole string of pseudobulbs could be
the sides. It is held erect, pointing upwards and produced (as seen in many epiphytic orchids),
forwards between the two lateral sepals and in practice only the two most recent are alive;
clasping the column at its base. There is no spur, the older, lower, pseudobulb is buried in the
and the very short, broad, green column projects moss and surrounded by the remains of the
horizontally from the centre of the flower, with previous year’s leaves. Unusually, the rhizome
the lid-like anther shrivelling to expose the of Bog Orchid grows vertically with the two
pollinia shortly after the flower opens. The two pseudobulbs one above the other. This vertical
pollinia are each made up of two thin plates of growth pattern presumably allows the orchid to
waxy pollen, and the minute rostellum is topped adjust to the changing level of the bog’s surface
by a small, sticky mass. The flowers are said to due to, for example, the growth of Sphagnum.
have a sweet, cucumber-like scent. There are no roots, merely root hairs, and
Bog Orchid is largely dependent on fungi
Subspecies throughout its life cycle; the rhizome and leaf
None. bases are ‘infected’ at all times. The pseudobulb,
Variation and varieties which acts primarily as a storage organ, is
None. separated from the older parts of the rhizome
003 systematic.indd 156 29/1/09 12:25:34

BOG ORCHID l 157

The specific name paludosa means ‘growing in
boggy ground’.
Bog Orchid is sometimes placed in Malaxis,
a genus of rather similar orchids. It is also
closely related to Fen Orchid and could easily
be united with it in the genus Liparis, but the
current evidence is that it is sufficiently distinct
to justify its continued separation in its own
genus, Hammarbya.
HISTORY aND
CONSERVaTION
The first British records, from Hertfordshire
and Kent, were published in John Parkinson’s
Theatrum Botanicum (‘The Theater of Plantes’)
of 1640: ‘Bifolium palustre…. In the low wet
grounds between Hatfield and S. Albones; in
divers places of Romney Marsh’.
Nationally Scarce in Britain and specially
protected in Northern Ireland under the
1985 Wildlife Order (NI) and in Eire under
the Flora (Protection) Order. Bog Orchid
is threatened and declining throughout its
European range due to loss of habitat, and some
of the largest populations are now found in
western Scotland and the New Forest.
Past and present occurrence of Bog Orchid in Britain and

M 13 July, New Forest.The lip, which is much smaller than Ireland (based on presence or absence in 10km squares
the sepals, is boldly striped with pale and dark green. of the National Grid; data from the New Atlas).
Britain Ireland
by a band of lignified tissue. This effectively cuts
off one year’s growth from the next and prevents 1500-1999
fungi from reaching the pseudobulb and its current range 118 (4%*) 15 (1.5%*)
store of nutrients. An internal root is produced
at the base of the new segment of the rhizome, % lost, 1500-1969 47% 59%
and this grows down through the lignified % lost, 1970-1986 14% 7%
barrier into the older segment, apparently % lost, total 61% 66%
reabsorbing nutrients and water from the
decaying tissue and, bypassing the pseudobulb,
carrying the fungal ‘infection’ into the new
Systematic searches have produced records
segment.
from many new localities in recent years but,
Life expectancy and the period between
despite this, Bog Orchid has vanished from
germination and flowering are unknown.
61% of its historical range in Britain and
Hybrids 66% in Ireland. It is now extinct in most of
None. England, including Kent, East and West Sussex
003 systematic.indd 157 29/1/09 12:25:35

O 29 June, New Forest. As

in Fen Orchid, the aerial
stem grows from a pseudo
bulb, a swelling in the stem
concealed by the base of
the leaves; a second, non
flowering plant is growing
alongside.
(where it was formerly found in several areas, Many of the remaining colonies are small,
especially Ashdown Forest, but was last seen with just a few plants. Even in its strongholds
in 1956), Surrey, Hertfordshire, Bedfordshire, Bog Orchid is uncommon to rare; in Cumbria
Cambridgeshire, Suffolk, Lincolnshire, there are around ten sites, with just two
Staffordshire, Cheshire, Lancashire and Co. populations regularly exceeding 100 spikes.
Durham. The dramatic decline in lowland In the New Forest there are around 30
Britain started with the Enclosure Acts of populations, with over 200 spikes appearing
the late 18th and early 19th centuries and the in good seasons at two or three of the largest,
consequent reclamation of bogs and wet heaths. and in Dorset there are nine recorded sites.
Habitat destruction has continued to the present In Norfolk, which holds the only remaining
day, and in the lowlands the remaining areas of population in eastern England, there is now just
mire and heath have also suffered from a lack of one site, and in recent years there have
grazing, necessary to maintain the open sward been only one to four flowering plants in an
that the Bog Orchid requires. Conversely, in the area where 117 were counted in 1910; to add
uplands, overgrazing may have caused suitable insult to injury all the flowering plants were
habitats to be degraded. stolen in 2000.
003 systematic.indd 158 29/1/09 12:25:37

Distribution
A genus of 11 species, exclusively found in North
and Central America with the exception of
Coralroot Orchid C. trifida, which extends to the
boreal zone in Europe and Asia. All Corallorhiza are
largely or totally dependent on fungi as a source of
nutrition throughout their life.
Name
The generic name Corallorhiza derives from the
Greek korallion ‘coral’ and rhiza ‘root’, thus ‘coralroot’;
there are no roots, however, only a rhizome, and this
does not resemble coral in all species.
Genus CORALLORHIZA
coralroot orchids
003 systematic.indd 159 29/1/09 12:25:40

160 l GENUS CORALLORhIZA
coralroot orchid Vulnerable

Corallorhiza trifida
Other names: Early Coralroot (North America)

This tiny orchid lacks green leaves and throughout its life is largely parasitic on its fungal
‘partner’, living entirely underground apart from the brief period during which the flower
spikes are produced. It is found throughout the Northern Hemisphere, and its generally
northerly distribution is reflected in Britain where it is confined to Scotland and northern
England. Both the English name and the generic name Corallorhiza refer to the shape of the
underground rhizome, which is said to be coral-like. Coralroot Orchid is the county flower
of Fife.
Identification
With no leaves (merely scale-like sheaths on
the stem) and tiny greenish-white flowers, this
species is very distinctive.
Similar species
None. Bird’s-nest and Ghost Orchids also lack
green leaves but are very different. Bird’s-nest
Orchid is usually taller and always more robust,
with large, honey-coloured flowers, and Ghost
Orchid has a proportionally much larger flower
with the lip uppermost and is only found, very,
very rarely, in southern England.
Habitat
Coralroot Orchid is found on permanently
damp ground with a good layer of peaty organic
matter or moss, including Sphagnum. It can
grow both in full sunlight and in the shade,
even heavy shade, and favours mildly acidic soils
which are low in nutrients. It is commonest in
wet willow and alder carr growing on raised
bogs and around lochs. Its other favoured
habitat is a damp dune slack with a carpet
of Creeping Willow but Coralroot Orchid
appears to have rather exacting requirements
regarding the level of the water table and does
not like prolonged flooding (for example, on
P 7 June, Northumberland. Among Creeping Willows in

a dune slack; Coralroot Orchid extracts nutrients from the
willow, via its fungal ‘partner’.
O 7 June, Northumberland. In dune slacks Coralroot
Orchid usually has a stout, mahoganybrown stem. Like all
orchids growing in dunes, it often has a ‘scorched’ look, in
this case with the tips of the sepals blackened.
003 systematic.indd 160 29/1/09 12:25:41

CORALROOT ORCHID l 161
Lindisfarne in Northumberland a slack that

regularly held 150 flower spikes now supports
only 20 after several wet years). Rather less
frequently, Coralroot Orchid is found in birch
and pine woodland, including plantations, and
in overgrown scrubby fens with a mixture of
Sphagnum, sedges and willows. It is found up to
365m above sea level (Braemar, Aberdeenshire).
Coralroot Orchid is parasitic on a group
of fungi that form mycorrhizal relationships
with birches and willows (also pines in North
America and presumably also in Scotland). The
presence of both the correct fungal ‘partner’ and
one of these trees is essential and may explain
the very local occurrence of Coralroot Orchid.
Flowering period
May to August. In dune slacks it flowers in
May and early June, increasingly in recent
years from early May onwards, and emerging
earlier in drier slacks compared to wetter ones.
Woodland plants are mostly in bloom from
early June to late July. M 6 June, Cumbria.
Range southernmost records come from Ribblesdale

Widespread in eastern and central Scotland, in Yorkshire. World range: This species has
from the Borders north to Caithness. However, a circumpolar distribution, occurring very
it is absent from the west, apart from Ayrshire widely in northern Europe, Asia and North
and Renfrewshire. In England it is very America, ranging southwards locally where
local, occurring in Northumberland and in mountains provide a suitable habitat. It occurs
Cumbria, where it is found in dune slacks at north to Greenland, Iceland and northernmost
Eskmeals, North Walney and Sandscale. The Scandinavia and south to the Pyrenees, Massif
Central, Alps, Apennines and Balkans, also
1987-99 Corsica, the Crimea and Caucasus. In Asia,
1970-86
pre 1970 primarily boreal but with isolated outposts in
the mountains of Central Asia and China. In
North America occurs from Newfoundland
east to Alaska and south in the mountains to
California, New Mexico and West Virginia.
How to find it
Small and leafless, this is an inconspicuous
orchid and very hard to spot. The number of
flowering spikes can also vary greatly from
season to season. As with all such diminutive
orchids, scanning from a low level can be
productive. Dune slack populations are the
easiest to localise, with one of the biggest at
Sandscale in Cumbria.
003 systematic.indd 161 29/1/09 12:25:42

DESCRIPTION
Height: 5-30cm, usually 10-13cm.
Stem: Usually yellowish-green in woodland
plants but tends to be mahogany-purple in
dune populations. Frequently found in small
groups and up to ten spikes may develop from
one rhizome.
Leaves: There are no green leaves, merely two
to four long, membranous, sheathing scales on
the lower half of the stem, and these may be
brown, whitish or green.
Spike: There are four to nine flowers
(occasionally as many as 13) held pointing
outwards and slightly drooping in an open,
lax spike.
Bract: Green, minute, triangular and pointed.
Ovary: Green or mahogany-purple, spindle-
shaped, six-ribbed, on an extremely short,
twisted stalk.
Flower: The sepals and petals are strap-shaped,
with the petals slightly smaller; both are
greenish-yellow, often tinged reddish-brown
around the fringes and tip and thus apparently
‘browned off ’. The interior of the petals may
also be blotched with reddish-brown. The
lateral sepals curve inwards and are held
forward and slightly drooping on either side
of the lip. The upper sepal and petals form a
loose hood. The lip is tongue-shaped, shorter
and broader than the petals and sepals, with
a ruffled margin. A central groove, which
may produce nectar, runs between two raised
longitudinal ridges and there is a very short
spur. The lip is kinked downwards towards
the base and, although technically ‘three-lobed’,
the tooth-like side-lobes near the base are very
small. It is white, spotted with crimson at the
base, and these spots may occasionally coalesce
into a larger blotch. The column is long, green
and curved, the lid-like anther lies on the top
and there are four waxy yellow pollinia, a small
rostellum and two distinct viscidia. The flowers
are slightly scented, with the perfume reported
to b e ‘musk-like’.
M In woodland Coralroot Orchid has a green stem and
ovaries and is relatively slender (Paul Sterry/Nature Subspecies
Photographers Ltd). None.
003 systematic.indd 162 29/1/09 12:25:44

CORALROOT ORCHID l 163

Plants growing in dune slacks tend to be
shorter than those in woodland, with reddish
rather than greenish stem and ovaries, but there
are no named varieties.
BIOLOGY
The flowers are routinely self-pollinated. The
rostellum is small and degenerates quickly,
and the pollinia crumble apart, falling onto
the stigma below; 85-100% of flowers set
seed. Small insects, including flies, wasps
and beetles, visit the flowers, but the pollinia,
although easily detached, do not readily stick
to the insects and any cross-pollination is
purely accidental; visiting insects are probably
more effective in nudging fragments of the
disintegrating pollinia onto the stigma below.
Vegetative reproduction may also occur,
via fragmentation of the rhizome, the side-
branches elongating and producing new plants.
Development and growth M The tiny flowers are usually selfpollinated and most
flowers set seed (Bob Gibbons/Natural Image).
The aerial stems grow from a creeping,
horizontal, underground rhizome. This is a fungus-tree partnership, receiving nutrients but
much-branched mass of cream-coloured, fleshy, giving nothing in return. It is therefore parasitic.
coral-like knobs. There are no roots and water (Coralroot Orchid is not a saprophyte; it gets
must be absorbed either through tufts of root its nutrients from a living fungus, not dead
hairs or via the fungal ‘partner’. organic matter.)
The rhizome is permanently ‘infected’ Coralroot Orchid does have a limited ability
with fungi and, throughout its life, Coralroot to photosynthesise as the stem, ovary and scale-
Orchid is almost completely dependent on like leaves on the stem contain chlorophyll.
its fungal associate for nutrients. Like Bird’s- Even in diffuse daylight such photosynthesis
nest Orchid, it is very fussy and only forms a can contribute in a small way to its overall
relationship with the Thelephora-Tomentella nutritional budget.
complex of fungi (family Thelephoraceae). This Seed germinates from the spring onwards
group of fungi, apart from its relationship with but may remain dormant until the spring of the
orchids, is exclusively ectomycorrhizal, forming second year, perhaps even longer. Germination
symbiotic relationships with the roots of trees will, however, only take place when the seeds
(see p.8). The fungi that Coralroot Orchid have been colonised by the appropriate fungi.
‘partners’ are species that simultaneously attach The seedling is initially a globular protocorm
themselves to the roots of willows, birches and which develops scattered root hairs. It then
pines. It has been shown in the laboratory that elongates and starts to branch; each time the
the orchid obtains carbohydrates from the trees rhizome branches, the main rhizome bends
via their mutual fungal ‘partner’, but the orchid in one direction and the side branch goes off
undoubtedly ‘cheats’ in its relationship with the in another, producing the ‘coralloid’ growth
003 systematic.indd 163 29/1/09 12:25:44

P 6 June, Cumbria. The

lip has a central groove
that may produce nectar;
this would attract insects
but despite this the flowers
are usually selfpollinated.
pattern. After as little as nine months, some (especially for the New Atlas scheme), and these
seedlings have developed into a branched have turned up many new sites. But, despite
rhizome 15-25mm long with the bud for the these new records, there has been a net loss
aerial shoot already well-developed. Flower of 46% of the total historical distribution and
spikes may be produced two to five years after many of the losses are relatively recent. Many of
germination but there is a little uncertainty its habitats seem relatively secure, however, and
about the exact timing. Some related species importantly many are not affected by changes in
of Corallorhiza are monocarpic and die after agricultural practices. It is possible that climate
flowering once. change could be responsible as many ‘northern’
plants are currently in retreat.
Hybrids
Woodland populations are usually small,
None.
scattered and hard to find but in some dune
Name and classification slacks there can be large numbers; at Sandscale
The specific name trifida comes from the Latin in Cumbria at least 3,000 plants were counted
for ‘split into three’, a reference to the shape of in five slacks in 1991, making it the largest
the lip. This is barely appropriate, as the lip is English population.
only very slightly three-lobed.
Past and present occurrence of Coralroot Orchid in Brit
HISTORY aND squares of the National Grid; data from the New Atlas).
CONSERVaTION
The first British record was published in Britain Ireland
1777 by the Rev. John Lightfoot, curate of total historical range, 102 0
Uxbridge. In Flora Scotica he noted: ‘Ophrys 1500-1999
Corallorhiza… In a moist hanging wood near current range 55 (1.9%*) 0
the head of Little Loch Broom on the western % lost, 1500-1969 17%
coast of Ross-shire’.
% lost, 1970-1986 29%
Nationally Scarce and classified as
% lost, total 46%
Vulnerable. This small, inconspicuous orchid
is hard to find, and the true picture of its * current range as a % of the total number of 10km squares
status and distribution is still emerging. Efforts

have been made to seek it out in recent years
003 systematic.indd 164 29/1/09 12:25:46

Genus GOODYERA
lady’s-tresses (i)
Distribution
A genus of 40-100 species, nearly worldwide in
distribution although most are found in South-East
Asia. Some are called ‘jewel orchids’ and are known
for their beautiful foliage. Only two species occur
in Europe, the widespread Creeping Lady’s-tresses
and G. macrophylla, endemic to Madeira.
Name
The genus Goodyera is named in honour of John
Goodyer (1592-1664), a manorial steward at
Petersfield, Hampshire, who has been dubbed the
‘first amateur naturalist’ and ‘the Cavalier botanist’.
003 systematic.indd 165 29/1/09 12:25:49

166 l GENUS GOODyERA
creepinG lady’s-tresses
Goodyera repens
Other names: Lesser Rattlesnake Orchid, Lesser Rattlesnake Plantain (North America)
This small, white-flowered orchid is almost always found growing under pine trees and is
locally common in Scotland, the home of the only native pine woodland in the British Isles.
A few colonies in old pine plantations in Norfolk represent an enigma. The lady’s-tresses
could have been introduced accidentally with the pine seedlings, could be indigenous to the
area and have simply moved into a congenial new home, or could have arisen from wind-
blown seed.
Identification
The spikes of small, densely hairy, white flowers
are distinctive. Creeping Lady’s-tresses is
evergreen and can be found and identified all
year. Indeed, it is often easier to locate in winter
when much of the other vegetation has died
down. It forms small patches of rosettes which
are composed of small, oval, dark-green leaves
rather like Garden Privet in size, shape and
colour. Notably, the veins on the leaves form
a faint net over the surface; almost all other
British orchids have veins that are parallel.
Similar species
The other species of lady’s-tresses belong to
the genus Spiranthes but are nevertheless rather
similar. They have small white flowers which
have a covering of glandular hairs, although
none of them are as densely hairy as Creeping
Lady’s-tresses. Their flower structure is similar
but the Spiranthes have the tip of the lip broadly
frilled or crimped, rather than being a simple
unadorned wedge shape.
Autumn Lady’s-tresses is very unlikely
to be found in the same pinewood habitat as
Creeping Lady’s-tresses (although it did occur
until 1979 in a grassy woodland ride within
a few metres of Creeping Lady’s-tresses at a
remarkable site at Holt in Norfolk). Once the
possibility of confusion is acknowledged, the
two species can be separated easily because
P 11 July, Norfolk. Typically grows under pines, some
times mixed with birch or, as here in Norfolk, oak.
O 10 July, Norfolk. Creeping Lady’stresses.
003 systematic.indd 166 29/1/09 12:25:51

CREEPING LADY’S-TRESSES l 167
Autumn Lady’s-tresses has its flowers arranged

1987-99
into a distinct row that is often spirally twisted. 1970-86
Irish and Summer Lady’s-tresses grow from pre 1970
a rosette of long, strap-shaped leaves which

have parallel veins, and are rather taller than
Creeping Lady’s-tresses; their flowers are also
arranged into a more definite pattern.
Habitat
Creeping Lady’s-tresses is found in mature
pinewoods with a damp, well-shaded forest
floor, growing in the deep humus formed by
the accumulation of dead pine needles. It
can be found in areas with a relatively open
understorey of grasses and small shrubs, such
as Heather and Bilberry, or in places where
there is merely a carpet of moss over the needles
(sometimes even bog mosses, Sphagnum), sufficiently mature to have a shaded, moist
indicating a suitably moist microclimate. The open, forest floor. A balance of light conditions
classic habitat is ancient ‘Caledonian’ woodland is required; too little light and the orchid
where Scots Pine is mixed with birches. It will not flower, too much sun and a dense,
has also spread into pine plantations that are overwhelming shrub layer develops. Rarely,
M 10 July, Norfolk. Growing in an old pine plantation on sand dunes.
003 systematic.indd 167 29/1/09 12:25:52

Creeping Lady’s-tresses is found on damp Holkham, although the colonies there are
dunes or among Heather and Bell Heather scattered and hard to find.
on moorland, sometimes far from woodland
(perhaps most often in coastal areas; for DESCRIPTION
example on a ‘peat moor’ at Auckengill Loch Height: 7-20cm, occasionally to 35cm.
in Caithness). Unlike most British orchids, Stem: Pale green and ridged, with dense
Creeping Lady’s-tresses is found on acid as well glandular hairs towards the tip.
as neutral soils. Most sites are in the lowlands Leaves: The flower spikes grow from a rosette
but it is found up to 335m above sea level of three to nine dark-green (almost blue-green)
(Morinsh, Banffshire). leaves which have a network of faint paler veins
(reticulations). The leaves are oval and taper
Flowering period
to a pointed tip and the leaf-stalk is short but
Late June to late August but often at its best in
broad (winged). They are held more-or-less flat
mid-July. In dry weather the individual flowers
to the ground, although there may be one or
are short-lived.
Range
Northern and eastern Scotland, with an
isolated population in Ayrshire, also northern
England in Cumbria, Northumberland and
Co. Durham, with an outpost in Norfolk (see
History and Conservation). World range:
This species has a circumpolar distribution
and occurs throughout Europe, northern Asia
and northern North America. Found north
to 70°N in Scandinavia and south to the
Pyrenees, Corsica, Alps, northern Apennines
in Italy, northern Greece, the Crimea and the
Caucasus, although confined to the mountains
in the south of the range. In Asia it ranges
across Siberia to the Kamchatka Peninsula,
Sakhalin and Japan and is also found in the
mountains of southern Asia in Turkey, Central
Asia, Afghanistan, the Himalayas and in a band
from southwest China to Korea. In northern
North America it occurs from Alaska east
to Newfoundland and south in the Rocky
Mountains to New Mexico and Arizona and in
the Appalachians to North Carolina.
How to find it
Often abundant where it is found, any site in
the ‘Caledonian’ pinewoods of Scotland could
yield this species. In Norfolk the well-known
population at Holt Country Park was largely
destroyed by forestry operations in 2002
(but may recover in time), and the species is M 14 July, Norfolk. The stem, bracts and flowers have
best looked for in the coastal pines at Wells- abundant glandular hairs.
003 systematic.indd 168 29/1/09 12:25:53

two leaves at the base of the flower spike, and BIOLOGY

several very small, long and narrow sheathing
leaves further up the stem. Sterile, non-
The flowers are thought to be pollinated by bees
flowering rosettes are also produced and the
which are attracted to the nectar at the base
leaves are evergreen.
of the lip. Bumblebees of the genus Bombus
Spike: The 5-25 flowers are arranged spirally
are usually mentioned as pollinators and have
around the stem but mostly face in the same
certainly been seen carrying pollinia, but
direction, and the spiral pattern is seldom
smaller bees of the genus Lasioglossum may be
obvious.
more important. The mechanism of pollination
Bract: Green, with scattered glandular hairs
is thought to be very similar to that of Autumn
towards the base. The bracts are lanceolate and
Lady’s-tresses and other orchids in the genus
slightly longer than the ovary, which they clasp.
Spiranthes. The flower tube is initially only wide
Ovary: Green, narrowly pear-shaped, tapering
enough to allow an insect’s proboscis to enter;
towards the base, three-ribbed and with
this trips the mechanism and removes the
glandular hairs. The ovary is held upright but
pollinia. The lip then moves slowly downwards,
bends at the tip, so that the flowers lie either
allowing access to the stigma, which becomes
horizontally or are slightly drooped, and is
sticky and receptive. Fragments of pollinia,
either stalkless and slightly twisted or has a
picked up from other flowers and attached
short stalk, in which case it is the stalk that is
to a visiting insect’s proboscis, can then effect
slightly twisted.
pollination. The mechanism is effective and
Flower: Small, creamy-white and densely hairy.
seed-set is good, with 77% of flowers setting
The sepals are oval and concave, with the upper
seed in one Scottish study.
sepal a little narrower than the lateral sepals;
Vegetative reproduction may be more
they may have a faintly greener midrib and
important than reproduction from seed. Buds
have many conspicuous long, glandular hairs
are produced on the tip of the rhizome in the
on their outer surface. The petals are similar
autumn, and these grow into slender runners
but slightly smaller and more spatulate. The
which grow horizontally through the cushion
upper sepal and petals form a tight hood and
of moss. At first these runners have merely
the lateral sepals are held drooped and slightly
a few short, sheathing scales, but eventually
spreading with their tips bent outwards. The
green leaves are produced from buds at or near
lip is white and is shorter than the sepals, oval
their tip (a period of five years was given by
in shape with a pointed tip; the basal hypochile
Summerhayes (1968) for the production of
is sometimes tinged pink and forms a deep,
green leaves but this seems excessively long).
rounded, bag-shaped pouch that contains
Once it has appeared above ground, the new
nectar. The epichile forms a narrow, blunt-
rosette will live for between two and eight years
tipped triangle, folded into a shallow groove and
before flowering and dying (and in Norfolk a
bent downwards towards the tip. The column is
period of at least six years has been recorded
creamy and the rostellum is formed into short,
between the first appearance of rosettes and
curved horns which enclose the single, roughly
the production of flowers). Each runner can
circular viscidium. The anther cap is ochre with
produce a separate plant; after flowering the
a reddish-brown margin and the two pollinia
central ‘mother’ plant dies off, leaving the
are yellowish. The flowers have a sweet scent.
surrounding rooted runners as separate entities.
Subspecies In this way large patches can form.
None.
Variation and varieties The aerial stem grows from a creeping rhizome
None. which is often only shallowly buried and which
003 systematic.indd 169 29/1/09 12:25:54

ground and assume an existence totally

dependent on fungi.
Seeds probably germinate in spring, but
the period between germination and the first
appearance above ground is not known.
Hybrids
None.
The specific name repens derives from the
Latin and means ‘creeping’. The American name
‘Lesser Rattlesnake Plantain’ probably derives
from it use, including a poultice of the chewed
leaves, in the treatment of snakebites.
HISTORY aND
CONSERVaTION
The first published British record dates from
1777, when Rev. John Lightfoot (1735-1785),
curate of Uxbridge, noted in his Flora Scotica:
‘We found it … in an old shady hanging birch
wood … about two miles from the head of
Little Loch Broom … Ross-shire’. (The species
had, however, been found a few years before by
a Scotsman, James Robertson (c. 1745-1796),
‘in a wood called Cregenon’, but this was not
published at the time.)
Nationally Scarce and the subject of a
Plantlife International ‘Back from the Brink’
project. It is common to locally abundant
in northeast and north-central Scotland,
especially in the relict ‘Caledonian’ pinewoods
M 10 July, Norfolk. The flower structure is very similar to of the Strathspey and Cairngorm regions,
the Spiranthes lady’stresses, but the lip forms a simple
spoutshape rather than being crimped or frilled. but it is scarcer in the west of Scotland and
northern England. Colonial, it may occur in
puts out a few short, thick, fleshy roots that large numbers, and we have counted 620 flower
have numerous hairs. Both rhizome and roots spikes in an area of only about 50m x 20m.
are heavily ‘infected’ with fungi, and the species Although still locally common, there has
is probably dependent on its fungal ‘partner’ to a been a significant reduction in the range, and
significant extent throughout its life. it has now gone from 44% of its historical
Creeping Lady’s-tresses is the only British distribution. It is extinct in Dumfries-shire,
orchid which is evergreen (some other species, West Lothian, Peebles-shire and Orkney
such as Autumn Lady’s-tresses and Early (where recorded in the 1950s). There are
Spider Orchid, bear leaves through the winter also old records from southeast Yorkshire
but are leafless and ‘dormant’ for at least part (1888, Houghton Hall Woods, near Market
of the year). Unlike most other orchids, this Weighton) and east Suffolk (1932-1935,
species shows no tendency to disappear below Stuston Common, a site with no pine trees).
003 systematic.indd 170 29/1/09 12:25:55

Past and present occurrence of Creeping Lady’stresses nurseries. The trees were, however, sometimes
in Britain and Ireland (based on presence or absence in
10km squares of the National Grid; data from the New packed in dry Bracken, Heather and moss
Atlas). ‘collected from the woods’ and this packing
could have been a source of seeds or plants.
Britain Ireland
An alternative theory is that Creeping
total historical range, 186 0 Lady’s-tresses was present in Norfolk before
1500-1999
the advent of pine plantations and merely
current range 104 (3.6%*) 0 moved in when they became suitable. There
% lost, 1500-1969 29.5% is some support for this, in that it was found
% lost, 1970-1986 14.5%
at Sheringham in 1909, concealed among
Heather on open heathland, and similarly it
% lost, total 44%
was found on heathland at Beeston Regis in
* current range as a % of the total number of 10km squares 1900. Alternatively, the Norfolk populations
may originate from wind-blown seed, either
With its requirements for a certain from Scotland or the Continent. The timing
minimum level of shade in order to thrive, of records is interesting, as Creeping Lady’s-
Creeping Lady’s-tresses is vulnerable to tresses were recorded for the first time from
woodland management. Colonies can survive some of the plantations many years after they
even if a wood is felled and replanted, but it were established; pines were first planted at
takes many years for the number of flowering Wells and Holkham in the 1850s, largely with
plants to recover in these circumstances and seedlings produced in a nursery at Holkham
the woods must be replanted with pines; itself, but the lady’s-tresses were not found
many sites have been lost when restocked until 1952.
with alien conifers such as spruce and fir. The At some sites there have been documented
extensive thinning of plantations can also have introductions by well-meaning naturalists,
a very adverse effect, although perhaps not a as at Horsford in Norfolk in 1955-58
permanent one. (although there were probably already ‘natural’
The status of the species in Norfolk has populations there) and Warwick Moor Wood
been the subject of controversy since its in Cumbria in 1931 (where the plantation
discovery in 1885 at Westwick. Subsequent was clear-felled in 1987). Other Cumbrian
records came from Holt and adjacent populations may well have been accidentally
localities from 1890, Beeston Regis from introduced with conifers when plantations
1900, Sheringham from 1909, Cawston from were established.
1910, Wells from 1952, Horsford from 1958, Whatever their origin, the populations
Holkham from 1962 and Cranwich from in Norfolk are extremely interesting from a
1965. scientific point of view, yet the controversy
Scots Pine is not native to Norfolk, but over their status has compromised their
there are plantations of conifers dating from conservation. After showing keen and active
at least the 1830s. It has been widely assumed concern in the 1950s and 1960s, the attitude
that Creeping Lady’s-tresses was accidentally of the Nature Conservancy and Nature
introduced with pine seedlings brought in from Conservancy Council (predecessors of English
Scotland, but there is little evidence to support Nature) to the largest and best-known
this. Investigations by W.H. Burrell in 1909 population at Holt changed, and in 1986 its
established that pines supplied by a selection status as part of an SSSI was removed. As the
of nurseries in Scotland came bare-rooted New Atlas maps all the British populations as
and thus free from ‘weeds’ and that Creeping ‘native’, attitudes may now change.
Lady’s-tresses was unknown as a ‘weed’ in these
003 systematic.indd 171 29/1/09 12:25:55

GENUS SPIRANTHES
LADY’S-TRESSES (II)
003 systematic.indd 172 12/2/09 16:56:48

GENUS SPIRANThES l 173
These small orchids have distinctive small, tubular white flowers and swollen, tuber-like
roots. Sadly, one member of the genus, Summer Lady’s-tresses, is the only orchid that is
extinct in Britain and Ireland.
Distribution period’ as a single tuber and the rhizome

The genus contains 45-300 species (the taxon- does not die off. New, swollen roots develop
omy is very uncertain), found in Europe, Asia, each year and these push the roots from the
Australia and South America, with the majority preceding season, which are shrunken and
in North and Central America. Four species depleted, out towards a horizontal position;
occur in Europe, with three in the British Isles. this may give the impression that there are
two types of roots: vertical, fleshy roots and
Floral structures
horizontal, fibrous roots.
The column is held horizontally within the
The aerial stem dies off once seed has been
tubular flower with the circular stigma on its
set, but a bud develops on the side of the base
underside, facing downwards. The rostellum
of the stem and this will go on to form the new
is well-developed and deeply forked with the
leaves and aerial stem.
single torpedo-shaped viscidium held between
the two prongs of the fork. There are two Vegetative reproduction
pollinia, each composed of two leaf-like plates, Additional lateral buds may develop in the late
and both are attached to the viscidium. Once summer or autumn at the base of the aerial
shed by the anther, the pollinia lie on top of stem. These can produce their own roots and
the column with the viscidium below their tips, leaves and may eventually become separate
waiting to be removed by a visiting insect. ‘daughter’ plants.
Growth pattern Name
The aerial stem grows from a very short The generic name Spiranthes means ‘twisted
rhizome which puts out two to six fleshy, flowers’ and derives from the Greek speira
tuberous roots. Unlike the tubers of Orchis meaning ‘twisted’ or ‘coiled’ and anthos meaning
and Dactylorhiza, the roots are heavily ‘infected’ ‘flower’. The English name probably refers to the
with fungi. And, again unlike Orchis and flower spikes of Autumn Lady’s-tresses, which
Dactylorhiza, the plant does not spend a ‘resting look like plaited hair.
remains
of
flower
new spike
tuberous rosette new
root dies off rosette
January March July August December

M Growth pattern of Autumn Lady’stresses (after Wells 1981).
O 18 August, Norfolk. Autumn Lady’stresses.
003 systematic.indd 173 29/1/09 12:25:58

174 l GENUS SPIRANThES
irish lady’s-tresses BaP

Spiranthes romanzoffiana
Other names: Hooded Ladies’-tresses (North America)

This delicate, white-flowered, late summer orchid has one of the most unusual distributions
of any British plant, occurring in North America, Ireland, western Scotland and southwest
England but nowhere else in Europe. It is also one of the hardest orchids to find, partly due
to the remote location of the colonies and partly to the unpredictable nature of its flowering.
The biology of this species is a mystery. It is fairly widespread, with many recorded sites,
but very rarely sets seed in Britain and Ireland and has no obvious method of dispersal or of
maintaining its numbers.
Identification
Distinctive, with long, grass-like leaves and a
compact spike of small white flowers, usually
arranged into three spirally twisted rows. The
upper stem, ovaries, bracts and sepals have
numerous glandular hairs. These features,
combined with its usual damp grassy habitat,
restricted range and late flowering, are unique.
Similar species
Creeping Lady’s-tresses is very occasionally
found on moorland and also has glandular
hairs on the flowers, but it is usually smaller,
with a different flower structure and very
different leaves.
Autumn Lady’s-tresses also flowers in August
and September but is much commoner, has a
predominantly southerly distribution and is
almost always found on short, dry, calcareous turf.
Its flowers are arranged into an obvious single row.
Summer Lady’s-tresses is now extinct in
Britain. Like Irish Lady’s-tresses, it has long,
lanceolate leaves on the stem and is found
in wet, acid grassland but the flowers, as
well as being arranged into a single row, are
significantly smaller; 6-8mm long rather than
10-14mm, with the bracts 6-9mm long rather
than 10-20mm, sometimes even 30mm.
Other white-flowered orchids, such as Small
White Orchid and some Heath and Common
Spotted Orchids, may be found in the same
habitats and in the same geographical area
O 7 August, Colonsay, Inner Hebrides (Richard Gulliver).
003 systematic.indd 174 29/1/09 12:25:59

IRISH LADY’S-TRESSES l 175
but flower earlier in the season. They all lack Flowering period
glandular hairs on the upper stem and flowers Early July to early September, depending on
and can also be excluded by looking at the weather conditions and water levels, but mostly
details of the flower structure. in late July and early August.
Habitat Range
Frequently found in open grassy areas where In Ireland, it is very local in Eire but there are
the soil is low in nutrients and permanently some good populations in the lakes area of
damp or wet, either because it is flushed with western Co. Galway and Co. Mayo, especially
ground water or, more usually, because it is around Loughs Conn, Cullin, Corrib and Mask.
close to a river, stream or lake and is flooded Otherwise, it occurs at a few sites in the far
from time to time. Often grows close to the southwest, in Co. Kerry and Co. Cork, and in
shore of a lough and has even been recorded the northwest, in Co. Donegal. In Northern
flowering with the leaves submerged. At most Ireland the main concentration of populations
sites the water and soil are mildly acidic to lies in Co. Antrim, around Lough Neagh and
neutral but Irish Lady’s-tresses is sometimes the streams and rivers that flow into it. It is
recorded from more alkaline, base-rich flushes. also present along the River Bann through
Suitable conditions are provided by damp Lough Beg and as far north as Coleraine in
meadows and rushy pastures, flushed grassy Co. Derry and around the southern shores of
slopes, wet heathland, and bogs, where it may Lough Neagh in Co. Armagh. There are also
grow amongst Sphagnum on the disturbed some scattered sites elsewhere in Co. Antrim,
ground around old peat workings. In the including Gortnagory on the Garron Plateau,
Hebrides it may particularly favour the band and it has recently been found at Aird on the
of marginal land (known locally as ‘blackland’) north coast near the Giant’s Causeway. There
that is found at the transition between the lime- are also a very few sites in Co. Fermanagh, Co.
rich machair and the acid moorland inland. Down and probably also still Co. Tyrone.
And, although often growing in damp habitats, In Scotland the Irish Lady’s-tresses is
it has also been found on heather moorland. widespread but scattered on Coll and Colonsay
‘Old lazy beds’ is often given as a prime habitat in the Inner Hebrides and has been recorded
for Irish Lady’s-tresses (lazy beds are used to from around 30 sites on each island. Other
grow potatoes). Recent studies on Coll and concentrations are found in the southern
Colonsay have, however, comparatively rarely
found the species in this habitat.
1987-99
Most of the sites in Britain and many in 1970-86
pre 1970
Ireland are subject to extensive grazing, which
helps to keep the sward comparatively short
and reduce competition from other vegetation.
Disturbance by grazing animals may also
stimulate dormant plants, and it can favour
areas where cattle are fed in the winter or where
the ground has been broken up by ditching or
fencing. Generally found at low altitudes but
recorded up to 240m above sea level in Ireland.
In North America the Irish Lady’s-tresses is
found in similarly damp habitats, such as bogs,
marshes, wet meadows and stream-sides, but in
California it is also found in coastal grasslands
and pine forests that are baked dry in summer.
003 systematic.indd 175 29/1/09 12:25:59

California, New Mexico and Arizona in the

west and Illinois and Pennsylvania in the east.
Irish Lady’s-tresses is found in North
America and on the western fringe of Europe
in the British Isles. This is an extreme example
of a pattern of distribution known as ‘amphi-
Atlantic’ and is shared by just a few other
plants, including Blue-eyed Grass, Pipewort
and American Pondweed. A variety of
theories have been advanced to account for
the distribution of Irish Lady’s-tresses but the
simplest and most likely is that it arrived in
Britain and Ireland either via the long distance
dispersal of wind-blown seed or by pieces
of root and rhizome being carried across the
Atlantic by birds.
How to find it
The grass-like leaves are very hard to see,
making this species effectively invisible when
not in flower. The number of plants flowering
varies from year to year and can be very low
following a dry spring, while many that do
flower are grazed off. A visit to the Hebrides,
Lough Corrib (Co. Galway) or Lough
Mask and Lough Con (Co. Mayo) would,
nevertheless, be worthwhile.
DESCRIPTION
Height: 10-35cm.
Stem: Yellowish-green with scattered glandular
hairs towards the tip.
M 31 July, Co. Londonderry (Sean Cole) The flowers are
arranged more or less obviously into three rows. Leaves: There are three to five (sometimes up
to eight) yellowish-green leaves. The lower are
Outer Hebrides on Vatersay, Barra, South long, narrow and parallel-sided with a hooded
Uist and Benbecula. Otherwise it is rare but tip and are held very erect. The upper leaves
has been recorded from Islay, Mull and Tiree. are short, pointed and loosely sheathe the
On the mainland the species is found in three stem. The margins of the leaves may be rolled
small areas of Inverness-shire close to its inwards, making them even narrower and more
Hebridean strongholds: Moidart, Morvern grass-like, especially in Northern Ireland. The
and Ardnamurchan (where small numbers shoots appear in the autumn, around October,
have been found around the western end of but the leaves may not expand until the spring;
Loch Shiel). There are also old records from in North America this species is wintergreen.
Kintyre. In England it is known only from Spike: Five to 40 flowers are arranged in three
one site in Devon, to the east of Tavistock rows up the stem (rarely only two rows) and
on the southwest fringe of Dartmoor. World each row is variably twisted; this arrangement is
range: Widespread in North America, from most obvious on plants with numerous flowers.
the Atlantic to the Aleutian Islands, south to Bract: Narrow and pointed, sheathing the
003 systematic.indd 176 29/1/09 12:26:01

ovary, with glandular hairs on its outer surface.

The lower bracts are as long as the flowers, the
upper bracts shorter.
Ovary: Cylindrical, three-ribbed, on a very
short stalk and with a few glandular hairs. It is
held vertically but bends at the tip so that the
flowers lie horizontally.
Flower: White and tubular. The sepals and
petals are creamy-white, washed green towards
the base. The sepals are narrowly triangular
and blunt-tipped, with glandular hairs on their
outer surface and three greenish veins; the
petals are narrower and more strap-shaped. The
sepals and petals form a hood that encloses the
column and the basal half of the lip, with the
tips of the sepals distinctly turned outwards.
The lip is creamy-white with fine green veining
and is fiddle-shaped, with the ‘waist’ nearer the
tip. The sides of the larger and broader basal
portion are turned upwards to form a trough or
gutter, with two small nectar-producing bosses
at the extreme base. The smaller distal portion
is tongue-shaped or square-ended, frilled
and toothed along the edges and sharply bent
downwards. The flowers have, at most, a faint
scent (earlier reports of a strong vanilla-like
scent are presumably errors).
Subspecies
None.
M 16 August, Colonsay, Inner Hebrides (Richard Gulliver).
Variation and varieties The lip is frilled and spoutlike as in all the Spiranthes; this
plant has rather widelyspaced flowers.
Plants in Northern Ireland are said to be
on average taller and ‘leggier’ than those in
southern Ireland. They have creamy rather that in large populations plants resembling
than white flowers, a looser spike, narrower both of these extremes as well as a range of
lip and leaves that are often in-rolled at the intermediates are present, and no subspecies
edges, giving them an especially grass-like or varieties are usefully recognised in the
appearance. Plants from Scotland and Devon British Isles.
are intermediate. At one time Northern Irish Genetic studies have shown that there is a
plants were treated as the same species as in split between a northern group of populations,
North America, although separated as a distinct from Coll, and from Barra and Vatersay in the
variety (S. romanzoffiana var. stricta), with Outer Hebrides and a southern group from
southern Irish plants separated as a distinct, Colonsay and Ireland (no plants from mainland
endemic, species (named ‘S. gemmipara’). Two Scotland were examined). There has been little
distinct subspecies are still recognised by some recent contact or gene flow between these two
authors, but examination of the variation across groups, but this division is not reflected in any
the entire range in North America has shown known differences in appearance or ecology.
003 systematic.indd 177 29/1/09 12:26:02

O 4 August, Benbecula,
Outer Hebrides (Richard
Gulliver). In a small
minority of cases two plants
grow together, presumably
the product of vegetative
reproduction. These are
very fewflowered spikes.
BIOLOGY prevents automatic self-pollination). But, given

the scattered distribution of Irish Lady’s-tresses
The reproductive biology of Irish Lady’s-tresses in the British Isles, wind-blown seed is the
in the British Isles is a mystery. In North only plausible mechanism for dispersal and
America, medium-sized, long-tongued bees subsequent propagation.
act as pollinators but until very recently bees Concerted research efforts have now
had only very rarely been recorded visiting the produced more records of bumblebees visiting
flowers in the British Isles. Even more curiously, the flowers in Scotland, and at the largest
mature seed capsules had never been found in colony in Ireland the orchids are frequently
Scotland and only once or twice in Ireland and it visited by medium-tongued and long-tongued
was assumed that little or no seed was produced bumblebees (Bombus pascuorum and B.
(studies in North America have shown that hortorum respectively), and occasionally by
the column structure of Irish Lady’s-tresses short-tongued honeybees. Only bumblebees
003 systematic.indd 178 29/1/09 12:26:03

were observed to pick up pollinia, however, and reason or another and relatively few develop.
presumably these act as the primary pollinators. Nevertheless, the incidence of ‘twinned’ orchids
Careful examination of plants from Colonsay in a population, presumably the product of
in the Inner Hebrides revealed that c. 40 % vegetative reproduction, varies from very low
of the flowers had their pollinia removed and to over 25% of plants. Genetic studies do not,
c. 70 % had pollen on their stigmas, while at however, indicate that groups of clones are at
the Irish colony pollinia were found on the all common. It is possible that new plants may
probosci/thorax of visiting bumblebees and develop from fragments of root, perhaps broken
on the stigmas of randomly checked flowers. off by cattle or sheep, but this yet to be recorded.
Not surprisingly, in view of these observations, Irish Lady’s-tresses is an enigma. Seed
it has been shown that a small but consistent is apparently rarely produced. Vegetative
proportion of flowers on Colonsay do set seed reproduction is uncommon and does not account
(although much lower than the proportion of for long-distance dispersal. Clearly, much more
flowers that are pollinated, and the number remains to be learned about this beautiful orchid,
of seeds per capsule is very low). What is especially if it is to be effectively conserved.
surprising is that examination of more than
1000 flowers at the Irish site found no ripe
Poorly understood. The aerial stem grows from
capsules. The reason for the very low levels of
a cluster of two to six thick, fleshy, tuberous
seed production is not clear, but the populations
roots, more-or-less vertical in the soil and
on Colonsay and in Ireland (the southern
connected at the top by a very short rhizome.
group) have been shown to have a low level of
A lateral bud develops at the base of the aerial
genetic diversity, perhaps indicating a ‘genetic
stem (or on the rhizome if the plant is ‘dormant’
bottleneck’ during which numbers fell to a very
underground) between July and October and
low level. High levels of inbreeding may cause
overwinters, going on to form the leaves in the
problems such as impaired stigma receptivity
spring of the following year and, if the plant is
and low pollen viability, but examination
to flower, the stem appears in early June. The
of flowers on Colonsay showed high levels
species may become ‘dormant’ underground,
of pollen germination on the stigma, with
with up to six year’s absence recorded.
pollen tubes growing down the style and some
Little is known about the development from
penetrating an ovule as normal. The northern
seed to flowering plant, but the species possibly
group of populations show a high level of
spends five years growing underground before
genetic diversity and it would seem logical that
the first leaves appear.
they produce seeds much more consistently.
Searches in 2007 failed to confirm this, but Hybrids
it was poor year for the species; when flowers None recorded in the British Isles.
are few and scattered, or the weather poor, the Name and classification
chances of insect pollination are much reduced. The specific name romanzoffiana honours
British and Irish populations can reproduce
Nicholas Romanzof, a Russian minister of state.
vegetatively through the development of an
The species was discovered in Alaska around
additional bud at the base of the stem (rarely
1828 when it was still a Russian territory.
two or very occasionally three). Two buds can
produce two aerial stems in the next growing
season, and these may eventually separate to HISTORY aND
form two plants. Extra buds are, however, only CONSERVaTION
produced by a small percentage of plants each Nationally Scarce in Britain and specially
year (less than 5% in the Hebrides), and it seems protected in Northern Ireland under Schedule
that many of these extra buds disappear for one 8 of the 1985 Wildlife Order (NI) and in Eire
003 systematic.indd 179 29/1/09 12:26:03

under the Flora (Protection) Order. It is the and the population trends are not clear. On
subject of a UK Biodiversity Action Plan and the one hand, it is known from an increasing
a priority for conservation because Britain and number of sites, both in Ireland and Scotland,
Ireland hold the only European populations. largely due to particular efforts being made to
The first Irish record was published in find it; it was found for the first time on Mull
1828 by Sir J.E. Smith in the English Flora: in 1990 and on Tiree in 2002. On the other
‘Near Castletown opposite to Bearhaven on hand, it tends to ‘vanish’ unpredictably and often
the northern side of Bantry Bay, County of rapidly from known localities. It is, however,
Cork, Mr Drummond…communicated to me effectively impossible to find unless in flower
in August, 1810’. Irish Lady’s-tresses was first and the most likely explanation for its erratic
found in Northern Ireland in 1892, at Brackagh appearances is that populations are relatively
Bog near Lough Neagh, Co. Armagh, on an old stable but grazing and, perhaps sometimes,
dug-out peat bog. the weather prevent many or most plants from
flowering. Sheep, cattle or rabbits often graze-off
Past and present occurrence of Irish Lady’stresses in Brit
ain and Ireland (based on presence or absence in 10km the flower spikes and whole colonies can appear
squares of the National Grid; data from the New Atlas). to vanish overnight if sheep are turned out in
the vicinity. As with many orchids, slugs can also
Britain Ireland
be a particular problem, grazing-off both the
total historical range, 21 44 flower spikes and the leaves. Recent experiments,
1500-1999
where sheep have been excluded from large
current range 17 (0.6%*) 17 (1.7%*)
populations, have produced a profusion of
% lost, 1500-1969 5% 36% flowers, and careful long-term monitoring on
% lost, 1970-1986 14% 25% Barra has indicated that the population there is
% lost, total 19% 61% comparatively stable and long-lived.
The impact of grazing on Irish Lady’s-
* current range as a % of the total number of 10km squares tresses is of conservation concern, but the
situation is not clear-cut. Colonies, sometimes
The first positive identification for Scotland
large, have been recorded on sites with a
came from Colonsay in 1930 (a lady’s-tresses
variety of grazing regimes, and indeed, it may
had been found on Coll in August 1921 but
be tolerant of heavy grazing. It is becoming
was not certainly identified until 1939). The
clear that many orchids do best when grazing
first record for the Scottish mainland was as
recent as 1954. reduces competition and breaks up the sward,
Irish Lady’s-tresses was found on the providing suitable sites for the establishment
southwestern edge of Dartmoor, Devon, in July of seedlings, even if this means that many or
1957, when there were seven plants on a heavily most of the flowers are grazed-off before setting
grazed lawn of Purple Moor-grass and on the seed. This may apply to Irish Lady’s-tresses.
adjacent ‘bog’. It has but not been seen since c. However, some well-established Irish sites are
1993, although the site is essentially unchanged. only lightly grazed or even ungrazed.
Most populations of Irish Lady’s-tresses The species is certainly vulnerable to
are small and scattered but some of the largest changes in management, such as drainage, and
produce from 100 to 200 spikes annually, relatively few sites are protected as SSSIs or
with 400 spikes recorded recently from one reserves.
area in Co. Mayo. The largest site in Scotland
was recently estimated to hold 1,100 plants
(flowering and vegetative combined).
Irish Lady’s-tresses is very hard to survey
003 systematic.indd 180 29/1/09 12:26:03

AUTUMN LADY’S-TRESSES l 181
autumn lady’s-tresses Near Threatened

Spiranthes spiralis
This delicate little orchid blooms in August and September and is the last species to flower
in the orchid season. It is confined to short turf, often near the sea, and is one of the orchids
that has taken happily to lawns. If conditions are right, hundreds or even thousands of
flower spikes may appear, but it can exist undetected for many years if the grass is cut too
often.
Identification
Distinctive. The delicate, slender spike is
decorated with a row of small, tubular, white
flowers that are usually arranged in a spiral
pattern. A few tiny, bract-like leaves clasp the
stem and there is a rosette of short, oval leaves
lying flattened to the ground a little to one
side.
Similar species
Irish Lady’s-tresses is confined to Ireland
and northwest Scotland with an outpost on
Dartmoor. When in flower it has prominent,
long, narrow leaves on the stem. Habitat is
also a good distinction, as it is found in wet
grassland and bogs.
Summer Lady’s-tresses is extinct in Britain
and, like Irish Lady’s-tresses, has long, narrow
leaves along the stem when in flower.
Creeping Lady’s-tresses is found in
coniferous woodland (rarely moorland or
dunes), usually on acid soils. It is found in
northern England and Scotland with a few
populations in Norfolk. Soil conditions can
change over a very short distance, however,
and prior to 1979 Autumn and Creeping
Lady’s-tresses could be found growing within
a few metres of each other at Holt in Norfolk.
Creeping Lady’s-tresses is easy to distinguish
as it grows from horizontal rhizomes that
form irregular patches of leaves with scattered
flower spikes. Its little bell-like flowers are
exceptionally hairy and its leaves are faintly
net-veined.
P 18 August, Norfolk. The flowers can be arranged into

either clockwise or anticlockwise spirals.
003 systematic.indd 181 29/1/09 12:26:04

003 systematic.indd 182 29/1/09 12:26:07

animals may knock-over or bite off the flower

spikes but ignore the leaf rosettes, and mowing
also leaves the rosettes unharmed. Conversely,
the lessening or abandonment of grazing or
‘improvement’ with fertilisers will promote
more vigorous vegetation that will swamp the
orchid. In the past, Autumn Lady’s-tresses
was also found in damp meadows in the drier
climate of eastern England, but these are almost
all now drained and ‘improved’ and the orchids
have gone.
Flowering period
Early August to the end of September or even
early October.
Range
Southern England from the Isles of Scilly
to Kent, now mostly south of a line from
Bristol to London and rare to the north of
this, although there are some scattered sites as
far north as Norfolk, southeast Lincolnshire,
Yorkshire and in Cumbria around the northern
shores of Morecambe Bay. Also found in
Wales, especially the southeast and coastal
M 18 August, Norfolk.The new rosette of leaves appears regions, the Isle of Man and Ireland, where
alongside the current flower spike but is hidden in the it is scattered in the west and on the south
grass.
and east coasts, and absent from the northern
O 30 August, Norfolk. third of the island. World range: Almost
confined to Europe, with outposts in North
Habitat Africa and around the Caucasus. Found north
Found on short, dry, nutrient-poor turf in
sunny situations, often near the sea. It usually
1987-99
grows on calcareous soils on chalk, limestone, 1970-86
dunes, shingle banks or in the grykes of pre 1970
limestone pavements. Ancient earthworks are

favoured, as are lawns; some old lawns were
made with turf from nearby pastures and
the orchids may have come with the turves,
but they can certainly also colonise lawns
via wind-blown seed. It is also sometimes
found on old tennis courts, road verges and
reservoir embankments and has occasionally
been recorded from grassy places on less-
acid heaths. The critical factor is the lack of
competition from taller and more vigorous
herbs and grasses, and the correct conditions
may be provided by grazing or mowing; grazing
003 systematic.indd 183 29/1/09 12:26:08

to Denmark and southernmost Sweden, east

to the Baltic States and western Ukraine
and south to the Mediterranean, including
most of the Mediterranean islands from the
Balearics to Cyprus. Also Algeria and Tunisia,
northern Turkey, Israel, Syria, the Caucasus and
northwestern Iran.
How to find it
The slender spikes of small greenish-white
flowers do not grab the attention and can easily
be missed from a walking height. They become
much more obvious from a low vantage point,
and squatting down to scan along the ground
is often the easiest way to see the plants. Non-
flowering lady’s-tresses are almost impossible
to find, however, as the rosettes are only 2.5-
7cm in diameter, lie hidden in the grass and
are easily overlooked as plantains. But, from
nothing, flower spikes may appear and start to
bloom in just a week.
The number of flowering spikes may
fluctuate very widely from year to year, but
the total number of plants present, including
non-flowering rosettes or those ‘dormant’
underground, is much more stable. The
proportion of plants producing flower spikes
is probably related to the weather, but the
optimum set of conditions is not known.
Some populations may not be able to flower
for several years but will then bloom en masse
when grazing or mowing ceases. In a dry
summer lawns may be left uncut, allowing
the orchids to flower, and until this happens
Autumn Lady’s-tresses may grow unseen for
many years; suitable lawns may be picked out
later in the autumn as they often support a
good variety of fungi, including the colourful
waxcaps.
DESCRIPTION
Height: 3-15cm, sometimes to 20cm high, but
in the wet summer of 2004 a population in
Norfolk averaged 15cm high, with the tallest
28.5cm.
Stem: Pale green, densely covered towards
M 22 August, Norfolk. The tiny flowers are arranged in a the spike with fine, white glandular hairs.
spiral around the stem, resembling a lady’s braided hair. The flower spike grows from the centre of the
003 systematic.indd 184 29/1/09 12:26:09

previous season’s rosette, with or without the whitish fringe. The bracts are lanceolate, taper
remains of the dead leaves at its base. The next abruptly to a fine point and are a little less than
season’s rosette grows beside it. twice the length of the ovary, which they clasp.
Leaves: The flowering spike has three to seven Ovary: Green, three-ribbed, stalkless, with fine
small, narrow, lanceolate, bract-like leaves that glandular hairs. The ovaries are held upright
tightly sheathe the stem. They are greenish with but bend at the tip so that the flowers are held
a narrow, whitish or translucent fringe. Just to more-or-less horizontally.
one side of the base of the spike there is a tight Flower: A small, white, trumpet-shaped tube.
rosette of up to ten leaves. These are around The sepals and petals are white, often washed
3cm long, dark, shiny green with a faint blue green towards the base. The sepals are oblong
tone, oval or elliptical in shape, taper to a point, but taper slightly to a blunt tip and have
have a thick keel and broadly sheathe the stem glandular hairs on the outer surface. The petals
at their base. This rosette of leaves emerges in are slightly shorter, rather narrower and more
August or September and overwinters, dying strap-shaped. The upper sepal and the petals,
off in late May or early June. together with the lip, form a long, narrow tube
Spike: 3-21 flowers (the average is 9-11) are with the tip of the upper sepal curved upwards.
arranged in a row up the stem. In most plants The lateral sepals are held slightly drooped and
this is twisted so that the flowers form a spiral spread horizontally away from the tube. The
pattern, but in some the twist is so slight that lip is pale green, becoming whiter towards the
the flowers simply form a line along one side of edges, with two small, globular nectaries at
the spike. The spiral can be either clockwise or the base. It can be oval or even heart-shaped
anti-clockwise and is mostly twisted through with a slightly squared-off tip, and the sides
less than 360°, although in some plants it may curve upwards to form a trough or gutter; for
be through three full turns. its entire length the lip also bends downwards
Bract: Pale green with scattered glandular hairs to resemble the lip of a china teapot’s spout,
towards the base and a narrow transparent- and the extreme tip is rolled downwards
and crimped. The greenish column projects
horizontally into the tube made by the petals
and lip. The flowers are honey-scented.
Subspecies
None.
None.
BIOLOGY
The flowers are pollinated by bumblebees.
When the flower opens the longboat-shaped
rostellum lies close to the lip at the bottom of
the tubular flower with the sticky viscidium
facing downwards. In this position the
rostellum blocks access to the stigma so that the
flower cannot be self-pollinated. A bumblebee
lands on the lip and inserts its proboscis in
M 22 August, Norfolk. The lip has a spoutlike tip with
crimped edges, and there are abundant glandular hairs search of the nectar that is produced at its base.
on the stem, ovaries and sepals. There is just enough room for the proboscis to
003 systematic.indd 185 29/1/09 12:26:11

all flowers. The production of abundant seed

is probably a factor is the lady’s-tresses success
in colonising new sites (although the capsules
contain an average of only 850 seeds each, a
relatively low figure).
The species also reproduces vegetatively
from lateral buds at the base of the stem. This
leads to the formation of clusters of two or
three plants (clusters of up to 12 have been
recorded). Vegetative reproduction is, however,
thought to play only a minor role in the
turnover of a population of Autumn
Lady’s-tresses.
The aerial stem arises from a very short
rhizome that is almost concealed by two or
three (rarely up to five) thick, fleshy, tuberous
roots that are very much like miniature parsnips
in shape and have a few short, transparent hairs.
The roots that supported the current year’s
leaves and flower spike eventually shrivel when
their store of food is exhausted and are replaced
M 22 August, Norfolk. Despite their small size, the flow by new ones. The protocorm that is produced
ers are pollinated by bumblebees. by the germinating seed is heavily infected with
fungus, but by the time the roots appear the
reach the nectaries but in the process it pushes
rhizome is free of ‘infection’. However, the roots
past the rostellum, and the pollinia are attached
are each infected in turn as they develop.
to the proboscis by the fast-drying glue of the
Plants may spend one or possibly more
viscidium. The bee then moves on, carrying the
years underground with no aerial leaves and
pollinia with it. Over the following 24 hours
still flower the following year. This indicates
or so the column and lip move apart, creating
that the fungal ‘partner’ plays a significant
enough space for a visiting bumblebee to insert
part in the nutrition of the mature plant. In a
its proboscis, with the pollinia attached, into
study in Bedfordshire the ‘half-life’ averaged
the flower, where they rub against the stigma
6.9 years and varied from 4.6-9.2 years (the
(which is now exposed and has become much
‘half-life’ is a measure of the life expectancy
stickier). The pollinia are brittle and break
of the orchid after its first appearance above
off in small pieces. A single bee can pollinate
ground. It marks the point at which 50% of
several flowers. Due to the 24-hour interval,
the population that emerged in any given year
older flowers are always pollinated with pollinia
have died).
from a younger flower. The lowest flowers in
Early researchers concluded that Autumn
a spike open first and, as bumblebees work
Lady’s-tresses took many years to go from seed
upwards from the bottom of a spike (i.e. from
to flower. However, in the laboratory green
older to younger flowers), they visit the older
leaves have been produced six months after
flowers first and cannot pollinate them with
germination, and flowering plants in five years.
pollinia taken from the same spike. In this way
cross-pollination is virtually guaranteed. The Hybrids
mechanism is efficient and seed is set by almost None.
003 systematic.indd 186 29/1/09 12:26:14


The specific name spiralis means ‘twisted’ or
‘spiral’ (as does the generic name).
HISTORY aND
CONSERVaTION
In 1548 William Turner wrote in his Names
of Herbes: ‘Satyrion is very commune in
Germany, and a certeyne ryghte kynde of the
same groweth besyde Syon [Sion, Middlesex],
it bryngeth furth whyte floures in the end of
harueste and it is called Lady traces’. With this
statement the Autumn Lady’s-tresses became
the first species of wild orchid to be recorded
in Britain, an honour it shares with Common
Twayblade.
Past and present occurrence of Autumn Lady’stresses

Atlas).
Britain Ireland

1500-1999
current range 302 (11%*) 34 (3.4%*)
% lost, 1500-1969 50% 62%

% lost, 1970-1986 5% 9%
M September, Norfolk. Autumn Lady’stresses can occur

In the 450 years since William Turner’s in large numbers and, although the number flowering
each year is very variable, the total population is relatively
account, the species has not fared too well in stable.
the British Isles and has disappeared from
55% of its historical range in Britain and 71% poor grassland the Autumn Lady’s-tresses
in Ireland; it is classified as Near Threatened. cannot tolerate any sort of ‘improvement’ by the
On the edge of its range in Britain and Ireland, addition of fertlisers. Ploughing of downland
Autumn Lady’s-tresses has retreated to its and pastures and their conversion to arable or
core habitats, and the losses are concentrated reseeding with vigorous grasses are even more
in the northern and eastern parts of its former directly destructive processes. Conversely, the
range and at inland sites in general. Sadly, abandonment of grazing leads to the invasion
it has almost gone from northern England, of grassland by scrub. These practices have been
the Midlands, East Anglia and Kent. It is taking their toll since the 19th century, but the
extinct in Northamptonshire, Warwickshire, losses are mitigated to a small extent by gains
Staffordshire, Nottinghamshire, Derbyshire, as the species is able to colonise new sites and
Cheshire and, needlees to say, Middlesex. has appeared in large numbers on the lawns of a
With its requirement for short, nutrient- few favoured housing estates.
003 systematic.indd 187 29/1/09 12:26:15

summer lady’s-tresses Extinct

Spiranthes aestivalis
The delicate, pure white flower spikes of Summer Lady’s-tresses once graced a few favoured
bogs on the Channel Islands and in the New Forest. Sadly, it is the only orchid to have
become extinct in the British Isles and was last recorded in about 1952. Changes to the
habitat were important factors in its decline, but it may have been given the final death knell
by collectors. It has also declined sharply throughout northwest Europe and the chances of
it reappearing naturally in England appear to be slim.
Identification
The small, white, trumpet-shaped flowers,
arranged on the stem in a spiral pattern,
identify this species as one of the lady’s-tresses.
When in flower it has several long, narrow
leaves at the base of the stem.
Similar species
Autumn Lady’s-tresses is relatively common on
short, dry grassland but has also been found,
albeit rarely, in damp meadows and in grassy
places on less acid heaths. When in flower it has
a few small, bract-like leaves on the stem but
the basal rosette has already died off; the new
rosette appears a little to the side and the leaves
are always much shorter and blunter than in
Summer Lady’s-tresses.
Irish Lady’s-tresses is largely confined to
Ireland and western Scotland but has been
found on Dartmoor and could conceivably
turn up elsewhere in southern England. It too
favours wet, boggy habitats but it has larger
flowers than Summer Lady’s-tresses, usually
arranged into three spiral rows. Each flower has
a longer bract, 10-20mm long, rather than just
6-9mm.
Habitat
In the New Forest the species was confined
to wet, peaty, valley bogs with bog mosses
(Sphagnum spp.). It favoured areas that were
slightly less acidic and where the vegetation
P The flowers are very similar in shape to Autumn Lady’s

tresses, but this species grows in acidic bogs, in this case
with Crossleaved Heath (Bob Gibbons/Natural Image).
003 systematic.indd 188 29/1/09 12:26:16

SUMMER LADY’S-TRESSES l 189
was relatively low and open. On the Channel Ovary: Yellowish-green, finely hairy, six-ribbed
Islands it was found in a Sphagnum bog on and slightly twisted. The ovaries are stalkless
Guernsey and on wet sandy ground on the and held upright but bend at the tip so that the
margin of St. Ouen’s Pond on Jersey. In Europe flowers lie horizontally.
it is also found on moist heathland, damp dune Flower: White and trumpet-shaped. The sepals
slacks and other damp, base-rich areas with and petals are glandular-hairy on their outer
short, open vegetation. surfaces and strap-shaped, with the petals a
Flowering period little shorter and narrower than the sepals.
Mid-July to mid-August. They form a tight tube around the column with
their tips splayed outwards at the mouth of the
Range tube. The lip is tongue-shaped with two small,
Extinct in the wild in Britain. Formerly found nectar-secreting glands at the base. Towards the
in the New Forest in Hampshire and on rear, the sides of the lip curve upwards to form a
Guernsey and Jersey in the Channel Islands. gutter shape and at the tip the lip turns sharply
World range: Central and southern Europe, downwards and has a conspicuously frilled
north to Germany and northwest France, or crimped margin. The flowers are slightly
east to the Czech Republic and the former scented.
Yugoslavia, and south to the Mediterranean,
including the Balearic Islands, Corsica and Subspecies
Sardinia. Also found in North Africa in None.
Morocco and Algeria, and possibly also in Variation and varieties
Turkey. It is rare and declining in much of None.
Europe due to habitat destruction, especially
towards the northern edge of the range, and is
extinct in Holland and Belgium.
BIOLOGY
How to find it
The structure of the flower is very similar to
On the surface there seems little chance of
Autumn Lady’s-tresses, and the flowers are
finding this species. From time to time there are
presumably pollinated in the same manner
rumours that it has appeared again in the New
but there is no specific information on this
Forest, but these have never come to fruition.
species. The flowers are said to be fragrant in
DESCRIPTION the evening, which may indicate that they are
Height: 10-20cm, sometimes to 40cm. pollinated by night-flying moths. The species
Stem: Yellowish-green, with fine glandular also reproduces vegetatively from additional
hairs towards the tip. lateral buds at the base of the stem.
Leaves: Yellowish-green, glossy, narrow and Development and growth
strap-shaped; three to six leaves are held erect at The aerial stem grows from a cluster of two to
the base of the stem and there are one to three six thick, fleshy roots. Little is known about the
smaller, bract-like sheathing leaves above these. period between germination and first flowering.
The leaves emerge in spring and are retained all Development may be relatively rapid, however,
summer. because, following germination, protocorms
Spike: Five to 20 flowers are arranged into a appear by the spring, the first root forms in
single row which is twisted spirally around the July and the first tiny leafy shoot appears above
stem, sometimes several times. ground in the late summer.
Bract: Lanceolate, finely hairy at the base. The
bracts are rather longer than the ovary, which Hybrids
they tightly clasp. None.
003 systematic.indd 189 29/1/09 12:26:17

003 systematic.indd 190 29/1/09 12:26:20

SUMMER LADY’S-TRESSES l 191
Name and classification of decline and extinction; rare and on the edge
The specific name aestivalis means ‘of the of its range, it was perhaps especially vulnerable.
summer’. In the 19th and 20th centuries all-too-
numerous collections were made of flowering
HISTORY aND plants, often complete with roots, both for
CONSERVaTION private and public herbaria.
Summer Lady’s-tresses was also found on
The first British record dates from 1840.
the Channel Islands. The first record was on
At a meeting of the Linnean Society on 17
24 July 1837, from the banks of St. Ouen’s
November of that year, ‘Mr. Janson exhibited a
Pond on Jersey. This was its only locality on the
specimen discovered in August last by himself
island and it was always scarce at this site. Due
and Mr. Branch near Lyndhurst, Hampshire’
to overcollecting it quickly became even scarcer
(Proceedings of the Linnean Society).
and was last recorded in 1926, when just a
In England this species was always confined
single plant was seen (and even this last plant
to the region southwest of Lyndhurst in the
was possibly later collected). On Guernsey it
New Forest. In this area it occurred regularly,
was found on boggy ground around the lake
sometimes in large numbers, in at least five sites
of Grande Mare in around 1841 and was
until the end of the 19th century. As with many
fairly common to start with, but collecting and
species of orchid, the number of flowering
drainage led to its demise by 1914.
plants varied tremendously from year to year.
There are occasionally rumours that
Thus, in 1901, E.D. Marquand noted that he
Summer Lady’s-tresses has been refound in
‘once saw half an acre of bog perfectly white
the New Forest or reports that it has been
with these flowers, but the following year only
discretely reintroduced, without official
a few spikes of bloom appeared’. At another site
sanction, to one or more of its former sites.
200 flowering spikes were noted around 1900.
Nothing has ever come of these stories, but the
There was, however, a marked decline in the
possibility of an undocumented reintroduction
20th century, and the species had vanished from
will cast a permanent shadow over any natural
most sites by 1940. It persisted in the original
reappearance.
1840 location, a bog north of the A35, just
north of New Forest Gate and east of Highland
Water, until 1952 (with a possible sighting
nearby in 1959).
The decline was partly due to habitat
destruction, especially drainage and
afforestation (thus a site west of Brick Kiln
Inclosure was unwittingly destroyed when
drained by the Forestry Commission). Some
of the original sites are still very wet, however,
and superficially unchanged, although shading
by trees and scrub and subtle changes to
the vegetation through a process of natural
succession may have made them unsuitable.
Summer Lady’s-tresses may also be one of the
few cases where collecting was a genuine cause
O Extinct in Britain, Summer Lady’stresses is also on

the decline throughout northwest Europe (Bob Gibbons/
Natural Image).
003 systematic.indd 191 29/1/09 12:26:20

Distribution
Primarily Asian, with 30 species in total, but just
one ranging west to Britain and Europe.
Name
The origin of the generic name Herminium is
uncertain. It may derive from the Greek and
means ‘buttress’ or ‘foot of the bed’. This is
supposedly an allusion to the ‘pillar-like’ tubers
but these are spherical in Musk Orchid, the first
species to be given the name. Alternatively, it may
derive from Hermes, the messenger of the gods.
Genus HERMINIUM
musk orchid
003 systematic.indd 192 29/1/09 12:26:24
MUSK ORCHID l 193
musk orchid Vulnerable, BaP

Herminium monorchis
Despite its name, this small orchid does not smell of musk. It is confined to a relatively
few sites in southern England, where it is found in very short turf on chalk or limestone,
occasionally in large numbers. Surprisingly, and despite its status as a Nationally Scarce
orchid that has been lost from around 70% of its former range, it seems to have received
remarkably little attention from scientists and conservationists.
Identification
This diminutive orchid is easily overlooked but
when found is not hard to identify. The flower
spike is crowded with tiny greenish-yellow
flowers. Unless examined closely, the lip is
hardly different in appearance to the petals and
sepals, and the flower therefore appears to be
made up of six almost identical narrow ‘petals’
that form a little bell.
Similar species
Bog Orchid is also very small and greenish-
yellow in colour but is strictly confined to acid,
boggy ground, and the structure of its flower is
completely different.
Habitat
Musk Orchid is found exclusively on short,
well-drained grassland on chalk or limestone
soils. Its small stature means that it cannot
compete if the vegetation is tall, so thin or
compacted soils that restrict plant growth
are favoured. It particularly likes the narrow
‘terracettes’ formed on steep downland slopes
by soil creep, as well as ancient earthworks,
abandoned quarries, chalk and lime pits, and
spoil heaps. It has been recorded up to 215m
above sea level.
Flowering period
Early June to early July, sometimes to early
August. There can be large variations from
O 29 June, Hampshire. The colour of the flower varies a

little, from slightly yellower to slightly greener.The smallest
plants may be just 2cm high.
P 29 June, Hampshire.The small, slender spikes of Musk

Orchid are often hard to see from walking height.
003 systematic.indd 193 29/1/09 12:26:26

194 l GENUS hERMINIUM
003 systematic.indd 194 29/1/09 12:26:29

MUSK ORCHID l 195
year to year in the number of flowering How to find it

spikes, with the temperature and amount of This small, greenish-yellow orchid is hard
rainfall over the previous summer being the to spot. This is especially so on anything but
determining factors. A hot dry summer can the shortest turf or if the spikes are few and
cause a big drop in the number of flowers the scattered. In those situations, scanning carefully
following year. from a low-level vantage point, sitting or
squatting, can help. Once one spike has been
1987-99 seen, however, others will usually be found
1970-86
pre 1970
nearby. It can be easier to tread on than to
see, so it is good practice to keep to whatever
paths are available. On the other hand, in
a few favoured sites Musk Orchid occurs
in large numbers and can form dense and
obvious stands. One of the best sites for this
species is Noar Hill (Hampshire), and others
include St Catherine’s Hill (Hampshire) and
Malling Down (Sussex), with small numbers
at Tottenhoe Knolls (Bedfordshire), Park Gate
Down (Kent), Box Hill (Surrey) and Ham Hill
(Wiltshire).
Range
Confined to the chalk of the North and
South Downs in Kent, Sussex and Surrey,
also Hampshire, Dorset (just two sites),
Wiltshire and Berkshire, the Cotswolds
in Gloucestershire and the Chilterns in
Buckinghamshire, Hertfordshire and
Bedfordshire. It is very local and is absent
from large areas of apparently suitable habitat.
World range: Widespread in Europe and
Asia. In Europe it is found north to c. 61°N
in Scandinavia, southern Finland and the
Baltic States, and it ranges south to southeast
France, Italy (the Apennines) and the Balkans,
with isolated populations in northwest Turkey
and the Caucasus. The species reaches its
western limit in northern and eastern France,
but there is an isolated population in central
Spain. It ranges in a narrow band across Siberia
to the Russian Far East, North Korea and
northeast China, and is also found in another M 29 June, Hampshire. In the tiny, belllike flowers the
narrow band across northern China, with lip appears similar to the petals and sepals, with only its
narrow central lobe visible.
scattered records from northern Japan and the
Himalayas. O 28 June, Hampshire.
003 systematic.indd 195 29/1/09 12:26:31

ridged towards the tip.

Leaves: There are two mid-green basal leaves
(rarely three or four), strongly keeled and
oblong to oval-oblong in shape, and one to
three small, lanceolate, bract-like leaves further
up the stem. The leaves emerge from early May
onwards and persist until mid-September but,
in a dry season, they may not appear until early
June and will wither early.
Spike: The flowers appear densely packed on
the spike due in part to the relatively large
ovary. There are 20-30 flowers on most plants
but over 70 on the very largest. The spike is
often one-sided.
Bract: Green, lanceolate and roughly the length
of the ovary.
Ovary: Relatively large and inflated, even
when the flowers are fresh, greenish-yellow,
prominently ribbed and slightly twisted. The
ovary is upright but narrows at the top into
a stalk-like base for the flower which is bent
through more than 90°, thus holding the flower
pendant.
Flower: The whole flower is greenish-yellow
and very small, about 2.5mm wide x 3mm from
front to back. It does not open widely and the
sepals, petal and lip all point forward and are
more-or-less parallel, giving the flower a spiky,
tubular or bell-like shape. The sepals are oval,
with the lateral sepals slightly smaller than the
upper sepal. The petals are slightly paler and,
rather unusually, longer than the sepals. They
are spear-shaped, with variable small side-lobes
towards the base and a narrower tip. The lip
has a long and narrow central lobe and rather
shorter side-lobes. To the rear the lip narrows
into a short, blunt chamber (a rudimentary
spur) which secretes nectar. There are two oval
pollinia, each attached by a very short, elastic
caudicle (stalk) to a saddle-shaped viscidium
M 29 June, Hampshire.The flowers are scented but smell that is almost the same size as the pollinia. The
of honey, not musk. viscidia each have a delicate skin but are not
enclosed in a bursicle (pouch). The flowers have
DESCRIPTION a sweet, honey-like scent.
Height: 2-30cm but usually 3.5cm-15cm and
rarely more than 20cm. Subspecies
Stem: Yellowish-green to dark green, distinctly None.
003 systematic.indd 196 29/1/09 12:26:32

MUSK ORCHID l 197
Variation and varieties this may be the major means of recruitment

None. to the population; two to five ‘daughter’ tubers
are produced at the ends of slender rhizomes
BIOLOGY up to 20cm from the parent tuber. In this way
Pollination and reproduction extensive clones can develop.
The flowers are pollinated by a variety of tiny Development and growth
insects such as flies, parasitic wasps, gnats and The leaves and flower spike grow from a single
beetles, typically just 1-1.5mm long. As it feeds spherical tuber which starts to wither away
on the nectar at the rear of the lip, the insect by flowering time, and there are a few short,
ruptures the skin of the relatively large viscidia, thin roots. The tip of the tuber and usually
and these stick the pollinia to the insect’s legs. also the roots are ‘infected’ with fungi. Two
Once the insect leaves the flower, the pollinia or more new tubers are formed each growing
rotate forward to be in the correct position to season, the larger of which provides for the
make contact with the stigma of the next flower next year’s leaves and flower spike, the smaller
to be visited; this can be on the same plant. ones are ‘daughter’ tubers. Musk Orchid is
There are conflicting reports on the possibility sometimes described as ‘migratory’ because the
of self-pollination. A study in Sweden found replacement tuber grows at the end of a short
that it did not take place, but it has also been rhizome and therefore the aerial shoot appears
stated that the flowers can be self-pollinated, in a slightly different place each season. After
the anther withering and the pollinia dropping flowering once, plants may appear again merely
onto the stigma immediately below. Whatever as vegetative rosettes or even remain ‘dormant’
the mechanism, 70-95% of flowers set seed. underground for one or two years before they
The species also reproduces vegetatively and flower again.
M 29 June, Hampshire. Musk Orchid can reproduce vegetatively to form extensive patches of clones.
003 systematic.indd 197 29/1/09 12:26:33

Musk Orchid is reported to flower after Not surprisingly, it is classified as Vulnerable.

a period of immaturity lasting several years, The ploughing of chalk grasslands from the
but in cultivation plants have flowered within late 18th century onwards caused the first
two years of seed being sown. Individuals can wave of losses, especially in East Anglia, and
be long-lived, with an age of 27 years being many sites had gone by 1930, including John
recorded. Ray’s in Cambridgeshire. Subsequently, the
‘usual suspects’ of agricultural intensification,
Hybrids
the scrubbing-over of grassland (especially
None.
following the outbreak of myxomatosis)
Name and classification and overgrazing have taken their toll. It is
The specific name monorchis means ‘one- now extinct in Somerset, Glamorganshire,
testicle’ (i.e. ‘one-tuber’). When flowering this Oxfordshire, Rutland, Cambridgeshire, Suffolk
species appears to grow from a single tuber, and west Norfolk, where three 18th or 19th
unlike members of genera such as Orchis and century records included a site at Heacham, the
Dactylorhiza, which have two obvious tubers northernmost locality for the species. However,
at the base of the stem. In fact, Musk Orchid it can, at least occasionally, colonise new areas.
has several, but the replacement and ‘daughter’ Many sites are now protected as SSSIs and
tubers are formed at the end of thread-like reserves, and a few colonies hold hundreds or
rhizomes and easily missed. The specific name even thousands of plants.
is therefore a misnomer. Despite its predilection for dry chalk
grassland, the species is vulnerable to
HISTORY aND summer drought, which can lead to the early
CONSERVaTION withering of the leaves and thus a reduction
Musk Orchid was first recorded in Britain in in the amount of carbohydrates stored in the
1663 by John Ray in his Catalogus Plantarum tubers for the following season. In some cases
circa Cantabrigiam nascentium (‘Catalogue there may then be a big fall in the number of
of plants found around Cambridge’): ‘Orchis flowering spikes, the plants appearing merely
pusilla odorata…In the chalk pit close at as non-flowering rosettes, but the population
Cherry Hinton’. can recover rapidly following a good growing
season. Prolonged droughts can, however,
Past and present occurrence of Musk Orchid in Britain devastate the species; the long, hot summer of
squares of the National Grid; data from the New Atlas). 1976, the worst drought in southern England
for 250 years, caused big losses in Sussex from
Britain Ireland which it has not yet recovered.
1500-1999
current range 32 (1.1%*) 0
% lost, 1500-1969 52%

% lost, 1970-1986 17%
% lost, total 69%
Musk Orchid has been lost from 69% of its

total historical range and is Nationally Scarce,
having been recorded from just 32 10km
squares in the New Atlas period of 1987-99. P 14 May, Kent, Lady Orchid.
003 systematic.indd 198 29/1/09 12:26:34

Genus ORCHIS
‘typical’ orchids
003 systematic.indd 199 29/1/09 12:26:38

200 l GENUS ORChIS
This genus contains some of the rarest and most spectacular orchids in the British Isles.
Many species have flowers that recall tiny human figures, with lips that are divided into
‘arms’ and ‘legs’. All grow from tubers and can spend part of the year underground in a safe
‘resting’ state. This allows the orchid to survive unfavourable periods, such as the summer
drought in the Mediterranean or sub-zero winter temperatures.
Distribution accurately termed ‘root-tubers’ or ‘root-stem

There are about 50 species, most of which are tubers’ and are essentially roots that have
found in Europe and adjacent areas in North been modified to become specialist storage
Africa and the Middle East. Just five species organs.
occur in the British Isles. To follow the pattern of growth through
Floral structures the year it is best to start in the late summer
There are two pollinia, each narrowing to a when the orchid is ‘resting’ and consists only of
caudicle (stalk) which is attached to one of a single tuber with one terminal bud; the leaves,
the two viscidia. The viscidia are each in turn rhizome and roots die off once the orchid has
contained within a bursicle, a pouch-like cavity flowered.
on the column. In the autumn the bud on the tuber
produces a short rhizome and from this a few
Pollination roots develop. The roots serve two functions.
Members of this genus are pollinated by insects, First, their ‘infection’ with fungi provides the
but only a few species produce nectar to reward plant with nutrients. Second, they supply the
their pollinators. The majority rely instead plant with water and this becomes particularly
on deceit; insects are attracted to the brightly important once the leaves have appeared. In
coloured and scented flowers but receive some members of the genus the leaves develop
no reward. in the autumn, in others they do not expand
Growth pattern until the spring.
At flowering time all Orchis species have two Once the roots have developed, they begin
almost spherical tubers side by side at the to produce nutrients. Some of these are stored
base of the aerial stem. The ‘tubers’ are more in a new tuber that has started to form on the
August October December February April May August
M Annual cycle of growth and replacement of tubers in the genus Orchis (after Wells, 1981).
003 systematic.indd 200 29/1/09 12:26:39

GENUS ORChIS l 201
rhizome, side by side with the old; due to its rhizome develops and the first tuber develops
relationship with fungi, the orchid can produce on this rhizome, the root and protocorm dying
nutrients even before its leaves appear. The away in the late summer to leave only the tuber.
rhizome goes on to form the aerial stem and In the autumn, a new rhizome grows from this
when the flowers open in the spring this has tuber and produces one or more roots and in
two tubers at its base. The older of these has the spring a leafy shoot is produced and a new
supplied the current season’s growth, including tuber forms at its base during the summer; the
the leaves, stem and flower spike. This tuber is pattern of growth is then similar to that of the
starting to shrink and will have vanished by the adult plant. Some species spend several years
late summer’s ‘resting period’. The newer tuber is as an underground seedling, producing a new
plump and swollen and continues to grow until tuber each year.
the leaves die down. It will go on to overwinter The pattern of development of the seedling
and form the flower spike in the following year. apparently varies slightly between species and
even between populations. Some may produce
Fungal partners
a tuber directly from the protocorm, without
The tuber does not have a fungal ‘infection’
any roots developing beforehand, whereas
although fungi are found in its epidermis (and
others develop an aerial stem and root in the
in hot, dry, climates the fungus may only be
first season after germination and then go on to
able to survive the summer drought in the
produce a tuber.
skin of the orchid’s tubers). Fungal activity is
concentrated in the roots and sometimes also Vegetative reproduction
the rhizome. All Orchis species are able to spend Additional tubers may be formed at the base
one or more years underground, presumably of the aerial stem and these will go on to form
sustained by fungi. separate plants as the connecting stem dies off
in the autumn.
In all species the seed germinates in late Name
summer or autumn and forms a protocorm. The generic name Orchis derives from the
This rapidly forms the first root, which is Greek orkhis and means ‘testicle’. The two
‘infected’ with fungi and supplies the protocorm rounded tubers have long been considered to
with nutrients. The following summer, a small resemble male genitalia.
protocorm
bud replacement
tuber
root tuber
spring summer autumn spring summer autumn

YEAR 2 YEAR 3
M Development from seed in the genus Orchis (after Rasmussen 1995).
003 systematic.indd 201 29/1/09 12:26:40

202 l GENUS ORChIS
man orchid Endangered, BaP

Orchis anthropophora
Formerly: Aceras anthropophorum

The narrow flower spikes of Man Orchid do not command attention from a distance but on
closer acquaintance they reveal how apt the English name is. Just why do they look so much
like tiny, hooded, human figures, and what benefit does the plant gain by having flowers of
this particular size and shape? As with so many questions about orchids, we do not know
the answer. Man Orchid is relatively widespread in southern England, occurring north to
Lincolnshire and west to Gloucestershire, but away from Kent and Surrey it is extremely
local and either uncommon or rare; in the long term the future of the few remaining isolated
populations must be in doubt.
Identification
The long, narrow spike and very man-like
flowers are distinctive. The sepals and petals
form a ‘hood’ or ‘cowl’ and the tiny figure faces
downwards, concealing its ‘face’. The lip is deeply
lobed to form the ‘arms’ and ‘legs’. The flowers are
yellowish to greenish, variably washed with red;
plants in full sun may be on average the reddest
and some may even have bright foxy-red lips.
Similar species
Frog Orchid may be similarly coloured but is
usually rather smaller, and the lip is not divided
into ‘arms’ and ‘legs’.
Common Twayblade is also vaguely similar
but the tiny green flowers are rather different in
shape and it has only two large, rounded leaves.
Habitat
Typically found on well-drained grassland
on chalk or limestone, often on or at the foot
of a slope, with a predilection for abandoned
quarries and pits. Roadside verges, churchyards,
field margins and stabilised dunes or shingle
can also provide suitable habitat. It frequently
grows in relatively long, rank grass and among
scrub and it will sometimes spread under the
eaves of nearby woodland. It is vulnerable to
O 2 June, Northamptonshire. The spikes are often tall
and very slender; the basal leaves may be hidden (or nib
bled by rabbits).
P 2 June, Northamptonshire. Although classified as
‘Endangered’, Man Orchid can still be found in good num
bers at a few favoured sites.
003 systematic.indd 202 29/1/09 12:26:43

003 systematic.indd 203 29/1/09 12:26:48
204 l GENUS ORChIS
competition, however, and dense scrub will

crowd it out; conversely, populations can be
eliminated by heavy grazing.
Flowering period
Early May to late June, sometimes from late April,
but usually at its best in late May. The flowers
open in slow progression from the bottom of the
spike, so flowering can be protracted.
Range
Southern and eastern England, with the bulk of
the population on the North Downs in Surrey
and Kent. It is rare and very local elsewhere
and confined to the region south and east of
a line from Bristol to the Humber. There are
four or five sites in Hampshire, around four in
Suffolk, mostly on protected roadside verges, M 9 June, Northamptonshire. The flowers are truly
manlike.
and between two and four sites in Essex. Two
each remain in Sussex, Northamptonshire Algeria and Morocco, and it also occurs in
and Lincolnshire, with just one in Wiltshire, southwest Turkey and Lebanon.
Gloucestershire, Oxfordshire, Bedfordshire and
Norfolk; there is a single site in Warwickshire How to find it
but it was apparently introduced there, in 1968. A very localised species, nowadays only likely to
World range: Western and southern Europe, be found on reserves or protected road verges.
North Africa and the Levant. It ranges north It can be hard to spot if it is growing among
to Holland, southern Germany, Switzerland, long grass. Two of the best sites for this species
the former Yugoslavia and Greece and south are Barnack Hills and Holes (Peterborough)
to the Mediterranean, including most of the and Wye (Kent). Box Hill (Surrey) is another
Mediterranean islands from the Balearics to classic locality, although it has declined there in
Cyprus. In North Africa it is found in Tunisia, recent years.
DESCRIPTION
1987-99 Height: 15-65cm but usually 20-30cm and
1970-86
pre 1970 rarely over 45cm.
Stem: Pale green with some membranous
sheaths at the extreme base.
Leaves: Green, dull or slightly bluish, distinctly
veined, keeled and narrowly oval-lanceolate to
strap-shaped. There is a basal rosette of three or
four leaves, some lying flat and some held at about
45°, and higher on the stem one or two smaller
and more lanceolate sheathing leaves. In some
areas the leaves appear in spring, in others in
November or December, becoming fully formed
by January or February. In all plants the tips of
the lower leaves are often scorched by May and
they all die off after flowering; this pattern of
003 systematic.indd 204 29/1/09 12:26:50

MAN ORCHID l 205
growth is probably an adaptation to the mild wet moderate to good, but despite this it is thought
winters and hot dry summers in the core area of that most reproduction is vegetative via the
its distribution around the Mediterranean. production of additional tubers.
Spike: Tall, narrow, more-or-less cylindrical Development and growth
and dense, with up to 50 or even 90 flowers. Once they have appeared above ground for
Bract: Green, lanceolate and half the length of the first time, plants may live for up to 14
the ovary. years although they may not flower every year
Ovary: Pale green, long, cylindrical, boldly or even appear above ground. Conversely,
ribbed and twisted. some may flower for five years in a row. Man
Flower: Green or yellow, variably tinged red, and Orchids only rarely die after flowering just
very man-like. The sepals are oval and various once, and in a study in Bedfordshire the
shades of yellowish-green, often with a distinct ‘half-life’ averaged 5.8 years and varied from
maroon fringe and midrib. The petals are pale 4.0-7.8 years (the ‘half-life’ is a measure of
green, slightly shorter, much narrower and more the life expectancy of the orchid after its
strap-shaped. Both the sepals and petals form first appearance above ground and marks the
a hood over the column, with the petals fully
concealed. The lip is variably green or yellow,
often strongly washed red or reddish-brown,
especially around the edges, but can be pure red
or yellow. It hangs almost vertically downwards
and has three lobes: two long, narrow side-lobes
at the base (the ‘arms’) and a terminal lobe that is
itself divided half way to the base into two lobes
(the ‘legs’). There is sometimes a tiny projecting
tooth between the ‘legs’. There is no spur, rather
there are two shiny, whitish swellings on either
side of the base of the lip which curve round to
join the column and enclose a shallow pit with
two small, nectar-secreting depressions. The
flowers have a faint, unpleasant smell.
Subspecies
None.
Var. flavescens lacks red pigments (anthocyanins)
and has a green hood and contrasting yellow lip.
It is rare.
BIOLOGY
Little is known about the pollination of this
species. However, the numerous hybrids with
Lady, Monkey and Military Orchids found
in Europe would suggest that it shares a suite
of pollinating insects with those species; in
England ants and hover flies have been seen M 8 May, Kent. Many flowers have largely red ‘arms’ and
with pollinia on their heads. Seed-set is ‘legs’.
003 systematic.indd 205 29/1/09 12:26:51

206 l GENUS ORChIS
point at which 50% of the population that have also been lost to scrub encroachment. Extinct
emerged in any given year have died). There in Somerset, the Isle of Wight, Hertfordshire,
is no information on the period between Buckinghamshire, Cambridgeshire, Leicestershire
germination and flowering. and also Derbyshire (where it was introduced at
Hybrids Ashover but not seen there recently).
In Europe, hybrids with Monkey, Military and
Past and present occurrence of Man Orchid in Britain and
Lady Orchid are common but in England such Ireland (based on presence or absence in 10km squares
hybrids have only been found twice. of the National Grid; data from the New Atlas).
O. x bergonii, the hybrid with Monkey Orchid,
Britain Ireland
was found in Kent in 1985, although it has
been suggested that this may be the result of total historical range, 109 (1.7%*) 0
1500-1999
inadvertent hand pollination.
O. x macra, the hybrid with Lady Orchid, current range 48 0
was found in Kent in 1998, when two plants % lost, 1500-1969 46%
were seen. % lost, 1970-1986 10%
Name and classification % lost, total 56%
The specific name anthropophora means ‘man- * current range as a % of the total number of 10km squares
bearing’.
Man Orchid was formerly placed in the
genus Aceras as Aceras anthropophorum. Indeed,
it was the only species in that genus, which was
distinguished from the genus Orchis by the lack
of a spur. Recent DNA studies have confirmed
that this difference is purely superficial and that
Man Orchid is a perfectly good Orchis. The
generic name Aceras derived from the Greek
and meant ‘without-a-horn’, a reference to this
absence of a spur.
HISTORY aND
CONSERVaTION
The first British record dates from 1690, when
John Ray published his Synopsis Methodica
Stirpium Britannicarum (‘Methodical synopsis
of British plants’): ‘Found by Mr. Dale in an
old Gravel-pit at Dalington (Ballingdon) near
Sudbury’.
Nationally Scarce and still decreasing in
numbers, Man Orchid has been lost from 56% of
its historical range and is classified as Endangered.
It was formerly much commoner in East Anglia
but vanished as pastures were ploughed from
the late 19th century onwards. Sites were also
destroyed when quarries and pits were used as
‘landfill’, field margins were sprayed or subject to
drift from nearby operations and road verges cut M 2 June, Northamptonshire. Some flowers are much
or sprayed unsympathetically. Conversely, sites yellower, with little or no red.
003 systematic.indd 206 29/1/09 12:26:53

MONKEY ORCHID l 207
monkey orchid Vulnerable, BaP

Orchis simia
One of our rarest orchids, the exotic, monkey-like appearance of the flowers, combined with
the inevitable secrecy that surrounded such a rarity, has excited botanists for generations. It
is confined to three sites in the Chilterns and Kent, two of which welcome visitors.
Identification
To see a Monkey Orchid you will almost
certainly have to visit one of its two ‘public’ sites,
so identification is straightforward. The flowers
are distinctive and definitely resemble a Spider
Monkey, with the hood forming the ‘head’ and
the lobes of the lip, the slender, curved ‘arms’
and ‘legs’.
Similar species
Military Orchid is also very rare and the two
species are not found together anymore in
England. However, should a new colony of
plants be discovered, identification would be an
issue. In Military Orchid the ‘legs’ are straight,
distinctly broader than in Monkey Orchid and
widen towards the tip, and the hood forms a
longer, neater ‘helmet’. The shape of the flower
spike is rather different, too, being taller and less
crowded, with the flowers opening in sequence
from the bottom of the spike upwards.
Habitat
Monkey Orchid prefers south-facing slopes
on open, grazed chalk grassland. It probably
favours the interface between grassland and
woodland or scrub, benefiting from the shelter
which scattered trees and shrubs provide from
desiccating winds and grazing animals. The
slightly moister conditions in light shade are
also likely to be beneficial and the bare ground
under scrub can provide suitable conditions
for seedling establishment. Should the shade
become too dense, however, it ceases to flower
or even to appear above ground. Monkey
Orchid may be badly affected by drought; for
example at Faversham in Kent the population
‘crashed’ after the hot, dry summers of 1975 P 28 May, Kent. The rosette of shiny, green basal leaves
and 1976 and was slow to recover. is usually hidden in the grass.
003 systematic.indd 207 29/1/09 12:26:54

208 l GENUS ORChIS
Range
Currently confined to Hartslock in Oxfordshire
and two sites in Kent (a confidential site near
Faversham and Park Gate Down, where it was
introduced). World range: Southern Europe
and the Mediterranean region, including
the Balearics, the Aegean islands, Crete and
Cyprus, north to Holland (very rare), southern
Germany, Hungary and Romania. Ranges east
to Syria, Turkey, the Crimea, Caucasus, Iran
and Turkmenistan, and also found in North
Africa in Algeria, Tunisia and Libya.
1987-99
1970-86
pre 1970
How to find it
Park Gate Down and Hartslock reserves are the
M 28 May, Kent. At Parkgate Down in Kent, Monkey places to go, and Monkey Orchids are easy to
Orchid grows on open downland, but it can also grow in
light scrub.
find at both.
Flowering period DESCRIPTION

Late May to early June; flowering peaks Height: 10-30cm tall, occasionally to 45cm.
around a week earlier in the Chilterns than Stem: Green, usually washed brownish-purple
in Kent. In most orchids the flowers open towards the tip, angled and with two or three
from the bottom of the spike upwards, but in sheaths at the base.
Monkey Orchid the flowers often open rapidly Leaves: There are three or four shiny green
from the top downwards, although they may basal leaves, oval-oblong, often keeled and
open synchronously (or even from the base blunt-tipped, and two or three sheathing leaves
upwards in Kentish plants). Overall, it is the higher on the stem. The leaves may appear
rapidity with which all the flowers open that is above ground as early as November but more
characteristic of Monkey Orchid, rather than usually do so in January or February.
the direction of opening. The spike is at its best Spike: Roughly globular in shape with the
for a rather brief period. flowers crowded together, and the whole thing
003 systematic.indd 208 29/1/09 12:26:56

MONKEY ORCHID l 209
looking a bit scruffy. Most spikes have ten

to 20 flowers but there are up to 30 on well-
developed plants and as many as 42 on the
most robust.
Bract: Very small, around a third of the length
of the ovary, triangular, chaffy and whitish.
Ovary: Green, heavily washed purple, boldly
six-ribbed, twisted and curved.
Flower: Monkey-like, white or pink, with
darker spots and variably redder extremities.
The sepals are lanceolate, and the petals are
slightly shorter, much narrower and more
strap-shaped; together they form a hood that
encloses the column with the sepals slightly
splayed outwards at the tips. The sepals are
whitish on their outer surface, variably washed
pink and with irregular violet-purple dots,
blotches and streaks. The inner surfaces are
more heavily blotched and streaked, sometimes
almost solidly so. The petals are similar but
sometimes more solidly washed purplish-pink.
The lip has two lobes forming the ‘arms’ and M 28 May, Kent. With long ‘arms’, ‘legs’ and a ‘tail’, it is
not hard to see the ‘monkey’ in a Monkey Orchid.
two lobes forming the ‘legs’, with an additional
small projection between the ‘legs’ (more surfaces, making the hood appear pale pink
phallic than tail-like); the tips of the ‘arms’ and rather than white on the outer surface, and
‘legs’ are curved forwards and upwards. The the spur is pale pink to purple (white to pale
lip is whitish in the centre, variably flushed pink in the Chilterns). However, there is much
violet-purple and spotted violet (the spots are overlap, and the differences do not warrant the
formed by tufts of papillae), becoming violet- recognition of distinct varieties. ‘Var. macra’,
purple towards the extremities of the ‘arms’ with a violet hood and purplish-red ‘arms’ and
and ‘legs’. The spur is pale pink, about half to ‘legs’, was recorded many years ago and may
three-quarters the length of the ovary, slightly have been more prevalent in large populations
down-curved and blunt-tipped. The column but probably falls within the normal range of
is reddish. The flowers have a faint scent of variation and may not be worthy of recognition.
vanilla.
BIOLOGY
Subspecies
None.
Some natural pollination does occur in
Variation and varieties England, with the flowers visited by flies,
The Kentish Monkey Orchids are on average bees and butterflies. Although the spur is not
taller than those in the Chilterns, with stouter thought to contain nectar, two swellings near
stems and, more significantly, more and bigger its mouth may contain sugars which can be
leaves; their largest leaf is up to twice as long. extracted by insects. Seed-set has been poor in
Their flowers have a more extensive area of England and some populations have been hand-
purple spotting on a more purple-washed pollinated. Pollination rates appear to improve
ground, and the spots are larger. The sepals significantly, however, once a population of
have larger and darker markings on their inner Monkey Orchids is above a certain threshold;
003 systematic.indd 209 29/1/09 12:26:58

210 l GENUS ORChIS
O 28 May, Kent. Kentish

Monkey Orchids are, on
average, taller and more
richlycoloured than those
in the Chilterns.
the spectacle of large numbers of flowers may not be entirely typical. A period of three or
be more attractive to potential pollinators. In a four years elapses between germination and the
study in Holland, vegetative reproduction was appearance of the first aerial leaf and a further
found to be rare and only to occur following three to six years before flowers are produced,
good growing seasons. by which time the plants have at least four basal
Development and growth leaves. A similar time scale has been recorded
Much of the information about the life cycle of for Kentish Monkeys. Young plants occasionally
the Monkey Orchid comes from observation disappear underground after they have
of a small population in the Netherlands. This produced their first leaves and then reappear
originated with a single founder plant and may again after one or two years.
003 systematic.indd 210 29/1/09 12:27:01

MONKEY ORCHID l 211
Individual plants can be long-lived, flowering Oxfordshire and East Yorkshire but there
for up to 19 consecutive seasons although this are now only three sites, one in Oxfordshire
may be exceptional. Plants often ‘rest’ between and two in Kent. A Red Data Book species,
bouts of flowering as vegetative rosettes and may it is classified as Vulnerable and is specially
even be ‘dormant’ underground for one or two protected under Schedule 8 of the Wildlife and
years. If absent for three years, however, they Countryside Act 1981.
are almost certainly dead. Severe winters will Monkey Orchid has apparently always
inhibit flowering and may result in many plants been very local in England, but in the 18th and
dying, as the tubers typically lie in shallow soil. early 19th centuries it was frequent in south
Similarly, if the leaves are grazed off the plant Oxfordshire, in the area between Wallingford,
will not flower the following season, presumably Reading and Henley. Most records came from
because without leaves it is unable to build up the slopes overlooking the north bank of the
sufficient resources. Thames between Goring and Caversham
Hybrids (the latter now on the northern outskirts of
O. x. beyrichii, the hybrid with Military Reading) where the river cuts through the
Orchid, occurred in the Thames Valley until the southern outliers of the Chilterns. The species
middle of the 19th century, after which the two then declined dramatically from about 1840,
parent species were not found together. due to the ploughing of downland and the
O. x. bergonii, the hybrid with Man Orchid, was collection of specimens for both herbaria
recorded in 1985 at Faversham in Kent (but it and gardens. However, the main cause of its
has been suggested that this may have been the demise may well have been a big increase in
result of hand pollination ‘gone wrong’). the number of rabbits. By the mid-1920s there
The hybrid with Lady Orchid has appeared was only one substantial colony remaining in
in small numbers since 2006 at the Hartslock
reserve.
The specific name simia means ‘of the ape’ or ‘of
the monkey’.
HISTORY aND
CONSERVaTION
In 1666 Christopher Merrett noted both
Monkey and Military Orchids in his Pinax
Rerum Naturalium Britannicarum (‘A picture of
British natural history’): ‘…on several Chalkey
hills neer the highway from Wallingford to
Redding on Barkshire side the river…’ This
reference to the Berkshire Downs was apparently
the first record of both species in England
and originated with information supplied by a
William Brown of Magdalen College, Oxford.
Old records of Monkey Orchid are often
difficult to assess because for a long time the
species was confused with Military or even
Lady Orchid. Monkey Orchid went on to M 28 May, Kent.The ‘monkeys’ often look as if they have
be recorded from Kent, Surrey, Berkshire, been thrown into a heap.
003 systematic.indd 211 29/1/09 12:27:02

212 l GENUS ORChIS
the Chilterns, at Hartslock, but such was the to around 60 plants in the late 1980s, with
secrecy surrounding this site that the species roughly a third flowering each year. In the early
was generally thought to have become extinct in 1990s the number of flower spikes actually
Britain. dropped, to as few as five in 1990, although
the total number of plants was more constant.
Past and present occurrence of Monkey Orchid in Brit Then, in 1994 the population at Hartslock
squares of the National Grid; data from the New Atlas). started to expand rapidly. Three factors may
have helped; first, a run of mild winters, which
Britain Ireland probably reduced mortality and encouraged
total historical range, 10 0 plants to flower. Second, systematic hand
1500-1999 pollination, which began in 1977; this has now
current range 2 (0.07%*) 0 ceased, as rates of natural pollination and seed
production are high. Third, in 1992 the main
% lost, 1500-1969 60%
colony was fenced against rabbits. Over the
% lost, 1970-1986 20% next three years the number of plants trebled
% lost, total 80% and by 1995 there were 123, including 47 new
* current range as a % of the total number of 10km squares individuals, of which 72 flowered. It is thought
that prior to the erection of the fence rabbits
The Hartslock colony was stable at over 100 ate most of the first orchids to emerge; they can
flowering plants in the 1920s and increased appear above ground as early as January, well
to 200 spikes in the years leading up to World before the seasonal warden puts wire netting
War Two; in 1933 over 30 were picked. But, cages on the orchids in April.
following the war, the increasing mechanisation By 1999 there were 200 plants with 100
of agriculture allowed steeper and steeper of them flowering, and from 2000 Hartslock
slopes to be cultivated and in 1949 and 1950 held around 300 plants, with one to two-thirds
the field at Hartslock that held the Monkey flowering each year. Despite this apparently
Orchids was ploughed. Fortunately, the upper healthy population, however, DNA studies have
part of the slope (the area where the orchids shown that the genetic variability at Hartslock
now grow) escaped as it was steeper, scrubbier is very low, as the population had gone through
and protected by a thick hedge. There may a genetic ‘bottleneck’ when it was reduced to a
already have been a few Monkey Orchids tiny handful of individuals. In the long term, it
growing in this refuge, and some tubers rescued may be better to introduce more variability by
from the lower part of the field were replanted hand pollinating again using pollen from other
there but with unknown results. Chiltern plants, Kent or even France. In the
Just one Monkey Orchid flowered at last few decades Monkey Orchid has appeared
Hartslock in 1950-52 and numbers remained sporadically and in very small numbers at other
painfully low for many years. It was not until sites in the Chilterns. In 1966 it was found
1968 that the population reached even the near Pangbourne and in 1965 and 1971-74 at
modest total of eight flower spikes, and even Aston Rowant.
in 1977 there were just eight plants. In 1975 In Kent, Monkey Orchid was first recorded
BBOWT bought Hartslock but, as on most in 1777 from near Faversham and then again
downland sites, scrub encroachment was a in the early 1800s but it was not seen again
problem. This led to a substantial programme until 1920-23 when a few plants flowered at
of scrub removal, and the area is now grazed in Bishopsbourne (near Canterbury). At the other
the autumn and winter by sheep. end of the county, in west Kent, from 1952
Despite the initial management efforts, onwards a single Monkey Orchid appeared
the orchid population increased very slowly on the rough grass of a disused tennis court
003 systematic.indd 212 29/1/09 12:27:02

MONKEY ORCHID l 213
at a vicarage at Otford. Every year, until 1955,

the vicar took the seed capsules and scattered
seed onto nearby downland. In 1956 there
was a single robust spike and a further six
non-flowering plants at the vicarage. However,
on the retirement of the botanically minded
vicar the new incumbent would not guarantee
to safeguard the colony. All the orchids were
moved to nearby private land, where the largest
flowered once only, in 1957, and then the
Monkeys vanished.
In 1955 a Monkey Orchid appeared near
Faversham in Kent (at the current native site),
but the single plant was eaten, perhaps by a
horse. It may well have been able to bloom
due to the outbreak of myxomatosis which
resulted in the mass-flowering of orchids in
Kent in that year. More were found in the
following years, with up to 38 plants, ten of
which flowered, in 1957-58, growing partly on
open downland and partly in a nearby hazel
copse. From 1958 until at least 1985 the plants
in this colony were hand-pollinated to make
sure that seed was produced. Numbers steadily
increased, with 246 plants and 162 flowering M 28 May, Kent.The colour of the flowers is very variable.
spikes in 1964 and 205 flowering plants in Some appear pale pink at a distance, others are darker
and redder.
1965. The 1975-76 drought badly affected
the colony, however, with none flowering but until 1983, after which the site was washed
in 1977 plants reappeared and by the mid- away in a storm.
1980s there were again 30-50 plants, with ten The Sainsbury Orchid Project organised by
or so flowering. The population here is now the Royal Horticultural Society has become
stable at over 200 plants, although only a small involved in the conservation effort. Each year a
proportion flower. few seedpods are sent to Kew from Hartslock.
From 1958 onwards seed from the Progress with propagating Monkey Orchid
Faversham site was scattered at several other has been slow because cultivation has been
places in Kent, and a population became restricted to sterile, asymbiotic techniques, and
established at Park Gate Down. The first three germination and growth rates are low. By 1996,
plants flowered in 1965, seven years after seed ten plants had been raised to a size where
was sown, but then not again until 1976 when they could be planted out, and in September
two spikes appeared. This population stabilised 1996 the tubers were placed in two clumps on
at about 100 plants in the mid-1990s but has the edge of the existing colony. In 1997 two
increased significantly since then. of these came up, but since then only one has
A few plants appeared in dune grassland at appeared.
Spurn Point in southeast Yorkshire in 1974,
over 250km from the nearest known source of
seed. This colony increased to a maximum of 25
plants, with nine flowering, but only persisted
003 systematic.indd 213 29/1/09 12:27:03

214 l GENUS ORChIS
military orchid Vulnerable

Orchis militaris
The Military Orchid combines two of the qualities which make orchids so alluring; great
rarity and great beauty. It is one of Britain’s most attractive species and is found regularly
at just three sites (two of which are, happily, open to the public). Thought to be extinct
in Britain by the early part of the 20th century, it was dramatically rediscovered in 1947,
although for a long time the site was kept a closely guarded secret. Like its cousin, the
equally rare Monkey Orchid, it has been monitored, managed and mollycoddled since its
rediscovery, but we are still far from unlocking all its secrets.
Identification
One of the so-called ‘manikin’ orchids in which
the flower resembles a tiny human figure,
Military Orchid brings to mind a soldier. The
sepals and petals form a ‘helmet’, purple-striped
on the interior, and the lip has four lobes, two
for the ‘arms’ and two for the ‘legs’. The rows
of purple spots down the centre of the lip are
reminiscent of buttons on a soldier’s tunic.
The allusion to the military was coined before
soldiers habitually wore red uniforms and may
refer to the resemblance of the hood to an
ancient ‘coal-scuttle’ helmet.
Similar species
Monkey Orchid resembles this species in the
general structure of the flower, but its ‘legs’ are
kinked, narrower and do not broaden towards
the tip, and the hood formed by the sepals and
petals is more open. Also, its flower spike is
not only shorter and more crowded but also
more jumbled and disarrayed, lacking ‘military
precision’, and all the flowers open at roughly
the same time.
Habitat
Military Orchid is found in grassland, scrub,
woodland glades, on woodland edges and,
formerly, rough fields. It always grows on chalk.
The species does best in light scrub on old
pastures and in the shelter of woodland edges.
It favours some shade and needs bare ground
for seedling establishment (rather than a closed
grass sward), but it does not do well if there is
too much shade. Recorded up to 183m above M 31 May, Suffolk. The colony at Mildenhall has thrived
sea level in the Chilterns. since the overshadowing trees were removed.
003 systematic.indd 214 29/1/09 12:27:04

MILITARY ORCHID l 215
Leaves: Bright shiny green, prominently keeled,

1987-99
1970-86
strap-shaped or, on more robust plants, broader
pre 1970 and more oval, and slightly hooded at the tip.
Two to five leaves are carried in a basal rosette
at around 45° above the horizontal, and there
are two or three sheathing leaves higher on
the stem, although the upper part of the stem
is bare. In Suffolk the tip of the shoot appears
above ground in late December or more usually
early January and the leaves start to unfurl by
early March.
Spike: Oval or conical, becoming cylindrical
as all the flowers open. There are two to 25
flowers but up to 57 have been recorded on a
particularly robust spike.
Bract: Triangular to oval, very short (much
shorter than the ovary) and green, strongly
Flowering period washed purple or rose.
Mid-May to mid-June, the flowers are at their Ovary: Green, washed purple, boldly ribbed
best in late May and early June. Once a flower and strongly twisted.
has been pollinated it usually shrivels within Flower: Man-like, whitish, with purple spots
a day. and purple extremities. The sepals are oval
Range with pointed tips, and the petals are narrower
Currently confined to two sites in the Chilterns, and more strap-shaped; they form a hood
in Buckinghamshire and Oxfordshire, and with the tips of the sepals swept upwards and
one in Suffolk. Plants have been introduced the mouth of the hood very open. The sepals
into industrial waste-ground at Bolton in are pale dove grey, their outer surfaces lightly
Lancashire (apparently from outside the United washed with lilac, becoming more purplish at
Kingdom) and to sites in Cambridgeshire and the base; the unopened buds are therefore pale
Kent. World Range: Europe and Siberia. In pinkish-grey. The inner surfaces have bold,
Europe occurs north to Holland, northern longitudinal purplish lines and an irregular
Germany, southeast Sweden and Estonia and purplish wash. The petals are more uniformly
south to northern Spain, central Italy, the pale purplish. The lip has two narrow, strap-
former Yugoslavia, Bulgaria, Romania and shaped lobes forming the ‘arms’ and two shorter
European Turkey. Confined to the mountains and broader lobes forming the ‘legs’, with an
in the southern parts of the range. In Siberia it additional small pointed projection between
extends east to the Altai Mountains and Lake the ‘legs’. It is whitish, flushed pink, with the
Baikal. ‘arms’ and ‘legs’ more-or-less solidly purple.
There are two rows of purple spots down the
How to find it centre of the lip (the spots are formed by tiny
Homefield Wood in Buckinghamshire and the papillae) and solid dark purple lines along the
Rex Graham Reserve at Mildenhall in Suffolk centre of the ‘arms’. The spur is purple, short
are the two public sites for the species. (about half the length of the ovary), cylindrical
DESCRIPTION and slightly down-curved. The flowers are
faintly vanilla-scented.
Height: 5-60cm but usually 20-45cm.
Stem: Green, variably tinged purple towards Subspecies
the tip. None.
003 systematic.indd 215 29/1/09 12:27:05

216 l GENUS ORChIS

Var. militaris is on average taller, with seven
to 42 flowers. It has a relatively lax, open spike
and paler flowers. The Suffolk colony has been
assigned to this variety which is widespread in
Europe.
Var. tenuifrons is on average shorter, with two
to 26 flowers, a denser spike, darker flowers
and narrower leaves. Endemic to England, the
Chilterns plants belong to this variety
BIOLOGY
Military Orchid is pollinated by hover flies
and bumblebees. It produces no nectar but
insects may be attracted to the sugary sap in
the wall of the spur. It has always been thought
that few flowers are pollinated in Britain; rates
of 3-11% are given for plants in the Suffolk
colony and 2-28% of flowers were recorded
as setting seed by Summerhayes (1968). At
Homefield Wood, however, 40% and 24% of
flowers were naturally pollinated in 1999 and
2000 respectively. Vegetative reproduction is
probably important for the British populations,
maintaining numbers when recruitment from
seed is low.

This is a relatively long-lived orchid. Many
plants will live for ten years after their first
appearance above ground and a significant
proportion live for at least 17 years. In a study
in Suffolk the ‘half-life’ varied from 2.2-7.8 years
(the ‘half-life’ is a measure of the life expectancy
of the orchid after its first appearance above
ground and marks the point at which 50% of
the population that emerged in any given year
has died).
The period between germination and the
appearance of the first aerial shoot is three to
five years and between 30% and 50% of plants
flower in their first year above ground. Once
they have appeared, most plants flower at least
once and many do so every year, but intervals
of 11 years between flowering have been noted.
M 31 May, Suffolk. Perhaps a third of plants retreat underground
003 systematic.indd 216 29/1/09 12:27:11

during their life, usually spending only one year

‘dormant’, but in some cases up to three years
and absences of up to eight years have been
recorded in Suffolk. In any given year, 5-15%
of the adult population can be underground.
However, ‘dormancy’ is not a good option for
the plants as underground plants have the
highest probability of dying.
Hybrids
O. x. beyrichii, the hybrid with Monkey
Orchid, occurred in Oxfordshire until the
middle of the 19th century, after which the two
parent species were not found together.
The specific name militaris means ‘military’.
HISTORY aND
CONSERVaTION
A Red Data Book species that is classified as
Vulnerable and specially protected under
Schedule 8 of the Wildlife & Countryside
Act 1981.
M 31 May, Suffolk. A fewflowered spike with very lightly
Gerard’s Historie of Plants, first published marked flowers.
in 1597, contains a description of ‘Souldier’s
Satyrion’ or ‘Soldier’s Cullions’ (literally (‘A picture of British natural history’): ‘…on
‘testicles’) which presumably pertains to the several Chalkey hills neer the highway from
Military Orchid: ‘Souldiers Satyrion bringeth Wallingford to Redding on Barkshire side the
forth many broad large and ribbed leaves, river…’ This reference to the Berkshire Downs
spread upon the ground like unto those of the was apparently the first record of both species
great Plantaine: among the which riseth up a fat in England and originated from information
stalke full of sap or juice, clothed or wrapped supplied by a William Brown of Magdalen
in the like leaves even to the tuft of flowers, College, Oxford.
whereupon doe grow little flowers resembling Military Orchid was often confused with
a little man, having a helmet upon his head, his Monkey and Lady Orchids, and many older
hands, and legs cut off; white upon the inside, records, unless supported by identifiable
spotted with many purple spots, and the backe specimens, are open to doubt. It was recorded
part of the flower of a deeper colour tending reliably from Belchamp Walter near Sudbury in
to redness. The rootes be greater stones than Essex in 1729, and there are also old records for
any of the kinds of Satyrions.’ Gerard gives, the North Downs of Surrey, including Box Hill
however, no localities for his Souldier’s Satyrion in the 1830s, and Kent (a herbarium specimen
and may have copied his description from a dated 1836 from Cobham near Rochester).
European herbal. Like Monkey Orchid, the stronghold of
The earliest localised record for the British the species was always the Chilterns, where
Isles was in 1666 when Christopher Merrett it ranged from ‘just west of the Thames in
noted both Military and Monkey Orchids in the region of Streatley and Basildon, along
his Pinax Rerum Naturalium Britannicarum the Chiltern Hills through Oxfordshire and
003 systematic.indd 217 29/1/09 12:27:12

003 systematic.indd 218 29/1/09 12:27:14
Buckinghamshire, to the neighbourhood of

Tring in Hertfordshire and to Harefield in
Middlesex. It seems to have been particularly
abundant around High Wycombe and between
this town and Great Marlow on the Thames’
(Summerhayes, 1968). Once fairly common
within this limited area, it declined dramatically
from around 1850 and was thought to be
extinct after 1929 when the last specimen was
collected from Hertfordshire. Several factors
contributed to the decline, including the
ploughing-up of downland and the collection
of specimens for both herbaria and gardens,
but the main cause may well have been a big
increase in the number of rabbits. There have
been no subsequent records from Berkshire,
Hertfordshire or Middlesex.
The Military Orchid became something
of a Holy Grail for British botanists. Indeed,
it was the title and main subject of Jocelyn
Brooke’s semi-autobiographical The Military
Orchid, published in 1948. In May 1947 it was
dramatically rediscovered by J. E. Lousley at
Homefield Wood in Buckinghamshire, an area
from which it had not previously been recorded.
‘The excursion was intended as a picnic, so M 31 May, Suffolk. A richlycoloured spike.
I had left my usual apparatus at home and
took only my note-book. But I selected our of all likely sites, the colony at Homefield Wood
stopping places on the chalk with some care, was found by Richard Fitter and Frances Rose.
and naturally wandered off to see what I could They dispatched a postcard to Lousley with
find. To my delight I stumbled on the orchid the simple but cryptic message ‘The soldiers are
just coming into flower’ (Lousley 1969). There at home in their fields’. The discovery was not
were 39 plants, with 18 flower spikes, although made public even then, but Homefield Wood
five had been bitten off, probably by rabbits. came to be managed by BBOWT in 1969
Lousley noted that the orchids growing in and they eventually announced to the press in
shade were either flowerless or had small spikes 1975 that the Military Orchid had returned.
and thought that the increase in available light This resulted in headlines like ‘The Beauty that
after trees were felled during World War Two must blossom in secret’. It was not until the end
had probably prompted the appearance of of the 1980s that the location was made truly
the plants. public, and visitors are now welcomed.
Lousley kept his discovery a closely guarded Homefield Wood was originally largely
secret, fearing that the colony would be wiped composed of Beeches with some areas of
out by collectors, and many others tried to track chalk grassland on a south-facing slope. The
down this hidden treasure but to no avail. At wood was largely clear-felled in the winter
last, in 1956, after a long and systematic search of 1947, and the Military Orchid was found
in the spring of that year on chalk grassland
O 31 May, Suffolk. with scattered trees and scrub. The Forestry
003 systematic.indd 219 29/1/09 12:27:16

220 l GENUS ORChIS
Commission has owned the wood since 1955, raked, and scrub is controlled. In addition,
and most of the felled areas were replanted up to 20% of the plants at Holmfield Wood
with conifers in the 1950s. The area around were hand-pollinated from 1986-98. Hand-
the colony was planted with Beech in 1960-61, pollination can result in almost 100% seed set
and other trees regenerated naturally to form a and does not ‘weaken’ the plants (as had been
mixed woodland; trees also invaded suggested); hand-pollinated individuals have
the grassland. flowered every year for a decade.
When first found, there were two distinct Military Orchid was found at a second site
colonies at Homefield Wood. Colony ‘A’ held 31 in the Oxfordshire Chilterns in 1970. The
plants in 1947 and peaked at 35 in 1949, but number of flowering plants did not exceed five
the numbers fell steadily until the last few were until 1999 and there were none in 1984-87.
seen in 1958, with a single plant reappearing Since 1999, however, there has been a rapid
in 1961. Myxomatosis had decimated rabbit increase, with 25 flowering plants in 2003,
populations in 1955, and this probably caused together with a further 25 vegetative rosettes.
a dramatic increase in scrub which, combined This population was also hand-pollinated for at
with disturbance during tree-planting and least 10 years from 1988.
the spread of Rosebay Willowherb, led to the As part of the conservation programme,
extinction of this colony. Colony ‘B’ presumably plants propagated asymbiotically at the
held just eight plants when found, but this Royal Botanic Gardens were planted out at
is the area where the orchid survives to the Homefield Wood and the nearby Warburg
present day. This colony was fenced off in 1968, reserve in 1996 (a total of 231 tubers, both
brambles invaded and a thick understorey one and two years old). Survival was poor but
developed. The population of Military Orchids those that remained first flowered in 2000 and
slowly fell to a low point of 28 plants in 1984, 2002 respectively. In addition, 25 wild plants
with just five flowering. were transplanted from Homefield Wood in
Active habitat management started in 2000 to a site around 25km away. Survival of
1981 with the removal of scrub and later on these mature plants has been better than young
large, overshadowing Yews. This may have led seedlings, with some flowering and setting
to a recovery in numbers in this area to over seed.
50 plants (with over 30 flowering) by 1989. On 2 June 1955 the Military Orchid was
In 1985 a small adjacent area was clear-felled found at Mildenhall in Suffolk, a region from
and the orchids eventually spread into this, which there were no previous records. The
flowering for the first time in 1995; this clearing colony was in an old chalk pit where there were
held half the flowering plants by 2003. Military at least 500 plants, mostly on a heap of pure
Orchids also appeared over 100m away through chalk among birch and Wild Privet. There were
the wood in a third open area in 1983. Overall, over 100 flowering spikes, but within a short
the number of plants at Homefield Wood time the majority had been nibbled off, perhaps
steadily increased to about 80 in 1995, with 45 by deer, and only 16 fruiting spikes remained
in flower. by mid-August. The colony was within a
Since 1995 there has been a dramatic Forestry Commission plantation and at the
upturn, with over 200 plants, at least half time of its discovery the surrounding pines
of which flowered, in 2003. In recent years were only 1.5m tall, thus the site was open
management has included fencing the colonies and sunny.
to exclude rabbits and deer during the growing By 1958 the number of plants in the Suffolk
season. These enclosures are grazed by sheep colony had risen to 2,854 and remained at this
in the autumn and winter, before the orchids level until the late 1960s, although only about
appear above ground. They are also mown and 10% flowered. However, the population then
003 systematic.indd 220 29/1/09 12:27:16

declined rapidly to 252 plants in 1971, with

perhaps only 100 in the following years. To
protect the orchids the Forestry Commission
had erected a tall wire fence around the pit in
the 1960s. This excluded both deer and people
and within a few years Sycamore and Wild
Privet had taken over. These were cleared in
autumn 1972 and the colony increased slightly
and stabilised at 300-400 plants, with about
100 flowering. Scrub clearance continued and
the overshadowing Corsican Pines were finally
removed in 1985-86, allowing in much more
light, and from 1987 there was a dramatic
increase in the population. In 1990 there were
279 flowering spikes and 1,115 plants and in
2000 there were 748 flowering plants and too
many non-flowering plants to count.
The story of the two Military Orchid
colonies highlights the need for the appropriate
management of rare orchids. For example,
in both cases the colonies were fenced in
the 1960s, and although this excluded deer
and rabbits it came close to destroying them
because scrub took over. The recent increases
in Military Orchid are likely to be directly
attributable to better management rather than
a more favourable climate. This species seems
well-adapted to hard winters and is therefore
unlikely to have benefited from recent mild
years, unlike Monkey Orchid.
Genetic fingerprinting has shown that the
three English colonies are distinct and may
represent independent colonisations from
Europe. Each colony is thus valuable and
worthy of conservation in its own right.
Past and present occurrence of Military Orchid in Brit

Britain Ireland

1500-1999
% lost, 1500-1969 79%

% lost, 1970-1986 5%
% lost, total 84%
M 7 June, Suffolk. A large spike, wellout. * current range as a % of the total number of 10km squares
003 systematic.indd 221 29/1/09 12:27:18

222 l GENUS ORChIS
lady orchid Endangered

Orchis purpurea
The sight of a group of Lady Orchids in a woodland glade in May is always a delight. Even
those lucky enough to see them regularly cannot fail to be impressed by the spectacular show
put on by this stately orchid. It is largely confined to Kent where it is relatively common in
chalky woodlands on the North Downs.
Identification
Straightforward. It is usually rather large
and statuesque, and the unopened buds are
dark reddish-purple. Lady Orchid is one of
the ‘manikin’ orchids, and the flowers form a
miniature human figure. The sepals and petals
form a dark ‘bonnet’ which, with the unopened
buds, contrasts strongly with the whitish, dark-
spotted lip. The lip itself is divided into several
lobes to form the ‘arms’ and the ‘skirt’ of the lady.
Similar species
Burnt Orchid is superficially similar, with dark
buds, a dark hood and a white, purple-spotted
lip but it is very much smaller, seldom more
than 15cm tall.
Habitat
Lady Orchid is found in woodland, both
ancient woodland and secondary woods, but
almost always on thin, well-drained chalky soils
(rarely also on limestone or other calcareous
substrates). It favours beechwoods and often
grows on south-facing slopes, frequently on
banks or on the ‘terraces’ formed by the root
plates of the trees, either among a carpet of
Dog’s Mercury or on bare leaf-litter. However,
its preferred habitat may be scrub or coppice,
and it does not flower so freely in shade, being
happier in open, well-lit situations, such as
along paths and rides, in clearings and along
the lower edges of woods. Indeed, it may cease
to flower and ‘disappear’ if the shade becomes
too dense, only spectacularly to reappear after
coppicing, tree falls or felling opens up the
canopy. Conversely, although it is often found
P 14 May, Kent. The leaves are shiny green and rather

flaccid.
003 systematic.indd 222 29/1/09 12:27:20

LADY ORCHID l 223
just outside a wood, it rarely occurs in the full

1987-99
sun of open downland. Lady Orchid also likes 1970-86
shelter from the wind, and in exposed situations pre 1970
the leaves and flowers may be scorched in a cold

spring, the leaves often turning yellow. It occurs
up to 200m above sea level.
Flowering period
Early May to early June, exceptionally from mid
to late April, varying from season to season,
but plants are generally at their best in mid
to late May.
Range
Lady Orchid is locally frequent on the
North Downs in Kent with more than 100
sites in two areas. First, the downs on either
side of the Medway Valley and sporadically
eastwards towards the Stour Valley. Second, Essentially Europe, with outlying populations
the downs between the eastern slope of the in North Africa, Asia Minor, the Crimea and
Stour Valley and Dover. Elsewhere, there are the Caucasus (the last sometimes treated as
two current sites in Oxfordshire (see History a distinct species). In Europe it occurs north
and Conservation) and it has recently been to Holland and Denmark, east to western
recorded in north Hampshire. World range: and southern Poland and Ukraine and south
M 14 May, Kent.There are around 100 sites in Kent, some with over 1,000 flowering plants.
003 systematic.indd 223 29/1/09 12:27:22

224 l GENUS ORChIS
to central Spain, Italy, northern Greece and

Bulgaria, also found on Corsica and Sardinia
and, in North Africa, in Algeria and Tunisia. In
the south of the range Lady Orchid is confined
to the mountains.
How to find it
This large and conspicuous species is easy to
spot when in flower, but as with all orchids,
non-flowering plants can be hard to find and
the proportion of a population in flower varies
from year to year. Especially good sites in
Kent include Yocklett’s Bank and it also occurs
at Wye.
DESCRIPTION
Height: 20-50cm, exceptionally to 100cm.
Stem: Green, becoming purplish-brown
towards the tip.
Leaves: Green and shiny or almost ‘greasy’.
There are three to five, sometimes seven, broad,
oval to oval-oblong leaves. The lower leaves
are blunter and form a basal rosette, the upper
leaves becoming successively more pointed,
keeled and clasping. They appear above ground
between mid-January and mid-February.
Spike: Oval to oblong in shape and lax or
densely flowered. Robust plants have up to
50 flowers.
Bract: Greenish to purple, tiny, elongated and
scale-like.
Ovary: Bright green, sometimes washed purple
along the six ribs, and distinctly twisted.
Flower: Whitish, finely spotted with reddish-
purple and with a contrastingly dark hood.
The sepals are oval, and the petals are shorter,
much narrower and more strap-shaped but
broaden to a spear-shaped tip; together they
form a hood (the ‘bonnet’). They are pale green,
irregularly blotched on both surfaces with
dark purple or purplish-brown, the blotches
becoming more numerous and coalescing
towards the tip, base and sides; the unopened
buds are therefore very dark. The lip is pale
pink to white, washed violet or rose around the
M 14 May, Kent. Although Lady Orchid is associated with
edges, variably spotted pink to reddish-purple
woodland, it does best where there is some sun. (the spots are formed by tufts of tiny papillae).
003 systematic.indd 224 29/1/09 12:27:23

LADY ORCHID l 225
hood may also vary from paler to darker shades

of purple.
Plants in west Kent differ slightly from
those in east Kent (from the eastern side of the
Stour Valley to Dover). They are, on average,
shorter (20-38cm tall rather than 30-76cm)
with a shorter, denser flower spike, shorter
ovaries (13-19mm rather than 19-25mm) and
lips that are more heavily spotted and washed
rose to purple (rather than salmon to brownish-
red); their lips are also blunter and less deeply
lobed. These two groups of populations have
not, however, been given names.
Var. albida lacks anthocyanin pigments and
thus has a pure white lip and a distinctive white
or straw-coloured hood with green veins. It is
scarce.
Var. pseudomilitaris has narrower and more
reddish lobes on the lip, resembling the ‘arms’
and ‘legs’ of a Military Orchid. The hood is,
however, the normal reddish-purple. It is rare.
M 14 May, Kent. The spots on the lip are composed of BIOLOGY

tufts of tiny papillae.
It is deeply lobed with two long, narrow ‘arms’ The flowers are pollinated by small flies and
and two broad terminal lobes, often with frilled bees, including small digger wasps. Seed-set
edges, which form the lady’s ‘skirt’ (the latter is variable, sometimes very low, with only
usually have a tiny tooth between them). The 3-10% of flowers producing ripe capsules.
spur is cylindrical, from a quarter to a half of However, in some years and at some sites it
the length of the ovary, curved and pale green can be good. Most reproduction is by seed
blotched with purple. The column is pale green although vegetative reproduction also takes
washed pinkish or purplish, and the pollinia places, and clumps of plants can be formed in
are blotched purple. The flowers are variably this way.
reported to smell of vanilla or bitter almonds or
to be unscented.
From germination to first flowering takes eight
Subspecies to ten years and the pattern of development
None. is typical of the genus Orchis. Plants may live
for at least another ten years, flowering at
least three times in that period but seldom
Relatively variable. Plants growing in the
every year; the remains of the previous year’s
open tend to be shorter and darker-flowered
dried, dead spike is not often seen next to the
than those in woods with a tendency towards
current flowers. There are therefore usually
brown rather than purple markings. There is
large numbers of non-flowering plants in any
also much variation in the shape, colour and
population.
markings of the lip; this may be white and even
unspotted in some plants and heavily washed Hybrids
pink with dark purple spots in others. The O. x wilmsii, the hybrid with Early Purple
003 systematic.indd 225 29/1/09 12:27:24

226 l GENUS ORChIS
O 14 May, Kent. A flower

with a relatively pale hood
and fine markings.
Orchid, has been reported very rarely in Kent. HISTORY aND

O. x macra, the hybrid with Man Orchid, was CONSERVaTION
found in east Kent in 1998. First recorded by Christopher Merrett in 1666
The hybrid with Monkey Orchid has in his Pinax Rerum Naturalium Britannicum (‘A
appeared in small numbers since 2006 at the picture of British natural history’), ‘On Gads-
Hartslock reserve. hill in Kent’.
Hybrids with Military Orchid have also Since its first discovery, the Lady Orchid has
been reported (Summerhayes, 1968). been largely restricted to Kent, where there are
Name and classification many sites. Currently, there are also two sites
The specific name purpurea means simply in south Oxfordshire. In the Thames Valley
‘purple’. area a single plant was discovered growing
003 systematic.indd 226 29/1/09 12:27:26

LADY ORCHID l 227
under Beeches in mixed woodland in 1961.

It flowered again in 1962 and 1964 but then
vanished. Seven plants were found at the same
site in 1986 (the Beeches and nearby Yews
having been felled in the late 1970s) and this
colony has subsequently flourished, with at least
46 plants and 29 flower spikes in 1997. Also in
south Oxfordshire, there have been up to three
plants (mostly non-flowering) at Hartslock
since at least 2000.
Past and present occurrence of Lady Orchid in Britain and

of the National Grid; data from the New Atlas).
Britain Ireland
1500-1999
% lost, 1500-1969 46%

% lost, 1970-1986 11%
% lost, total 57%
Classified as Endangered, Lady Orchid is

Nationally Scarce and has been lost from 57%
of the historic range. It was formerly found
in Surrey, mostly prior to 1930, but the last
site was at Coulsdon, where it was present
until 1959. Also extirpated in West Sussex,
where there were at least five sites, the most
recent record being in 1976; there is an old,
unconfirmed record for East Sussex. Elsewhere, M 14 May, Kent. The spike is pyramidal at first but
becomes cylindrical as more flowers open.
recorded from Herefordshire in a disused
quarry in 1967, at Leigh Woods in the Avon
Gorge, Somerset, in 1990 (although this plant others are declining, and it does not easily
quite possibly originated from Lady Orchids in colonise new areas. It has long been noted that
cultivation at the University of Bristol Botanic Lady Orchids are subject to the depredations of
Gardens a short distance away), and formerly rabbits and deer which nip off the flowers and
occurred in the Channel Islands. A widely attack the leaves. Slugs are also attracted to the
quoted 1738 record from Essex has been shown species. Deer are now commoner in England
to have involved Military Orchid. than at any time for a thousand years, making
Past declines were due to the loss of this more of an issue. Feral wild boars are also a
woodlands and the cessation of coppicing. The potential problem as they can root out and eat
population in Kent is now stable and some the tubers.
colonies are large (over 3,000 plants were
counted at one site in recent years) although
003 systematic.indd 227 29/1/09 12:27:28

228 l GENUS ORChIS
early purple orchid

Orchis mascula
As its name suggests, this is the first orchid to appear in the spring in most of Britain and
Ireland. It heralds the coming season and gladdens the heart on an early spring day, whether
as splashes of purple among a carpet of Bluebells or scattered among Cowslips and violets on
a roadside bank. One of the commonest and most widespread orchids, Early Purple Orchid
is nevertheless very local in many areas and has largely vanished from farmland habitats in the
lowlands where it is now confined to woods, roadsides, churchyards and nature reserves.
Identification
Early-flowering, purple flowers and spotted
leaves are a distinctive combination. Unspotted
leaves are not uncommon, however, and there is
a scarce white-flowered variant; these could be
more problematic but a look at the structure of
the flower should prevent any confusion.
Similar species
Green-winged Orchid is easily separated by the
parallel green stripes on its sepals. The hood of
its flower is formed by all the sepals and petals,
so that it lacks the erect ‘wings’ of Early Purple
Orchid. In addition, Green-winged Orchid
always has unspotted leaves and usually also a
smaller, fewer-flowered spike, and it does not
grow in woodland.
Marsh orchids have flowers of various
shades of purple and some have spotted leaves,
but the flowers are usually more extensively and
more heavily marked with black dots, lines and
squiggles. Their spur is short and often sack-
like and is either straight or curves downwards,
whereas it curves slightly upwards in Early
Purple Orchid. The spots on the leaves tend to
be regular and often elongated sideways rather
than irregular and often elongated lengthwise as
in Early Purple Orchid.
Habitat
Very variable. It can occur in both grassland
and woodland and on a variety of soils,
although it does have a definite preference
for calcareous soils on chalk, limestone or
M 23 April, Norfolk. In most places this is the first orchid to boulder clay and avoids acid conditions. It
bloom, with rich purple flowers; wellnamed indeed. is found in a wide variety of old grasslands,
003 systematic.indd 228 29/1/09 12:27:29

EARLY PURPLE ORCHID l 229
both dry chalk downland and damp hill most are in flower from late April to late May.
pastures, as well as meadows, rocky mountain Flowering is on average a little later in upland
ledges, railway embankments and cuttings, areas and in Scotland, the season occasionally
road verges, grass-covered dry-stone walls lasts until early July. Plants in sunny, sheltered
and limestone pavements. It also grows in spots flower earliest, whereas those in cool,
deciduous woodland, usually in the better-lit shaded, wet areas will be last. Once it has
areas along rides, tracks and woodland edges. set seed, the tall spikes with numerous dark
Early Purple Orchid is particularly associated purplish-brown capsules are conspicuous well
with coppice woodland, where there is usually a into the summer.
great increase in the number of flowering plants
Range
in the second or third year after coppicing but
Found throughout the British Isles, including
a decline thereafter as the canopy closes again.
the Channel Islands, Isle of Man, Hebrides,
The species does not colonise new sites easily
Orkney and Shetland. Although generally
and is usually found in ancient woodland rather
fairly common or even abundant, it may be very
than relatively recently established plantations
local in areas of acid soils and is largely absent
or secondary woodland (unless, of course,
from some regions, such as the Fens, south
woodland and scrub have invaded old orchid-
Lancashire and mid-west Wales. It is also very
rich grassland). Conversely, many colonies on
scattered in the Borders, northeast Scotland,
road verges and banks may be relicts of long-
Orkney, Shetland, the Outer Hebrides and
gone woods. Recorded up to 880m above sea
southeast Ireland. World range: Western
level (Caenlochan, Angus).
Europe, extending to North Africa and Asia
Flowering period Minor. Occurs north to the Faeroe Islands, c.
Early April to early June in the south, 70°N in Norway, central Sweden and the Baltic
exceptionally from mid-March, although States. The southern and eastern limits are
M 24 May, Co. Clare. Early Purple Orchid is very conspicuous in The Burren and sometimes forms beautiful ‘rock
gardens’ with Primroses and other flowers.
003 systematic.indd 229 29/1/09 12:27:31

230 l GENUS ORChIS
003 systematic.indd 230 29/1/09 12:27:35

Spike: Oval or cylindrical but often rather

1987-99
1970-86
irregular in shape and rather open, especially
pre 1970 in the lower half, with ten to 50 flowers,
occasionally more.
Bract: Green, usually strongly washed purple.
The bracts are lanceolate and about as long as
the ovary, which they clasp.
Ovary: Green, variably washed purple and
clearly ribbed.
Flower: Various shades of purple, occasionally
pale rose-pink or white. The sepals are
lanceolate, the lateral sepals asymmetrical, and
the petals are smaller and more arrow-shaped.
The upper sepal and petals form a hood over
the column and the two lateral sepals are held
erect and pushed backwards, resembling ‘angel’s
wings’ (they may almost touch at the rear).
poorly known due to confusion with closely- The lip points downwards and outwards and
related species, but it is found south and east is three-lobed with the central lobe usually the
to Italy, Greece, Turkey and Bulgaria and is
probably absent from much of Central Europe.
Also present in the Canary Islands, Balearics,
Corsica, Sicily and Malta, and Tunisia in
North Africa.
How to find it
Generally common and easy to find. In some
areas, such as the Peak District, Yorkshire
Dales and The Burren in western Ireland,
Early Purple Orchid occurs in large, extensive
colonies but it is more usually found in small
and often scattered groups.
DESCRIPTION
Height: 10-45cm, occasionally to 60cm.
Stem: Stout, pale green, angled and usually
flushed purple towards the tip.
Leaves: The three to eight basal leaves are
variably oblong-lanceolate in shape and often
blunt-tipped. They are glossy green, usually
marked with large, irregular, rounded or
elongated blackish-purple spots on the upper
surfaces and rarely also on the undersides. They
are held close to the ground, either spreading
upwards and outwards or in a flatter rosette.
There are two or three rather smaller and more M 2 May, Norfolk. The shape of the lip and, to a lesser
extent, the colour of the flower, is variable.
pointed sheathing leaves higher on the stem and
these may have a few spots or a purple wash. O 2 May, Norfolk.
003 systematic.indd 231 29/1/09 12:27:37

232 l GENUS ORChIS
largest. The side-lobes are folded downwards,

and the entire lip is sometimes folded or
creased along its centre line. The terminal lobe
is shallowly notched and the edges of all three
lobes are crinkled (crenate). The base of the
lip and mouth of the spur are much paler and
whiter, sometimes with some yellow tones,
and usually spotted with purple, the spots
composed of dense tufts of short papillae. The
spur is long and narrow, at least as long as the
ovary, broadens a little towards a blunt tip
and curves upwards. The column is variably
greenish or purple, and the pollinia are dark
green. The flowers are initially sweet-smelling,
recalling honey or Lily-of-the-valley, but this
quickly changes to a rank smell, usually said
to be like tomcat’s urine and to be especially
pungent at night.
Subspecies
None recognised. In the Outer Hebrides, far
north of mainland Scotland, Shetland and a
few places in western Ireland plants are just
5-10cm tall and flower from early June to early
July. These populations are sometimes named
O. m. ebudium.
As in many orchids, plants growing in the open M 25 May, Co. Clare.The white flowered variety alba has
are often shorter and stockier than those in an unmarked lip and yellow pollinia. It is scarce.
woodland. These are sometimes referred to as
the ‘grassland’ and ‘woodland’ forms.
Var. alba lacks anthocyanin pigments and has
white, unmarked flowers with yellow pollinia,
an entirely green stem and bracts, and no leaf
spots. It is rather scarce. An even scarcer variant
has white flowers which retain purple spots at
the base of the lip and sometimes has spotted
leaves too. A ‘broken-coloured’ variant has also
been recorded very rarely and this has a pale
pink lip, copiously flecked with tiny purplish
markings.
BIOLOGY
The flowers are pollinated by bumblebees and
M 12 May, Norfolk.The long spur contains no nectar; the
bright colours of the flower are therefore a ‘deceit’ to lure to a lesser extent cuckoobees and a variety
potential pollinators. of solitary bees. A visiting insect touches the
003 systematic.indd 232 29/1/09 12:27:40

column and the pollinia are stuck to its head

by the sticky viscidia. Once the bee has left
the flower, the pollinia on the insect rotate
forward in about 30 seconds so that they will
make contact with the stigma in the next flower
visited. Early Purple Orchid is self-compatible
and is sometimes self-pollinated.
The flowers of Early Purple Orchid have
no nectar and offer no reward to a pollinator,
although it has been suggested that a sugary sap
is produced inside the wall of the spur. Rather,
it is thought that the orchid takes advantage of
the naivety of the bees which, newly emerged
from hibernation in the spring, have yet to learn
which flowers are genuine sources of nectar.
These naive bees are attracted, at least for a
while, by the bright colours and scent of the
flowers. The various species of bee emerge from
hibernation at different times, and therefore
the orchid can take advantage of a succession of
pollinators as the season progresses. Seed-set
is variable, with the lowest, earliest-opening
flowers most likely to be pollinated. Vegetative
reproduction may occur occasionally via the
production of additional tubers.
The first aerial leaves are usually produced in
the fourth year after germination, and more
and bigger leaves appear in successive seasons
until enough reserves have been accumulated
to produce a flower spike; up to eight years
may elapse between the first appearance above
ground and the first flowers. Around 60%
of plants will flower in successive years; the
remainder either appear as a rosette of leaves
or, in about 17% of cases, spend a year ‘dormant’
underground before appearing again. Up to
12 years dormancy has been noted in Europe.
A maximum lifespan of 13 years after the first
appearance of leaves has been recorded.
Hybrids
O. x wilmsii, the hybrid with Lady Orchid, has
been reported very rarely in Kent.
P 22 May, Co. Clare.The influence of the different flower

colours on potential pollinators is unknown.
003 systematic.indd 233 29/1/09 12:27:42

234 l GENUS ORChIS

The specific name mascula means ‘male’ and may
be a reference to its vigour, early flowering or is
perhaps another allusion to the masculinity of
the tubers.
HISTORY aND
CONSERVaTION
The first British record dates from as long ago
as 1562 when William Turner noted in his
Herball, ‘There are divers kindes of orchis …
one kinde … hath many spottes in the leafe and
is called adder grasse in Northumberland’.
Locally common in many areas, Early
Purple Orchid is the third most widespread
species of orchid in the British Isles (after
Common Spotted and Heath Spotted
Orchids). However, it has vanished from 28%
of its historical range in Britain and 21% in
Ireland, and the decline appears to be ongoing
in Britain. Losses are due to the destruction
or ‘coniferisation’ of woodland and, perhaps
more importantly in recent years, the loss of
permanent grasslands as pastures and meadows
have been ploughed and reseeded. In most of
lowland Britain, away from reserves, the species
is now largely confined to ancient woodland
and to marginal sites, such as road verges and
churchyards, which have escaped agricultural
improvement. In the north and west
overgrazing may be a problem as the species
is tolerant of light grazing only.
Past and present occurrence of Early Purple Orchid in

Atlas).
Britain Ireland
total historical range, 1,971 475

1500-1999
M 24 April, Norfolk. Hazel coppice is the classic habitat. current range 1,416 (50%*) 377 (37%)
Intergeneric hybrids % lost, 1500-1969 17% 17%

‘O. x morioides’, the hybrid with Green- % lost, 1970-1986 11% 4%
winged Orchid, has been recorded rarely and
sporadically in England and Wales. As Green-
winged Orchid is no longer in the genus Orchis,
a new name is needed for this hybrid.
003 systematic.indd 234 29/1/09 12:27:43

The single species in this genus is
found in Europe and Greenland,
with just a toe-hold in North
America, in eastern Canada.
Taxonomy
Sometimes Pseudorchis is united
with the fragrant orchids in the
genus Gymnadenia. It differs in
having a shorter spur, all three
sepals arranged with the petals to
form the hood of the flower (not
just the upper sepal) and a tuber
that is so deeply divided that it may
appear to be several cylindrical
tubers.
Name
The generic name Pseudorchis
derives from the Greek ‘pseud’
meaning false and is a reference to
its relationship to the genus Orchis.
GenuS PSEUDORCHIS
Small white Orchid
004 systematic.indd 235 29/1/09 12:31:31
236 l GENUS PSEUDORCHIS
Small white Orchid Vulnerable, BAP

Pseudorchis albida
Other names: Leucorchis albida, Gymnadenia albida; in North America Pseudorchis straminea
Newfoundland Orchid
This delicate and unassuming orchid is a boreal species whose generally northern distribution
worldwide is reflected in Britain and Ireland. In this respect it is similar to Lesser Twayblade
and Creeping Lady’s-tresses. Like many other northern plants, it has declined drastically in
the southern parts of its range and disappeared from much of England, Wales and Ireland.
Identification
The combination of its small stature, dense
spike of creamy-white flowers and deeply three-
lobed lip is distinctive. The individual flowers
are bell-shaped and very small, just 2-4mm
across (smaller even than the ovary). The spike
is carried on a long stem with a cluster of shiny
green leaves at the base.
Similar species
Creeping Lady’s-tresses is superficially similar
but flowers later in the year, on short, dry
turf (although occasionally on moorland). Its
flowers also have glandular hairs but its lip is
rather different, being spout-shaped rather than
three-lobed.
Irish Lady’s-tresses flowers even later in
the summer than Creeping Lady’s-tresses and
barely overlaps with Small White Orchid. It
has bigger flowers arranged in three columns
around the spike. The lip is also formed into a
spout.
Dense-flowered Orchid is superficially
similar to Small White Orchid, and the
flowering periods may just overlap, but the lip is
very different and the hood is tightly closed.
White-flowered varieties of Pyramidal,
spotted and fragrant orchids have been
mistaken for Small White Orchid, but their
flowers are larger and differ in many other
details.
Habitat
M 6 June, Cumbria. Despite its white flowers, this petite Small White Orchid grows in rough grassland
orchid can be hard to see in long grass.
on poor, well-drained soils, both mildly acidic
O 6 June, Cumbria. and base-rich. It is found on hill pastures, hay
004 systematic.indd 236 29/1/09 12:31:32

SMALL WHITE ORCHID l 237
meadows, road verges, banks, streamsides and

grassy ledges. It will grow in the partial shade
of shrubs and bushes and is sometimes found
on recently burnt moorland among short
Heather (disappearing again once the Heather
regenerates to form a closed community).
Very rarely, the species has been found in oak
woodland on acid soils and on stabilised coastal
dunes. Generally it occurs in the uplands at over
165m above sea level and has been recorded as
high as 500m at Alston Moor in Cumbria and
550m at Ben Chaisteil in Argyll, but it is also
found near sea level in western Scotland.
Flowering period
Late May to mid-July, depending on altitude
and latitude. It is latest at higher altitudes in the
north and earliest in Ireland but is generally at
its best around mid-June. The flowers tend to
wither quickly.
Range
Small White Orchid is commonest and most
widespread in northern and western Scotland,
including the Inner Hebrides and Orkney. It is
now almost absent from the central lowlands
and southern uplands of Scotland, merely
clinging on at a few isolated sites in Dumfries-
shire, Ayrshire and Roxburghshire. The
species is similarly much reduced in northern
England and is now found rather rarely in
1987-99
1970-86
pre 1970
M 7 June, Perth and Kinross.The sepals are whiter than the

petals and lip, which tend to be washed greenish-yellow.
004 systematic.indd 237 29/1/09 12:31:34

over the Urals into northwest Siberia). In

the south it is found in the Pyrenees, Massif
Central, Corsica, Alps, Apennines, Balkans,
Carpathians and Transylvanian Alps east
to Ukraine. Between and around these two
centres of distribution there are scattered
records, e.g. in Bulgaria, northern Greece and
recently from Crete. But, as in the British Isles,
the species has vanished from many lowland
areas and is extinct in Belgium, Luxemburg
and Holland. It is also found in southern
Greenland and in North America, where
it is restricted to a small area in northwest
Newfoundland and western Quebec.
How to find it
Small and inconspicuous, this orchid can be
hard to spot, especially as it is often found
singly or in small numbers. Sites in Scotland
include Keltneyburn (Perth & Kinross), Glen
Cova (Angus) and Feoch Meadows (Ayrshire).
DESCRIPTION
Height: 8-40cm but usually less than 20cm and
M 6 June, Cumbria. The three-lobed lip can only be seen only very exceptionally to 40cm.
on close inspection.
Stem: Greenish, slightly angled towards the tip,
Cumbria, Northumberland, Co. Durham with two or three whitish or brownish sheaths
and mid-west and northwest Yorkshire. In at the base.
Wales it is found at a few scattered sites in Leaves: There are four to six shiny, green, oval
Breconshire, Merionethshire, Denbighshire to oval-lanceolate, keeled sheathing leaves at the
and Caernarvonshire. In Ireland it is similarly base of the stem and one or two narrower and
very scattered, with odd sites in Co. Kerry, more bract-like leaves above them.
Co. Limerick, Co. Clare, Co. Cavan and Co. Spike: Dense, cylindrical and often rather one-
Donegal. There are small clusters of sites in sided, with 20-40 flowers (exceptionally as few
northern Co. Tipperary and central Galway but as ten or as many as 70).
the main concentration lies in the northwest, Bract: Green, lanceolate with a pointed tip, and
in Co. Sligo, Co. Leitrim and Co. Fermanagh. as long as or just longer than the ovary, which
Elsewhere there is another cluster of sites in it clasps.
Co. Tyrone, Co. Derry and Co. Antrim in Ovary: Green, slightly twisted, with three
Northern Ireland. World range: Small White obvious ridges. The ovary is strongly curled over
Orchid has an ‘amphi-Atlantic’ distribution towards the tip so that the flower faces more-
and is found in Europe and northeast North or-less downwards.
America. In Europe it occurs in two discrete Flower: The flowers are small (rather smaller
areas. To the north it is found in Iceland than the ovary and bracts) and very pale, with
(where it is abundant), the Faeroe Islands, the sepals whitish or creamy and the petals, lip
Denmark, Scandinavia (north to northernmost and spur washed more greenish or yellowish.
Norway) and northern Russia (just creeping The sepals are elliptical and blunt-tipped and
004 systematic.indd 238 29/1/09 12:31:36

SMALL WHITE ORCHID l 239
form a loose hood that encloses the similarly taper gradually to a long, pointed tip and are
shaped petals and the column. The lip is short, often deeply divided into several long ‘fingers’
broader than long and deeply three-lobed. The that diverge widely. There are also long, fleshy
central lobe is triangular, usually longer, wider roots that lie horizontally close to the surface
and blunter than the side-lobes, which are of the soil. The roots and the slender tips of the
narrower and more lanceolate in shape. The tubers have a heavy fungal ‘infection’. The first
spur is short (2-3mm), tubular or sack-shaped, aerial stem is reported to appear four years after
blunt-tipped and down-curved and contains germination.
abundant nectar. The flowers have a delicate
scent of vanilla.
Subspecies
Small White Orchid is divided into two
subspecies. The nominate subspecies P. a. albida
is found in the British Isles, central Europe and
lowland Scandinavia. Subspecies straminea,
with larger and yellower flowers, is found in the
mountains of central Europe and Scandinavia,
Faeroe Islands, Iceland, Greenland and North
America. The latter is often treated as a distinct
species, Pseudorchis straminea.
British plants belong to the nominate
subspecies, P. a. albida, which in turn is
divided into two varieties.
Var. albida is found on more acid soils and has
the lateral lobes of the lip clearly shorter than
the central lobe.
Var. tricuspis favours calcareous soils and has
the lateral lobes almost as long as the central
lobe. The distribution and abundance of the
two varieties in the British Isles has not been
studied.
BIOLOGY
The flowers produce nectar and are visited by
butterflies, day-flying moths and solitary bees.
The specific pollinator has not been identified,
but the narrow entrance to the spur suggests
that it may be butterflies. Some self-pollination
also occurs, as the pollinia eventually fall onto
the stigma if an insect has not removed them.
Seed may be set by over 90% of flowers.
M 6 June, Cumbria.The flowers are small – smaller than
Development and growth the ovary. The specific pollinatior is unknown but seed-set
The aerial stem grows from paired tubers that is good.
004 systematic.indd 239 29/1/09 12:31:37

Intergeneric hybrids Berwickshire, the Lothians, Fife and

X Pseudadenia schweinfurthii, the hybrid with Stirlingshire. It is extinct in Shetland, where
fragrant orchid (presumably Heath Fragrant there has only been one record, from Bressay
Orchid), has been recorded from several places prior to 1845.
in Yorkshire and Scotland, where it is fairly In England, Small White Orchid is now
frequent in the northwest. rare even in Cumbria, a former stronghold that
X Pseudorhiza bruniana, the hybrid with still holds the bulk of the English populations,
Heath Spotted Orchid, was recorded from although most colonies comprise just a handful
Orkney in 1977 and Skye in 1994. of plants. It is now extinct in Kent (with just
one old record, from near Lyminge), East
and West Sussex (where present until at least
The specific name albida means ‘white’.
1913, with three localities in West Sussex
HISTORY AND and ten in East Sussex, notably Ashdown
Forest), Gloucestershire (last recorded 1899),
CONSERVATION Herefordshire, Staffordshire, Shropshire
The first British record was in 1670 when it was (last recorded 1856), Derbyshire, Cheshire,
discovered by John Ray on Mount Snowdon Lancashire, southwest and northeast Yorkshire
in North Wales: ‘This we found on the back (where it has recently gone from the North
of Snowdon-hill by the way leading from York Moors National Park). The situation
Llanberis to Carnarvan’ (Catalogus Plantarum in Wales is probably even worse, as numbers
Anglicum et Insularum adjecentium). there have collapsed in recent years, with no
Classified in Britain as Vulnerable, the records from Monmouthshire, Glamorganshire,
species is specially protected in Northern Carmarthenshire, Cardiganshire, Montgomery-
Ireland under Schedule 8 of the 1985 Wildlife shire and Flintshire. In total, Small White
Order (NI) and in Eire under the Flora Orchid has been lost from 66% of the historical
(Protection) Order. range in Britain. In Ireland there has also been
Past and present occurrence of Small-white Orchid in Brit-
a very sharp drop in numbers. It has been
ain and Ireland (based on presence or absence in 10km lost from 70% of the historical range and is
squares of the National Grid; data from the New Atlas). extinct in Co. Cork, Co. Waterford, Laois,
Co. Wicklow, Co. Dublin, Co. Mayo, Co.
Britain Ireland
Monaghan and Co. Down.
total historical range, 385 110 Populations tend to be small and scattered
1500-1999
and even in Scotland many vice-counties have
current range 132 (4.6%*) 33 (3.3%*) just one or two sites for the species. Colonies
% lost, 1500-1969 52% 63.5% were lost throughout the 20th century due
% lost, 1970-1986 13.5% 6.5%
to habitat destruction, forestry, agricultural
improvement and overgrazing. As with many
% lost, total 65.5% 70%
grassland orchids it requires both a short
* current range as a % of the total number of 10km squares sward and areas of bare soil so that seedlings
can become established. It therefore needs a
Small White Orchid is commonest in certain level of grazing to thrive; too little and
northern and western Scotland and becomes it is swamped by coarse grasses and scrub, too
rarer further south; in the remainder of much and it can never flower and is eventually
Scotland it has almost vanished from the area eliminated.
south of a line from the Clyde to Aberdeen
and has not been recorded recently from
Kirkcudbrightshire, Lanarkshire, Selkirkshire, P 14 June, Kent. Greater Butterfly Orchid.
004 systematic.indd 240 29/1/09 12:31:38

GENUS
PLATANTHERA
BUTTERFLY
ORCHIDS
004 systematic.indd 241 12/2/09 17:01:41

242 l GENUS PLATANTHERA
Distribution separate plants as the connecting stem dies off

There are 80-200 species in this genus. They are in the autumn.
scattered almost worldwide, although most are
Name
found in the temperate regions of the Northern
The generic name Platanthera derives from
Hemisphere, with eight in Europe and two in
the Greek plat meaning ‘broad’, ‘wide’ or ‘flat’
the British Isles. and antherus meaning ‘flowery’ or ‘anther’. It is
Floral structures usually taken to mean ‘flat-anthers’, a reference
The column is short and there are two pollinia. to the shape of the anther, which has a hollow
They taper into slender caudicles (stalks), each at its base to hold the pollinia.
attached to one of the two viscidia, which are
naked (there is no pouch-like bursicle). There is
column
a single flat stigmatic zone.
Growth pattern
pollinium
The aerial stem grows from a pair of spindle-
shaped tubers which taper to a long, narrow
point, and there are also a few slender roots
that spread into the surface layers of the soil. As
with the genus Orchis, one of the tubers forms stigma
the summer before and has supplied the current
year’s growth, while a new tuber, complete
with a bud ready for next year’s flower spike,
develops beside it.
Fungal partners
All the roots are ‘infected’ with fungi. mouth of the spur
viscidium
Development from seed lip
Seed probably germinates in the spring, usually Greater Butterfly Orchid
not far below the surface of the soil. The
protocorm produces a large bud in the first column
autumn and this develops into a leafy shoot
the following spring. Following the emergence
of this shoot, a root is produced. These die off,
presumably in the late summer, and the seedling pollinium
overwinters as a short, two-segmented rhizome.
Similarly, a leafy shoot and one or two roots
are produced the following summer, and it is
not until the third year that the first tuber is stigma
produced and the protocorm finally vanishes. viscidium

A full-sized tuber appears after the fourth
summer. (This chronology is based on work mouth of
lip
in the 1920s and, as with many other orchids, the spur
development may be rather faster.)
Lesser Butterfly Orchid
Additional tubers may be formed at the base M Columns of butterfly orchids showing position of
of the aerial stem. These will go on to form pollinia (sepals and petals removed).
004 systematic.indd 242 29/1/09 12:31:41

LESSER BUTTERFLY ORCHID l 243
leSSer Butterfly Orchid Vulnerable, BAP

Platanthera bifolia
The more delicate and daintier of the two butterfly orchids, this species has a northerly
and westerly bias to its distribution and has vanished from much of central and eastern
England. It is found on rough pastures, damp heathland, bogs and, rather less frequently, in
woodland. The delicate flowers are highly scented at night in order to attract hawkmoths;
as the moth sips nectar from the orchid’s long spur the pollinia are glued to its proboscis and
it becomes the unwitting servant of the orchid.
Identification
The two butterfly orchids are distinctive. They
have two oval, shiny leaves, placed at the base
of the stem and often hidden away in the grass,
and exquisite white flowers. The lip of the
flower is long, narrow and undivided, and the
spur is extremely long and slender and projects
prominently to the rear. There are two subtly
different forms of Lesser Butterfly Orchid,
‘heathland’ and ‘woodland’ (see Variation and
varieties).
Similar species
Greater Butterfly Orchid can easily be separated
by the size and shape of the pollinia. In Lesser
Butterfly Orchid the two pollinia are placed
close together and parallel, while in Greater
Butterfly Orchid the bases of the pollinia are
well separated and they lean inwards towards
the tip. Other differences between the two
species are subtler and less consistent. Lesser
Butterfly Orchid is generally smaller and
daintier (averaging around two-thirds the size
in most dimensions), with fewer, smaller flowers
in a narrower spike; the mouth of the spur is
smaller and the spur tends to be straighter.
Habitat
The ‘heathland’ form of Lesser Butterfly Orchid
is by far the commoner and grows on heathland
in the south and east and on moorland and
damp pastures in the north and west. On both
heathland and moorland it usually occurs in
the damper areas, and these are frequently
P 19 June, Norfolk. A robust plant with numerous flow-

ers in a compact spike.
004 systematic.indd 243 29/1/09 12:31:43

open deciduous woodland and scrub, often

beechwoods, on calcareous soils, especially
‘clay-with-flints’. It can also occur on open chalk
downland and among Bracken on the less acid
areas of dry grassy heaths. Lesser Butterfly
Orchid occurs up to 365m above sea level
(Glenfeshie, Inverness-shire).
Flowering period
Late May and June in woodland populations,
June to July for heathland forms. On average it
flowers latest in the north of Scotland.
Range
Lesser Butterfly Orchid occurs throughout
Britain and Ireland, including the Isle of Man,
Inner and Outer Hebrides and Orkney. There
is, however, a distinct western and northern
bias to the distribution, and the species was
always scattered and local in much of England,
eastern Scotland and in southern and southeast
Ireland. It is now extinct in most of East Anglia,
the Midlands and northern England, although
it is still widespread in Cumbria. World range:
Widespread in Europe and Asia and also found
in North Africa. In Europe it occurs north
to the Faeroe Islands and c.70°N in northern
Scandinavia. It ranges south to Spain, Italy,
northern Greece, the Crimea and Caucasus.
It is found on the Balearic Islands, Corsica,
Sardinia, Sicily and the Aegean islands. In
1987-99
1970-86
pre 1970
M 20 June, Norfolk. A rather small, few-flowered plant in
the early morning dew.
P 19 June, Norfolk.
marked out by the dusty grey foliage of Cross-

leaved Heath. It can often be found around
the margins of valley bogs or growing on drier
tussocks and other raised areas amidst sodden,
boggy ground. It grows on neutral or mildly
acidic soils; on moorland and around bogs it
favours the areas where springs and flushes
buffer the general acidity to some extent. The
‘woodland’ form is much scarcer and largely
restricted to southern England. It is found in
004 systematic.indd 244 29/1/09 12:31:45

004 systematic.indd 245 29/1/09 12:31:48
North Africa it is found in Algeria and Tunisia. Bract: Green, narrow, pointed and slightly
In Asia it occurs in northwest and northeast shorter than the ovary.
Turkey and northern Iran. The range extends Ovary: Pale green, long, narrow, clearly ribbed
across southern Siberia to at least Lake Baikal. and twisted and also curled into a C-shape to
hold the flowers pointing outwards and slightly
How to find it
downwards.
Often rather small and slender, this can be a
Flower: Whitish, with a long, strap-shaped
hard plant to find wherever the vegetation is tall.
lip and extremely long spur. The sepals are
However, in other areas it can be very obvious,
white, washed greenish towards the tip and
especially when found in large numbers.
bluntly lanceolate in shape. The upper sepal
DESCRIPTION is slightly broader and more triangular and
Height: 15-30cm, occasionally to 45cm. may be bent upwards at the tip. The petals are
Stem: Pale green, more-or-less triangular creamy and variably washed greenish. They are
and ribbed towards the tip, with two or three smaller, narrower and more strap-shaped than
whitish or brownish sheaths at the extreme base. the sepals. The upper sepal and petals form a
Leaves: The two pale green, slightly shiny or loose hood over the column. The lateral sepals
‘greasy’ looking leaves are held opposite each are held spreading and slightly drooped. The
other, one just above the other, at the base of lip is creamy, sometimes greener towards the
the stem. They are variable in shape, from oval tip, narrow, strap-shaped and 6-12mm long. It
to narrower and more strap-shaped, and taper projects forwards and downwards. The spur is
at the base to a whitish, winged stalk. There long and slender (1mm wide x 13-23mm long,
are also one to five small, lanceolate, bract-like occasionally to 27mm), sometimes slightly
leaves higher on the stem. curved and it may be washed with green.
Spike: Variable, the five to 25 flowers The pollinia are whitish, about 2mm tall and
(occasionally more) form a compact cylindrical lie parallel, 1mm apart, at the front of the
spike in moorland plants but are more widely column. The flowers emit a heavy, sweet scent,
spaced in woodland plants. sometimes likened to carnations, especially
at night.
Subspecies
None.
In both Greater and Lesser Butterfly Orchids
plants growing in the open tend to be shorter
and more compact than those in shaded
woodland localities. In Lesser Butterfly Orchid
the ‘heathland’ form has leaves that are egg-
shaped and a relatively dense flower spike
whereas the ‘woodland’ form has narrower
and less pointed, more tongue-shaped leaves.
The heathland form is found on acid soils
throughout Britain and Ireland whereas the
woodland form is most frequent in southern
England, often growing on calcareous soils,
and becomes rare to the north, although it has
M 20 June, Norfolk.The pollinia lie parallel to each other, been found in southern Scotland. Intermediates
above the mouth of the spur. occur and the differences between the two
004 systematic.indd 246 29/1/09 12:31:49

LESSER BUTTERFLY ORCHID l 247
forms may be caused solely by the different

environments in which the plants are growing.
Var. trifolia has three main leaves rather than
two. It is not rare.
Var. quadrifolia has four leaves.
BIOLOGY
The flowers are pollinated by insects attracted
by the copious nectar in the spur (nectar can
be seen filling the tip of the spur, which is
translucent). However, the nectar can only
be reached by an insect with a suitably long
proboscis and Lesser Butterfly Orchid is
pollinated by night-flying moths, especially
hawkmoths such as Elephant, Small Elephant
and Pine Hawkmoths. The flowers’ scent,
which is particularly pungent around dusk,
and white coloration (they almost ‘glow
in the dark’) help the moths to find them.
Hawkmoths hover in front of the flower to
feed, resting their forelegs on the lateral sepals,
whereas other moths land on the flower itself.
The two pollinia, which are relatively small,
extremely short-stalked and placed parallel
to each other at the mouth of the spur, have
small sticky pads, the viscidia, at their base. As
the moth inserts its proboscis into the spur,
the viscidia glue the pollinia to it. The moth
continues on its way, visiting other flowers,
and after a short while the pollinia rotate M 19 June, Norfolk. The flowers are pollinated by moths,
forwards and in this new position will make especially hawkmoths, and a long proboscis is required to
access the nectar in the spur, visible here.
contact with the stigma in the next flower
visited. The mechanism is fairly effective and
seed-set is moderate to good. length. The differences in habitat, flowering time
and flower structure make hybridisation less
Development and growth likely than it seems; in Lesser Butterfly Orchid
Grows from a pair of underground tubers.
the pollinia are attached to the proboscis of
There is no information on the period between
a visiting moth, whereas in Greater Butterfly
germination and flowering in the wild but in
Orchid they are typically attached to the eyes.
cultivation plants may flower in three or
In these different positions the chances of the
four years.
pollinia making contact with the stigma of the
Hybrids ‘wrong’ species are greatly reduced. Hybrids
P. x hybrida, the hybrid with Greater Butterfly have the pollinia in an intermediate position,
Orchid, has been reported from scattered reducing the effectiveness of the pollination
localities but is rare; hybrids are intermediate mechanism and making such hybrids less likely
in the positioning of the pollinia and in spur to reproduce successfully.
004 systematic.indd 247 29/1/09 12:31:50

Past and present occurrence of Lesser Butterfly Orchid

Atlas).
Britain Ireland

1500-1999
current range 342 (12%*) 159 (16%*)
% lost, 1500-1969 48% 36%

% lost, 1970-1986 16% 12.5%
% lost, total 64% 48.5%
range in Britain and 48.5% in Ireland. It is

classified as Vulnerable and is one of the most
rapidly declining wild flowers in the British
Isles. It has vanished from much of southern
and eastern England and is now extinct in
Essex, Suffolk, Hertfordshire, Middlesex,
Berkshire, Northamptonshire, Worcestershire,
Lincolnshire, Nottinghamshire, Derbyshire,
Cheshire, Denbighshire, Roxburghshire and
the Lothians. It is also seriously reduced in
many other areas. For example, in Norfolk it
was ‘locally common’ on wet heaths in 1914 and
‘fairly frequent on bog and wet heath, always on
acid soils’ in 1968, when 12 localities were listed,
M 13 July, Norfolk.The parallel pollinia are clearly visible. but by 2008 was present at only two sites, with
just a handful of plants at each. It has now gone
Name and classification altogether from some classic heathland area in
The specific name bifolia means ‘two leaves’. The southern England such as Woolmer Forest in
two butterfly orchids, although easily separated Hampshire and Ashdown Forest in Sussex.
in the field, are extremely close genetically. Some losses are due to the outright
This suggests that they have only very recently destruction of heathland, with both agriculture
separated into two species. and urban development being responsible. Even
where heathland remains, the losses continue,
HISTORY AND probably due to the lack of grazing and resulting
CONSERVATION transformation of heathland into scrub and
The first reference to the Lesser Butterfly woodland. Notably, Lesser Butterfly Orchid is
Orchid was in the second edition of John Ray’s still common in the New Forest, which continues
Synopsis Methodica Stirpium Britannicarum to support large numbers of grazing animals.
(‘A methodical synopsis of British plants’), Away from heathland sites the ‘improvement’ of
published in 1696, which simply noted its pastures and hay meadows and the clearance or
presence i n ‘pastures’. ‘coniferisation’ of woodland are responsible for the
Lesser Butterfly Orchid has declined decline. Paradoxically, overgrazing is a problem in
greatly, with a loss of 64% of the historical the uplands of western and northern Britain.
004 systematic.indd 248 29/1/09 12:31:51

GREATER BUTTERFLY ORCHID l 249
Greater Butterfly Orchid

Platanthera chlorantha Near Threatened
Found locally throughout mainland Britain and Ireland but with a distinctly southern
bias, this species favours both grassland and woodland. The exquisite white flowers, often
held on a tall, stately spike, have evolved to ‘glow in the dark’ as the pollinators are night-
flying moths.
Identification
The two butterfly orchids are distinctive, with
a pair of oval, shiny green leaves at the base of
the stem and an open spike of beautiful ‘waxy’
white or greenish-white flowers. The lip is long,
narrow and undivided, and the extremely
long slender spur projects backwards from
the rear of the flower across the width of the
flower spike.
Similar species
Lesser Butterfly Orchid is distinguished by
the shape and position of its pollinia, which lie
close together and are parallel for their entire
length. In Greater Butterfly Orchid the pollinia
are well-separated at the base but lean inwards
so that their tips almost touch. There are other,
subtler differences between the two species
but none of these can be taken as diagnostic.
Greater Butterfly Orchid is on average taller
and sturdier, with a broader flower spike (the
ovaries are longer, holding the flowers further
away from the stem). It has greener flowers with
a larger and more obvious mouth to the spur,
and the spur itself is usually slightly expanded
at the tip.
Habitat
Rather variable but it almost always grows on
calcareous soils: chalk, limestone and base-
rich clays. It is found in deciduous woodland
(where it is strongly associated with ancient
woodland) and has a preference for hazel
coppice. It grows in light, dappled shade
and is usually found in the more open areas
around the edge of a wood and in clearings and
P 14 June, Kent. In woodland the spike is often very
attenuated.
004 systematic.indd 249 29/1/09 12:31:53

populations may be relicts of deciduous

woodland previously on the site.
In the north and west of Britain and Ireland
Greater Butterfly Orchid is most frequently
found in grassland such as old pastures and
hay meadows, again usually on calcareous
soils; it has been found on rare occasions on
mildly acidic soils on moorland, wet heath
and pastures but is clearly less tolerant of acid
conditions than Lesser Butterfly Orchid. It was
probably frequent in pastures and meadows in
southern England prior to their ‘improvement’
but in the south, away from woodland sites, is
now only likely to be found on chalk grassland,
often in long grass with variable amounts of
scrub. It is sometimes also found in calcium-
rich dune slacks and on railway embankments.
Occurs up to 460m above sea level (Harwood
Dale, Northumberland).
Flowering period
Late May to late July, being earliest in the south,
where it is sometimes in flower from mid, or
even early, May. Usually it is at its best in early
June in much of England but it may not be
in full flower in parts of Scotland until July.
Typically it flowers one or two weeks earlier
than Lesser Butterfly Orchid.
1987-99
1970-86
pre 1970
M 15 June, Kent. Greater Butterfly Orchid will grow on

open grassland as well as in woodland. This plant has
three leaves rather than the usual two.
along rides. It is tolerant of both dry and wet

conditions but in woodland on chalk may have
some preference for the heavier and wetter soils
found at the foot of slopes. Greater Butterfly Range
Orchid has occasionally been found growing Found throughout Britain and Ireland except
in rides through conifer plantations but such Orkney and Shetland, although rather scattered
004 systematic.indd 250 29/1/09 12:31:55

in the southern half of Ireland and absent Bract: Green, narrow, pointed and as long as
from large areas of the Midlands, northern the ovary.
England, southern and eastern Scotland and Ovary: Pale green, long, narrow, clearly six-
the Outer Hebrides. The centre of gravity ribbed and twisted. It is also curved to hold
of the distribution lies in southern England the flowers pointing outwards and a little
and Wales, in contrast to the northern and downwards.
western bias shown by Lesser Butterfly Orchid. Flower: White, washed green, with a long
World range: Almost confined to Europe, strap-shaped lip and extremely long curved
occurring north to c. 63°N in Scandinavia, spur. The sepals are white, washed greenish
southern Finland and the Baltic States. It towards the tip; the lateral sepals are oval-
ranges south to the Mediterranean, Crimea and triangular, asymmetrical and even sickle-shaped
Caucasus, including Corsica, Sicily, the Aegean whereas the upper sepal is rather shorter,
Islands, Cyprus and possibly Sardinia. Occurs broader and more triangular or heart-shaped.
eastwards to central European Russia. It is also The petals are white, variably washed greenish
found in North Africa in Tunisia and possibly
Morocco and in Turkey, northeast Syria and
northwest Iran.
How to find it
Woodland plants are often easy to spot,
being rather tall, but those in the open
among grass and scrub can be well hidden.
A large proportion of most populations are
non-flowering, and when heavily shaded in
overgrown coppice or very dark woods plants
can remain in a vegetative state for decades.
They ‘reappear’ and flower again following
coppicing, tree-falls or other changes that let
more light in. In woods with a regular coppice-
cycle, flowering is most prolific two or three
years after coppicing.
DESCRIPTION
Height: 20-40cm, sometimes to 65cm.
Stem: Pale green, more-or-less triangular
and ribbed towards the tip, with one to three
brownish sheaths at the extreme base.
Leaves: Two, oval to elliptical, keeled, pale
green (often slightly bluish) and rather shiny.
They lie opposite each other at the base of the
stem, one just above the other, and are held
variably erect. There are also one to six small,
lanceolate, bract-like leaves higher on the stem.
Spike: Variable. The ten to 30 flowers
(occasionally up to 40) form a loose, open spike
in woodland plants, but in full sun the spike M 14 June, Kent. The flower spike is extraordinarily
tends to be more compact. graceful.
004 systematic.indd 251 29/1/09 12:31:56

252 l GENUS PLATANTHERA.
in the open tend to be shorter and more

compact and to have shorter leaves than those
in shaded woodland localities. It seems likely
that these differences are produced by the
local environment rather than by any genetic
difference. Plants may occasionally have just
one leaf or sometimes three or four.
BIOLOGY
The pollination mechanism is very similar to
that of Lesser Butterfly Orchid, and night-
flying moths are the primary pollinators. But
although hawkmoths are involved, members
of the large family of Noctuid moths are
more important as pollinators. In the Greater
Butterfly Orchid the sticky viscidia at the base
of the pollinia face inwards on either side of,
M 14 June, Kent. The extraordinarily long spur contains and just above, the entrance to the spur. Due to
nectar that can only be reached by insects with an equally this more widely spread position, the pollinia
long proboscis.
usually become attached to the large compound
eyes of the visiting moth rather than its
and are shorter, narrower and more strap-
proboscis. The mechanism is effective and seed
shaped than the sepals. The upper sepal and
is set in 70-90% of flowers.
petals form a loose hood over the column, and
the lateral sepals are spread horizontally and
are often a little twisted or wavy-edged. The
lip is creamy but becomes greener towards the
tip and is narrow and strap-shaped, 10-16mm
long and projects forwards and downwards. The
spur is long and slender, 19-35mm long x 1mm
wide, broadens towards the tip and is often
strongly curved; it is washed green. The pollinia
are conspicuously yellow and relatively large,
3-4mm tall including the long caudicle (stalk).
The viscidia at the base of the pollinia lie about
4mm apart on either side of the foot of the
column, and the pollinia lean inwards towards
each other. The flowers emit a heavy scent,
especially at night, and it is said that people with
a sensitive nose can smell them from several
hundred metres on a still summer’s evening.
Subspecies
None.
M 8 July, Perth and Kinross. The lip is longer than wide
Variation and varieties and often only obscurely lobed, and the ‘wings’ are oval,
As in Lesser Butterfly Orchid, plants growing pointed and usually held more or less drooped.
004 systematic.indd 252 29/1/09 12:31:59

Vegetative reproduction can occur via the meadows, have been responsible for the decline.
formation of additional tubers but is of little The species will also ‘vanish’ from woodland if it
importance in the dynamics of a population. becomes too intensely shaded but may reappear
if the canopy is opened again.
The process of development from seed to
flowering plant is very similar to Lesser
Butterfly Orchid. The adult plant is reported
to be entirely independent of fungi, but this
seems highly unlikely given that it can become
‘dormant’ in heavy shade for long periods.
Hybrids
See Lesser Butterfly Orchid.
The specific name chlorantha means ‘green
flowers’. The two butterfly orchids, although
readily separated by the shape of the pollinia,
are genetically almost indistinguishable,
suggesting that they have only very recently
separated into two species.
HISTORY AND
CONSERVATION
The first British record dates from 1597 when
John Gerard noted this species in his Herball:
‘…in the wood belonging to a worshipfull
gentleman of Kent named Master Sedley of
Southfleete.’
Past and present occurrence of Greater Butterfly Orchid in

Britain and Ireland (based on presence or absence in 10km
Britain Ireland

1500-1999
current range 626 (22%*) 117 (11.6%*)
% lost, 1500-1969 32% 41.5%

% lost, 1970-1986 14% 12%
% lost, total 46% 53.5%
It has been lost from 46% of the historical

range in England and 53.5% in Ireland and is
classified as Near Threatened. The destruction
and coniferisation of woodland, as well as M 28 May, Norfolk. The spike may be relatively few-
agricultural improvements to pastures and hay flowered and can be inconspicuous.
004 systematic.indd 253 29/1/09 12:32:01

GENUS GYMNADENIA
FRAGRANT ORCHIDS
004 systematic.indd 254 12/2/09 17:03:12

Distribution Development from seed
Europe and Asia, with about 14 species, Early observations suggested that seeds
five of which occur in Europe and three in probably germinate in the spring. The
Britain. Until recently all three were treated resulting protocorm produces roots in the
as subspecies of a single species, the ‘Fragrant second autumn, the first leaves in the third
Orchid’. spring (i.e. when it is two years old) and the
first tuber the following season. Given the dry
Floral structures
habitats, however, in which fragrant orchids
The column is short and erect with a long
often grow, it seems unlikely that protocorms
rostellum that projects forward between the
could survive for over two years, and the
two viscidia. The two club-shaped pollinia
period prior to the development of the first
each narrow into a caudicle (stalk) that is
tuber may be very much shorter. The sequence
attached to one of the two long, narrow
protocorm-root-aerial stem-tuber is shared
viscidia, but there is no bursicle (the flap or
with other genera, including some of the
pouch-like structure found in many orchids
Orchis and Dactylorhiza orchids.
which covers and protects the viscidia). The
two stigmas lie on the side-lobes of Vegetative reproduction
the column. Additional tubers may be formed at the base
of the aerial stem. These will go on to form
Growth pattern
separate plants as the connecting stem dies off
The aerial shoot grows from a pair of
in the autumn.
flattened, deeply divided tapering tubers.
Roots develop in the autumn and penetrate Name
the upper layers of the soil. The roots and the The generic name Gymnadenia originates from
root-like tips of the tubers are ‘infected’ with the Greek gymnos ‘naked’ and aden ‘gland’ and
fungi. The annual cycle of growth and the means quite literally ‘naked-gland’, a reference to
replacement of tubers are similar to those of the lack of a bursicle (see Floral structures).
the genus Orchis (see p.200).
004 systematic.indd 255 12/2/09 17:03:17

256 l GENUS GYMNADENIA
cOmmOn fraGrant Orchid

Gymnadenia conopsea
Other names: Chalk Fragrant Orchid

This attractive orchid is found in species-rich grassland on chalk and limestone soils
throughout England and Wales but is rather rare in Scotland and Ireland. It sometimes
occurs in large numbers, and the fragrance emitted by a big colony can be quite overpowering,
especially in the evening.
Identification Intermediates also occur, and it is best to

The fragrant orchids are easy to separate from examine several plants in a population to gain
other orchids. The pink flowers are held in a an overall impression. The criteria used to
tall, narrow, spire-like spike. Each flower has separate the three fragrant orchids are still
two ‘wings’ (the lateral sepals) that are held being developed, and some plants or even whole
roughly horizontally, a flat, three-lobed lip and a populations may not yet be identifiable.
long, slender, down-curved spur. In Common Fragrant Orchid the flowers are
medium sized, the lip is about as wide as long,
Similar species
lacks ‘shoulders’ and is distinctly three-lobed.
Pyramidal Orchid has flowers that are a similar
The central lobe is the longest. The ‘wings’ are
colour, size and shape to those of Common
angled downwards and are pointed, narrow and
Fragrant Orchid and also have a long slender
parallel-sided (with the lateral sepals rolled into
spur. Pyramidal Orchid has, however, an
a tube to produce this linear shape). See also
obviously shorter, more conical and more
p.258.
closely-packed flower spike. Its flowers are a
deeper pink with a more markedly three-lobed Habitat
lip that has two diagnostic parallel raised ridges Found in dry, species-rich grassland on
at the base. Pyramidal Orchid also flowers later calcareous soils, mostly chalk downland in
in the summer, and there is usually relatively the south and limestone pastures in northern
little overlap between the two species. England but sometimes also on stabilised
Fragrant orchids are difficult to separate dunes, road verges, railway banks and in old
from each other, although they are usually quarries on suitable calcium-rich soils. It
found in distinct habitats. Common Fragrant occasionally occurs in base-rich fens (together
Orchid favours old, species-rich grassland on with Marsh Fragrant Orchid) or on alkaline
chalk or limestone and is just occasionally Leblanc waste in Lancashire. Recorded up to
found in fens. Its scent is sickly-sweet but has 365m above sea level (near Whitely Shield,
a slightly acid, rancid, or musty background. Northumberland).
Its main flowering period is June and early July. Flowering period
However, even in its preferred dry grassland Late May (sometimes mid-May) to late July,
habitat, Marsh Fragrant Orchid or Heath but most are in flower during June.
Fragrant Orchid could also occur, and a
careful examination of the flower is necessary Range
to be reasonably certain of the identification. Widespread in England, although confined
to areas with suitable habitat, such as
O 14 June, Kent. At some sites Common Fragrant Orchid the North and South Downs, the chalk
occurs by the thousand.
districts of Hampshire, the Berkshire
P 14 June, Kent. Downs and Chilterns, Dorset, and the
004 systematic.indd 256 29/1/09 12:32:06

COMMON FRAGRANT ORCHID l 257
004 systematic.indd 257 29/1/09 12:32:09

Separation of Common, Marsh and Heath Fragrant Orchids.
Common Marsh Heath

Fragrant Orchid Fragrant Orchid Fragrant Orchid
Width of florets* 10-11mm (7-13mm) 11-13mm (10-14.5mm) 8-10mm (7-12mm)
Lip: width 5.5-6.5cm (4.5-7mm) 6.5-7mm (5.5-8mm) 3.5-4mm (3-5mm)
Lip: length 5-6mm (4-6.5mm) 3.5-4mm (3-4.5mm) 4-4.5mm (3.5-5mm)
Lip: shape fractionally broader much broader longer than
than long than long broad
Lip: lobes distinctly 3-lobed, distinctly 3-lobed, obscurely lobed,
central lobe longest side-lobes largest longer than wide
Lip: shoulders** absent distinct narrow
Lateral sepals: width c. 1mm c. 1mm c. 2mm
Lateral sepals: length c. 5-6mm c. 6-7mm c. 4-5mm
Lateral sepals: shape linear, pointed at tip linear, blunt at tip oval-lanceolate,
pointed at tip
Lateral sepals: deflexed at c. 30° horizontal deflexed
position
Length of spur 12-14mm (11-17mm) 14-16mm (13-17mm) 11-14mm (8-15mm)
Height 20-40cm, sometimes 30-60cm, less in 15-25cm, rarely
more dry habitats more
Fragrance sickly-sweet with ‘acid’ spicy sweet, clove- spicy sweet,
or ‘rancid’ overtones like; no acid overtones recalling cloves
Flowering period early June-mid July early July-mid August June-August
Habitat chalk & limestone fens; rarely chalk mildly acidic to
grassland grassland on north- base-rich grassland;
facing slopes base-rich heathland
flushes; very rarely
chalk grassland
Largely after Francis Rose in Rich & Jermy (1998).
* The width of the floret is measured from tip to tip of the lateral sepals
** The ‘shoulders’ are formed by the abrupt angle at the base of the lip where it narrows towards the mouth of the spur.
limestone areas of the Cotswolds, Mendip

1987-99
Hills, Northamptonshire, Lincolnshire, the 1970-86
Derbyshire and Yorkshire Dales, Cumbria and pre 1970
Co. Durham. Widespread in Wales, but in Common, Heath

and Marsh
Scotland it is rare and local and information Fragrant Orchids
combined
about its distribution is incomplete; it may
have been recorded from Dumfries-shire,
Roxburghshire, Banffshire and a few areas in
the far north, in Sutherland, Caithness and
the Outer Hebrides. Its status in Ireland is
also obscure due to confusion with the other
fragrant orchids and with hybrids between
them. It may be most frequent in the central
counties, but it is probably scarce or rare.
World range: The ‘Fragrant Orchid’, i.e. an
aggregate of all three species, is found in Europe
004 systematic.indd 258 29/1/09 12:32:10

and Asia east to Japan and Korea. In Europe

it ranges north to northernmost Scandinavia
and south to the Mediterranean (although it
avoids the Mediterranean lowlands), Turkey
and the Caucasus. Further east it occurs in
the Himalayas and in central China. Records
seldom distinguish the three species of fragrant
orchid, and their exact ranges have yet to be
defined. ‘Fragrant Orchid’ has been found once
in North America, in Connecticut in 1887.
How to find it
This is usually an easy species to find at suitable
sites, now often reserves or SSSIs. It sometimes
appears in large numbers but the number of
flowering spikes can vary widely from year
to year.
DESCRIPTION
Height: 10-60cm but usually15-30cm and
seldom over 40cm.
Stem: Green, becoming more purplish towards
the flower spike.
Leaves: Mid-green. The three to five narrow,
strap-shaped basal leaves, keeled and with
pointed tips, are held loosely erect. These grade
into two or three narrow, lanceolate, bract-like
leaves higher on the stem.
Spike: More-or-less cylindrical and moderately
densely packed with 20-50 flowers, rarely more.
The spike becomes looser as more flowers open.
Bract: Green, sometimes tinged purple, strap-
shaped, narrowing around the mid-point into
a finer pointed tip and roughly as long as
the ovary. M 9 June, Northamptonshire. A full, densely-flowered spike.
Ovary: Green, variably washed purple, long,
narrow, prominently three-ribbed and twisted. the lateral sepals are held to the side, about 30°
Flower: Pink with a hint of purple, varying below the horizontal and pressed backwards.
in the exact shade. The sepals are elongated- The lip is flat and about as wide as long, with
oval in shape, the lateral sepals being slightly three well-developed, rounded terminal lobes;
irregular but with their upper and lower the central lobe is longer than the lateral lobes
margins rolled backwards so that face-on they and often broader. The spur is a darker, more
appear as parallel-sided oblongs with a short purplish-pink, very slender, down-curved and
pointed tip. The petals are a little shorter, more rather long, around twice the length of the ovary.
oval-triangular in shape and asymmetrical, It is filled with nectar and this can sometimes
with one side squared off. The upper sepal and be seen through its semi-translucent walls. The
petals form a hood over the column, whereas flowers have a strong sickly-sweet scent.
004 systematic.indd 259 29/1/09 12:32:12

Subspecies
None.
Var. albiflora has white flowers and is fairly
frequent.
Var. crenulata has broad lateral lobes to the
lip which are narrowly serrated at the edges.
It is rare.
BIOLOGY
Nectar is produced in the bottom of the
long spur, and only insects with a sufficiently
long proboscis can reach this. Pollinators
include butterflies and both day- and night-
flying moths, including Large Skipper, Six-
spot Burnet and hawkmoths. Night-flying
moths may, however, be the most important
pollinators; the scent becomes more pungent
towards dusk, and, with its white flowers, var.
albiflora may only be attractive to nocturnal
moths. As the visiting insect advances to sip
nectar from the spur, the pollinia are fixed by
their sticky viscidia (which lie just above the
mouth of the spur) to the insect’s proboscis.
The pollinia are then carried to another flower
or another plant, having in the meantime swung
forward into a position ready to make contact
with the stigma. Pollination is very efficient and
seed is set in large quantities.
Vegetative reproduction may also occur via
the production of additional tubers.
The interval between germination and flowering
is usually around five years but may be as short
as three.
Hybrids
Opposite is a list of all the hybrids listed
officially for ‘Fragrant Orchid’ by Stace (2004),
which does not separate the three fragrant
orchids into distinct species. The names of these
hybrids may well need to be revised to reflect the
new status of the fragrant orchid parent:
O 14 June, Kent. The white-flowered variety albiflora is

fairly frequent.
004 systematic.indd 260 29/1/09 12:32:14

P 9 June, Northampton-
shire. The lip is distinctly
three-lobed and about as
long as it is wide, while
the ‘wings’ formed by the
lateral sepals are narrow,
parallel-sided and held
slightly drooped.
X Gymnanacamptis anacamptis, the hybrid X Dactylodenia wintonii, the hybrid with

with Pyramidal Orchid, has been recorded Southern Marsh Orchid, has been found in
rarely (presumably x Common Fragrant Devon and Hampshire.
Orchid). X Dactylodenia varia, the hybrid with
X Pseudadenia schweinfurthii, the hybrid Northern Marsh Orchid, has been found in
with Small White Orchid, has been recorded northern England, Scotland and Co. Down in
from Yorkshire and Scotland, where it is fairly Ireland.
frequent in the northwest (presumably x Heath Common Fragrant Orchid can also
Fragrant Orchid). hybridise with Marsh Fragrant Orchid and
X Dactylodenia jacksonii, the hybrid with Frog Heath Fragrant Orchid, but these hybrids do
Orchid, has been noted rarely but widely. not yet have names.
X Dactylodenia st-quintinii, the hybrid with
Common Spotted Orchid, is uncommon but Name and classification
has been found widely through Britain and The specific epithet conopsea means ‘gnat-like’
Ireland; all three species of fragrant orchid may or ‘cloudy’ but the reason for this reference is
be involved. obscure.
X Dactylodenia legrandiana, the hybrid Since 1997 the ‘Fragrant Orchid’ has been
with Heath Spotted Orchid, has been found separated into three distinct species, Common,
widely scattered through Britain and Ireland Marsh and Heath Fragrant Orchids. Long
(presumably x Heath Fragrant Orchid). treated as separated varieties or subspecies, new
X Dactylodenia vollmannii, the hybrid with molecular and genetic evidence has shown that
Early Marsh Orchid, has been recorded from they merit specific status, even though they can
Cornwall and Cumbria. be hard to identify with certainty.
004 systematic.indd 261 29/1/09 12:32:15

HISTORY AND
CONSERVATION
The first British record of ‘Fragrant Orchid’
dates from 1634 when Thomas Johnson
published the Mercurius Botanicus (‘Botanical
Mercury’) and recorded the ‘Orchis palmata
minor calcaribus oblongis…. In montosis’
(‘lesser palmate Orchis with oblong spurs… In
mountains’). In 1660 John Ray stated that the
‘Fragrant Orchid’ could be found ‘in meadows
everywhere’ in Cambridgeshire.
Although the species is still common or
even abundant at suitable sites, it declined
significantly in the 19th and 20th centuries due
to the conversion of downs and pastures into
arable land and the ‘improvement’ of grazing
land. Even where the habitat still remains,
like all grassland orchids this species requires
a specific level of grazing; too little and scrub
invades, too much and the diversity of species
is lost.
The identification and recording of the
three fragrant orchids is still in its infancy, and
it is hard to identify trends for the individual
species. But, taken together, the ‘Fragrant
Orchid’ has vanished from 39.5% of the total
historical range in Britain and 30.5% in Ireland.
Past and present occurrence of the fragrant orchids (all

three species combined) in Britain and Ireland (based on
presence or absence in 10km squares of the National
Grid; data from the New Atlas).
Britain Ireland
total historical 1,341 365
range, 1500-1999
current range 810 (28%*) 253 (25%*)
% lost, 1500-1969 26.5% 24.5%

% lost, 1970-1986 13% 6%
% lost, total 39.5% 30.5%
* current range as a % of the total number of 10km squares M 29 June, Hampshire. A rather open, few-flowered spike.
004 systematic.indd 262 29/1/09 12:32:17

HEATH FRAGRANT ORCHID l 263
heath fraGrant Orchid

Gymnadenia borealis
Formerly: Gymnadenia borealis subspecies densiflora

Until very recently this species was, like Marsh Fragrant Orchid, considered to be merely
a subspecies of the composite ‘Fragrant Orchid’. It favours a variety of heathy and grassy
habitats and is the commonest fragrant orchid in northern and western Britain with scattered
outposts in southern England. It may yet be discovered elsewhere.
Identification
This is a relatively small, delicate, few-flowered
cousin of Common Fragrant Orchid. The
conical spikes of pink flowers, each with a
vaguely tri-lobed lip and long slender spur,
should nevertheless identify it as one of the
three fragrant orchids. These can be difficult
to separate from each other and, although
they are usually found in distinct habitats, can
occur together. Unlike Common and Marsh
Fragrant Orchids, Heath Fragrant Orchid does
not require alkaline soils and can be found in
neutral grassland in the uplands of northern
and western Britain and also very locally in
boggy hollows on grassy heaths in southern
England. Its status in Ireland is not certain
due to confusion with the other two fragrant
orchids. The scent is very sweet and carnation-
or clove-like and its main flowering period is
from mid-June to late July.
Similar species
Marsh and Common Fragrant Orchids are
similar, and where there is a possibility that
these could also occur a closer look is required
to be certain of the species. In Heath Fragrant
Orchid the flowers are relatively small, around
8-10mm across. The lip is longer than wide
and often rather obscurely lobed, with small
side-lobes and a longer central lobe. The ‘wings’
(lateral sepals) are oval, taper to a pointed
tip and are held angled downwards to about
four and eight o-clock. Intermediates occur,
however, and it is best to examine several
plants in a population and take an average. M 15 June, New Forest. Genetic evidence has recently
See table on p.258. confirmed that Heath Fragrant Orchid is distinct.
004 systematic.indd 263 29/1/09 12:32:18

Habitat Flowering period

In the north and west it is found on unimproved June to August in Scotland and Ireland and
hill pastures, roadside verges and grassy early June to early July in the New Forest
moorland, often in areas flushed with ground (around the same time as Common Fragrant
water. In such places it grows on tussocks of Orchid or just a little later); anomalously, the
grass and heathers, sometimes with Lesser population on chalk grassland in Sussex flowers
Butterfly Orchid and Heath Spotted Orchid. from late July to early August.
It is also found on machair, dunes and in old
quarries. It is tolerant of a wider range of pH Range
than Common and Marsh Fragrant Orchids Poorly understood as it has only recently been
and will grow on soils that are both mildly acidic separated from the other fragrant orchids. So
and alkaline, on sands, limestones and clays. In far it is known to occur in southern England
the New Forest and Ashdown Forest it is found in Cornwall (including the Lizard Peninsula
in ‘marl bogs’ (flushes and hollows on grassy and Bodmin Moor), Devon, Dorset, the New
heaths on base-rich clays). Very rarely it has been Forest in Hampshire (about 12 relatively small
recorded from chalk grassland, as near Lewes in populations of 50-100 flowering plants), East
Sussex. Occurs up to 610m above sea level (Ben and West Sussex (Ashdown Forest and three
Lawers, Perthshire). sites on the South Downs), and in Shropshire
O 15 June, New Forest. Growing

with Heath Spotted Orchid on a
large tussock.
004 systematic.indd 264 29/1/09 12:32:20

HEATH FRAGRANT ORCHID l 265
(one site), Derbyshire, Northumberland and

Cumbria (just two sites). In Wales it is known
only from Breconshire and Cardiganshire.
Scattered through much of Scotland, including
the Inner and Outer Hebrides, Orkney and
Shetland (although very rare there and now
found only on Unst). Occurs in western
and northern Ireland but there is very little
information on the distribution of Heath
Fragrant Orchid in Ireland. World Range:
Apparently more widespread in Europe than
Common Fragrant Orchid and found in France
as far south as the Dordogne. See Common
Fragrant Orchid.
How to find it
Widespread and locally common in Scotland
but very local in England and Wales. The M 8 July, Perth and Kinross. The lip is longer than wide
and often only obscurely lobed, and the ‘wings’ are oval,
pointed and held more or less drooped.
New Forest, where it is locally frequent,

is undoubtedly the best area to look for
this species in southern England. It can be
surprisingly elusive among tussocks of long
grass, and the number of flowering spikes can
vary widely from year to year.
DESCRIPTION
Height: 10-25cm, rarely to 30cm.
Stem: Green, becoming more purplish-brown
towards the tip.
Leaves: Mid-green, the three to five narrow,
strap-shaped basal leaves are keeled and have
pointed tips. They are held loosely erect, often
in two ranks, and grade into two or three
narrower and more lanceolate bract-like leaves
higher on the stem.
Spike: More-or-less cylindrical, although often
slightly irregular in shape, with 20-30 or more
flowers in a fairly lax spike.
Bract: Green, tinged purple (especially on
the edges), strap-shaped, narrowing abruptly
around the mid-point into a finer pointed tip,
and about one-and-a-half times the length of
the ovary.
Ovary: Green, variably washed purple, long,
M 8 June, Dunbartonshire.The spike can be relatively squat. narrow, three-ribbed and twisted.
004 systematic.indd 265 29/1/09 12:32:21

Flower: Dark pink to lilac. The sepals are Subspecies

oval or drop-shaped with the margins of the None.
lateral sepals rolled back to give them an oval-
lanceolate shape with a pointed tip. The petals
Var. albiflora has white flowers and is fairly
are shorter, more oval-triangular but less regular
frequent.
in shape. The upper sepal and petals form a
hood over the column, whereas the lateral sepals BIOLOGY
are held to the side, a little below the horizontal
and pressed backwards. The lip is flat and longer
No specific information, although the
than wide, lobed at the tip with the side-lobes
differences in the size and shape of the flower
shorter than the central lobe; the lobes are
suggest that a different suite of pollinators is
usually poorly developed with the incisions
involved for each of the three fragrant orchids.
between them reduced or absent. The spur is a
See Common Fragrant Orchid.
darker, more purplish-pink, very slender, down-
curved and rather long, around twice the length Development and growth
of the ovary; it is filled with nectar. The flowers No specific information.
have a powerful, sweet scent, recalling cloves,
Hybrids
which is strongest when they are newly opened
Hybrids have been recorded with both
and lacks the rancid overtones of Common
Common Fragrant and Heath Fragrant
Fragrant Orchid. See also table on p.258.
Orchids. See also Common Fragrant Orchid
(p.260).
The specific name borealis means ‘northern’, a
reference to the generally northerly distribution.
This species was originally described as
a variety of ‘Fragrant Orchid’ by the British
botanist G.C. Druce in 1918. It was raised to
the status of a subspecies in 1991 before finally
achieving specific status (yet to be universally
recognised) a few years later. Despite the
difficulties in identification, DNA evidence
strongly supports the separation of Heath
Fragrant Orchid from Marsh and Common
Fragrant Orchids.
HISTORY AND
CONSERVATION
This species is the most poorly known of the
three fragrant orchids and has only really been
brought to the attention of botanists since
1988 (when it was treated in the Botanical
Society of the British Isles’ Plant Crib). Its
history and conservation is obscure, but there
has undoubtedly been a considerable decline,
M 8 June, Dunbartonshire. The spikes are variable in with overgrazing of upland grasslands being a
shape and in the density of flowers. particular problem.
004 systematic.indd 266 29/1/09 12:32:22

MARSH FRAGRANT ORCHID l 267
marSh fraGrant Orchid

Gymnadenia densiflora
Formerly: Gymnadenia conopsea subspecies densiflora

Until recently this orchid was treated as a subspecies of ‘Fragrant Orchid’, a composite
that encompassed Common, Marsh and Heath Fragrant Orchids, but genetic evidence
has recently confirmed that all three are ‘good’ species. It has been recorded from scattered
localities throughout the British Isles and is rather scarce, being mostly confined to alkaline
fens, a habitat that has suffered badly from drainage and water abstraction. Although
coverage was undoubtedly incomplete due to confusion with its cousins, the New Atlas
mapped it from just 58 10km squares in Britain during the period 1987-99, and it probably
qualifies for the status of Nationally Scarce.
Identification
With spire-like spikes of pink flowers, each
with a distinctly three-lobed lip and a long,
slender, curved spur, this species is easy to
identify as one of the three fragrant orchids.
Similar species
The fragrant orchids can be hard to separate
from each other, although they usually grow in
different habitats. Marsh Fragrant Orchid is
found in meadows and fens where the ground
water is distinctly alkaline and sometimes
occurs in large numbers, often together with
Marsh Helleborine. It is occasionally recorded
from chalk downland and can sometimes occur
together with either Common or Heath Fragrant
Orchids. Marsh Fragrant Orchid is nevertheless
relatively distinctive, being a tall, robust plant
with numerous broad basal leaves that are
usually held noticeably erect and a good-sized
spike of relatively large, dark pink flowers. To
clinch the identification, a careful examination,
preferably of several plants, is necessary. In
Marsh Fragrant Orchid the flowers are around
11-13mm across and the lip is broader than long,
prominently lobed (with the side-lobes larger
than the central lobe) and has distinct ‘shoulders’.
The ‘wings’ (lateral sepals) are long and narrow
with parallel sides and a blunt tip and are
held roughly horizontal. See table on p.258.
P 11 July, Norfolk. The robust fragrant orchids of fenland

habitats have long been recognised as distinct.
004 systematic.indd 267 29/1/09 12:32:23

Intermediates and hybrids do occur, however, Flowering period

and some populations do not ‘fit’ any of the Late June to mid-August.
three species (see photograph on p.270).
Range
Habitat Rather poorly known because until recently
Usually found in fens and meadows flushed many records were given merely as ‘Fragrant
with calcareous, base-rich water, also dune Orchid’. In England found in a belt of counties
slacks and, on the Isle of Wight, slumped clay along the south coast from Cornwall to Kent
cliffs. Occasionally occurs on chalk grassland (including the Isle of Wight), at a handful
on north-facing slopes, as at Ditchling Beacon of sites in Berkshire and Hertfordshire, in
and Heyshott Down in Sussex. Recorded up to Norfolk and adjacent parts of Suffolk, and
310m above sea level (Tomintoul, Banffshire). then through the Midlands and northern
England north of a line from Cambridgeshire
to Herefordshire, although very scattered.
In Scotland the species is similarly local and
has been recorded from Peebles, Midlothian,
Banffshire, a few islands in the Inner and Outer
Hebrides, Ross & Cromarty, Sutherland and
Orkney. Similarly scattered in western and
northern Wales. Apparently the commonest
of the three fragrant orchids in Ireland but
there are few specific records. World range:
Widespread in Europe and especially common
in the southeast, particularly the Balkans.
How to find it
The tall plants in fens and calcareous meadows
are easy to find, although this habitat is very
local. Chalk grassland populations can very easily
be overlooked, however, and any population of
fragrant orchids is worth careful examination.
DESCRIPTION
Height: 30-60cm, exceptionally to 90cm; plants
in chalk grassland are the smallest.
Stem: Green, washed purple towards the tip.
Leaves: There is a rosette of several erect,
lanceolate and relatively broad leaves at the base
of the stem and several smaller, narrow and
more bract-like leaves higher up.
Spike: Up to 100 flowers form a rather tall,
narrow spike.
Bract: Green, washed purple (especially towards
the edges) and narrowly oval, tapering to a point;
M 17 July, Norfolk. The deep, purplish-pink colours of the bracts are roughly the length of the ovary.
the fragrant orchids are extremely hard to reproduce in Ovary: Green, variably and sometimes heavily
print.
washed purple; the ovary is long, narrow,
P 11 July, Norfolk. A typical tall, spire-like spike. curved, prominently three-ribbed and twisted.
004 systematic.indd 268 29/1/09 12:32:24

004 systematic.indd 269 29/1/09 12:32:27

Flower: Deep pink, variably tinged purple

and often whiter around the base of the lip.
The sepals are elongated-oval in shape with
the upper and lower margins of the lateral
sepals rolled back so that they appear to be
parallel-sided with a blunt tip. The petals are
a little shorter and more oval-triangular in
shape. The upper sepal and petals form a hood
over the column whereas the lateral sepals are
held horizontally to the side. The lip is flat,
broader than long and has distinct ‘shoulders’
(abrupt angles on each side at the base where
it narrows towards the mouth of the spur).
It has three well-developed rounded lobes,
with the side-lobes larger than the central
lobe. The spur is relatively dark, very slender,
down-curved and long. The scent is spicy M 11 July, Norfolk. The lip is wider than long, with large
sweet, recalling cloves, and lacking the rancid side lobes and distinct ‘shoulders’, and the ‘wings’ are long,
overtones of Common Fragrant Orchid. narrow, blunt and usually carried horizontally.
See also table on p.258.
Subspecies
None.
The dune-slack population at Kenfig in south
Wales has recently been assigned to ‘var.
friesica’, described from Dutch plants and said
to intermediate in appearance between Marsh
and Heath Fragrant Orchids. The validity of
‘var. friesica’ and its relationship to the three
species of fragrant orchid has, however, yet to
be clarified.
BIOLOGY
No specific information but the differences in the
size and shape of the flower, notably the long spur
and distinct scent in Marsh Fragrant Orchid,
suggest that a different suite of pollinators is
involved for each of the three fragrant orchids.
See also Common Fragrant Orchid.
No specific information.
Hybrids
Hybridises with Heath Fragrant Orchid in
Ireland and sometimes with Common Fragrant
M 9 July, Norfolk.The shape of the flowers is variable. Orchid. See also Common Fragrant Orchid.
004 systematic.indd 270 29/1/09 12:32:30


The specific name densiflora refers to the
densely packed flower spike, although the spike
is not necessarily more crowded than Common
Fragrant Orchid.
Originally described as a distinct species
in 1806, Orchis densiflora was quickly reduced
to being merely a variety of ‘Fragrant Orchid’
(although it did appear as a distinct species in
Francis Rose’s excellent Wild Flower Key as
long ago as 1981). Recently, DNA evidence has
shown that it, together with Heath Fragrant
Orchid, merits full specific status.
HISTORY AND
CONSERVATION
Confusion with Common Fragrant Orchid
means that its past history in Britain is obscure.
Although ‘var. densiflora’ warranted some
attention, the identification criteria used were
sometimes dubious, notably the width of the
lower leaves and the length of the flower spike,
and many records merit re-examination.
Marsh Fragrant Orchid has undoubtedly
undergone a serious decline in much of Britain.
The direct drainage and destruction of fens
and marshes has caused some losses. However,
subtler effects, including eutrophication and
the lowering of water tables, have also caused
declines which reserve or SSSI status could
not protect against. The few chalk grassland
populations are also subject to losses due to
‘improvement’ or abandonment. The species
is now much reduced in Norfolk, one of its
strongholds, and has withdrawn to just five
sites in Suffolk. There are no recent records
from Somerset, Wiltshire, Gloucestershire,
Bedfordshire, Lincolnshire, Co. Durham,
Lanarkshire, Berwickshire, Perthshire or
Angus.
P 3 July, Norfolk.The flowering period can be somewhat

variable – in this small population by two to three weeks
– and is of limited use in identification or in delineating
the various forms.
004 systematic.indd 271 29/1/09 12:32:31

GenuS DACTYLORHIZA
marSh OrchidS and
SpOtted OrchidS
004 systematic.indd 272 29/1/09 12:32:39

GENUS DACTYLORHIZA l 273
This genus, which includes the marsh and spotted orchids, presents the toughest identification
challenge among British and Irish orchids. The species are closely related and individually
quite variable. Indeed, there is disagreement as to how the various populations should be
classified into species, subspecies and varieties. Hybridisation is also relatively common,
making identification even more difficult. The situation can be particularly complex in
Ireland, west Wales and western Scotland due to the number of possibilities that have to
be considered. If in doubt it may be necessary to take a ‘statistical’ approach and carefully
measure a selection of plants in order to reach a satisfactory identification. On the other
hand, a field full of Southern Marsh Orchids or the scatter of Heath Spotted Orchids across
a heather moor are straightforward to identify and, indeed, spectacular.
Distribution become reproductively isolated from its parents.

Approximately 75 species are found in This hybridisation event occurred several times
Europe and temperate Asia. The range of one and with slightly differing parents, producing the
species extends into Alaska via the Aleutian closely similar species we see today.
Islands, and Frog Orchid has a circumpolar On genetic evidence the Frog Orchid has
distribution. recently been moved into Dactylorhiza. It is
thought to be a ‘primitive’ member of the genus
Classification
and differs from the other British species not
Dactylorhiza is one of several closely related
only in its general appearance but also in having
genera that all have well-developed tubers. It
a rudimentary spur and bursicle, by producing
is probably most closely related to the fragrant
nectar, and by having the ‘hood’ of the flower
orchids Gymnadenia and butterfly orchids
made up from all of the sepals and petals (in
Platanthera and only more distantly to the
the other species the lateral sepals are held away
‘typical ‘ orchids in the genus Orchis. This is
from the hood).
despite the fact that until the 1930s and 1940s
the marsh and spotted orchids were placed in Identification
the genus Orchis. The most useful tools in the identification of
The Dactylorhiza orchids can be divided marsh and spotted orchids are a small ruler, a
into two groups depending on the number of notebook and pencil. The ruler is useful because
chromosomes they possess. The diploid group for many species the width of the flower’s lip
has a chromosome count of 2n = 40. This is a critical measurement, as well as its length.
includes Early Marsh Orchid and Common Sometimes the width of the widest leaf and
Spotted Orchid. The tetraploid marsh orchids the total height of the plant are important too.
have a chromosome count of 2n = 80. This The notebook and pencil are useful because it
group includes Southern, Pugsley’s, Northern, is almost always necessary to take the details of
Irish and Hebridean Marsh Orchids. The several plants. The following are worth noting:
chromosome count is significant for two reasons. 1. Lip size: the width across the widest point of
The first is hybridisation as it has some influence the flattened lip and also the length (from the
on the fertility of hybrids. Second, the tetraploid mouth of the spur to the tip of the central lobe).
marsh orchids are thought to have evolved from 2. Lip shape: the lip is often folded or has the
a cross between the ancestors of two members sides turned down. To judge its shape it is
of the diploid group, Early Marsh Orchid necessary to flatten it. This can easily be done
and Common Spotted Orchid, followed by a by sliding the ruler or a finger below the lip and
doubling of the chromosomes (in which 2n = gently pushing upwards. The degree to which
40 became 2n = 80). This allowed the hybrid to the side-lobes are turned downwards in its
004 systematic.indd 273 29/1/09 12:32:43

274 l GENUS DACTYLORHIZA
lateral upper
lateral
sepal sepal
sepal
petal
petal
mouth of
the spur
side-lobe
bract
non-sheathing double loop

leaf markings
sinus
central lobe
non-sheathing sheathing
leaf natural position may also be important. The
leaf
lip is almost always lobed but it may also have
small cuts or incisions (sinuses) separating the
lobes; see figure above.
3. Lip markings.
sheathing
leaf 4. Flower colour: only useful in some cases,
as the tetraploid marsh orchids (Southern,
Northern, Pugsley’s, Irish and Hebridean) are
all similar shades of purplish-pink.
basal 5. Position of lateral sepals: these may be held
leaf horizontally at the side of the flower, vertically
over the flower or at any angle in between.
6. Lateral sepal markings.
7. The shape of the spur.
basal 8. The number of flowers: surprisingly hard to
sheath
count and only really useful in the identification
of Pugsley’s Marsh Orchid. If there are too
many to count, it cannot be that species.
9. The number of leaves: the leaves are divided
into sheathing and non-sheathing. Sheathing
leaves are found on the lower part of the stem
tuber and at their base they completely encircle the
(divided into stem with a short, unbroken, tubular sheath.
finger-like lobes) Non-sheathing leaves are found higher on the
stem, are narrower and more bract-like. They
have a clearly defined base that may encircle the
M A typical Dactylorhiza orchid.
stem but never have an unbroken sheath.
O 1 June, Norfolk. Southern Marsh Orchid: The Dacty- Importantly, the basal leaf is not included
lorhiza may produce spectacular displays, but the colours
of the marsh orchids are extremely hard to reproduce in in any counts. This can range from a very short
print. sheath at ground level with a green tip to a
004 systematic.indd 274 29/1/09 12:32:44

GENUS DACTYLORHIZA l 275
fully-formed leaf up to half the length of the As in the genus Orchis, the next season’s
sheathing leaf immediately above it (see figure tuber develops alongside the current one and
opposite). the system of annual replacement is similar (see
10. Leaf markings. p.200).
11. Leaf hooding: the tips of the leaves in some Fungal partners
species are described as hooded but this is a In the adult plant only the roots and sometimes
subtle and very variable character and is of the tips of the tubers are ‘infected’ with fungi.
limited use (see figure below). Members of this genus have well-developed
foliage, and fungi may play a relatively minor
role in their nutritional budget.
hooded leaf-tip Development from seed
Seed probably germinates in the autumn
to produce a tiny, conical or turnip-shaped
protocorm about 2mm long. The first root and
leafy shoot are produced the following spring,
and the first tuber develops from a bud at the
base of this leafy shoot. This tuber is rod-
shaped with just one finger-like extension and
in the late summer the protocorm, root and
leafy shoot disappear, leaving just the tuber.
In the autumn a short rhizome grows from a
Floral structures bud at the tip of the tuber. This produces roots
There are two pollinia, each narrowing into a
and, in the spring, elongates to produce the
caudicle (stalk) and attached by a basal disc
next leafy shoot. The sequence of events is very
to one of the viscidia. A two-lobed bursicle
similar to that of the genera Orchis, Gymnadenia
is present and encloses the two viscidia. The
and Platanthera (see p.201). A regular annual
stigma is more-or-less two-lobed and situated cycle now begins. The tuber that produced
at the roof of the entrance to the spur. the current year’s growth gradually shrivels
Pollination away, and a new tuber forms beside it from
Dactylorhiza are cross-pollinated by bees but a bud on the short rhizome. This swells over
most members of the genus produce no nectar. the summer as it stores up nutrients produced
The insects receive no reward and are perhaps by photosynthesis. The period between
just attracted to the mass of brightly coloured germination and flowering is four or five years
flowers which are often found in large numbers. in most species of Dactylorhiza.
The attraction is therefore a deceit. The exception Vegetative reproduction
is Frog Orchid, which does produce nectar. Additional tubers may develop at the end of
Growth pattern short shoots growing from the base of the stem,
The aerial stem grows from a pair of tubers. and these become separate plants when the
These are flattened and each divided into two to central stem dies away in the autumn.
five finger-like lobes which taper to a fine point. Name
They may be long in dry habitats, extending The generic name Dactylorhiza originates
well down into the soil, but in wet areas may from the Greek daktulos ‘finger’ and rhiza ‘root’
bend upwards towards the surface of the soil, and means ‘finger-like root’, a reference to the
perhaps to avoid becoming waterlogged. There distinctive shape of the tubers.
are also several long, fleshy roots growing near
the surface of the soil.
004 systematic.indd 275 29/1/09 12:32:44

early marSh Orchid

Dactylorhiza incarnata
Formerly: Orchis latifolia, Dactylorchis latifolia

This is the most widespread of the marsh orchids, occurring throughout Britain and Ireland
in a wide variety of habitats. However, it is always rather local, often uncommon and indeed
has disappeared from many areas. It is also the most variable marsh orchid with five named
subspecies and flowers that range in colour from deep red to creamy. Because the subspecies
are so different and occur in different habitats, they are discussed separately in the text.
Identification
The five subspecies in brief:
1. D. i. incarnata has flowers that are usually
very pale pink but sometimes purplish-pink. It
is found in alkaline fens and marshy meadows
throughout the British Isles.
2. D. i. coccinea has flowers that are deep red
and often looks like a fat little hyacinth. It
occurs scattered on both the east and west
coasts in coastal dune slacks. There are also
some inland sites, especially in Ireland.
3. D. i. pulchella has flowers that are purplish-
pink. It is largely confined to bogs on acid
heathland in southern England.
4. D. i. cruenta Flecked Marsh Orchid has
flowers that are mid to dark pink. It also
frequently has bold spots on the leaves and
bracts. It is confined to alkaline fens in western
Ireland and northwest Scotland.
5. D. i. ochroleuca has unmarked creamy
flowers and is found in alkaline fens in East
Anglia but is now very rare.
Early Marsh Orchid is probably the most
variable species of orchid in the British Isles,
both in flower colour and in stature. It may
be very petite and just 5cm tall or a robust
giant at 60cm. Nevertheless, it is relatively
distinctive because the individual flowers have
a characteristic size and shape: They are always
M 6 July, Lancashire. Sub- small, indeed disproportionately so on robust
species incarnata. A very plants compared to the stem, bracts and ovaries.
stout plant.
The lip is no more than 9mm wide and usually
O 13 June, Norfolk. Sub- rather less (push a finger gently upwards from
species incarnata. A slen-
der, petite plant with few below to flatten the lip before measuring it).
flowers. Furthermore, the flowers appear even narrower
004 systematic.indd 276 29/1/09 12:32:45

EARLY MARSH ORCHID l 277
M 19 June, Norfolk. Subspecies coccinea.

This subspecies appears in many different
shades of red, from orange-red to purplish-
red.
O 27 June, Norfolk. Subspecies coccinea.

Grows in dune slacks and also on slumped
clay cliffs.
because the sides of the lip are often folded D. i. coccinea is characteristic, but the various
downwards, sometimes sharply so, especially as purple-flowered forms can be more confusing.
the spike matures. This narrow appearance is Southern Marsh Orchid always has larger
further accentuated by the lateral sepals, which flowers with a broader lip. The sides of the
are held vertically above the flower as if it is lip are quite frequently folded downwards
holding its ‘arms’ up in surrender. And, whatever but never as sharply as in some Early Marsh
their coloration, the flowers have the same Orchids, and the lip is often held flat or even
basic pattern of markings (with the exception dished. It typically lacks prominent double loop
of the unmarked creamy flowers of subspecies markings and although Leopard Marsh Orchid
ochroleuca). The centre of the lip has a scatter (var. junialis of Southern Marsh Orchid) does
of wriggly lines of varying length and these are have loop markings it also has ring-shaped
all bounded by a solid dark line that forms a spots on its leaves.
double loop, with few, if any, dark markings Northern Marsh Orchid is quite similar
outside this loop. to purple-flowered Early Marsh Orchids but
Similar species its flowers are on average slightly larger with
The pink-flowered subspecies D. i. incarnata a broader, distinctively diamond-shaped lip
is distinctive as no other British marsh orchid which is usually held flat. It is often an intense
shows this coloration. Similarly the deep red reddish-pink, lacks the double loop markings,
004 systematic.indd 277 29/1/09 12:32:48

extend to the edge of the lip and it often lacks

the double loop of Early Marsh Orchid. In
Scotland, ‘Lapland Marsh Orchid’ (subspecies
lapponica of Pugsley’s Marsh Orchid) has been
found in the same localities as D. i. cruenta,
the spotted-leaved subspecies of Early Marsh
Orchid, but although they share spotted leaves
and bracts and heavily marked lips, they can be
separated as above.
Irish Marsh Orchid is usually stouter
than Early Marsh Orchid with the leaves
neither as rigidly upright nor hooded. It has a
proportionally larger flower spike and a flatter,
broader lip that has more evenly rounded lobes.
Flowering period
Late May to late June, occasionally from
mid-May or to late July. There is considerable
variation both within individual colonies and
between sites but late May and early June is
probably the peak for many. Beyond a tendency
for plants in the south and west to flower
earlier and plants in wetter sites and at higher
altitudes to flower later, there seems to be no
real difference between the flowering times of
the various subspecies.
Habitat
Subspecies incarnata is found on calcareous
or neutral soils on damp to wet grassland.
It is especially characteristic of unimproved
wet meadows on floodplains. It is also found
in spring-fed fens and flushes, even in the
mountains of Scotland. Paradoxically, although
M 15 June, New Forest. Subspecies pulchella. Grows in it is usually found in wet areas, the species
acid bogs with bog-mosses Sphagnum.
cannot survive a prolonged submergence.
and its lateral sepals are not usually held as Other habitats include dune slacks (where it
near to the vertical. Subspecies cambrensis of may be found with the red-flowered subspecies
Northern Marsh Orchid has boldly spotted coccinea), old fly-ash tips, occasionally old chalk
leaves and a slightly more three-lobed lip than quarries where soil compaction may cause
typical plants, but the lip is still broader than in seasonal water-logging and, very rarely, chalk
Early Marsh Orchid. grassland. Recorded up to 440m above sea level
Pugsley’s Marsh Orchid has narrower leaves (Ben Lawers, Perthshire; plants not referred to
than Early Marsh Orchid, a rather few-flowered any subspecies have been found at 610m in the
spike and relatively large flowers. The lip is Atholl District in Perthshire and also at 610m
usually over 9.5mm wide and distinctly three- at Caenlochan in Angus). Subspecies coccinea
lobed, often with a prominent projecting central is found in damp dune slacks, the machair
lobe. The dark markings are variable but usually grasslands of the Hebrides, the flushed slopes
004 systematic.indd 278 29/1/09 12:32:49

vegetated areas. Suitable sites were created

when peat was cut for fuel and the workings
subsequently flooded. Such ‘turf ponds’ slowly
fill with vegetation which eventually forms a
floating mat or ‘hover’ but in the long run the
continued build-up of material dries them out
and they become unsuitable.
Range
Subspecies incarnata is locally frequent in
England and Wales but is very scattered in
Scotland and Ireland. Subspecies coccinea
is very locally common on the west coast
from Devon north to the Inner and Outer
Hebrides and Shetland (South Mainland
only; it is absent from Orkney). On the east
coast it is scattered in southeast Scotland,
northeast England and north Norfolk. It has
been recorded inland in Hertfordshire and
Derbyshire, and in Ireland is very scattered but
occurs both inland and on the coast. Subspecies
coccinea is often found in very large numbers.
Subspecies pulchella occurs in Cornwall,
Dorset, the New Forest, the heaths around
the Surrey/Hampshire border, Ashdown
Forest in Sussex and perhaps also Dartmoor
in Devon. Similar plants, perhaps subspecies
pulchella (rather than subspecies incarnata with
pinkish-purple flowers), may be found in acid
M 15 June, New Forest. Subspecies pulchella. The flower
habitats elsewhere in Britain, especially Ireland,
is purplish-pink. but the exact status of these populations is
of slumped clay cliffs in east Norfolk and damp

1987-99
lake shores inland in Ireland. It is also recorded 1970-86
from alkaline Leblanc waste in Lancashire and pre 1970
pulverised fly-ash waste. Subspecies pulchella

is found in valley bogs and acid marshes on
heathland, often growing among Sphagnum and
sundews. Subspecies cruenta Flecked Marsh
Orchid is found in the limestone districts of
Ireland in calcium-rich fens around loughs and
turloughs, frequently with Black Bog-rush;
indeed, it often grows in the hollows amongst
slabs of limestone. Its only three Scottish sites
are on roughly neutral flushes and mires up to
450m above sea level. Subspecies ochroleuca
is found in calcareous spring-fed fens where it
grows on a mossy carpet in the more sparsely
004 systematic.indd 279 29/1/09 12:32:51

through the mountains of Central Asia to the

Himalayas. Subspecies coccinea and subspecies
pulchella are endemic to Britain. Subspecies
cruenta is widespread in Scandinavia and the
Alps and is also found in Poland, the Baltic
States and through central and northern Russia
to eastern Siberia. Subspecies ochroleuca is
local and rare but occurs widely in central
Europe from Haute-Savoie in southeast France,
Switzerland, Austria, Hungary and Romania
north to Denmark, southern Scandinavia,
Poland, the Baltic States and perhaps northwest
Russia; it has also been recorded from Spain.
But as in Britain, confusion with pale-flowered
variants of the other subspecies has obscured its
precise distribution.
How to find it
With the exception of dune populations, Early
Marsh Orchid tends to occur in small numbers,
unlike the massed ranks of Southern, Northern
or Irish Marsh Orchids, and is easily overlooked.
A search of any rough-looking riverside
meadows, with a good growth of rushes, is
recommended. In the London area, the Lee
Valley Park is an excellent site for this species.
The red-flowered form can be found in good
numbers at most of the larger protected dune
systems, and the purple-flowered subspecies
pulchella is common in many of the New Forest
bogs and also at Thursley Common in Surrey.
A visit to western Ireland is necessary to see
M 23 May, Co. Clare. Subspecies cruenta. Grows in cal-
careous fens. Not all plants have spotted leaves. Flecked Marsh Orchid, subspecies cruenta.
unclear. Subspecies cruenta occurs in western DESCRIPTION (incarnata)

Ireland in Co. Galway, Co. Mayo and the Height: 7-65cm but typically 20-40cm and
southeastern portion of The Burren in Co. rarely more than 20cm in Scotland.
Clare. In Scotland there are a couple of tiny Stem: Bright yellowish-green or apple green
populations near Ullapool in Ross & Cromarty and usually hollow; in larger plants the thick,
and one in Assynt in Sutherland. Subspecies hollow stem is conspicuous.
ochroleuca is confined to East Anglia. World Leaves: Bright yellowish-green or apple green.
range: Europe and Asia. Subspecies incarnata There are three to five sheathing leaves, often
is found throughout Europe away from the crowded towards the base of the stem and
Mediterranean lowlands, north to northern rather broad (the largest more than 2cm wide,
Scandinavia and south to Spain, Italy, Greece, sometimes as much as 3.5cm), with up to two
the Crimea and Caucasus. In Asia it occurs non-sheathing leaves higher on the stem. The
from northern Turkey to northern Iran and leaves are typically held rather erect at c. 45°, are
in Siberia east to Lake Baikal and south strongly keeled and often hooded at the tip.
004 systematic.indd 280 29/1/09 12:32:52

Spike: Crowded, with ten to 70 flowers. Marsh Orchids can be found growing together.
Bract: Apple green, sometimes flushed rose- This variation in flower colour appears to be
pink or purplish, lanceolate and fairly long, up commoner in Wales, Cumbria, Scotland and
to twice as long as the ovary and projecting Ireland, but even in these areas ‘mixed’ colonies
well beyond the flowers in the lower part of are still mostly or always found in neutral or
the spike. base-rich habitats.
Ovary: Green, six-ribbed and twisted, also bent (Note that ‘D. i. gemmana’ has been recorded
through approximately 45°. from east Norfolk and east Galway with similar
Flower: The flowers are usually pale pink but plants found elsewhere in England, Wales and
at some sites there may also be plants with Ireland. Ignored or forgotten for many years
flowers of various shades of purplish-pink. it has recently been ‘resurrected’ but it may be
The sepals and petals are off-white, variably best to treat it as a large, late-flowering variant
washed rose-pink. The lateral sepals are oval of subspecies incarnata. It is robust, growing to
to strap-shaped, rather like donkey’s ears, 50cm (sometimes 80cm), has six or more leaves
and slightly asymmetric; they are held erect and is large flowered, with the lip more than
and often have dark pink spots or sometimes 8.5mm wide x 7mm long and the spur usually
ring-shaped markings. The upper sepal and longer than 7.5mm. The flowers are either pink
petals are more oval in shape (the petals a little or purple and the lip is marked with fine dots
smaller) and form a tight hood. The lip is off- rather than a double loop, suggesting perhaps a
white washed rose-pink, especially towards hybrid origin.)
the edges, with irregular dark pink dots and D. i. coccinea is often relatively small at 5-20cm
lines in a central zone enclosed within two but may grow to 30cm; it can appear stout,
complete or near-complete loops. It is usually squat and apparently stemless. The flower is a
less than 8.5mm wide and 7mm long and is
slightly to moderately lobed, with the sides
usually strongly turned downward; indeed,
this deflexing may be so marked that the
lip is almost folded in two. The spur is pale
pink, stout, slightly tapering and slightly to
moderately decurved (rarely straight); it ranges
from half as long to nearly as long as the ovary
(but less than 7.5mm long).
Subspecies
Each of the five subspecies is sometimes treated
as a distinct species by European authors,
but recent genetic analyses show that they
are very similar indeed, with the exception
of subspecies cruenta, which does show some
genetic differences. The dividing lines between
the various subspecies are not hard and fast
and intermediates occur. See also Habitat and
Range.
D. i. incarnata is the ‘typical’ subspecies and is
described above.
In southern England the vast majority
of plants have pale pink flowers but at a few M 23 May, Co. Clare. Subspecies cruenta. The flower is
sites both pale pink and purplish-pink Early mid to dark pink.This is a heavily marked plant.
004 systematic.indd 281 29/1/09 12:32:53

004 systematic.indd 282 29/1/09 12:32:58

distinctive deep red with even darker markings it differs from subspecies incarnata in having
although these are less obvious than in other thicker, bolder and more complete double loop
subspecies due to the reduced contrast with the markings on the lip, which is less obviously
dark red ground colour; the markings on the lip lobed. The sides of the lip either do not fold
are rather narrow and appear quite faint. The downwards at all or do so only as the flower
bracts are rather long and are washed and edged gets older, and the lip may therefore be flat
purple, and the upper stem and ovaries are also or even slightly dished. On average the leaves
strongly washed purple. The spur is stout and are narrower and a darker and deeper green.
bag-like or conical. The leaves are broad-based The bracts are shorter and less prominent.
but sharply tapering, dark green and unspotted. There is some variation in flower colour and
In most dune colonies there are also plants with in some colonies very pale yellowish flowers
flowers which are paler and pinker, intermediate are relatively common; these are subspecies
with subspecies incarnata, and often some pulchella, var. ochrantha.
typical pale pink incarnata too. The subspecific name pulchella is often
D. i. pulchella has purplish-pink flowers that used willy-nilly for any purple-flowered Early
fade to white around the mouth of the spur; Marsh Orchid wherever it grows. We prefer
they are rather similar in colour to Southern to reserve the name pulchella for the heavily
Marsh Orchid although slightly more purple. marked, purple-flowered populations growing
The upper stem, bracts and ovaries are variably in acid bogs and to treat the purple Early Marsh
washed purple, and as in subspecies incarnata Orchids that grow with pale pink-flowered
the lateral sepals may have ring-shaped plants in alkaline habitats as colour variants of
markings. Apart from the colour of the flower subspecies incarnata.
M 6 July, Lancashire. Subspecies incarnata among Creeping Willow. On the edges of dune slacks subspecies incarnata
and subspecies coccinea can be found growing together.
O 25 May, Norfolk. Subspecies incarnata.
004 systematic.indd 283 29/1/09 12:32:59

1 2 3
4 5 6
M 1. 19 June, Norfolk. Subspecies incarnata. 2. 19 June, Norfolk. Subspecies incarnata. 3. 17 June, Norfolk. Subspecies
coccinea. 4. 15 June, New Forest. Subspecies pulchella. 5. 23 May, Co. Clare. Subspecies cruenta. 6. 14 June, Suffolk.
Subspecies ochroleuca.
D. i. cruenta ‘Flecked Marsh Orchid’ has more strongly washed purple than in subspecies
mid to dark pink flowers that lack the purple pulchella and are sometimes spotted or flecked
tones of subspecies pulchella. Around 30-65% darker. There are sometimes ring markings
of plants have dark purplish-brown spots on on the bracts, too (almost never spotted in
one or both surfaces of the leaves, the spots pulchella), and the lateral sepals more often
becoming denser towards the tips of the leaves have ring markings. The lip is more distinctly
and sometimes merging. If present, the spots three-lobed, about 4.5-7.5mm long x 4.5-9mm
on the underside of the leaves are paler, smaller wide, the side-lobes are moderately reflexed
and sparser. The stem, bracts and ovaries are and the margins of the lip are often slightly
004 systematic.indd 284 29/1/09 12:33:10

wrinkled (crenate). The lip has broader, bolder

loop markings but few dashes within the loops
and these markings often spread over most of
the lip (extending over no more than two-thirds
of the lip in pulchella). The stem is slender,
the leaves are held stiffly erect, and the spike
is looser and less crowded than in subspecies
incarnata, especially in Scottish plants.
Controversy surrounds this subspecies and
doubts are sometimes expressed as to whether
British and Irish plants are ‘true’ cruenta as
found in Europe. Bizarrely, it is sometimes
suggested that while Scottish plants are the
‘real thing’, Irish plants are not, being merely
spotted-leaved variants of subspecies pulchella.
D. i. ochroleuca is often relatively large,
sometimes as much as 70cm tall, with a broad
stem and large leaves that are usually held very
erect. The bracts are large, the lowest usually
larger than 30mm x 20mm. The flowers are
creamy to pale yellow with a relatively large lip,
around 9mm wide x 7mm long, that is deeply
three-lobed with notches on the side-lobes.
This subspecies has often been confused with
pale-flowered variants of other subspecies, i.e.
with var. ochrantha, but it has larger and more
distinctly lobed flowers.

As well as the various subspecies, several
varieties have been named. In principle, each
different subspecies could be found as each
M 13 June, Norfolk. Subspecies incarnata. Early Marsh
different variety, giving around 15 different Orchid can be robust, with many proportionally small
combinations. flowers.
Var. punctata is usually rather small with a
it has also been recorded among incarnata,
few small dots on the leaves towards the tip. It
coccinea and cruenta.
is rare but has been recorded from Yorkshire,
A hyperchromic variant with an excess of
the New Forest and the Isle of Coll, among
pigmentation and a solidly dark lip has also
subspecies incarnata and pulchella.
been recorded, albeit rather rarely.
Var. leucantha has unmarked pure white
flowers but is very rare.
Var. ochrantha has unmarked pale creamy- BIOLOGY
yellow flowers with the lip pale yellow or rarely Pollination and reproduction
green at the base (it is similar to subspecies There is no nectar, and pollination is by
ochroleuca but has smaller and less distinctly bumblebees that are ‘duped’ into visiting the
three-lobed flowers). It is scarce and usually flowers. The process is efficient and seed-set
found among subspecies pulchella, although is good.
004 systematic.indd 285 29/1/09 12:33:11

Hybrids
Most or all of these hybrids are sterile and
only found as isolated individuals or in small
groups.
D. x kernerorum, the hybrid with Common
Spotted Orchid, has been found scattered
throughout Britain and Ireland. It is sterile.
D. x carnea, the hybrid with Heath Spotted
Orchid, has been recorded scattered in Britain
and Ireland but is rare. It is probably sterile.
D. x wintoni, the hybrid with Southern Marsh
Orchid, has been found scattered throughout
the range of the latter but is rare. It is sterile.
D. x latirella, the hybrid with Northern Marsh
Orchid, has been recorded throughout the
range of the latter but is scarce.
D. x dufftii, the hybrid with Pugsley’s Marsh
Orchid, has been found in northwest Wales,
Yorkshire and Co. Wicklow. It is rare.
D. x aschersoniana, the hybrid with Irish Marsh
Orchid and Hebridean Marsh Orchid, has
been recorded in Co. Limerick and the Outer
Hebrides respectively. This name is no longer
valid for both species due to changes in the
classification.
Intergeneric hybrids
X Dactylodenia vollmannii, the hybrid with
fragrant orchid (in the broad sense), has been
recorded twice from Cornwall.
The specific name incarnata means ‘flesh-
coloured’. Most of the subspecific names also
M 14 June, Suffolk. Subspecies ochroleuca. Grows in cal- refer to the flower colour; pulchella is Latin for
careous, spring-fed fens.
‘beautiful’, coccinea means ‘crimson’ or ‘scarlet’ (as
Development and growth in the dye produced from galls on the Kermes
Seed probably germinates in the spring, and the Oak), cruenta is Latin for ‘blood-coloured’ and
first leaves appear a year later. Early estimates ochroleuca comes from the Greek for ‘yellowish-
of the period between germination and first white’.
flowering were as long as 16 years, but these The Early Marsh Orchid and its relatives are
were made by counting growth marks in tubers often known as the ‘diploid marsh orchids’. This
and other structures that were thought to is because their chromosome count is 2n = 40
represent a year’s development. This technique compared with a count of 2n = 80 in the
is probably unreliable, and in cultivation plants ‘tetraploid marsh orchids’, a group that includes
flower aged four or five years. Individual plants all the other British and Irish marsh orchids.
have been recorded living for up to 25 years Although of little use to the orchid-lover in the
after their first appearance above ground. field, it represents a fundamental division.
004 systematic.indd 286 29/1/09 12:33:12

HISTORY AND Past and present occurrence of Early Marsh Orchid (all
subspecies) in Britain and Ireland (based on presence or
CONSERVATION absence in 10km squares of the National Grid; data from
For a long time this species was amalgamated the New Atlas).
with the other marsh orchids. The first British Britain Ireland
record that distinguished the Early Marsh
Orchid was in the Flora of Shropshire published 1500-1999
in 1841.
current range 671 (23.5%*) 202 (20%*)
Early Marsh Orchid is frequently found
in small numbers and is often the first of the % lost, 1500-1969 29.5% 30.5%
marsh orchids to vanish if its habitat dries out % lost, 1970-1986 14% 8.5%
due to drainage or other changes. Overall, the % lost, total 43.5% 39%
species has gone from 43.5% of its historical * current range as a % of the total number of 10km squares
range in Britain and 39% in Ireland. In lowland
areas the species declined significantly as Similar factors have led to the near
riverside meadows were ploughed, drained extinction of the creamy-flowered subspecies
or otherwise ‘improved’, the water table fell ochroleuca, which is treated in the British Red
due to abstractions, or indeed meadows were Data Book and is a BAP priority species. It was
abandoned to scrub. first recorded from Roydon Fen (near Diss) in
Norfolk in 1936 and was subsequently found
at several fens in the Waveney Valley on the
Norfolk-Suffolk border and at Chippenham
Fen in Cambridgeshire. It is now extinct at all
sites with the exception of Chippenham Fen,
where it has declined from around 30 plants in
1990 to a single clump, and one of the Waveney
Valley fens in Suffolk, where it was rediscovered
in the late 1990s, and there are currently
around a dozen flowering plants.
The red-flowered dune subspecies
D. i. coccinea has suffered declines due to the
scrubbing-over of sand dunes and also coastal
development but still occurs in very large
numbers at some favoured sites. The spotted-
leaved Flecked Marsh Orchid D. i. cruenta is
also treated in the British Red Data Book and
is classified as Endangered. It is only found
at three sites in Scotland which, although
remote, may suffer from overgrazing. It is
rather commoner in Ireland but with the rapid
changes occurring there, including The Burren
region, it could be vulnerable to development
and ‘improvement’.
M 14 June, Suffolk. Subspecies ochroleuca. A well as

being a pale cream colour, the lip is very distinctly three-
lobed.
004 systematic.indd 287 29/1/09 12:33:14

frOG Orchid Vulnerable, BAP

Dactylorhiza viridis
Formerly: Coeloglossum viride; Other names: Bracted Green Orchis (North America)
The English names of many orchids make sense, and it is easy to see the ‘man’ in a Man
Orchid or the ‘monkey’ in a Monkey Orchid. Not so this species, for although the name has
been in use since the 17th century it is hard to see the ‘frog’ in a Frog Orchid; the two-lobed
lip could resemble hind legs, and the hood may look like a frog’s body, but to anyone except
the most imaginative any real resemblance to a frog is fanciful. Frog Orchid grows in short
grassland, frequently on chalk or limestone, but is inconspicuous and often hard to find. It
has undergone a serious decline and is now missing from much of its former range.
Identification
The small size, generally greenish or reddish-
purple flowers, tight hood and rather plain,
strap-shaped lip are distinctive.
Similar species
Of the vaguely similar species, Man Orchid has
long and narrow ‘arms’ and ‘legs’ on its lip and
Common Twayblade has a smaller flower, again
with ‘arms’ and ‘legs’.
Habitat
In southern Britain this species is confined
to well-drained short grassland on chalk or
limestone, especially the slopes of ancient
earthworks, abandoned quarries, old chalk
and lime pits and spoil heaps. It is tolerant of
grazing and trampling but cannot compete
with rank vegetation. In the past it was also
found in damp or wet permanent pastures and
meadows in southern England, but this habitat
has almost entirely vanished, taking the Frog
Orchid with it. In the north and west it is found
in a wider range of short-grass habitats, often
damp, and on both calcareous and neutral soils,
including limestone pavements, rocky ledges,
road verges, railway embankments, upland
flushes, mountain pastures, coastal grassland,
machair and dune slacks. It occurs from sea
level to 915m (Glen Doll, Angus).
O 29 June, Hampshire. Frog Orchid is ‘shy’ and the flow-

ers typically face downwards without showing their ‘face’.
004 systematic.indd 288 29/1/09 12:33:15

FROG ORCHID l 289
Flowering period
Early June to early August, sometimes from
late May or even until early September, but
usually peaking in late June and early July. Once
pollinated, the flowers persist for a long time,
although the lip will have withered.
Range
Frog Orchid is widely distributed in Britain
and Ireland, including the Inner and Outer
Hebrides, Orkney and Shetland, but has
declined severely in much of lowland Britain.
In southern England it is now almost confined
to the chalk districts of Hampshire, northeast
Dorset and Wiltshire, extending very locally
into the Berkshire Downs, Chilterns and the
South Downs in Sussex. In the Midlands it is
more-or-less restricted to the Peak District. It
is very local in Wales, lowland Scotland and
the southern half of Ireland (and absent from
southernmost Ireland from Co. Cork to Co.
Wicklow) but more widespread in northern
England, upland Scotland and the northern
half of Ireland. World range: Frog Orchid
has a circumpolar distribution and is found in
Europe, Asia and North America. It is mostly
confined to the temperate regions but extends
north to Iceland, the Faeroes, northernmost
Scandinavia and Alaska. It ranges furthest
south in the mountains, to Spain, Italy, Greece,
the Crimea and Caucasus (and adjacent
1987-99
1970-86
pre 1970 M 13 July, Dorset. Frog Orchid varies from overall green
to generally very reddish.
northeast Turkey); in Asia to Central Asia and

the eastern Himalayas; in North America to
Washington, New Mexico, Iowa and North
Carolina.
How to find it
Although variable, this species is often small,
green, rather inconspicuous and easy to tread
on. It can easily be overlooked; the first record
in Cornwall for over 50 years was when over
4,000 flower spikes were found near Colliford
Reservoir in 1986. It is also rather local in
004 systematic.indd 289 29/1/09 12:33:17

004 systematic.indd 290 29/1/09 12:33:20

FROG ORCHID l 291
occurrence, and the number of flowering plants washed brown or reddish-purple, in some the
can vary greatly from year to year. As with whole flower may be reddish-purple. The sepals
Musk Orchid, another short-turf specialist, it are dark green variably washed reddish-purple,
can be easiest to spot when scanning from a roughly oval to oval-triangular in shape. The
low-level vantage point. In southern England upper sepal is a little smaller than the lateral
small numbers are found at several chalk sepals. The petals are pale green, washed
grassland reserves from Dorset east to Sussex. reddish-purple around the edges and strap-
Notable sites in Scotland include North shaped, much smaller and narrower than the
Berwick and Crail golf courses which hold sepals. The sepals and petals form a tight hood
populations of many thousands. over the column, with the petals often showing
as much paler slots between the sepals. The lip
DESCRIPTION is usually paler and greener than the rest of the
Height: 4-45cm but mostly 5-15cm and only flower and is even paler and yellower towards
very exceptionally over 25cm. the base, with reddish tones usually confined
Stem: Green, on some plants washed reddish- to the tip and edges. It is tongue-shaped and
purple towards the flower spike. There are may broaden slightly towards the tip, with two
one or two brown, leafless sheaths at the small terminal lobes that have a notch between
extreme base. them and a third smaller lobe within this
Leaves: There are three to five dark green, notch. There is also a thickened ridge down the
strap-shaped sheathing leaves, sometimes rather centre of the lip, and nectar is secreted from
broad, with the two lowest leaves being the two hollows formed by the raised and curved
largest and bluntest. They are held at around margins of the lip at its base. The lip may hang
30°-45° above the horizontal. Higher on the downwards or be folded backwards below the
stem there are also several narrower and more ovary. The spur is very short (1-2mm), almost
lanceolate non-sheathing leaves. The rosette
appears in the autumn and overwinters, and
there may be a relatively large number of non-
flowering plants; these have fewer leaves.
Spike: Rather loose and often irregular in
shape, with the ovaries variably twisted and
the flowers therefore pointing in different
directions. Mostly there are 5-25 flowers but
sometimes only two, and particularly large
plants may have up to 50.
Bract: Green, variably washed reddish-purple
and lanceolate in shape; the lower bracts may
be rather longer than the flowers (up to a
maximum of twice the flower’s length), but
towards the tip of the spike the bracts
become shorter.
Ovary: Green, variably washed reddish-purple,
spindle-shaped, six-ribbed and twisted.
Flower: Green or yellowish-green, variably
O 29 June, Hampshire.
P 29 June, Hampshire. It is not easy to see the ‘frog’ in a

Frog Orchid.
004 systematic.indd 291 29/1/09 12:33:21

hemispherical in shape and rather colourless. first year above ground. Frog Orchid is short-
The anthers are washed purple, and the pollinia lived with a rapid turnover of the population.
are club-shaped and pale yellowish with a Many plants are monocarpic and die after just
very lobed surface. The flowers are faintly one year above ground, although some may live
honey-scented. and flower for at least seven years. In a study in
Holland the ‘half-life’ of a population averaged
Subspecies
1.5 years and varied from 1.0 to 2.4 years (the
None.
‘half-life’ is a measure of the life expectancy
Variation and varieties of an orchid after its first appearance above
On average shorter, stouter and more reddish- ground and marks the point at which 50% of
brown in the dry grassland habitats of southern the population which emerged in any given year
England and taller and greener in the north. have died). Occasionally, plants may be ‘dormant’
Some plants, including the flower spike, may be underground but not for more than one year.
entirely yellowish-green.
Var. longibracteatum has unusually long bracts Hybrids
and is rather taller and more robust. It has been D. x mixtum, the hybrid with Common
found in northern England. Spotted Orchid, is rare but has been found at
widely scattered localities.
BIOLOGY D. x conigerum, the hybrid with Heath Spotted
Orchid, has been found rarely (the two parent
Pollination and reproduction species do not occur together as frequently).
Unlike the other Dactylorhiza orchids, the D. x viridella, the hybrid with Northern Marsh
flowers produce nectar, and this is secreted Orchid, has been recorded from Co. Durham
into the short spur. A variety of insects visit and the Inner and Outer Hebrides.
the flowers, including small beetles and wasps,
especially ichneumons. These alight on the lip Intergeneric hybrids
and are directed by its central ridge to either X Dactylodenia jacksonii, the hybrid with
side; as they approach the spur, the pollinia fragrant orchid (in the broad sense) has been
are stuck to their head by the viscidia which lie noted sporadically but widely in England.
on either side of the spur. Once on the insect’s Name and classification
head it takes about 20 minutes for the pollinia The specific name viridis means ‘fresh-green’ or
to rotate forward. They are then in position to ‘youthful’.
make contact with the stigma of the next plant Until recently the Frog Orchid was named
to be visited. Self-pollination is also reported Coeloglossum viride, and indeed the genus
to occur. Seed-set is variable, with the capsules Coeloglossum contained just this one species.
maturing rapidly and containing around 1,250- However, genetic and biochemical techniques
5,000 seeds. This species only occasionally have shown that Coeloglossum is sufficiently
reproduces vegetatively. similar to the genus Dactylorhiza for the Frog
Development and growth Orchid to be transferred to it. This explains
The aerial stem grows from a pair of tubers why the Frog Orchid has such a propensity
that are forked into three or four finger-like to form hybrids with the other members of
lobes. They are typical of the genus Dactylorhiza the genus Dactylorhiza, the spotted orchids
although rather small. and marsh orchids. The chromosome number
The period between germination and is 2n=40, and Frog Orchid is one of the
flowering is short. The first green leaves appear more primitive Dactylorhiza. (The old name
one to three years after seed has germinated Coeloglossum means ‘hollow-tongue’, a reference
and a flower spike is usually produced in that to the short conical spur at the base of the lip.)
004 systematic.indd 292 29/1/09 12:33:21

FROG ORCHID l 293
Generally commoner in the north, especially

in Scotland, Frog Orchid is abundant at a
few favoured localities with hundreds or even
thousands of flower spikes. It is rather local,
however, even in Scotland and very much so in
the rest of Britain.
Past and present occurrence of Frog Orchid in Britain and

of the National Grid; data from the New Atlas).
Britain Ireland

1500-1999
current range 381 (13%*) 99 (9.8%*)
% lost, 1500-1969 49% 47%

% lost, 1970-1986 11.5% 6.5%
% lost, total 60.5% 53.5%
This species has declined throughout

almost the entire range in the British Isles
and in much of Europe too. The biggest losses
have been in England, especially the Midlands
and East Anglia, where the ploughing-up of
old pastures on neutral or chalky boulder-clay
soils appears to have been a major factor as
well as the development of scrub and rank
M 13 July, Dorset. Frog Orchid has recently joined the vegetation on what had been short grassland.
spotted and marsh orchids in the genus Dactylorhiza but Frog Orchid is extinct in Essex, Norfolk,
any similarity is not obvious.
Middlesex, Huntingdonshire, Worcestershire,
Warwickshire, Lancashire, Monmouthshire,
HISTORY AND Breconshire, Radnorshire, Pembrokeshire,
CONSERVATION Montgomeryshire, the Isle of Man,
Despite its inconspicuous nature, the Frog Dumfries-shire, Wigtownshire, Berwickshire,
Orchid was known to early naturalists and Kincardineshire and Dunbartonshire. It was
was first recorded in Hertfordshire in 1650 last seen in Kent in 1988, and just one site
by William How in his Phytologia Britannica each remains in Suffolk, Cambridgeshire,
natales exhibens Indigenarum Stirpium sponte Lincolnshire and Herefordshire. Not
emergentium (‘A British botany presenting surprisingly, it has recently been classified
the origins of native wild plants’): ‘Orchis as Vulnerable by the UK government’s
Batrachites…By Barkway, Dr. Johnson’s MS.’ conservation advisers.
The species was then listed in John Ray’s flora
of Cambridgeshire in 1660 and noted by
Christopher Merrett growing near Lewes in
Sussex and, ‘in many places about Oxford’ in his
Pinax of 1666.
004 systematic.indd 293 29/1/09 12:33:22

cOmmOn SpOtted Orchid

Dactylorhiza fuchsii
Formerly: Orchis fuschii, Dactylorchis fuschii

Aptly named, this species is both common and has boldly spotted leaves. Indeed, it is the
commonest and most widespread orchid in the British Isles and is found in a very wide
variety of habitats from open grassland to woodland and fen. An opportunist, it can colonise
new areas relatively easily and quickly and can sometimes appear in very large numbers;
the massed ranks of thousands of Common Spotted Orchids on a road verge or in an
abandoned quarry or chalk pit has to be one of the most spectacular sights of the summer.
Common Spotted Orchid is the county flower of West Lothian and Linlithgowshire and
the ‘Hebridean Spotted Orchid’ (subspecies hebridensis) is the county flower of the Western
Isles.
Identification
Identification is usually straightforward. The
leaves are marked all over with solid dark
spots or bars, the flowers are various shades of
pink or lilac (and are often very pale or almost
white), and the lip is divided into three roughly
equal lobes and decorated with bold dark lines
and loops. It is, however, very variable: the
leaves may be heavily or lightly marked; plants
with unspotted leaves are not uncommon;
and the flowers vary from white to rather dark
pinkish-purple. It tends to be tall, attenuated
and pale-flowered in shady woodland sites
but shorter and more compact in the open. It
thrives best in damp or wet habitats; plants on
dry chalk grassland can be very petite while
those in exposed, windswept sites may be short
and squat.
Similar species
Early Purple Orchid has spotted leaves, but
the lip is a very different shape and does not
have bold darker lines and loops. It also flowers
rather earlier in the spring.
Heath Spotted Orchid can be very similar
to Common Spotted Orchid but is separated
by its fewer, rather narrower, more obviously
keeled and more pointed leaves, with the lowest
leaf not significantly shorter or blunter than
the remainder. The spots on its leaves are often
M 6 July, Norfolk.The leaves are typically well-spotted. small and rounded, compared to the larger and
004 systematic.indd 294 29/1/09 12:33:23

COMMON SPOTTED ORCHID l 295
bolder marking of Common Spotted Orchid,

and plants with unspotted leaves are more
frequent. The lip of Heath Spotted Orchid
is broader and the lobes are very unequal in
size. The central lobe is rather small and barely
longer than the side-lobes, and the incisions
(sinuses) between the lobes cut much less than
halfway to the base of the lip. The lip is often
more faintly marked, with dots and fine dashes
rather than bold loops and lines, and the spur is
longer and rather finer.
Marsh orchids mostly have darker and
more purple flowers, although there is some
overlap, and often have unspotted leaves. Both
Common and Heath Spotted Orchids can
be distinguished from the marsh orchids by
having a spur that is slender and parallel-sided
rather than short, fat and bag-like (but beware
hybrids). The stem of the spotted orchids is
usually solid and ‘unsquashable’, whereas that of
some of the marsh orchids is obviously hollow.
However, Common Spotted Orchids growing
in wet habitats may have hollow stems, so this
is not always a safe distinction. The number of
non-sheathing leaves on the upper part of the
stem may also be helpful; often two to six in
Common Spotted (but sometimes just one) and
one or two in most marsh orchids.
Hybrids are common, particularly with
Southern and Northern Marsh Orchids.
Identification of hybrids requires a good
knowledge of the range of variation of both
species and even then is not always certain. M 22 June, Norfolk. A petite, few-flowered plant.
First generation hybrids usually show ‘hybrid
vigour’ and may be obviously large, sometimes conditions in wet meadows, the machair of
ridiculously so. Hybrids with marsh orchids northwest Scotland, marshes, fens and dune
often have the fat, conical spur of the marsh slacks, including alkaline flushes in otherwise
orchid parent rather than the thin spur of the unsuitable moorland. The species is also found
spotted orchid but in subsequent generations in woodland, usually in the better-lit areas
back-crossed with the parent species, or in hybrid along rides, clearings and edges, although it
swarms, these distinctions will break down. can tolerate quite deep shade where it may
persist in a non-flowering state. Common
Habitat Spotted Orchid is usually found on alkaline or
Common Spotted Orchid favours a very wide neutral soils but it can grow in slightly acidic
range of habitats. It occurs on dry grassland, conditions, such as among grass and Heather
from heavily grazed downland swards to ranker on less acid areas of heathland. An opportunist,
and scrubbier sites, and also thrives in damper it is capable of colonising new sites, including
004 systematic.indd 295 29/1/09 12:33:24

man-made habitats such as abandoned gravel

and chalk pits and industrial waste sites (e.g.
fly ash tips and alkaline Leblanc waste). It is
often found on roadside verges and cuttings,
including motorways, on railway embankments
and sometimes on lawns, and can quickly build
up in numbers to form substantial populations.
Recorded up to 530m above sea level in
Cumbria (Garrigill) and 600m in Perthshire
(Loch na Lairige).
Flowering period
Mid-May to early August, being earliest in
open, sheltered localities in the south (where
generally best around mid-June) and later in
woodland sites and in the north.
Range
Found throughout Britain and Ireland,
including the Isle of Man, Inner and Outer
Hebrides and Shetland, and also the Channel
Islands. However, it is absent from most of
Cornwall, much of north Devon and from
large areas of northern and northeast Scotland.
Formerly recorded from Orkney and the Isles
of Scilly. World range: Widespread in Europe
away from the Mediterranean lowlands, south
to Spain, northern Italy and Greece, north to
Scandinavia and Finland and east to Russia
and Siberia. The precise boundaries of the
distribution are obscure due to confusion with
Heath Spotted Orchid. M 20 June, Norfolk. ‘Typical’ coloration.
How to find it
1987-99
1970-86 In much of Britain and Ireland this is the
pre 1970
commonest orchid, and it is often found in large
numbers. Nevertheless, there are large tracts of
countryside that are not graced by its presence.
DESCRIPTION
Stem: Pale green, sometimes lightly washed
with purple towards the tip, grooved and
usually solid (but can be hollow, especially in
larger plants in wetland habitats).
Leaves: Green, usually marked all over with
elongated solid dark spots and blotches but
sometimes unmarked. There is a relatively
small, oval, blunt-tipped leaf at the base of the
004 systematic.indd 296 29/1/09 12:33:27

green, often with a violet tinge.

Flower: Various shades of pale lilac, pink or
purplish-pink or sometimes almost white. The
sepals are oval-lanceolate with the lateral sepals
asymmetrical, marked with lines and spots
and usually held spreading horizontally (but
they may be closer to 45°). The upper sepal
and the petals, which are a little shorter, form
a hood over the column. The lip is a flattened
oval in shape, 8-12mm wide, divided into three
lobes by deep wedge-shaped incisions (sinuses)
that extend about halfway to the base. The
side-lobes have broad rounded tips that are
sometimes toothed with roughly parallel sides.
The central lobe is slightly narrower and more
triangular in shape but as long or longer than
the side-lobes. The lip is held flat or with the
side-lobes lightly depressed. It is usually paler
and whiter towards the centre and is marked
with bold dashes and broken dark pink or dark
reddish-purple lines which often form a pattern
of concentric double loops (occasionally just
dots and dashes). The spur is about half as
long as the ovary to roughly the same length,
narrow, cylindrical, slightly tapering and either
straight or very slightly curved. The pollinia are
pale brownish-pink to purple, or sometimes
yellowish. The flowers are faintly scented.
Subspecies
M 6 July, Norfolk. A relatively pale spike.
D. f. fuchsii is the commonest and by far the
stem and the three to six (sometimes two to most widespread subspecies.
seven) sheathing leaves are crowded together D. f. hebridensis ‘Hebridean Spotted Orchid’ is
above this; they are broad, up to 4cm or even characterised by dark flowers and a broad lip.
5.5cm wide, and lanceolate. The two to six It is found on the machair and similar coastal
(sometimes one to nine) non-sheathing leaves habitats in the Outer Hebrides, northwest
higher on the stem become narrower and more Scotland, Shetland and western Ireland (Co.
bract-like towards the spike. Donegal, Co. Galway and Co. Kerry). It tends
Spike: Variable in shape but often pyramidal or to flower relatively late and often occurs in large
conical as the flowers begin to open, becoming numbers. In some areas overlaps and mixes
longer and more cylindrical later on, with 20-70 with subspecies fuchsii.
flowers, exceptionally 150. Hebridean Spotted Orchid is small and
Bract: Green, sometimes washed purple, stocky, 8-20cm tall (range 6-40cm), with
lanceolate and finely pointed; the lowest are the upper stem usually washed purple and
longer than the ovaries but higher on the spike the leaves heavily spotted. The spike is often
they are about equal in length. pyramidal or conical and densely packed, the
Ovary: Cylindrical, six-ribbed, twisted and lip is usually over 9.5mm wide (8-15mm), and
004 systematic.indd 297 29/1/09 12:33:28

004 systematic.indd 298 29/1/09 12:33:31
the flowers are often deep rose-pink or reddish-

purple (rarely white) with the spur relatively
long at 7-8.5mm. It is sometimes treated as
a separate species by European authors but
its distinctiveness has been exaggerated and,
conversely, it is also sometimes treated as merely
‘var. hebridensis’.
Woodland and grassland plants are sometimes
separated as distinct varieties but the
differences between the tall leafy plants found
in the shade of woodland and scrub and the
shorter plants of sunny grassland (the so-called
var. ‘trilobata’) seem to be purely a product of
their different growing conditions.
Var. alpina is a dark-flowered variant of
subspecies fuchsii. The dark flowers recall
those of subspecies hebridensis but they are
smaller, with the lip usually less than 9.5mm
wide and with narrower side-lobes. It is found
in Scotland, both at inland sites and on the
machair on the Hebrides and northwest coast
(including some populations in the Inner
Hebrides traditionally called ‘hebridensis’). It is
also present at inland sites in northern England
and possibly also Wales and western Ireland.
Var. cornubiensis is very similar to subspecies
hebridensis and essentially differs only
statistically. It is a little bigger and more
robust, with a larger and looser flower spike,
longer bracts (often over 8mm long) and a
narrower lip. The lateral sepals are rarely held
horizontally, and the spur is stouter. Confined M 10 June, Norfolk. A particularly richly-coloured spike.
to Cornwall, where it is found on the north O 11 July, Norfolk.
coast on stabilised dunes at Lelant golf course
and cliffs near St Ives and at Tintagel. It is found along the western seaboard of
Var. albiflora has unmarked white flowers Ireland, on the Isle of Man and in northern
(rarely washed cream or pink) and unspotted Britain; individual white-flowered plants within
leaves. It is widespread but rather uncommon. these populations are well-nigh identical to var.
Var. okellyi has narrow leaves and flowers albiflora (and vice-versa).
which are sometimes a little smaller or broader A lot of controversy surrounds okellyi with
than typical plants and often almost white or some European authors treating it as a distinct
creamy with very faint pink or lilac markings, species and other botanists treating it merely
although in some areas a substantial proportion as a poorly defined variety. The classic okellyi
(often the majority) have darker and heavier of the literature always has white, almost un-
markings; the leaves are spotted or unspotted. marked flowers and is said to be confined to
004 systematic.indd 299 29/1/09 12:33:33

western Ireland (especially The Burren region of BIOLOGY

Co. Clare and Co. Galway), the Isle of Man and
the west coast of Kintyre in western Scotland Pollination and reproduction
(with single records for Tiree and Sutherland). A wide variety of insects has been recorded
However, in The Burren and elsewhere these carrying off pollinia from this species, but
classic white-flowered okellyi are just part of a studies in Austria and Poland have identified
population of plants with a variable flower colour. beetles, especially longhorn beetles, as the major
Var. rhodochila is an attractive hyperchromic pollinators. In common with other members
variant with an excess of pigmentation. The of the genus Dactylorhiza, Common Spotted
lip is solidly reddish or purple, usually with a Orchid does not produce ‘nectar’. Insects may
narrow paler pink or white border. The leaves therefore be deceived into visiting the flowers,
may be more heavily spotted or in extreme cases although Charles Darwin suggested that
entirely washed purple. It is widespread but rare. visiting insects feed on sap sucked from inside
the wall of the spur. This hypothesis has since
been disputed, and it has been suggested that
they feed instead on tiny sugar-rich papillae
that are present in a zone from the middle of
the lip down into the spur. A third option is
that bumblebees receive no reward and are
deceived into pollinating the flowers but that
honeybees are able to access a sugary liquid
produced on the surface of the stigma.
Whatever the mechanism, pollination is
efficient and around 50-90% of flowers produce
seed. Notably, twice as many of the lower
flowers, which open first, produce ripe capsules,
compared with the upper flowers, which open
later on. This suggests that as the season
progresses insect pollinators become scarcer
or lose interest as there is little or no reward.
Vegetative reproduction is also possible.
The length of the period between germination
and flowering in the wild is unknown but
in cultivation it can be as little as two years
although it is more often four or five years.
It is thought that the plant can flower for
several years in a row or remain as a dormant
non-flowering rosette if conditions become
unsuitable.
Hybrids
Sterile hybrids are likely to be found either
singly or in very small numbers, whereas even
partial fertility can result in large numbers of
hybrids which in turn can show a great deal of
M 6 July, Norfolk. A few-flowered spike. individual variation.
004 systematic.indd 300 29/1/09 12:33:34

D. x transiens, the hybrid with Heath Spotted

Orchid, is widespread but scarce and can be
hard to identify with certainty. It is sterile. See
also Name and classification.
D. x kernerorum, the hybrid with Early Marsh
Orchid, has been found scattered throughout
Britain and Ireland. It is sterile.
D. x mixtum, the hybrid with Frog Orchid, is
rare but there are widely scattered records.
D. x venusta, the hybrid with Northern Marsh
Orchid, occurs throughout the range of the
latter and can be common. It is partially fertile
and can back-cross with either parent species
to produce a range of characters in a hybrid
swarm.
D. x grandis, the hybrid with Southern Marsh
Orchid, can be found not only where both
species occur together but also in the absence
of one or even both parents. The pollen is
highly sterile and seed production is very low.
Nevertheless this partial fertility allows back-
crossing with both parents, creating hybrid
swarms which may be very persistent. It is
probably the commonest orchid hybrid in
southern Britain.
D. x silvae-gabretae, the hybrid with Pugsley’s
Marsh Orchid, has been recorded from
northwest Wales, Yorkshire and Ireland. The
pollen is highly sterile.
‘D. x braunii’, the hybrid with Irish Marsh
Orchid, has been recorded in Co. Clare.
(The name may be changed following the
elevation of Irish Marsh Orchid to the status
of a full species.)
X Dactylodenia st-quintinii, the hybrid with
fragrant orchid, is uncommon and scarce but
has been found widely through Britain and
Ireland. All three species of fragrant orchid may
be involved.
The specific name fuchsii commemorates
Leonard Fuchs (1501-66), a German botanist
who is also celebrated in the name ‘Fuchsia’.
M 10 June, Norfolk. Presumably Common Spotted Orchid
Common Spotted Orchid is closely related x Southern Marsh Orchid. Such hybrids can sometimes
to Heath Spotted Orchid and although they occur where one of the parents is absent.
004 systematic.indd 301 29/1/09 12:33:36

between the two are not absolute. Common

Spotted Orchid is generally found on calcareous
or neutral soils and Heath Spotted Orchid
on acid soils. However, there is some evidence
that spotted orchids growing on intermediate,
neutral or slightly acidic soils do tend to be
intermediate in appearance.
HISTORY AND
CONSERVATION
Although spotted orchids had been known
from the time of Turner’s Herball of 1562,
this species was not distinguished from the
Heath Spotted Orchid until 1915 when it
was described by the English botanist George
Claridge Druce.
This is the most widespread species of
orchid in the British Isles and is currently
present in 67% of the total available 10km
squares in both Britain and Ireland. There
has been some decline but Common Spotted
Orchid is so versatile that it can usually ‘hang-
on’ somewhere within any particular 10km
square. The New Atlas figures are thus likely
to conceal a significantly greater fall in the
number of actual populations. Notably, much
of the decline in Britain appears to be recent
and it could be that a fall in the number of
populations is increasingly expressing itself as
a decline in the overall range. At least some of
the losses are compensated for, however, by the
colonisation of new areas, especially in recent
man-made habitats.
M 30 June, Norfolk. A densely-flowered spike; as all the Past and present occurrence of Common Spotted Orchid
flowers open the spike typically becomes more cylindrical. in Britain and Ireland (based on presence or absence in
are clearly distinct in Britain, France and Atlas).
Scandinavia they are rather similar in central
Britain Ireland
Europe and are considered to be the same
species by many European authors. They are total historical range, 2,219 774
usually quoted as differing in the number 1500-1999
of chromosomes, with Common Spotted current range 1,913 (67%*) 673 (67%*)
Orchid having 2n=40 and Heath Spotted % lost, 1500-1969 7% 12%
Orchid 2n=80, but various studies have given
% lost, 1970-1986 7% 1%
chromosome counts of 2n=40, 2n=60 and
2n=80 for both species (i.e. diploid, triploid
and tetraploid). Even in Britain the differences * current range as a % of the total number of 10km squares
004 systematic.indd 302 29/1/09 12:33:37

HEATH SPOTTED ORCHID l 303
heath SpOtted Orchid

Dactylorhiza maculata
Formerly: Orchis ericetorum, Dactylorchis ericetorum

Closely related to Common Spotted Orchid, this species replaces it in more acid, heathy
habitats. It is similarly very widespread but has a northern and western bias to its range and
is now absent from much of the Midlands and southeast England. The flowers are usually
rather pale pink with delicate markings, and the lip is broad, resembling a frilled skirt. It can
occur in large numbers and gives a real splash of colour to otherwise sombre tracts of moor
or bog.
Identification
A dainty and very attractive orchid, variable in
stature but often just 10-20cm high, especially
in exposed or relatively dry sites. The flowers
are very variable but are often a pale shade of
pink or pinkish-lilac and marked with fine dots
and dashes, the ground colour frequently fading
to whitish as they age. The broad lip resembles
a voluminous petticoat and its side-lobes are
much larger than the small, tooth-like central
lobe and often have serrated or frilly edges.
The leaves are usually marked with numerous
dark spots.
Similar species
Common Spotted Orchid is rather similar
but can be separated by its broader and flatter
leaves with the lowest leaf usually significantly
shorter than the rest with a broad, rounded tip.
In both species the leaves are usually spotted
but in Common Spotted the spots are often
larger, bolder and usually elongated into short
bars rather than being rounded. In Common
Spotted the lip is more deeply lobed with the
incisions (sinuses) between the lobes cutting
around halfway to the base of the lip and the
three lobes more equal in size; the central lobe
is at least half the width of the side-lobes and
usually a little longer too. Finally, in Common
Spotted Orchid the lip is marked with loops
and lines rather than dots and the spur is
shorter, thicker and tapers slightly towards
the tip.
P 5 July, Norfolk.
004 systematic.indd 303 29/1/09 12:33:39

Marsh orchids are usually easily separated

as most have darker and more boldly marked
flowers than is typical in Heath Spotted
Orchids. Sometimes, however, Heath Spotted
may have darker and more purplish flowers
and, especially if the leaves are unspotted,
may superficially resemble a marsh orchid.
In case of doubt check the spur: long, slender
and roughly parallel-sided in Heath Spotted
and squat and sack-like in marsh orchids.
Also check the number of non-sheathing
leaves: often two to five in Heath Spotted (but
sometimes just one) and one or two in most
marsh orchids. Finally, the stem of the spotted
orchids is usually solid and ‘unsquashable’ and
that of at least some of the marsh orchids is
obviously hollow.
Hybrids are common, particularly with
Northern and Irish Marsh Orchids. First
generation hybrids usually show hybrid vigour
and may be obviously large. Hybrids with
marsh orchids often have the fat, conical spur
of the marsh orchid parent rather than the thin
spur of the spotted orchid. Later generations
of hybrids, if back-crossed with one or other of
the parent species, become harder and harder
to determine.
Habitat
Heath Spotted Orchid is found in a wide
variety of habitats. It favours heathland and
moorland, especially the damper, more peaty
areas on the margins of valley bogs, flushes and
mires and the raised and slightly drier areas M 15 June, New Forest. A petite but richly marked plant.
within Sphagnum bogs; suitable areas are often
picked out by the greyish foliage and dusty-pink Orchid is not as tolerant of shade as Common
flowers of Cross-leaved Heath. It is also found Spotted and is much less commonly found in
on acid grassland (for example rhos pastures light woodland and even then usually on the
in Wales and culm grassland in southwest edge or along rides rather than in dense shade.
England), damp unimproved meadows and hill Recorded from sea level up to 915m (Ben
pastures. It usually grows on mildly acidic soils Lawers, Perthshire).
and in areas of chalk or limestone bedrock;
suitably acidic conditions may be provided by Flowering period
pockets or blankets of peat or areas of drift Mid-May to July, sometimes from the second
(superficial deposits of sands and gravels). week of May or rarely to early August. Typically
It may sometimes grow in neutral or even it flowers earliest in drier situations and later
slightly alkaline marshes and fens and in such in wetter habitats. The flowering time relative
habitats can be very robust. Heath Spotted to Common Spotted Orchid varies in different
004 systematic.indd 304 29/1/09 12:33:40

leaves crowded towards the base of the stem.

1987-99
1970-86
These are long, narrow, keeled and pointed,
pre 1970 the largest up to 20mm wide (more rarely
to 25mm) with the tips sometimes hooded.
There are also two to five (one to seven) non-
sheathing leaves and these become more bract-
like towards the flower spike. All the leaves are
green, variably marked with solid dark spots
which are rounded or only slightly elongated;
the leaves are fairly frequently unspotted.
Spike: Usually rounded or pyramidal, even
when all the flowers are open, with 5-20
flowers, sometimes up to 60 on robust plants.
Bract: Green, often washed purple around the
edges and towards the tip, lanceolate, tapering
to a fine point, one-and-a half to two times
longer than the ovary on the lower flowers but
parts of the country and can be later or earlier, around equal in length higher up.
perhaps depending on their respective habitat Ovary: Green, cylindrical, six-ribbed and
in the area. twisted.
Range
Found throughout Britain and Ireland,
including the Scottish Islands, Isle of Man and
the Channel Islands, but with a predominantly
northern and western bias to the distribution.
World range: Western Europe, north to
Iceland, the Faeroes, Scandinavia, Finland and
the Baltic States, east to Russia and south to
northern Portugal and Spain, Italy, the former
Yugoslavia and Bulgaria. As with Common
Spotted Orchid, dispute over the exact
definition of the species makes it hard to be
precise about the boundaries of the range.
How to find it
Usually common in suitable habitats, the delicate
pale pink blooms stand out among the grasses
and heathers that typically surround it.
DESCRIPTION
Height: 4-50cm but usually 10-25cm. Plants
have been noted up to 75cm tall but such
robust individuals are probably hybrids.
Stem: Pale green, washed purple towards the
tip and slightly ridged.
Leaves: There are one or no basal leaves (or M 21 July, Norfolk. A relatively few-flowered spike, with
sheaths) and two to four (one to five) sheathing sparse but bold markings.
004 systematic.indd 305 29/1/09 12:33:43

that are usually rather fine and form a pattern

of concentric loops or circles but are sometimes
bolder and form the clear double loops more
typical of Common Spotted Orchid. The spur
is slender, cylindrical or slightly tapering, about
half as long as the ovary to roughly the same
length and straight or slightly decurved. The
flowers are faintly scented.
Subspecies
British and Irish plants are conventionally
treated as belonging to subspecies ericetorum
which also occurs along the Atlantic seaboard
of Europe from northern Portugal to northern
Scandinavia and Finland. The typical
subspecies, D. m. maculata, is found in the
remainder of continental Europe.
Within the British Isles there is much
variation but it does not show any consistent
pattern, and no other subspecies can usefully be
recognised, even though some European authors
state that subspecies maculata and/or subspecies
elodes occur here (and some split the spotted
orchids into many different species and treat
M 21 July, Norfolk. The flower often has a ‘blousy’ look subspecies maculata and ericetorum as different
and the central lobe of the lip is much smaller than the
side lobes.
species, stating that both occur in Britain).
Plants from Rhum in the Inner Hebrides
Flower: Usually whitish to pale pink or with dark flowers intermediate in shape
pale lilac, sometimes purplish-pink, with
dark reddish markings. The lateral sepals
are lanceolate, asymmetrical, marked with
darker spots and lines and held horizontally
or drooping. The upper sepal and petals are
slightly shorter and form a hood over the
column. The lip is shorter than wide (around
5-9.5mm long x 6.5-13mm wide), roughly
circular or circular-diamond shaped. It is
divided into three lobes by relatively shallow,
triangular notches (sinuses). The side-lobes
are much larger than the central lobe and their
edges may be wavy or toothed. The central lobe
is small, triangular, rather tooth-like and does
not usually project beyond the side-lobes (if
it does, it rarely projects by more than 1mm).
The lip is held flat, slightly dished or with
the side-lobes slightly deflexed. It is whitish
to pink, sometimes deep lilac-pink, variably
marked all over with darker dots and dashes M 21 July, Norfolk. A boldly marked plant.
004 systematic.indd 306 29/1/09 12:33:44

between Heath and Common Spotted Orchids

were described as subspecies rhoumensis. But
the population on Rhum is very variable and
such dark, three-lobed flowers occur elsewhere.
It is not worthy of recognition.
Var. concolor has an excess of pigmentation.
It has a solidly dark reddish-purple lip and
the leaves may also be washed purple. It is
extremely rare, with just one or two records.
Var. leucantha has pure white flowers. It is
widespread but rare.
BIOLOGY
A variety of insects have been observed carrying
pollinia, especially bees and flies, and seed
production is good (45% of flowers set seed
in one Scottish study). The mechanism is
probably similar to Common Spotted Orchid,
although the pollinia are relatively small,
possibly an adaptation to smaller and less
robust pollinating insects.
The pattern of development is probably similar
to Common Spotted Orchid, with seedlings
appearing above ground less than two years
after germination and the first flower spike
three years later.
Hybrids M 20 June, Norfolk. The white flowered var. leucantha is
widespread but rare.
Sterile hybrids are likely to be found either singly
or in very small numbers, whereas even partial can therefore create hybrid swarms.
fertility can result in large populations of hybrids D. x formosa, the hybrid with Northern
displaying a great deal of individual variation. Marsh Orchid, occurs throughout the range
D. x transiens, the hybrid with Common of Northern Marsh Orchid. It is fertile and
Spotted Orchid, is widespread but scarce. It is frequently back-crosses with the parents to
highly sterile. form hybrid swarms. This is probably the
D. x carnea, the hybrid with Early Marsh commonest hybrid orchid in northern Britain
Orchid, has been recorded scattered in Britain and Ireland.
and Ireland but is rare. It is probably sterile. D. x jenensis, the hybrid with Pugsley’s Marsh
D. x conigerum, the hybrid with Frog Orchid, Orchid, has been recorded from northwest
has been found rarely but widely. Wales, Yorkshire and Ireland. It appears to be
D. x hallii, the hybrid with Southern Marsh sterile.
Orchid, occurs infrequently as the parent D. x dinglensis, the highly fertile hybrid with
species favour different habitats. It is fertile and Irish Marsh Orchid, has been found widely.
004 systematic.indd 307 29/1/09 12:33:46

of disagreement over the number of species

involved. Some European authors treat them
both as the same species, and there is some
support for this argument as the differences
between the two are not as clear cut in parts
of Europe as they are in the British Isles. Even
in Britain the differences are not as absolute as
some field guides suggest (see also discussion
under Common Spotted Orchid).
HISTORY AND
CONSERVATION
The first British record of a spotted orchid
dates from 1562 when the second part of
William Turner’s Herball noted: ‘There are
divers kindes of orchis…one kinde…hath many
spottes in the leaf and is called adder grasse in
Northumberland’.
One of the commonest and most
widespread orchids in the British Isles. There
has been some decline, however, especially in
lowland areas in England, and the species now
occupies only around three-quarters of its
historic range (and many of the losses in Britain
are relatively recent). Agricultural changes,
leading to the ploughing and ‘improvement’
of meadows and pastures and the destruction
of heaths have caused some losses, as has
the abandonment of heathland causing it to
M 20 June, Norfolk. Hybrid Heath Spotted Orchid x scrub-over and turn into woodland; extensive
Common Spotted Orchid. Surprisingly infrequent, such heath fires can also be damaging. The species
hybrids are typically very robust. tolerates, and even benefits from, some grazing
Intergeneric hybrids but in northern Britain it has, like much native
X Pseudorhiza bruniana, the hybrid with vegetation, suffered from overgrazing.
Small White Orchid, was recorded from Past and present occurrence of Heath Spotted Orchid in
Orkney in 1977. Britain and Ireland (based on presence or absence in 10km
X Dactylodenia legrandiana, the hybrid with squares of the National Grid; data from the New Atlas).
fragrant orchid (presumably x Heath Fragrant Britain Ireland
Orchid), has been found widely scattered
through Britain and Ireland. total historical range, 2,025 687
1500-1999
Name and classification current range 1,587 (56%*) 531 (53%*)
The specific name maculata means ‘spotted’ or
% lost, 1500-1969 12.5% 20.5%
‘blotched’.
% lost, 1970-1986 9% 2%
Heath and Common Spotted Orchids are
members of a complex group of Dactylorhiza % lost, total 21.5% 22.5%
orchids which have generated a great deal * current range as a % of the total number of 10km squares
004 systematic.indd 308 29/1/09 12:33:47

SOUTHERN MARSH ORCHID l 309
SOuthern marSh Orchid

Dactylorhiza praetermissa
Formerly: Orchis praetermissa, Dactylorchis praetermissa

This species often occurs in large numbers and can produce a spectacular show in riverside
meadows and pastures. Locally common and widespread in much of England (south and
east of a line from the Ribble to the Humber) and in south and southwest Wales, it can
tolerate a wide variety of conditions, not always ‘marshy’, and is even found on certain
favoured brownfield sites where a variety of chemical wastes have been dumped. In size and
flower colour it is rather variable, and it also hybridises readily with the spotted orchids,
making identification difficult at times.
Identification
In its typical form Southern Marsh Orchid is
distinctive. It is usually a robust orchid with a
stout stem, numerous rather broad, unmarked
green leaves and a spike of purplish-pink
flowers. The lip is broad and rounded with a
tooth-like central lobe; to judge the shape of the
lip it needs to be flattened out from below, using
a finger or a ruler. In its natural pose the lip may
be held flat, slightly dished or with the sides
turned downwards, and it is marked in a central
zone with fine dots and short dashes.
Southern Marsh Orchid is, however, rather
variable. Most plants have flowers that appear
a little ‘dusty’ or washed-out, but they may be a
darker and more intense colour or have bolder
and darker markings on the lip. In addition,
two varieties of Southern Marsh Orchid are
markedly different: var. junialis, the so-called
Leopard Marsh Orchid has large ring-shaped
spots on the leaves and a boldly marked flower
and var. bowmanii has a very prominently three-
lobed lip in various shades of pink with bold,
dark dashes and loops (recalling Common
Spotted Orchid in lip shape and pattern).
Similar species
Early Marsh Orchid is usually easily
distinguished by flower colour alone (very pale
pink, deep red or creamy). The various purple-
flowered forms of Early Marsh Orchids can be
very similar in colour to Southern Marsh but M 22 June, Norfolk. Southern Marsh Orchid can be tall,
always have smaller flowers with a narrower slender and stately.
004 systematic.indd 309 29/1/09 12:33:48

lip that usually has the side-lobes clearly this, too, has spotted leaves (see below for more
folded downwards and prominent double-loop details). Southern Marsh Orchid also freely
markings. hybridises with Pugsley’s Marsh Orchid, and as
Northern Marsh Orchid usually has darker these hybrids are fully fertile, back-crossing can
and redder flowers (deep magenta), a smaller, lead to a range of plants intermediate between
more angular, diamond-shaped lip with heavier, the two parent species.
more blotched or looped dark markings and
often fewer non-sheathing leaves. Habitat
Pugsley’s Marsh Orchid has a more slender Found in a variety of habitats on calcareous
stem, fewer, narrower leaves, and a rather loose, to neutral or even acidic soils that are usually
one-sided spike composed of a few, relatively (but not always) moist or wet. Typical habitats
large flowers. include damp meadows, fens, marshes, the less
Hybrids are a major identification headache. acid parts of bogs and wet heathland, dune
Southern Marsh Orchid frequently hybridises slacks, marshy gravel pits, road verges and also
with Common Spotted Orchid, and the hybrids old industrial sites, especially in northwest
are usually tall and robust with the sheathing England and the West Midlands (waste alkali,
leaves shorter and blunter than Southern colliery and fly-ash tips). It is also found in old
Marsh Orchid and the non-sheathing leaves chalk quarries where compacted ground may
narrower and more pointed. Importantly, lead to water-logging, and it occasionally grows,
the leaves are usually faintly spotted, and often in a dwarf form, on dry chalk grassland
the flowers are intermediate between the and downs. Perhaps, surprisingly, Southern
two parents. These hybrids can be harder to Marsh Orchid cannot tolerate being submerged
distinguish from Leopard Marsh Orchid as for long periods, and winter floods lasting more
M 21 June, Suffolk. Often occurs in large numbers.

P 6 June, Norfolk.
004 systematic.indd 310 29/1/09 12:33:50

l 311
004 systematic.indd 311 29/1/09 12:33:54

O 17 June, Norfolk. Southern Marsh

Orchid can be short and squat.
than a month may cause a severe decline or even North America, where it has been present for
kill all the plants. many years at Tilt Cove, Newfoundland. It was
probably accidentally imported with mining
Flowering period
equipment from Europe in the early 1900s.
Late May to early July, occasionally from
mid-May to mid-July or even to early August. How to find it
Southern Marsh Orchid is fairly common and
1987-99 sometimes occurs in large numbers, although
1970-86
pre 1970 in the modern agricultural landscape it is
necessarily local. Big colonies stand out but
odd plants in marshy meadows may be easy to
miss; the presence of quantities of rushes is a
good clue.
DESCRIPTION
Height: 20-50cm, occasionally to 70cm or even
95cm (the largest plants possibly show some
hybrid influence). Dwarfed plants, just 10cm
high, may occur in marginal habitats such as
chalk grassland.
Stem: Stout (usually over 5mm in diameter),
hollow and green.
Leaves: Mid-green, sometimes slightly tinged
Range greyish-green. There are three to nine sheathing
Widespread in England roughly south of a leaves (typically four to six), held erect at 45° or
line from the Ribble to the Humber with a few more and often slightly more crowded towards
scattered colonies a little to the north of this. It the base of the stem. Oblong-lanceolate, the
is also found in south and southwest Wales, the leaves are more-or-less flat, lack a marked keel
Isles of Scilly and the Channel Islands. World and may be slightly hooded at the tips. The
range: A European endemic found along the largest are usually 2-3.5cm wide but they can
Atlantic fringe of the continent from northern be as narrow as 1.5cm. There are one to three
France through Belgium, Holland, Germany narrower and more pointed non-sheathing
and Denmark to southwest Norway (and leaves higher on the stem.
possibly also Sweden and Finland); reported Spike: Rather crowded and occupies around
recently from northern Italy. Introduced to 20% of the stem with 20-60 flowers, although
004 systematic.indd 312 29/1/09 12:33:55

M 22 June, Norfolk. The markings on the lip are very variable but tend to be relatively fine, short dashes and dots
concentrated in the central zone.
large plants may have over 100. Conical as the lobe which usually projects slightly; there are
first flowers open, becoming cylindrical later on. shallow incisions (sinuses) between the lobes.
Bract: Green, often washed purple. The bracts The side-lobes turn downwards at the base but
are around twice the length of the ovary and are often level or turn upwards towards the
project beyond the flowers (although usually tip of the lip, which therefore appears drop-
not as prominently as in Early Marsh shaped or like a well-rounded triangle in its
Orchid and less obviously so once the spike natural position. It is purplish-pink, becoming
is fully open). paler towards the base and then white around
Ovary: Green, variably washed purple, the mouth of the spur, and is marked with
cylindrical, six-ribbed and twisted. fine dark spots and short dashes, usually
Flower: Purplish-pink but variable in its exact concentrated in the centre. The spur is two-
shade; sometimes a deep, intense colour but thirds to four-fifths the length of the ovary,
usually a little ‘washed-out’. The lateral sepals stout, tapering slightly to a blunt tip and may
are lanceolate, asymmetrical and occasionally be slightly down-curved.
blotched darker. They are variably positioned,
Subspecies
from nearly vertical over the flower to nearly
None.
horizontal but usually held at around 45°. The
upper sepal and petals are lanceolate, the petals Variation and varieties
slightly shorter than the sepals, and together Var. junialis ‘Leopard Marsh Orchid’
they form a loose hood over the column. The has leaves marked with large oval spots or
lip is 9-12mm long x 8.5-14mm wide (but sometimes bars which are hollow, and its
mostly more than 10mm wide). Overall it is lip is marked with unbroken dark loops or
roughly circular in shape (if slightly ‘squashed’) horseshoe-shaped marks rather than just dots
and subtly three-lobed, with broad, rounded and dashes. It is found most often in the south
side-lobes and a small, often tooth-like central and southeast but is uncommon.
004 systematic.indd 313 29/1/09 12:33:57

nearby, suggests, however, that junialis is a

variety of Southern Marsh, and this has been
supported by genetic evidence.
Both Leopard Marsh Orchid and hybrids
can have hollow, ring-shaped spots on the
leaves, and the presence of such annular spots
is not diagnostic of Leopard Marsh Orchid,
although hybrids are more likely to have solid
dark spots. The following characters are useful
in distinguishing Leopard Marsh Orchid from
ring-spotted hybrids:
Size: Relatively small and slender, with fewer
leaves (averaging smaller and more slender than
normal Southern Marsh Orchids). Hybrids are
often robust, with many leaves, especially many
non-sheathing leaves.
Lip size: On average narrower, up to 10mm
wide. Hybrids have broader lips, 10mm or more
– sometimes much more.
Lip shape: Often more obviously three-lobed
than typical Southern Marsh Orchids but not
as distinctly three-lobed as the hybrid.
Lip markings: Confined to the central zone,
within the heavy double loop, with just one or
two faint markings outside this area. In hybrids
heavy markings extend over most of the lip.
Var. bowmanii is relatively slender, with slightly
fewer, narrower leaves than typical Southern
Marsh Orchid (three or four sheathing leaves
and one to three non-sheathing leaves). The
flowers have a prominently three-lobed lip with
a very long central lobe and heavy line and loop
markings. Recorded from marshy areas or dry
grassland on acid soils in Devon, Dorset and
Hampshire. Initially described as a variety of
M 22 June, Norfolk. A ‘Leopard Marsh Orchid’ with bold
Pugsley’s Marsh Orchid but clearly outside the
loop markings on the lip and ring-spotted leaves; it is easy range of that species and genetic evidence places
to confuse this variety with hybrid Southern Marsh Orchid it with Southern Marsh Orchid.
x Common Spotted Orchid.
Var. macrantha has a lax flower spike and large
At one time, Leopard Marsh Orchid was flowers (the lip usually much more than 7.5mm
thought to be a variation of the hybrid between long x 9.5mm wide) with an obvious central
Southern Marsh Orchid and Common Spotted lobe that exceeds the lateral lobes by more than
Orchid which is commonly found whenever 1mm. It is rare and found mostly at sites where
the two species grow together. The occurrence Southern Marsh and Pugsley’s Marsh Orchids
of relatively uniform populations of Leopard occur together; it may be an intermediate
Marsh Orchid, lacking hybrid vigour and often between the two.
without Common Spotted Orchids growing Var. albiflora has white flowers but is very rare.
004 systematic.indd 314 29/1/09 12:33:58

BIOLOGY Name and classification

The specific name praetermissa means
Pollination and reproduction ‘overlooked’ and refers to the fact that the
Pollination is efficient and large quantities of
Southern Marsh Orchid was not recognised
seed are produced. Vegetative reproduction,
until 1914.
although far less important, may sometimes
This species is one of the tetraploid marsh
produce groups of plants.
orchids and the chromosome number is
Development and growth 2n = 80.
The period between germination and flowering
is two or three years. HISTORY AND
Hybrids CONSERVATION
D. x grandis, the hybrid with Common Spotted This species was overlooked (due to confusion
Orchid, is rather frequent and can usually be with Early Marsh Orchid) until 1914 when it
found where both species occur together (even was described by the eminent English botanist,
on dry chalk grassland and in the absence George Claridge Druce.
of one or both parents). It is probably the
Past and present occurrence of Southern Marsh Orchid
commonest hybrid orchid in southern Britain. in Britain and Ireland (based on presence or absence in
The pollen is highly sterile, and seed production 10km squares of the National Grid; data from the New
very low but even such partial fertility allows Atlas).
back-crossing and the creation of long-lived Britain Ireland
hybrid swarms.
D. x hallii, the hybrid with Heath Spotted 1500-1999
Orchid, occurs much less frequently than
D. x grandis as the two parent species favour
different habitats. It is fertile and can form % lost, 1500-1969 12%
hybrid swarms. % lost, 1970-1986 8%
D. x wintoni, the hybrid with Early Marsh % lost, total 20%
Orchid, has been recorded north to Lancashire
but is rare. It is sterile.
D. x insignis, the hybrid with Northern
Southern Marsh Orchid has inevitably
Marsh Orchid, is rare and has only been
suffered quite significant declines as agricultural
recorded from Wales. Despite their superficial
changes and development, especially the
similarity and ‘complementary’ distributions
draining and ploughing of flood-plain meadows
in northern and southern Britain, Northern
and pastures, have destroyed its habitats. It has
and Southern Marsh Orchids may not be very
vanished from 20% of the former range with
closely related.
much of the loss relatively recent. This statistic
x Pugsley’s Marsh Orchid has been recorded
is likely to conceal a much greater decline,
from Norfolk, Suffolk and Cambridgeshire.
however, as the number of populations within
Importantly this hybrid is fully fertile and
each 10km square can fall significantly without
through a process of back-crossing with its
it registering as a ‘loss’. This problem applies to
parents can form a full range of intermediates
all orchids but is perhaps most acute with those
(see also var. macrantha opposite).
species found in meadows and unimproved
Intergeneric hybrids pastures, habitats that were once so much a part
X Dactylodenia wintoni, the hybrid with of the British landscape. The colonisation of
fragrant orchid (in the broad sense), has been derelict industrial sites is small compensation
found in southern England. for such losses in the wider countryside.
004 systematic.indd 315 29/1/09 12:33:59

puGSley’S marSh Orchid

Dactylorhiza traunsteinerioides
Formerly: Narrow-leaved Marsh Orchid Dactylorhiza traunsteineri; also Dactylorhiza majalis

subspecies traunsteineri
The scarcest, most enigmatic and most localised of the marsh orchids (with the exception
of Hebridean Marsh Orchid), this species is endemic to Britain and Ireland. It occurs at
scattered localities throughout the British Isles, almost always in wet fens that are formed
by the upwelling of lime-rich but nutrient-poor springs. It hybridises freely, especially with
Southern Marsh Orchid, making certain identification difficult at times. ‘Lapland Marsh
Orchid’, a heavily marked plant found at a few localities in Scotland, has recently been shown
to be a variant of Pugsley’s Marsh Orchid.
Identification
This delicate marsh orchid can be identified by
a combination of features which taken together
give it a subtle but distinctive appearance. It
is typically slender with a thin, weak stem,
and some plants even appear to need to be
supported by the surrounding vegetation. It
has a few narrow leaves that are often held
erect. The spike is made up of a few relatively
large flowers and is usually rather open and
one-sided, with all the flowers facing roughly
the same way. The upper stem and especially
the bracts are washed purple. The flowers
themselves are variable in colour but often
purplish-pink. The lip is usually obviously
three-lobed with the side-lobes turned
downwards and the central lobe projecting as
a prominent ‘tooth’ (itself also sometimes
turned downwards). It is well-marked with
dark dots, loops and squiggles, the markings
often extending right to the edge of the lip.
Habitat is also a good clue, as the species is
almost always found in alkaline fens, often in
relatively low or sparse vegetation.
In order to confirm the identification it is
necessary to look carefully and in detail at the
orchids. It is best to examine a selection of
plants to get a feel for the range of variation
and work out some average measurements. It is
M 25 May, Norfolk. Typically slender, with narrow leaves
and a few, relatively large flowers (in this plant there are
11).
004 systematic.indd 316 29/1/09 12:34:00

PUGSLEY’S MARSH ORCHID l 317
unlikely that a Pugsley’s Marsh Orchid would

occur as a single isolated individual among a
colony of commoner species such as Southern
or Northern Marsh Orchid, and there should
always be a few candidates to look at. To be
Pugsley’s Marsh Orchids, plants must have the
following characteristics:
1. Number of flowers: Relatively few, usually
just five to 14, but sometimes as many as 18,
particularly on the robust plants found in some
parts of Ireland.
2. Width of lip: At least 7mm wide and usually
wider, up to a maximum of around 13mm
(the average of several plants should be at least
9mm). The individual flowers are relatively
large.
3. Lip shape: Distinctly three-lobed, usually
with prominent incisions (sinuses) between
the lobes and with the central lobe longer and
rather narrower than the rounded side-lobes;
flatten the lip from below with a finger in order
to measure the width and accurately judge the
shape. In a natural position the lip often has
a characteristic triangular or deltoid shape,
narrower at the base and broadest at the tip; a
shape accentuated by the side-lobes being more
strongly reflexed at the base and flatter towards M 28 May, Norfolk. The lip is usually marked with dots,
the tip. loops and squiggles, the markings often extending right to
the edge, and is typically three-lobed with the central lobe
4. Total number of leaves: Three to five, projecting as a little ‘tooth’.This plant has 12 flowers.
including up to one, rarely two, bract-like non-
sheathing leaves; do not count the short basal Early Marsh Orchids in similar habitats will
leaf hidden at the bottom of the stem. The
have pale pink flowers. The purple-flowered
leaves are more-or-less evenly spaced, rather
forms of Early Marsh Orchid are, however,
than being gathered into a rosette.
much more similar in colour and sometimes
5. Width of leaves: Relatively narrow, with
have few flowers on a spike. Also, in Scotland
the second leaf from the bottom, which is
the rare spotted-leaved subspecies cruenta of
usually the widest, 6-15mm across (rarely to
Early Marsh Orchid has been found in the
18mm and usually averaging no more than
same localities as the similarly spotted ‘Lapland
12mm wide).
Marsh Orchid’. But, whatever the colour of
6. Leaf spotting: Mostly unmarked but at a few
the flower, Early Marsh Orchid always has a
sites in Yorkshire and Ireland the plants have a
more ‘crowded’ flower spike and the individual
scatter of fine dark spots, and in Scotland the
flowers have a rather smaller lip, less than
subspecies lapponica (‘Lapland Marsh Orchid’)
9mm wide. The markings are also different on
has very boldly marked leaves and bracts
Early Marsh Orchid, with dots and dashes in
Similar species a central zone enclosed by bold double loops
Early Marsh Orchid is usually easy to separate and few if any markings outside this area. Their
from Pugsley’s Marsh Orchid because most leaves are broader and their spur is slightly
004 systematic.indd 317 29/1/09 12:34:01

tapering and slightly to moderately decurved. and more crowded at the base of the stem. The
Southern Marsh Orchid is typically taller spike usually has more than 20 flowers and
and more robust than Pugsley’s Marsh Orchid. the flowers themselves often have a slightly
It has a rather thicker stem and more leaves smaller lip (sometimes less than 8mm wide
which are broader and frequently more crowded and averaging less than 9mm), which is usually
towards the base of the stem; there are often roughly diamond-shaped and held flat or
two or more non-sheathing leaves. The spike slightly dished. The flowers are also usually a
is larger and usually has many more flowers, dark, ‘velvety’ reddish-pink, a rather different
and these have a less obviously three-lobed colour to those of Pugsley’s Marsh Orchid.
lip which is often held flat or slightly dished. Irish Marsh Orchid can be of a similar
Small, petite Southern Marsh Orchids are more size to Pugsley’s Marsh Orchid and has a
similar to Pugsley’s Marsh Orchid but still three-lobed lip (although not so deeply lobed),
have relatively unmarked flowers with less often with similar markings. It has, however,
prominent lobes. more leaves which are often broader and more
Northern Marsh Orchid is rather closer crowded at the base of the stem and a denser
in size and stature to Pugsley’s Marsh Orchid spike with more flowers.
but again has more leaves which are broader Hybrids with Common and Heath Spotted
Orchids and Early Marsh Orchid should show
clearly intermediate characters. Much more
problematical is the hybrid Pugsley’s Marsh
Orchid x Southern Marsh Orchid which is
apparently fully fertile. Through a process of
back-crossing this can form a complete range
of intermediates between the two species, and
at some localities in southern England there are
hybrid swarms and it is difficult to find any plants
that are completely ‘pure’ Pugsley’s Marsh Orchid.
Habitat
This species has very specific habitat
requirements and is only found where these are
met. It needs wet fens and flushes where the
ground water is ‘base-rich’ due to the influence
of chalk or limestone. It grows in the wettest
areas among a relatively open community of
sedges, rushes and scattered reeds, rooting
into the mossy layer at the base of the taller
vegetation. It is almost always associated with
Black Bog-rush. Some of the sites are fairly
extensive but at others the correct conditions
are restricted to small patches within a larger
area of marshland. In Ireland it has been
recorded rarely from dune slacks. Pugsley’s
Marsh Orchid is only exceptionally found in
very slightly acid conditions and in Scotland
‘Lapland Marsh Orchid’ may spread from its
M 30 June, Norfolk. Tends to flower early, from mid-May
to mid-June, but as with all orchids, this can vary (12 base-rich flushes into nearby wet heathland,
flowers). which is slightly more acid. It is found from
004 systematic.indd 318 29/1/09 12:34:02

sea level to 370m (Carperby, Wensleydale,

northwest Yorkshire), with subspecies lapponica
recorded up to 310m (Knapdale, Kintyre).
Flowering period
Plants come into flower in late May in the south
and west, exceptionally in early May, and some
may still be in flower in late June. The onset
of flowering can be a little later, towards early
June in the north, with ‘Lapland Marsh Orchid’
sometimes in flower until July.
Range
Found across the British Isles but the colonies
are very scattered and it is absent from most
areas. The strongholds are in Norfolk (and
formerly adjacent parts of north Suffolk),
northern Yorkshire, Anglesey and the Llyn
Peninsula in Caernarvonshire. Elsewhere
there are a few sites (often just one) in north
Somerset, north Hampshire, Berkshire,
Cambridgeshire, Co. Durham, Cardiganshire
and Pembrokeshire. Pugsley’s Marsh Orchid
is similarly scarce and local in Ireland: in Co.
Carlow, Co. Wicklow and Co. Kildare in the
east and from Co. Limerick north to Co. Mayo
in the west, with just three sites in Northern
Ireland, all in Co. Antrim. In Scotland it is rare, M 28 May, Norfolk. Although it had 16 flowers and a
and both the typical form and ‘Lapland Marsh total of 5 leaves, this well-marked plant was part of a
larger population of classic Pugsley’s Marsh Orchids.
Orchid’ have a broadly similar distribution
in the west, in Kintyre, west Inverness-shire, Ross & Cromarty, west Sutherland, the Inner
Hebrides (Rum, Raasay, Tiree) and Outer
1987-99 Hebrides (Harris). World Range: Endemic to
1970-86
pre 1970 Britain and Ireland.
How to find it
Pugsley’s Marsh Orchid is easily overlooked.
This species is small, slender and often partially
concealed among relatively long grasses and
sedges. At some localities it can be abundant
but it may also be scarce and ‘hidden’ amongst
hybrids. At many sites, especially in Scotland,
there may only be small numbers of flower
spikes. Two good clues for locating the species
are flowering period and habitat. Pugsley’s
Marsh Orchid is one of the first marsh orchids
to come into flower and is almost always found
with Black Bog-rush. This is a rather large,
004 systematic.indd 319 29/1/09 12:34:03

tussock-forming member of the sedge family

that through the winter and into the spring
forms stands of distinctively grey vegetation.
Many of the sites for this species are rather
small and vulnerable to excessive disturbance
or, as at Beeston Common in Norfolk, have be-
come dominated by hybrids. The best areas to
see it include Buxton Heath in Norfolk and
the Anglesey fens. In Ireland sites include
Slyne Head (Co. Galway) and Blacksod Bay
(Co. Mayo).
DESCRIPTION
Height: 6-40cm but usually 10-30cm with the
most robust plants in Ireland.
Stem: Green, usually washed purple towards
the flower spike, slightly hollow and rather
slender. The stem is often rather floppy or
bendy, averaging only 2.3-3.5mm in diameter
and is rarely more than 5mm across.
Leaves: Green, unspotted or occasionally with a
few, well-scattered, faint, purplish-brown spots
or transverse bars about 1mm in diameter;
spotting is only common in Yorkshire and
Ireland (see also Subspecies). The uppermost
M 28 May, Norfolk.The spike is often very one-sided (11
non-sheathing leaf may be washed purple. flowers).
There is one short basal leaf and two to four
(rarely five) sheathing leaves. These are narrow, Bract: Lanceolate and long (the lower bracts
strap-shaped, moderately keeled, sometimes longer than the flowers), the bracts are green,
slightly hooded at the tip, well spaced along variably but often strongly washed reddish-
the stem and often held erect at around 45°. purple.
The longest leaf is rarely more than 12cm long Ovary: Green, strongly washed purple,
(17cm in robust Irish populations) and the cylindrical and six-ribbed.
broadest leaf is usually 6-15mm wide (rarely Flower: Often purplish-pink, similar in colour
to 18mm). There are also up to one (rarely to Southern and Irish Marsh Orchids, but
two) bract-like non-sheathing leaves on the variable in intensity from rather pale to dark
upper part of the stem, and the total number of and deep. The sepals are oval-lanceolate and
sheathing and non-sheathing leaves is three to the petals slightly smaller and more oval.
five (this count excludes the short basal leaf ). The lateral sepals are asymmetrical, very
Spike: Short and distinctly loose or ‘lax’ with occasionally have darker markings and are
rather few flowers, usually just six to 14, held at an angle that varies between 45° and
occasionally as few as two or as many as 18 on vertical but is usually closer to the latter. The
the largest plants. The spike is rather irregular upper sepal and petals form a hood over the
in shape but roughly cylindrical or tapering and column. The lip is relatively large, 6.5-9.5mm
the flowers all face in more-or-less the same long x 7-13mm wide (usually at least 7.5mm
direction (the spike is thus described as secund long x 9.5mm wide). It is very variable in
or subsecund). shape but is usually a flattened oval divided
004 systematic.indd 320 29/1/09 12:34:04

into three obvious lobes with the central lobe Hebrides on Harris. It has also been identified
rather narrower and variably longer than the in Northern Ireland, on the Garron Plateau in
side-lobes. The lobes are separated by incisions Co. Antrim. In at least one site ‘Lapland Marsh
(sinuses) of various depths but sometimes Orchid’ is found with typical Pugsley’s Marsh
there are no obvious sinuses, just a prominent Orchids, and intermediates have been recorded.
central projection. The side-lobes are folded ‘Lapland Marsh Orchid’ was first discovered
downwards to a variable extent (and the tip at Knapdale (Kintyre) in 1967 and initially
of the central lobe may also turn downwards), identified as Pugsley’s Marsh Orchid. This
giving the lip a rather drop-shaped outline, identification was then revised and it became
narrowest at the base. The lip is a variable ‘Western Marsh Orchid’ (the form cambrensis,
shade of purplish-pink, becoming whitish at now itself a subspecies of Northern Marsh
the mouth of the spur, with dark lines, dots and Orchid). Later still, in 1988, the plants
blotches in an irregular pattern that extends were identified as ‘Lapland Marsh Orchid
more-or-less to the edges (sometimes forming D. lapponica’ a species found in the Alps,
a ‘double loop’ as in Early Marsh Orchid). Scandinavia and northern Russia which was
There is no correlation between the ground new to the British Isles. Ironically, the wheel
colour of the lip and the extent and colour of has now turned full circle and Scottish ‘Lapland
the markings; dark plants in Yorkshire may Marsh Orchids’ have become a subspecies
have relatively small dot and dash markings of Pugsley’s Marsh Orchid, confirming in a
whereas paler flowers on Anglesey may have roundabout way the initial identification.
long, heavy and contrastingly dark lines and ‘Lapland Marsh Orchid’ may, in fact, be best
blotches. Conversely, pale flowers in southern treated as a variety of Pugsley’s Marsh Orchid,
England may have relatively light markings.
The spur is long, straight and thick but tapers
slightly to a blunt tip.
Subspecies
D. t. lapponica ‘Lapland Marsh Orchid’ has
bold, dark purplish-brown spots, blotches, bars
and rings on the upperside of the leaves and
bracts, which may also be edged with purple
(and the bracts are often spotted below). There
are up to two non-sheathing leaves which may
also have a few small marks on the underside.
The lip has intense dark purple or crimson
lines, rings and spots on the lip, sometimes
merging to form a dark patch in the centre, and
the lateral sepals are marked with dark rings,
elongated spots and dots. It averages slightly
shorter than the typical form at 6-18cm tall,
sometimes to 24cm, but Scottish populations
of typical plants are also small.
Occurs very locally in western Scotland,
in west Sutherland, Ross & Cromarty, west
Inverness-shire (including Ardnamurchan and
M 11 June, Kintyre. ‘Lapland Marsh Orchid’. A beautifully-
Morvern), Kintyre and, in the Inner Hebrides, marked plant, with bold loops and blotches on the lip and
on Skye, Rum and Raasay and on the Outer lateral sepals.
004 systematic.indd 321 29/1/09 12:34:06

O 8 June, Kintyre.‘Lapland
Marsh Orchid’. In classic
plants the leaves, bracts
and flowers are boldly
marked, but may occur in
‘mixed’ colonies with more
typical Pugsley’s Marsh
Orchids; there appears to
be a north-south gradation
in the degree and depth
of markings, from rela-
tively lightly-marked plants
with unspotted leaves in
southeast England to bold-
marked plants in northern
and western Scotland.
rather than a subspecies; the Lapland Marsh of this variety and as spotted leaves are also
Orchid of Scandinavia remains, however, a found in Irish populations it does not seem
distinct species, D. lapponica. worthwhile to use this name. Rather small
Variation and varieties Scottish plants from near Loch Maree (Ross
Pugsley’s Marsh Orchid is rather variable. & Cromarty) with very pale flowers and
The size and shape of the leaves, presence or contrasting dark markings have been named
absence of leaf spotting, number of flowers var. francis-drucei but may represent just one
and their colour and markings all vary, both small aberrant population. (Note that var.
between colonies and within them. Yorkshire bowmanii, originally described as a variety
plants with spotted leaves are sometimes of Pugsley’s Marsh Orchid, is treated under
named var. eborensis. However, as there is Southern Marsh Orchid.)
disagreement about the other characteristics Var. albiflora has white flowers. It is very rare.
004 systematic.indd 322 29/1/09 12:34:08

BIOLOGY Orchids. Pugsley’s Marsh Orchid is intriguing

in that the same parent species have crossed on
Pollination and reproduction several occasions, in the British Isles, the Alps,
There is little specific information. Pollination
Scandinavia and perhaps elsewhere, and each
is probably undertaken by bees and although
hybridisation event has given rise to a similar
other insects such as flies may remove the
plant. After much debate, it was generally
pollinia they do not go on to pollinate other
accepted that British and Irish plants were the
orchids successfully. In studies of Pugsley’s and
same species as those in Europe and should
‘Lapland’ Marsh Orchids in western Scotland,
be known as Narrow-leaved Marsh Orchid
only 17-35% of flowers set seed, reflecting low
D. traunsteineri. However, recent genetic
levels of pollination, probably due to a lack of
research has shown that, although very
suitable pollinators and the poor weather. Each
similar, they each have a distinct ancestry
ripe capsule that was produced contained about
and represent separate lineages and are best
3,000 viable seeds. Vegetative reproduction may
treated as separate species. British and Irish
also be possible.
plants therefore become an endemic species, D.
Development and growth traunsteinerioides, and should generate greater
No specific information. interest among scientists and conservationists.
Note that Delforge (2005) gives the range of
Hybrids
this species as Britain, Ireland and northern
D. x silvae-gabretae, the hybrid with Common
France but his concept of Pugsley’s Marsh
Spotted Orchid, has been recorded from
Orchid appears to be rather different; for
Ireland, Anglesey and Yorkshire. The pollen is
example, he gives the number of flowers as
highly sterile.
10-25, with extremes of 5-30.
D. x jenensis, the hybrid with Heath Spotted
Orchid, has been recorded from northwest HISTORY AND
Wales, Yorkshire and Ireland. It is probably CONSERVATION
sterile.
This species was first found in Ireland by H.W.
D. x dufftii, the hybrid with Early Marsh
Pugsley, who realised that it was something
Orchid, has been found in northwest Wales,
different from herbarium specimens and plants
Yorkshire and Co. Wicklow but is rare.
sent to him from Co. Wicklow. But, from
x Southern Marsh Orchid has been recorded
the start, the exact status of Pugsley’s Marsh
from Norfolk, Suffolk and Cambridgeshire.
Orchid was the subject of much debate. In
Fully fertile, it can be common where the two
1936, Pugsley described his plants as a new
species occur together.
subspecies of a widespread European marsh
Name and classification orchid and called it D. majalis subspecies
The specific name traunsteinerioides is derived traunsteinerioides. In 1940, however, Pugsley
from traunsteineri, the scientific name of changed his mind and declared it to be an
Narrow-leaved Marsh Orchid, a European entirely new species, D. traunsteinerioides. Then,
species that is, in turn, named in honour of in 1953, the controversial British botanist
Traunsteiner, a pharmacist who lived in Austria J. Heslop-Harrison demonstrated that it
from 1798-1850. belonged with another European species, D.
This species is a tetraploid marsh orchid traunsteineri. This determination generated a
with the chromosome number 2n = 80. All large and sometimes controversial literature,
of the tetraploid marsh orchids are thought but the consensus eventually emerged that it
to have originated from ancient hybridisation was indeed the same as the European plant. Just
events involving the ancestors of Early Marsh as the question seemed to have been settled,
Orchid and Common or Heath Spotted however, genetic evidence showed that Pugsley
004 systematic.indd 323 29/1/09 12:34:09

had been right all along and that it should to drainage and abstraction) can cause a
be treated as a separate, endemic species, D. degradation of the habitat. This has been
traunsteinerioides. For more details of ‘Lapland especially noticeable in East Anglia where the
Marsh Orchid’ see Subspecies. species has declined sharply in the Waveney-
Pugsley’s Marsh Orchid is Nationally Ouse fens along the Norfolk-Suffolk border. A
Scarce in Britain and is specially protected in more insidious threat is posed by hybridisation.
Northern Ireland under Schedule 8 of the 1985 The hybrid with Southern Marsh Orchid is
Wildlife Order (NI). In addition, ‘Lapland fully fertile and at some sites hybrids have
Marsh Orchid’ (until recently treated as a come to dominate the population and Pugsley’s
distinct species) is in the Red Data Book and is Marsh Orchid has been ‘hybridised out’ (e.g. at
considered Near-threatened in Great Britain Beeston Common in Norfolk there were several
and fully protected under Schedule 8 of the hundred Pugsley’s Marsh Orchids in the late
Wildlife and Countryside Act 1981. 1980s but by 2000 it was difficult or impossible
to find a single ‘pure’ plant). At other sites,
Past and present occurrence of Pugsley’s Marsh Orchid hybrids apparently involving this species are
(including ‘Lapland Marsh Orchid’) in Britain and Ireland
(based on presence or absence in 10km squares of the found where Pugsley’s Marsh Orchid has never
National Grid; data from the New Atlas). been recorded, indicating a more extensive
Britain Ireland
distribution in the past. Similar threats face
the species throughout Europe as its highly
1500-1999 specialised habitat is degraded or destroyed.
current range 63 (2.2%*) 13 (1.3%*) In the uplands, sheep or deer graze most
sites. This can keep the vegetation open and
% lost, 1500-1969 8.5% 37.5% prevent a succession to scrub but it limits
% lost, 1970-1986 17.5% 30% the number of flowers produced, sometimes
% lost, total 26% 67.5% severely so. When sites are not grazed the
* current range as a % of the total number of 10km squares number of flowering plants has increased,
although this may not reflect the number of
‘Lapland Marsh Orchid’ is mapped in the seeds that are able successfully to germinate.
New Atlas for 18 10km squares (seven of which More direct threats include forestry and
it shares with typical Pugsley’s Marsh Orchid). drainage, and only a few of the 30 or so
Pugsley’s Marsh Orchid has declined populations of ‘Lapland Marsh Orchid’ are
by 26% in Britain, with most of the protected in any way at all.
decline relatively recent and concentrated Any changes in the overall status of
in East Anglia. It is probably extinct in Pugsley’s Marsh Orchid are hard to assess. It is
Huntingdonshire and Northumberland. Losses a difficult species to identify with certainty and
have been even greater in Ireland, at 67.5%, also easy to overlook and is therefore still being
and its numbers appear to have fallen there found and identified at new sites. These new
more than almost any other orchid. There are records do not indicate, however, an expansion
no recent records from Co. Cork, Co. Kerry, or increase as the species has presumably
Co. Tipperaray, Co. Offaly, Westmeath, Meath, always been present at these sites. This applies
Co. Leitrim, Co. Cavan, Co. Louth and Co. particularly to Lapland Marsh Orchid, which
Fermanagh. has been actively searched for since its discovery.
In the lowlands, Pugsley’s Marsh Orchid Conversely, Pugsley’s Marsh Orchid is probably
has disappeared due to the destruction and recorded from some sites by mistake (i.e. it is
drainage of its habitat. Even where wetlands ‘overrecorded’).
remain, often protected as SSSIs and reserves,
a general lowering of the water table (due
004 systematic.indd 324 29/1/09 12:34:09

HEBRIDEAN MARSH ORCHID l 325
heBridean marSh Orchid Vulnerable

Dactylorhiza ebudensis
Formerly: Dactylorhiza majalis subspecies occidentalis

This is one of four species of orchid endemic to the British Isles (the others being
Lindisfarne Helleborine and Pugsley’s and Irish Marsh Orchids). It is known only from
Berneray and North Uist in the Outer Hebrides where it is locally abundant in the flower-
rich machair.
Identification seldom bend upwards at the edges in the

Usually rather short and squat with a manner that gives the lip of Northern Marsh its
distinctive purple tone to the flowers. The leaves characteristic diamond shape.
vary from more or less plain to solidly blackish l Flowers usually a clearer, purer purple (when
purple, but most are marked with rings or fine freshly opened the flowers of Northern Marsh
blotches. In general, plants with darker leaves have a distinct crimson tone; on older flowers
have a darker ground colour to the flower. the colour become more purple, but the dark lip
Similar species markings retain a deep crimson lustre).
l Leaves often (but not always) more heavily
Only likely to be confused with Northern
Marsh Orchid, but distinguished as follows: marked, with ring-spots and blotches, and
l Flower spike looser, less symmetrical and also finely edged with purple (especially the
often one-sided (typically symmetrical and uppermost leaf ), even when not spotted.
compact in Northern Marsh). Northern Marsh may have fine, regular spotting
l Lip relatively larger, more spreading, and on the leaves, but never extensive dark markings
more obviously three-lobed; the side lobes (subspecies cambrensis of Northern Marsh
M 14 June, North Uist.The flowers are distinctly purple, and the leaves variably spotted.
004 systematic.indd 325 29/1/09 12:34:11

Orchid, which has heavily spotted leaves, has stretching for c. 4 km south-westwards from
been recorded elsewhere in the Outer Hebrides, Newtonferry (Port nan Long), with a couple of
but the spots tend to be smaller, more even in very small satellite populations a few km to the
size and more evenly distributed over the leaf west, and to the north, on Berneray.
surface than in Hebridean Marsh).
l Whole plant averages smaller and more petite. DESCRIPTION
Height: 4.5-20cm.
Habitat
Stem: Green, heavily washed with dark purple
Found on the machair, a unique, species-rich,
towards the tip and hollow.
coastal grassland habitat which develops on
Leaves: There are two or three, sometimes
low calcareous dunes in the wet and windy
four, keeled, lanceolate sheathing leaves that
environment of western Scotland and Ireland.
are relatively broad (10-14mm wide) and are
Flowering period crowded towards the base of the stem with
Late May to late June. the lower leaves tending to be held nearer the
horizontal than the vertical; the lowest leaf
Range
is often rather shorter than the rest and the
Confined to North Uist and Berneray in the
single non-sheathing leaf is lanceolate. On most
Outer Hebrides. World range: Endemic to
plants the leaves are green, moderately marked
Scotland.
with brownish-purple rings or fine blotches
How to find it (especially the non-sheathing leaves). Markings
Only found on the north coast of North Uist, are concentrated towards the tips of the leaves
where there is effectively a single metapopulation and on more heavily marked plants they begin
M 14 June, North Uist.Two plants, showing some of the range of variation in leaf markings.
004 systematic.indd 326 29/1/09 12:34:13

HEBRIDEAN MARSH ORCHID l 327
to coalesce; there is a continuous gradation Hybrids

from plants with unmarked green leaves to x Northern Marsh Orchid? Plants showing
those with leaves that are solidly blackish- characters intermediate between these two
purple; spotting is occasionally also present species have been found but may be merely
on the underside of the leaves. The leaves are variant Northern Marsh Orchids; they are
also finely rimmed with purple (especially the fertile.
uppermost), even when not spotted. Name and classification
Spike: Short, compact and cylindrical, with The specific name ebudensis is from the Latin
5-20 flowers. ebudes, ‘Hebridean Islands’.
Bract: Lanceolate, the lower slightly longer than The taxonomy of this species is confusing.
the flowers but becoming shorter towards the Until recently Hebridean Marsh Orchid was
top of the spike. They are green, heavily washed grouped together with Irish Marsh Orchid
purple or sometimes entirely purple and often and the majaliformis form of Northern Marsh
spotted darker. Orchid into one species named ‘Western Marsh
Ovary: Ribbed, twisted and very heavily Orchid D. majalis’. Thus, in many recent books,
washed purple. the populations on North Uist will be found
Flower: Deep purple-magenta with darker under the heading ‘Western Marsh Orchid
markings. The sepals and petals are elliptical, Dactylorhiza occidentalis’ and either listed as
with the lateral sepals slightly asymmetric and subspecies ‘scotica’ or combined with Irish
held at 45° above the horizontal, or even higher, plants as subspecies ‘occidentalis’. Unexpectedly,
whereas the upper sepal and petals (which genetic studies have shown that Hebridean
are shorter) form a hood over the column. and Irish Marsh Orchids should be treated
The lip is broader than long (6-8.5mm long x as distinct species (and thus Western Marsh
8-12mm wide) and distinctly three-lobed with Orchid D. occidentalis does not occur anywhere
the central lobe longer than the side-lobes. It is in Britain or Ireland). Hebridean Marsh Orchid
flat or dished when the flower first opens but is one of the tetraploid marsh orchids and has
the side-lobes may become slightly deflexed a chromosome number of 2n = 80. Genetic
with age. The lip is rich purple-magenta, evidence suggests that it originated in a cross
whiter towards the throat of the spur, and between Early Marsh Orchid, subspecies
heavily marked with dark purple spots and coccinea, and Common Spotted Orchid,
dashes which are bounded within one or two subspecies hebridensis.
concentric dark loops. The spur is a little
longer than the ovary, thick, cylindrical and HISTORY AND
slightly tapering. CONSERVATION
Subspecies This distinctive orchid first attracted attention
None. in the 1930s, but it was not until 1976 that
it was given a name. The first to be used was
Variation and varieties ‘scotica’ but this was not published in the correct
A hyperchromic variant has occasionally been manner according to the International Code
found. This has an excess of pigmentation of Botanical Nomenclature and was eventually
producing a solidly dark lip with a paler border. replaced by ebudensis.
BIOLOGY Much of the range of the species lies within
the Machairs Rabach and Newton SSSI and
Pollination and reproduction there seems to be no immediate threat to the
No specific information. species, but such a limited distribution means
Development and growth that Hebridean Marsh Orchid will inevitably
No information. always be vulnerable.
004 systematic.indd 327 29/1/09 12:34:13

nOrthern marSh Orchid

Dactylorhiza purpurella
Formerly: Orchis purpurella, Dactylorchis purpurella

This is the most commonly encountered magenta-flowered orchid of damp, marshy ground
in northern Britain and can sometimes be found in large numbers, giving a ‘wow’ factor to
the landscape, even sometimes to derelict industrial sites. As its name suggests, this species
has a northerly distribution and occurs widely in northern England, northern Ireland
and Scotland, as well as in Wales. Although the intense reddish-pink flowers are usually
distinctive, the presence of hybrids with the spotted orchids can be a complication.
Identification
Relatively easy to identify. The flowers are
a deep ‘velvety’ magenta with a distinctive
crimson tone that is most obvious when they
are freshly opened, while the dark markings
on the lip are always a lustrous dark crimson.
The lip typically appears diamond-shaped
with straight sides, especially to the base of the
‘diamond’ (the lip is actually three-lobed but
the side lobes are folded upwards at the edges
to form the diamond). The leaves are either
unspotted or have a few small spots, except
in parts of west Wales, northern England
and northwest Scotland where the subspecies
cambrensis has dark spots all over the leaves.
Similar species
Early Marsh Orchids with purple flowers are
quite similar to Northern Marsh Orchid, but
their flowers are, on average, slightly smaller,
with the lip rather narrower and usually
obviously three-lobed but with the side-lobes
sharply turned downwards (it may be unlobed,
but is never the distinctive diamond shape
of Northern Marsh); their lateral sepals are
usually held vertically over the flower.
Southern Marsh Orchid has more purplish-
pink flowers that are usually a little ‘washed out’
and lack the deep crimson tones of Northern
Marsh. Its lip is usually roughly circular, with
rather broad rounded side-lobes and a small
tooth-like central lobe. The dark markings are
M 6 June, Cumbria. As with all the marsh orchids,
Northern Marsh Orchid is variable in size and structure often small, fine and confined to the centre of
and can look quite stocky. the lip. Leopard Marsh Orchid (var. junialis
004 systematic.indd 328 29/1/09 12:34:18

NORTHERN MARSH ORCHID l 329
of Southern Marsh) has bolder, more looped

markings on the lip but also has heavily spotted
leaves, with the spots usually in the form of
hollow rings.
Irish Marsh Orchid has a distinctly
rounded, three-lobed lip that is on average a
little broader than Northern Marsh. It is more
purplish-pink and often also paler and less
intensely coloured. Up to 50% of Irish Marsh
Orchids have spotted leaves.
For distinctions from Hebridean Marsh
Orchid, see p. 325.
Hybrids with both Heath Spotted and
Common Spotted Orchids are relatively
common and can be abundant where the two
parent species grow together. Both hybrids
are very variable. They can have either spotted
or unspotted leaves and flowers intermediate
between the two parent species.
Habitat
Northern Marsh Orchid is found in damp or
wet sites on calcareous, neutral or slightly acid M 6 June, Cumbria. The leaves may have some fine dark
soils, such as marshy fields, roadside verges, spots.
lake margins, fens, marshes, flushes, seepages N 7 June, Northumberland. Northern Marsh Orchid can
along coastal cliffs, dune slacks, machair and be common in dune slacks.
004 systematic.indd 329 29/1/09 12:34:27

sometimes in less acidic peat bogs or in open,

damp woodland. It is an opportunist and has
colonised old quarries and urban ‘brownfield’
sites, including derelict industrial areas and
old waste tips, such as alkaline Leblanc waste
in Lancashire. Northern Marsh Orchid is
sometimes found on drier substrates such
as rubble and has been found in gardens.
Recorded up to 610m above sea level (Creag
Dhuba, Loch Ericht, west Inverness-shire).
Flowering period
Late May to late July (exceptionally from mid-
May) but mostly early June to mid-July.
Range
Widespread in north and west Wales, northern
England and Scotland, including the Isle of
Man, Inner and Outer Hebrides, Orkney and M 6 June, Cumbria.The lip is usually a distinctive diamond
shape.
Shetland. Occurs north of a line from Swansea
to Hull, but the population is sparser and more and in Hampshire it was discovered in 1955 at
scattered towards this southern boundary. Moorgreen near Southampton where two small
Also found in Northern Ireland, especially in colonies held up to 100 flowering plants at
Co. Fermanagh, Co. Tyrone, Co. Derry and times. It was thought that a cutting for the M27
Co. Antrim, and in adjacent Co. Donegal, but in 1986 had destroyed these colonies but plants
otherwise rare and local in Eire, with almost were found again in the area in 1999. (It has
all the current sites on the east coast or on the been suggested, however, that this species was
coasts of Co. Waterford, Co. Cork, Co. Galway introduced to Hampshire.) World range: A
and Co. Mayo. There are a few isolated records European endemic found along the northwest
from southern England. In Oxfordshire a single fringe of the continent in western Norway
specimen was found near Wychwood in 1981 (north to around Trondheim), Sweden,
Denmark and the Faeroe Islands.
1987-99
1970-86
How to find it
pre 1970 Quite common and sometimes found in large
numbers.
DESCRIPTION
Stem: Green, washed purple towards the tip,
ribbed, often stout and slightly hollow.
Leaves: The four to eight lanceolate sheathing
leaves are slightly crowded towards the base of
the stem, broad (the largest 1.5-2.5cm wide),
hooded at the tip and held at up to 45° above
the horizontal. There are also one or two non-
sheathing leaves higher on the stem (sometimes
none, rarely up to four) and a small basal leaf.
004 systematic.indd 330 29/1/09 12:34:29

O 13 June, North Uist. De-

spite their names, North-
ern and Southern Marsh
Orchids are not each
other’s closest relatives and
are not analogous. Perhaps
as a consequence, hybrids
between these two species
are surprisingly scarce.
The leaves are mid-green to dark green, oval and marked with irregular dark reddish-
unspotted or sometimes with a few very small purple rings and lines. The petals are a little
spots near the tip. shorter and unmarked. The lateral sepals are
Spike: Dense, with ten to 40 flowers (some- held at around 45° and the upper sepal and
times as many as 80), oval to cylindrical in petals form a loose hood over the column. The
shape and often flat-topped when fully open. lip is relatively small, 5-8mm long x 6-10mm
Bract: Green, often flushed purple, especially wide. Although variable in shape, it is usually
towards the edges. The bracts are narrow and shallowly three-lobed with a small central lobe
lanceolate, the lower longer than their flowers, and and held flat or dished, with the margins
the upper shorter. of the side lobes folded upwards to give the
Ovary: Green, washed purple (and sometimes lip a diamond shape. Heavy dark crimson
spotted on the ribs), cylindrical, six-ribbed and lines, dots and swirls cover much of the lip in
twisted. a concentric pattern. The spur is thick, conical,
Flower: Deep ‘velvety’ magenta, whiter around slightly downward pointing and shorter than
the mouth of the spur. The sepals are roughly the ovary.
004 systematic.indd 331 29/1/09 12:34:33

lobes slightly reflexed. It tends to be taller than

typical Northern Marsh Orchids with longer
and narrower leaves but with the spike smaller
in proportion to the whole plant.
This subspecies grows near to the sea in
western Scotland, often on damp grassy slopes,
in the Outer Hebrides, Kintyre, Ross and
Cromarty, Sutherland, Caithness and Orkney.
In west Wales it has been found rather rarely
in dune slacks and flood plain meadows on the
coast from Cardiganshire to Anglesey. It has
also been recorded from southeast Yorkshire
and recently from several sites in Cumbria, but
is otherwise only known from Denmark. It
has a chequered history and both Scottish and
Welsh plants started life as forms of ‘Western
Marsh Orchid’ (a conglomerate of cambrensis
with Irish and Hebridean Marsh Orchids).
Var. albiflora has white flowers.
Var. atrata is a hyperchromic variant with an
excess of pigmentation. The lip is solidly dark
magenta, lacking spots or lines but sometimes
with a well-defined paler margin. The leaves are
heavily spotted (rarely there is an overall purple
wash) with a few small spots on their lower
surfaces too. It is known only from damp fields
near an industrial area at Hartlepool which may
M 6 June, Cumbria. The lip markings are variable but be contaminated with heavy metals.
there tend to be heavy lines, dots and swirls more or less Var. crassifolia has broad, fleshy leaves and a
all over the lip, often in a concentric pattern.
large lip. It is rare.
Var. maculosa has many small dots all over the
Subspecies
D. p. cambrensis (formerly ‘majaliformis’) has leaves and sometimes a paler lip. Rare, it has
leaves that are boldly spotted dull purple, been recorded in southeast Scotland.
and bracts that are also spotted or washed Var. pulchella has the lip marked with spots
purple. The spots are distributed over the and dashes rather than bold loops. It has been
whole surface of the leaf and can be sparse recorded from Scotland.
or occasionally so numerous that they
coalesce into a solid dark patch; spotting BIOLOGY
is absent in a small minority of plants. The Pollination and reproduction
flowers average slightly paler and pinker than No specific information, but it is probably
normal and therefore the dark markings are pollinated by bees, as bumblebees, especially
more contrasting. The lip is a little larger at queens of Bombus terrestris, have been recorded
6-8.5mm long x 9-11mm wide and is rather as frequent visitors. Pollination is efficient and
distinctly three-lobed, often with incisions seed-set is good (52% of flowers setting seed in
(sinuses) between the lobes and with the side- one Scottish study).
004 systematic.indd 332 29/1/09 12:34:37

Development and growth HISTORY AND

No specific information. CONSERVATION
Hybrids Northern Marsh Orchid was not distinguished
Hybrids are frequently found, especially with from the other marsh orchids until 1920
Common and Heath Spotted Orchids. when it was described from plants found near
D. x formosa, the hybrid with Heath Spotted Aberystwyth by T. and T.A. Stephenson.
Orchid, occurs throughout the range and is Inevitably there have been losses due to
probably the commonest orchid hybrid in habitat destruction (drainage, ploughing-up
northern Britain and Ireland. of pastures, etc.) but the overall boundaries of
D. x venusta, the hybrid with Common the range have remained stable. It has declined
Spotted Orchid, occurs throughout the by 21% in Britain, with many of the losses
range and can be common. It is variously comparatively recent, and by 46% in Ireland.
noted that the pollen is highly sterile and Past and present occurrence of Northern Marsh Orchid in
little seed is produced or that this hybrid is Britain and Ireland (based on presence or absence in 10km
sometimes partially fertile. Whatever the case, squares of the National Grid; data from the New Atlas).
it is certainly capable of backcrossing with Britain Ireland
either parent species or crossing with itself to
produce a range of intermediate characters in a 1500-1999
hybrid swarm. current range 946 (33%*) 81 (8%*)
D. x latirella, the hybrid with Early Marsh
Orchid, has been recorded throughout the % lost, 1500-1969 11% 33%
range but is rare. % lost, 1970-1986 10% 13%
D. x insignis, the hybrid with Southern % lost, total 21% 46%
Marsh Orchid, has only been recorded in * current range as a % of the total number of 10km squares
Cardiganshire and Merionethshire. Despite
the relatively extensive area of overlap between
the two species in Wales and northern England
they tend to maintain a separate identity, even
in mixed colonies.
D. x viridella, the hybrid with Frog Orchid, has
been recorded from Co. Durham and the Inner
and Outer Hebrides.
x Hebridean Marsh Orchid? Plants showing
characters apparently intermediate between
the two species have been recorded but these
may be variant Northern Marsh Orchids; they
are fertile.
X Dactylodenia ‘varia’, the hybrid with one
of the fragrant orchids, has been found in
northern England, Scotland and Ireland.

The specific name purpurella means simply
‘purple’. This species is a tetraploid marsh orchid M 6 June, Cumbria.The lip shape can vary.This plant has
(2n = 80). a projecting central lobe.
004 systematic.indd 333 29/1/09 12:34:41

iriSh marSh Orchid

Dactylorhiza occidentalis
Formerly: Dactylorhiza majalis subspecies occidentalis; Dactylorhiza comosa subspecies

occidentalis; Orchis occidentalis; Dactylorchis occidentalis
An Irish endemic, this species is fairly widespread and locally common in marshy or grassy
habitats although it has suffered some decline, especially more recently. As with all the marsh
orchids, it can be tricky to identify with certainty but its relatively early flowering season is
a great help.
Identification
The compact spike of purplish-pink, well-
marked flowers identifies this species as one
of the marsh orchids. It is rather variable, and
the dwarf plants of coastal grasslands can look
rather different to the robust populations in
damper and more sheltered spots. The leaves
may be spotted or unspotted. The lip has three
rounded lobes and is usually heavily marked,
often with double loops.
Similar species
Separation of the marsh orchids can be difficult
at times, and in Ireland three species need to
be eliminated: Early, Northern and Pugsley’s
Marsh Orchids.
Early Marsh Orchid is very variable and
includes the purple and pink-flowered subspecies
pulchella and cruenta, the latter often showing
bold spots on the leaves. But, whether their
leaves are spotted or unspotted, all Early Marsh
Orchids have rather smaller flowers with a
narrower lip that is usually less than 9mm wide
and narrower leaves that are held stiffly erect.
Northern Marsh Orchid tends to flower a
little later, has unspotted leaves or only a few
small spots, and flowers that are a darker and
deeper reddish-pink. Characteristically it has
an unlobed, straight-sided, diamond-shaped
lip that, on average, is a little narrower, no more
than 10mm wide.
Pugsley’s Marsh Orchid is reasonably
distinctive, with a slender stem, a few relatively
M 22 May, Co. Clare. Some have relatively paler and
pinker flowers and at least 50% of plants have unmarked
large flowers held in an open, one-sided spike
leaves. and a lip that appears longer than wide.
004 systematic.indd 334 29/1/09 12:34:46

IRISH MARSH ORCHID l 335
Early Purple Orchid flowers at the same

1987-99
time as Irish Marsh Orchid, sometimes in 1970-86
the same grassy habitats, and also has heavily pre 1970
spotted leaves. It is generally tall and slender,

however, with the leaves arranged in a floppy
rosette at the base of the stem. Its flowers are
much less heavily marked, with just a few dark
spots in the centre of the lip.
Habitat
Varied but mostly on neutral or slightly alkaline
soils and including wet meadows and pastures,
road verges, lough shores, the edges of acid
bogs, dune slacks, damp hollows on short,
closely grazed grassland and dry grassy slopes
near the sea.
Flowering period
Mid-May to mid-June, sometimes to July. How to find it
Range Generally fairly common and sometimes found
In Eire there are concentrations in Co. Donegal in large numbers.
in the northwest, from Co. Galway south to
Co. Cork in the west and in Co. Wexford. DESCRIPTION
Otherwise, it is absent from much of Eire, Height: 10-30cm, rarely to 40cm.
especially the east, apart from a cluster of Stem: Green, variably washed purple, ridged
records from Co. Longford. Although mapped towards the tip and slightly hollow.
for Northern Ireland, its occurrence there is Leaves: The four or five (sometimes only three)
disputed. World range: Endemic to Ireland. lanceolate sheathing leaves are fairly broad,
M 25 May, Co. Clare. Irish Marsh Orchid can still be found in large numbers, even on roadside verges.
004 systematic.indd 335 29/1/09 12:34:49

004 systematic.indd 336 29/1/09 12:34:52

The lip is purplish-pink, often rather deep but

becoming paler towards the centre and white
around the mouth of the spur (near to Southern
Marsh Orchid in colour but slightly deeper,
more intense and closer to purple). It is usually
wider than long, 7-11mm long x 9-14mm wide
and broadest around the middle. It has three
distinct rounded lobes with noticeable incisions
(sinuses) between them. The side-lobes are
variably reflexed, especially as the flower gets
older, and sometimes notched or with wavy
edges. The central lobe is smaller and sometimes
tooth-like and often projects a little beyond the
side-lobes. The lip is marked with heavy dots
and dashes, often enclosed within double loops
M 21 May, Co. Clare. Some plants (perhaps especially in (sometimes concentric double loops), and there
coastal grassland) are short and squat. may also be markings outside the loops. The
O 24 May, Co. Clare. spur is roughly conical and straight.
usually 15-25mm wide (extremes 12-30mm). Subspecies

They are grouped towards the base of the stem None.
and are usually held rather spreading, closer to Variation and varieties
the horizontal than the vertical. There are also Var. kerryensis is found in southwest and
two, sometimes three, non-sheathing leaves western Ireland, sometimes in ‘pure’ colonies,
higher on the stem. The leaves are green and in
about 50% of plants spotted with brownish-
purple. The spots are generally concentrated
onto the outer half of the leaf and are variable
in shape; they may be rings or elongated into
transverse bars. Rarely, there are spots on
the lower surface of the leaf, too, or it has a
purple rim.
Spike: Crowded, with about 20 flowers
occupying 25-50% of the stem length.
Bract: Fairly long, green, variably washed
purple (sometimes strongly so) but only
occasionally spotted.
Ovary: Slender, twisted and ridged, green,
often heavily washed purple.
Flower: Purplish-pink with a well-marked
three-lobed lip. The lateral sepals are oval,
asymmetrical, variably marked darker with dots,
squiggles, or rings (sometimes unmarked) and
usually held closer to the horizontal than the
vertical. The upper sepal is oval, and the petals
M 24 May, Co. Clare. The lip has three well-defined
are slightly shorter, narrower and more pointed. rounded lobes marked with heavy dots and dashes, often
Together they form a hood over the column. both within and without double loop markings.
004 systematic.indd 337 29/1/09 12:34:54

Marsh Orchid, which it resembles in flower

colour and pattern although the dark spots on
the lip are more numerous and more extensive.
A hyperchromic variant with an excess of
pigmentation and a solidly dark lip has, rarely,
been recorded.
BIOLOGY
Hybrids
‘D. x aschersoniana’, the hybrid with Early
Marsh Orchid, has been recorded in Co.
Limerick. This name may be changed following
taxonomic changes.
D. x dinglensis, the highly fertile hybrid with
Heath Spotted Orchid, has been found widely.
D. x braunii, the hybrid with Common Spotted
Orchid, has been found in Co. Clare.
The specific name occidentalis derives from
‘occidental’, i.e. ‘western’, while the older name
majalis means ‘in May’.
Until recently this species was part of a
conglomerate of marsh orchids that took the
name ‘Western Marsh Orchid Dactylorhiza
majalis’. The conglomerate included D.
majalis from mainland Europe as well as Irish
Marsh Orchid, Hebridean Marsh Orchid
and the cambrensis subspecies of Northern
Marsh Orchid. Recent research, including the
M 24 May, Co. Clare. The coloration varies from a deep analysis of DNA, has suggested that Irish and
magenta to paler purple or deep pink. Hebridean Marsh Orchids are unique endemic
species and that cambrensis from west Wales
but is scarce. It always has unspotted leaves and
belongs with Northern Marsh Orchid. This
bracts. Its flowers are, on average, paler and less
means that D. majalis of Europe does not occur
intense and the lip is often flat and marked with
anywhere in the British Isles.
dots and dashes that do not form a pattern of
This species is a tetraploid marsh orchid and
loops. It also tends to be slightly shorter and
the chromosome number is 2n = 80.
stockier and is said to flower a little later, into
July.
This form is sometimes treated as a distinct HISTORY AND
species, ‘D. kerryensis’, due to its floral characters CONSERVATION
and later flowering, and allied with Southern Irish plants were recognised as being distinctive
004 systematic.indd 338 29/1/09 12:34:56

O 25 May, Co. Clare. The

sides of the lip can be
sharply folded downwards,
and the markings are very
variable.
in the 1930s and were first given the name Past and present occurrence of Irish Marsh Orchid in Brit-
occidentalis in 1935 by the British botanist squares of the National Grid; data from the New Atlas).
H.W. Pugsley.
Britain Ireland
Irish Marsh Orchid is vulnerable
to agricultural changes, especially with 1500-1999
European Union money funding agricultural
current range 0 130 (13%*)
‘improvement’ in Eire, but so far losses appear
to be modest. The difficulties of marsh orchid % lost, 1500-1969 9%
identification and the relatively low level of % lost, 1970-1986 9%
botanical recording in Ireland may be obscuring % lost, total 18%
any changes in its status. * current range as a % of the total number of 10km squares
004 systematic.indd 339 29/1/09 12:34:58

Distribution Growth pattern
Six species, found in central and southern The aerial stem grows from a pair of tubers
Europe, North Africa and the Middle East. and the annual cycle of the replacement of
tubers is similar to that of the genus Orchis
Classification
(see p.200).
Until recently Dense-flowered Orchid was
the only species in this genus, but recent Vegetative reproduction
genetic studies have seen Neotinea enlarged Additional tubers may be formed at the base
to embrace five more from the genus Orchis, of the stem and these will form separate plants
including Burnt Orchid. Paradoxically, when the connecting stem dies off in the
Neotinea is probably as close to the bee autumn.
orchids Ophrys, tongue orchids Serapias and
Name
Lizard Orchid as it is to the genus Orchis.
The genus is named neo (= new) plus tinea,
Floral structures after Vicenzo Tineo, a Sicilian botanist
There are two pollinia, each narrowing to (1791-1856). An alternative origin has been
a short stalk which attaches it to one of the suggested, ‘New-Tinnea’, due to its supposed
viscidia, which in turn are enclosed in a similarity to Tinnea, a genus of shrubby herbs
bursicle. There are two stigmas, placed on from Africa named after Dutch explorers
each side of the mouth of the spur but joined Henrietta Tinne and her daughters.
at the base.
GenuS NEOTINEA
denSe-flOwered
and Burnt OrchidS
004 systematic.indd 340 29/1/09 12:35:01
DENSE-FLOWERED ORCHID l 341
denSe-flOwered Orchid
Neotinea maculata
This small and rather inconspicuous orchid is confined to western Ireland, although for a
while it also occurred on the Isle of Man. It has a very peculiar world range, being essentially
a Mediterranean species adapted to the mild wet winters and hot dry summers of that
region and seemingly ill-fitted to the damp, windy, oceanic climate of western Ireland. It
is not unique, however, as several other Mediterranean species, such as Strawberry-tree,
occur in Ireland. Various theories have been advanced to explain this distribution, including
land-bridges between Ireland and the Continent or glacial refuges off western Ireland, and
the subject has generated a good deal of controversy. On balance, colonisation of Ireland via
wind-blown seed would seem the most likely explanation.
Identification
The tiny off-white flowers are distinctive.
Similar species
Superficially similar to Small White Orchid,
but the lip is a very different shape.
Habitat
Typically found on short turf on pastures,
road verges, limestone pavements and around
loughs and turloughs. It has occasionally been
recorded on dunes or from ash and hazel
woods in the hills. Although mostly confined
to calcareous soils on limestone it has been
found growing on gravels and also on light,
peaty soils overlying more acidic rocks. Occurs
up to 300m above sea level but most are found
below 100m.
Flowering period
Late April to early June, usually peaking in mid-
May. The flowers go over very quickly.
Range
Confined to Ireland, where most populations
are in The Burren region of Co. Clare and
adjacent Co. Galway, including the Arran
Islands. Otherwise there are a few very scattered
sites in eastern Co. Cork, Co. Limerick and
O 29 May, East Sussex. Burnt Orchids.
P 21 May, Co. Clare. Var. alba is commonest in Ireland

and has unspotted leaves and unmarked creamy flowers.
004 systematic.indd 341 29/1/09 12:35:03

342 l GENUS NEOTINEA
How to find it
1987-99
1970-86
Small and inconspicuous, this is a hard orchid
pre 1970 to spot, especially from walking height. It is
usually found in small scattered colonies with
1-20 flowering plants. Locally frequent in The
Burren, a mid-May excursion to this beautiful
area is well worthwhile.
DESCRIPTION (var. alba)

Height: 4-40cm but mostly 6-15cm and very
rarely over 30cm.
Stem: Green, with one or two brown
membranous sheaths at the extreme base.
Leaves: There are three to six rather dark green
leaves, of which two or three (sometimes four)
are broadly oblong-elliptical in shape and held
rather spreading at the base of the stem; the
Co. Donegal and in Northern Ireland in remainder are narrower and more pointed and
Co. Fermanagh. Extinct in the Isle of Man. loosely sheathe the stem with the uppermost
World Range: The Mediterranean basin even smaller and more bract-like. The leaves
from Portugal and Spain east to Cyprus, appear in October and are wintergreen.
Turkey, Syria and Israel; ranges north to Spike: Cylindrical and dense, composed of 15-
southwest and southern France, the former 20 tiny flowers (occasionally as many as 35), all
Yugoslavia, Greece, Romania and the Ukraine, facing more-or-less in the same direction.
and south to Morocco, Algeria and Tunisia Bract: Green, becoming whitish towards the
in North Africa. Also found on most of the tip. The bracts are lanceolate with a pointed
Mediterranean islands, the Canary Islands and tip and sometimes a tooth at the side and are a
Madeira. little shorter than the ovary, which they clasp.
M 21 May, Co. Clare.With Bird’s-foot Trefoil in the rocky landscape of The Burren.
004 systematic.indd 342 29/1/09 12:35:04

DENSE-FLOWERED ORCHID l 343
M 21 May, Co. Clare. The tiny flowers have a distinctly

three-lobed lip and vaguely resemble human figures.
Ovary: Pale green, fat, cigar-shaped and

twisted, with three diffuse ribs. The ovary is
rather larger than the flower and stands upright.
It narrows and bends at the tip, however, to
hold the flower facing outwards.
Flower: Small and whitish, resembling a tiny
man with a rather oversized helmet. The sepals
are white, washed greenish and with green
veins. They are oval-lanceolate to lanceolate,
pointed at the tip and fused at the base to
form a tight elongated hood. The petals are
greenish-white with green veins, very narrow,
strap-shaped and pointed; they are enclosed
within the hood. The lip is small in relation to
the hood and is held at an angle forwards and
downwards. It is three-lobed with the central
lobe strap-shaped and notched or shallowly
forked at the tip (sometimes with a small tooth
in the notch). The side-lobes are positioned
near the base of the lip and are rather shorter
and narrower. The spur is very short and
conical. The flowers are reported to smell faintly
of vanilla.
Subspecies
None.
Var. alba has unspotted leaves and whitish or
creamy flowers (rarely more greenish). This is
M 21 May, Co. Clare. Like most ‘Mediterranean’ orchids,
the common variety in Ireland. the leaves appear in October and are wintergreen, dying
Var. maculata has spotted leaves with the off after flowering.
004 systematic.indd 343 29/1/09 12:35:07

produced in good quantities (c. 1350 seeds per

capsule).
Hybrids
None known.
The specific name maculata means ‘spotted’ or
‘blotched’.
HISTORY AND
CONSERVATION
First found in Ireland in May 1864 by Miss
Frances More at Castle Taylor in Co. Galway.
It was discovered on the Isle of Man in 1966
in an area of dunes on the north coast at The
Ayres, Ballaghennie. This colony persisted
until 1986 but Dense-flowered Orchid is now
extinct there.
In Ireland the species is scarce and local
and has been lost from 46% of the historical
range. Many of the losses are comparatively
M 24 May, Co. Clare. The ‘typical’ variety, which is scarce recent, although some of the decline has
in Ireland, has spotted leaves, purple-veined flowers and been offset by the discovery of new sites.
often spotted ovaries, too.
It is extinct in Co. Offaly, Co. Roscommon
reddish or purple spots arranged in parallel and perhaps Co. Mayo (where noted from
longitudinal lines. The lip is pale pink or Lough Mask by Summerhayes, 1968). The
marked with a longitudinal pink stripe and, ‘improvement’ of pastures and overgrazing are
the sepals and petals have pink or brown likely causes for the losses. With the current
veins. Occasionally the stem is spotted and rapid pace of development in Eire there is
sometimes also the bracts, ovaries and even cause for concern about the future of such a
sepals and petals. This is the ‘typical’ variety (in localised orchid.
that it takes its name from the species) but it is
Past and present occurrence of Dense-flowered Orchid
relatively scarce in Ireland. in Britain and Ireland (based on presence or absence in
Var. luteola has primrose-yellow flowers. It is 10km squares of the National Grid; data from the New
very rare and perhaps extinct in Ireland. Atlas).
Britain Ireland
BIOLOGY
Pollination and reproduction 1500-1999
The spur contains traces of nectar, suggesting current range 0 13 (1.3%*)
that the flowers are attractive to small insects.
% lost, 1500-1969 0% 21%
Although cross-pollination is possible, however,
% lost, 1970-1986 100% 25%
most or all plants are self-pollinated, sometimes
even before the buds have opened. Whether % lost, total 100% 46%
self-pollinated or cross-pollinated, seed is * current range as a % of the total number of 10km squares
004 systematic.indd 344 29/1/09 12:35:09

BURNT ORCHID l 345
Burnt Orchid Endangered, BAP

Neotinea ustulata
Formerly: Orchis ustulata; Other names: Burnt-tip Orchid

This delicate species is one of our most attractive and delightful orchids but, sadly, is also
probably the orchid that has suffered the greatest decline in the last 50 years or so. It is now
confined to around 75 sites in England and just one in Wales, mostly on well-managed
chalk grassland reserves in southern England but with a few colonies still surviving on the
limestone of northern England. The unopened buds are deep purple, giving the top of the
spike a scorched or burnt appearance, hence the name. Burnt Orchid is the county flower
of Wiltshire.
Identification
Burnt Orchid is one of the group of orchids
in which the flower resembles a tiny human
figure, with the lip divided into ‘arms’ and
short, stumpy ‘legs’. It is easy to identify, with
a distinctive combination of small size, dark
reddish-purple buds and a white lip marked
with fine reddish-purple spots. The contrast
between the dark buds at the top of the spike
and the white lips of the lower flowers is very
striking. There are two forms of Burnt Orchid,
early and late flowering, but they do not differ
significantly in appearance.
Similar species
Lady Orchid resembles Burnt Orchid in general
flower structure and colour but is very much
larger, and its lip differs in the details of shape
and coloration.
Habitat
Burnt Orchid is found on ancient short
grassland on chalk and limestone soils, often on
south- or west-facing slopes, although the late-
flowering populations are not so fussy about
the aspect. In southern England the species
favours the narrow ‘terracettes’ that follow the
contours of the slope on the chalk downs and
also Bronze and Iron Age earthworks where
the ground has been undisturbed for centuries.
Only on rare occasions will it colonise new sites.
For example, at Martin Down in Hampshire
P 13 July, Hampshire. In the late flowering var. aestivalis

the hood does not fade after the flower has opened.
004 systematic.indd 345 29/1/09 12:35:11

004 systematic.indd 346 29/1/09 12:35:14
BURNT ORCHID l 347
it appeared in the 1980s on grassland that

1987-99
had been under the plough until 1957. Burnt 1970-86
Orchid is also sometimes found in alluvial pre 1970
water meadows where the silt is derived from

chalk, e.g. in Lincolnshire and formerly at sites
in Hampshire and Oxfordshire. This habitat
was always uncommon in southern England but
in the past Burnt Orchid was found relatively
frequently in riverside pastures in the north. It
is predominantly a lowland species but has been
recorded up to 275m above sea level.
Flowering period
There are two varieties that differ in their
flowering period. The early-flowering form
appears from the second or third week of May
to mid-June in southern England and may, on
average, be just a few days later in the north; it Wiltshire (four sites), Hampshire (five sites,
is usually at its best in the last ten days of May. plus one now destroyed) and East Sussex (14
The late flowering form flowers from the end of sites, mostly between Lewes and Eastbourne,
June through July to early August. one holding nearly 1,000 spikes in 2002).
World range: Confined to Europe, although it
Range extends just slightly into western Siberia (to c.
Once found throughout the chalk and 70°E). It occurs north to Denmark, southern
limestone areas of England, the species has Sweden, Estonia and the St Petersburg region
undergone a major decline and is now extinct of Russia and south to northern Spain, Italy,
in many areas. In southern England, Burnt northern Greece, Bulgaria and the Ukraine. It
Orchid can still be found in Dorset, the Isle of is also found in the Caucasus. In the south of
Wight and West Sussex but the strongholds are the range it is mostly found in the mountains
the downs of Wiltshire, Hampshire and East and is absent from the Mediterranean lowlands.
Sussex. Outlying relict populations cling on Extinct in Holland.
in Gloucestershire, Berkshire, Hertfordshire,
Bedfordshire and Kent. The decline has How to find it
been even more marked in the Midlands and Sadly, the chances of coming across this lovely
northern England, and the species is now only little orchid unexpectedly have become slim but
found in Derbyshire, Lincolnshire, northern it is nevertheless still worthwhile looking for in
Yorkshire and at a single site on the coast of suitable undisturbed short swards. Otherwise,
Co. Durham (the most northerly in Britain). In a visit to a good chalk grassland reserve is
1993, bucking the trend of range contraction necessary to see it but even then it can be hard
and decline, Burnt Orchid was recorded for the to find. The colonies can be very localised, with
first time in Wales, on limestone grassland at hundreds of plants in one area and few or none
Llanmadog, Glamorganshire. elsewhere, even on seemingly suitable ground.
The late-flowering variety aestivalis (see Being so small, it can be surprisingly hard to
Variation and varieties) is only found in spot among the various short downland herbs
such as thyme and milkwort. As with many
southern England. It has been recorded from
orchids, the number of flowering spikes varies
O 29 May, East Sussex. In the early flowering form the greatly from year to year and in extremely dry
hood fades once the flower has opened. seasons there may be none at all. One of the
004 systematic.indd 347 29/1/09 12:35:16

to form a spike that is initially conical but

lengthens and becomes cylindrical when all the
flowers have opened.
Bract: Reddish-purple, lanceolate and rather
short, at around two-thirds the length of the
ovary.
Ovary: Green, twisted and obscurely six-ribbed.
Flower: The sepals are oval, the lateral sepals
asymmetrical with their outer surfaces dark
reddish-purple (thus the unopened buds are
very dark) and their inner surfaces greenish.
The petals are paler and more strap-shaped.
The sepals and petals form a compact hood that
embraces the column; this is reddish-purple
when the flower is fresh although it quickly
fades (and see var. aestivalis below). The lip
is white with a few scattered reddish-purple
spots (formed by minute papillae) and is deeply
lobed to form two broad chubby ‘arms’ and two
short stumpy ‘legs’, sometimes with a tiny tooth
M 13 July, Hampshire. The flowers look like tiny human
figures; this is the late flowering var. aestivalis. between them. There is a groove in the centre
of the lip towards the base that leads into the
best sites is Mount Caburn (East Sussex). short, conical, down-curved spur. The column
Others include Seven Sisters (also East Sussex), is short and whitish and the pollinia are yellow.
Martin Down (Hampshire) and Pewsey The flowers of the early-flowering variety have a
Downs, Parsonage Downs and Clearbury strong, sweet, honey-like scent said to be similar
Rings (Wiltshire). to Heliotrope, and var. aestivalis has a weak,
citron-like scent that is vaguely unpleasant.
DESCRIPTION
Subspecies
Height: 2.5-15cm, very exceptionally to 30cm.
None.
In southern England it is usually around 4-10cm,
with the very shortest plants found at heavily Variation and varieties
grazed sites. In northern England it is a little Var. ustulata and var. aestivalis Very unusually
taller, usually around 9-12cm, even to 15-25cm there are two varieties of the Burnt Orchid
when growing among taller vegetation. which differ in their flowering time. The early-
Stem: Yellowish-green, slender and ridged flowering var. ustulata is much the commoner
towards the tip, with two or three membranous and blooms in late May and early June. The
white sheaths at the very base. late-flowering var. aestivalis is only found in
Leaves: Green with a faint blue tone, fading to southern England and flowers around July. The
pale green by flowering time. There is a rosette two varieties are rarely if ever found together,
of two to five elliptical-oblong keeled leaves but at a site in East Sussex a colony of 200 late-
at the base of the stem and one or two bract- flowering plants merges with a much smaller
like leaves higher up. The leaves appear in the group of early-flowering plants on a north-
autumn, are wintergreen and start to wither facing chalk slope.
as the plant comes into flower (by which time The two varieties differ slightly in
non-flowering plants have already vanished). appearance. In the late-flowering aestivalis the
Spike: The 15-50 flowers are densely packed reddish-purple of the hood is on average darker
004 systematic.indd 348 29/1/09 12:35:17

BURNT ORCHID l 349
and the colour does not fade once the flower

has opened. There is also a distinct rose-purple
wash to the edges of the lip and this wash
may spread across the lip. The early-flowering
variety never shows the persistent dark hood or
the coloured fringe to the lip. Other differences
are rather subtler. On average, var. aestivalis is
taller, although this may simply be a product
of the plants having to compete with the taller
vegetation found later in the season. It has a
more open spike of slightly smaller flowers and
the lip is also shorter, with a slightly narrower
‘waist’ and shorter ‘legs’ which have a deeper
cleft between them. Also, the spots on the lip
are larger.
Similar late-flowering populations in
Europe have been described as a distinct
subspecies, N. u. aestivalis, and it has even
been suggested that they should be treated as
a different species. Genetic analysis shows that
there is far too small a difference between the
early- and late-flowering varieties to justify this.
Var. albiflora has unmarked white flowers. It is
very rare but has been recorded in Derbyshire,
Kent, Berkshire, Hampshire and Wiltshire.
BIOLOGY
Poorly known. Early-flowering plants are
pollinated by flies that feed on nectar and M 13 July, Hampshire. The late flowering var. aestivalis
sugary plant juices. Presumably the flies are differs slightly in appearance from typical plants as well
initially attracted by the combination of as in its flowering period, but genetic research shows no
appreciable difference between the two.
colour and scent, but this species does not
produce nectar to reward them and they must flowers but have not been recorded carrying
be satisfied in some other way. Once on the pollinia. With its different scent and later
flower, the flies work from the uppermost, flowering, var. aestivalis may attract a different
unopened buds downwards and insert their suite of species and, in Europe, beetles have
proboscis in a head-down position. They been recorded as pollinators. Whatever its
‘taste’ with their feet beforehand and there mechanism, pollination is not very efficient in
may be some sugary secretion from the flower England and seed-set is relatively poor, with
to guide them into the ‘correct’ position; the around 20% of flowers producing ripe capsules.
groove at the base of the lip may in addition Nevertheless, most new plants are recruited to
act as a ‘leading line’. The mouth of the spur is the population from seed.
narrow and rather like a keyhole; this shape Vegetative reproduction takes place but is
and the design of the column may be related thought to be relatively unimportant. Groups of
to the unconventional ‘upside-down’ position four to six spikes are not uncommon, however,
of the pollinating fly. Butterflies also visit the and we have seen a compact cluster of 12 (see
004 systematic.indd 349 29/1/09 12:35:18

photo); such groups may well be the product of of development). Both in cultivation and in the
vegetative reproduction. wild Burnt Orchids can flower within about
Development and growth three years of germination, but older estimates
The aerial stem grows from a pair of tubers. of up to 16 years are still quoted. This figure
The degree of fungal ‘infection’ is unclear, is based on the pioneering work of Fuchs and
with both high and low rates being reported. Ziegenspeck in the 1920s. They estimated the
‘Dormancy’ is common, however, and Burnt length of time the protocorm spent developing
Orchids are able to spend up to three years underground by examining it for ‘constrictions’,
(sometimes even four) underground without each of which was thought to represent a
producing aerial parts. During this period the year’s growth, like annual rings on a tree. This
fungal ‘partner’ must play a major role (although technique is now known to be unreliable.
dormant plants have a significantly reduced Hybrids
chance of surviving). Most plants flower for one None are known.
to four seasons in succession (rarely for up to
seven successive years) and then either die or Name and classification
retreat into a period of dormancy underground The specific name ustulata means ‘scorched-
or as non-flowering rosettes. looking’. Until recently Burnt Orchid was placed
Seed germinates to produce a protocorm in the genus Orchis, but genetic studies have
which grows to a length of 20-30mm before the resulted in its transfer to the genus Neotinea.
first root is produced (the longest protocorm There it joins the superficially rather different
known among orchids with a similar pattern Dense-flowered Orchid.
M 13 July, Hampshire.This group of 12 plants (var. aestivalis). is probably the product of vegetative reproduction.
004 systematic.indd 350 29/1/09 12:35:20

BURNT ORCHID l 351
HISTORY AND
CONSERVATION
The first British record was in 1634 in Thomas
Johnson’s Mercurius Botanicus (‘Botanical
Mercury’): ‘...in montosis pratis’ (mountain
pastures). The first specific locality was given in
1650 in William How’s Phytologia Britannica
natales exhibens Indigenarum Stirpium sponte
emergentium (‘A British botany presenting the
origins of native wild plants’): ‘On Scosby-lease,
Mr Stonehouse’. Therefore the first localised
record was from near Doncaster in northern
England rather than the chalk downs of the
south where it was presumably much commoner.
Formerly locally frequent over much of
England, the Burnt Orchid has suffered one
of the most catastrophic declines of our wild
orchids. It is now greatly reduced, both in terms
of numbers and distribution, and is classified
as Endangered. Out of a total historical range
covering 265 10km squares, the New Atlas
records a post-1987 presence in just 55,
representing a 79% decline. It is now Nationally
Scarce. M 29 May, East Sussex. The reddish-purple buds and
hood give the flowers a ‘scorched’ or ‘burnt’ appearance.
Past and present occurrence of Burnt Orchid in Britain
and Ireland (based on presence or absence in 10km through the 19th and 20th centuries due to
the continued loss of downland and pastures
Britain Ireland (perhaps especially during World War Two),
total historical range, 265 0 while in the latter half of the 20th century
1500-1999 the ‘improvement’ of grasslands with artificial
current range 55 (1.9%*) 0 fertilisers and pesticides continued to take a toll.
The remaining grassland sites are vulnerable
% lost, 1500-1969 72%
to scrub invasion due to a lack of grazing,
% lost, 1970-1986 7%
especially in southern England, where first
% lost, total 79% rabbits were decimated by myxomatosis and
* current range as a % of the total number of 10km squares latterly livestock has become less economically
viable. Paradoxically, overgrazing is also a
It is possible that subtle changes in climate problem, especially in northern England, where
in the 19th and 20th centuries may have losses due to building and other development
caused some decline, but most losses can be have also occurred. Sites were even wilfully
directly attributed to agricultural changes. The destroyed, as at Cottondale in southeast
species vanished from some counties at the Yorkshire, ploughed out in 1965 ‘to counter the
edge of the range long ago (e.g. last recorded attention shown it by botanists’.
in Norfolk at the end of the 18th century), There were around 350 historical localities
mainly due to the ploughing of grasslands in southern England but only around 75
following the Enclosures. The decline continued colonies survived into the 1990s. Most sites
004 systematic.indd 351 29/1/09 12:35:21

have fewer than 50 flowering plants and,

indeed, at some just one or two spikes make
ExtINCt
sporadic appearances; fewer than ten regularly
the Burnt Orchid is extinct in the following vice-
hold more than 200 plants. The strongholds counties (the number of recorded sites, followed
are now in Wiltshire, Hampshire and East by the date of the last record, is given in paren-
Sussex, and notable concentrations include theses). Source: Foley (1987, 1990, 1992).
Parsonage Down in Wiltshire where there
may be 30,000 flower spikes spread over 95ha South Devon (four sites, last seen in 1932)
in one continuous ‘mega-colony’, probably North Somerset (seven, 1961)
West Kent (three, late 19th century)
the largest surviving population in northwest
Surrey (twelve, 1966)
Europe. In other counties where Burnt Orchid
North Essex (two, 19th century)
still survives, many or most of the sites have Middlesex (one, pre 1737)
been lost. Thus in Dorset only two out of 12 Oxfordshire (two, 1982)
known sites remain and the equivalent figures Buckinghamshire (five, 1961)
for the Isle of Wight are one of 12, for West East Suffolk (but this 19th century site might
Sussex one of six, for Kent seven of 18, for have been in Norfolk)
Berkshire one of 17, for Hertfordshire one of West Suffolk (three, 1939)
11, for Bedfordshire one of ten and for east West Norfolk (one, 18th century)
Gloucestershire one of 21. Cambridgeshire (twelve, 1955)
Of over 120 sites recorded from northern Huntingdonshire (one, 1890)
Northamptonshire (four, 1956)
England around 30 survive. There are a few
West Gloucestershire (eight sites)
in Derbyshire and northern Yorkshire; it
Herefordshire (two, 19th century)
reappeared in mid-west Yorkshire in 1988 but Worcestershire (five, early 20th century)
the single plant was promptly dug up. In the Staffordshire (two, early 20th century)
north and west of Lincolnshire there are three Shropshire (six, 19th century)
small colonies and just one remains in Co. Leicestershire (two doubtful old records)
Durham. Only two of the sites in the north, in Rutland (three, around 1915)
Derbyshire and northwest Yorkshire, hold more Nottinghamshire (three sites)
than 200 spikes, and the total population in West Lancashire (two, 1940s)
northern England is probably fewer than 1,000 Southeast Yorkshire (eleven, 1973)
flowering plants. Southwest Yorkshire (four, prior to 1888)
Fortunately many sites are now protected South Northumberland (one, prior to 1868)
Westmorland (seven, 1970s)
as SSSIs, nature reserves or are on Ministry
Cumberland (ten, early 1970s)
of Defence land and managed for the benefit
of their chalk flora and fauna. As the chalk
landscape was created and maintained by
hundreds or thousands of years of grazing, it is
important to continue this practice. For Burnt
Orchid the ideal regime is light spring grazing
until late April with the return of sheep from
late July until late September (when the new
leaves start to appear); cattle are unsuitable as
they can trample and destroy the plants and
also damage the turf.Distribution
004 systematic.indd 352 29/1/09 12:35:22

● 353
GENUS HIMANTOGLOSSUM
LIZARD ORCHID
Distribution
There are five closely related
species in Europe, North Africa
and in the Middle East from
Turkey and Lebanon to Iran.
Perhaps surprisingly, their closest
relatives include the bee and
spider orchids Ophrys, tongue
orchids Serapias and pyramidal
orchids Anacamptis.
Name
The generic name
Himantoglossum derives from
the Greek himas ‘leather strap’ or
‘thong’ and glossa ‘tongue’, thus
‘strap-tongued’, a reference to the
long, narrow lip.
004 systematic.indd 353 12/2/09 17:06:31

354 l GENUS HIMANTOGLOSSUM
lizard Orchid Near Threatened

Himantoglossum hircinum
The bizarre flowers of this robust orchid bear a fanciful resemblance to a lizard and smell
strongly of billy goat. It has always been rare in Britain but its range and numbers have ebbed
and flowed, probably due to subtle changes in climate; it once occurred north to Yorkshire
but retreated southwards and is now confined to a few sites in southern England, although it
is currently on the increase again and could turn up almost anywhere in the south and east.
Identification
Very distinctive. The tall spikes of large greyish-
green flowers have an untidy, straggly, ragged
appearance and on close inspection their
structure is unique. The long central lobe of
the lip resembles a lizard’s tail, and the shorter
side-lobes form the back legs. The hood of the
flower is said to recall the head and body of
the lizard, but it seems much more reasonable
to say that the fore-quarters of the lizard have
been swallowed and vanished into the throat of
the flower.
Similar species
None.
Habitat
The Lizard Orchid usually grows in open sunny
situations among fairly rank grass, although
sometimes it also occurs on shorter and more
closely-cropped swards. It can appear among
scrub or on the edge of woodland but would
eventually be shaded out if the canopy developed
too much. It is always found on well-drained
soils, usually on chalk but sometimes also on
limestone and occasionally on boulder clays,
sands or gravels; it has even been recorded
growing through broken tarmac. Whatever the
soil, it will probably be calcium-rich, although
some recent records have been on neutral
soils. Suitable sites include road verges, railway
embankments, ancient earthworks, field margins,
M 30 June, East Sussex. By flowering time the lower old chalk pits and dunes, and it even grows on
leaves, which appear in the autumn, have withered.
lawns on one favoured housing estate. At least six
O 25 June, Cambridgeshire. Smelling of billy goat and of the 19 recent populations are on golf courses,
with such extravagant flowers, this is a truly bizarre orchid.
and it has been suggested that golfers may have
P 27 June, Kent. carried the seeds from one course to the next on
004 systematic.indd 354 29/1/09 12:35:27

LIZARD ORCHID l 355
004 systematic.indd 355 29/1/09 12:35:32

their shoes or equipment. Found from sea level

1987-99
to about 200m in Gloucestershire and on the 1970-86
North and South Downs. pre 1970
Flowering period
Early June to late July, sometimes from late May.
Traditionally peaking in the first week of July it
has, like many other orchids, tended to flower
earlier in the last few years, in mid-June.
Range
The distribution has ebbed and flowed, but
all the records have been southeast of a line
from the Severn to the Humber apart from
one old site further north in Yorkshire and a
doubtful record from Lancashire. In recent
years there has been a very large established
colony at Sandwich Bay in Kent and a smaller
permanent colony at Newmarket in Suffolk, as Balkans are often treated as separate species.)
well as small colonies in East Sussex, Dorset The species is commonest in France, where it is
and Suffolk. Other recent records, either of a roadside weed in the wine-growing regions.
sporadic appearances or tiny populations,
come from north Somerset, north Hampshire, How to find it
West Sussex, Surrey, Oxfordshire and west Despite their size, the greyish-green spikes can
Gloucestershire. It has also been recorded from be hard to see among long grass. Undoubtedly
Jersey and Guernsey in the Channel Islands. the best site is at Sandwich Bay in Kent where
World range: Southwest and southern Europe, there have been up to 3,000 flower spikes in
north to Holland, Germany, Austria and the recent years, with 5,000 in 2000 (when the total
Czech Republic, south to northern Spain (and population was estimated at 27,500 plants over
very scattered in central and southern Spain), an area of 0.5 square km). Another well-known
Italy and Sicily and east to Greece, also North colony is on the Devil’s Dyke that runs across
Africa in Morocco, Algeria and Tunisia. (The Newmarket’s July racecourse, with around 200-
lizard orchids further to the east and in the 250 flowering plants.
M 27 June, Kent. Lizard Orchid is an adaptable species and a prestigious colonist on a lawn.
004 systematic.indd 356 29/1/09 12:35:34

LIZARD ORCHID l 357
M 28 June, Cambridgeshire. As wih many orchids, the function of the elaborate lip is obscure.
DESCRIPTION are oval and are fused at the base to form

Height: 25-90cm but usually below 60cm and a loose hood. They are pale greyish-green,
commonly around 30cm. variably flushed or rimmed purple with a paler
Stem: Pale green with faint purple blotches, interior that has lines of reddish-purple spots
ridged towards the tip and with two or three and dashes along the veins (which may show
scale leaves at the extreme base. through to the exterior). The petals are very
Leaves: Numerous, greyish-green and narrow, strap-shaped and spotted reddish-
sometimes blotched with purple. There are purple. The lip is divided into three lobes.
four to ten large, oval-oblong, keeled basal The central lobe is 2.5-6cm long, narrow,
leaves forming an untidy rosette and three to parallel-sided and ribbon-like, with a deeply
five smaller, narrower and more pointed leaves notched tip. It is twisted through two or three
which loosely clasp the stem up to the flower turns and is whitish at the base, becoming pale
spike. The leaves are wintergreen, and the basal lilac-brown for the outer 80%. The side-lobes
leaves have yellowed and often withered by are very much shorter and a little narrower,
flowering time, especially in dry years. pointed and pale lilac-brown. The sides of the
Spike: Tall, occupying almost 50% of the stem, lip at the base are boldly folded or corrugated,
cylindrical and rather dense, with 15-80 flowers. and this corrugation continues on to the outer
Bract: Pale green to off-white, variably but edges of the side-lobes. The centre of the base
often strongly washed reddish, narrow and of the lip is trough-like with a dense ‘fur’ of
pointed, and up to twice the length of the ovary. tiny white papillae and with variable bright
Ovary: Pale green, cylindrical, tapering both purplish-red tufted spots and blotches. The
towards the flower and at the base into a short lip is coiled like a spring in bud and slowly
stalk, twisted and with three ridges. unrolls to project outwards and downwards at
Flower: Overall greyish-green. The sepals about 45° (sometimes nearer the horizontal on
004 systematic.indd 357 29/1/09 12:35:36

few-flowered spikes). The spur is short, curved tuber, as well as the leaves, are often withering.
downwards and bluntly conical. The anther is At flowering time, therefore, the plant has
greenish-white, and the two greenish pollinia two large, egg-shaped oval tubers (one old and
are fixed to a single viscidium that is concealed withering, one new). After flowering and setting
in the purple bursicle. The stigmatic zone is seed the plant will disappear, leaving only the
purple. The strong scent recalls a billy goat and new underground tuber to survive the summer.
is most pungent in the evening. Individual plants can be long-lived, surviving
for up to at least 19 years after first emergence,
Subspecies
but they may not flower every year and can
None.
remain ‘dormant’ as underground tubers for
Variation and varieties a year or appear merely as a rosette of leaves,
None. sometimes for many years. An individual plant
in Sussex flowered in 1984 and then not again
BIOLOGY until 1995, and a colony in Suffolk of between
Pollination and reproduction 16 and 40 plants did not produce any flowers
The flowers are visited by a variety of insects, over a ten-year period.
including hover flies, various bees, wasps, ants, Seed germinates in the autumn up to
beetles, butterflies and moths. Of these, bees two years after the seeds have ripened and
are probably the main pollinators. There is no dispersed. Although some of the seed may blow
nectar (although glucose has been detected away, it is common for mature flowering plants
in the spur), and the flower seems to offer its to have large numbers of seedlings close by, and
pollinator no reward; despite this the pungent many seeds stay in the pods until the dead spike
scent may serve to attract flies and night- falls over. With the aid of fungi, the seedling
flying moths. Self-pollination is physically initially develops into a protocorm the size of a
possible but is thought not to occur. The small pea. In the spring following germination,
species is self-compatible, however, and solitary the protocorm produces the first tuber and
isolated plants do set seed; for this to happen then withers away, leaving just the tuber to
either self-pollination (autogamy) or cross- oversummer. In the autumn, a short rhizome
pollination from other flowers on the same grows from this tuber and develops a root and
spike (geitonogamy) must occur. Whatever a second tuber which is fully developed by the
the mechanism, it does not seem to be very following spring. This establishes the annual
efficient and only around 30% of flowers are cycle of replacement tubers. When the seedling
pollinated, although each capsule contains up is more than two years old the first leafy shoot
to 1,200 seeds, perhaps more. It is possible for appears above ground. The seedling usually
the species to reproduce vegetatively by forming remains in the one-leaf stage for two or more
extra tubers, but this seems to be a rare event. years and then moves on to two leaves, again for
several years, and then three leaves. Although
Development and growth in very favourable conditions a plant with two
The Lizard Orchid spends the summer ‘resting or three leaves can flower, it is only when it
period’ underground as a tuber. The aerial has grown four leaves that it can be considered
shoot appears from late August onwards, with mature. Even then it may not flower every
the lower leaves unfolding and some short and year. The period between germination and first
rather thick roots growing from the base of the flowering may therefore be a minimum of six
stem. These leaves will overwinter and by early years.
spring may start to blacken at the tips. By then a
new tuber has started to form from a bud at the Hybrids
base of the stem and by May the roots and older None.
004 systematic.indd 358 29/1/09 12:35:36

LIZARD ORCHID l 359
P 28 June, Cambridge-
shire.The long central lobe
of the lip uncoils like a
spring.
Name and classification species that is classed as Near Threatened and

The specific name hircinum means ‘of goats’, a fully protected under Schedule 8 of the Wildlife
reference to the scent. and Countryside Act 1981. Despite this it has
been targeted by collectors, and plants have
HISTORY AND been dug up at Sandwich Bay in Kent (where
CONSERVATION a 24-hour guard was maintained in the 1970s
The first British record dates from 1641 when and 1980s) and as recently as 2003 from a road
Thomas Johnson published the second part of verge in Dorset.
his Mercurius Botanicus (‘Botanical Mercury’) The Lizard Orchid has an intriguing history
and recorded it ‘Nigh the highway betweene in England. From its discovery until about 1850
Crayford and Dartford in Kent’. the species was restricted to the Dartford area
The Lizard Orchid is a Red Data Book of Kent, although the original 1641 colony
004 systematic.indd 359 29/1/09 12:35:38

Past and present occurrence of Lizard Orchid in Britain nine to 11, although some sites were lost and
squares of the National Grid; data from the New Atlas). others were gained. During this period all the
colonies were small and even the most thriving
Britain Ireland lasted only about 20 years. A further increase
started around 1994 with plants appearing west
1500-1999 to Somerset and Gloucestershire. There were
16 sites in 1996 and 19 by 2000. As usual some
of the new sites are ephemeral with the orchid
% lost, 1500-1969 76.5% flowering once and vanishing, e.g. by 2000 four
% lost, 1970-1986 6% of the 1996 sites had disappeared but there
% lost, total 82.5% were seven new ones (thus a net increase of
* current range as a % of the total number of 10km squares three). With protection, some colonies, such as
the very large population at Sandwich Bay, have
had been destroyed due to road-widening. expanded.
There was then a decline in that area and just Lizard Orchid is on the edge of its range
odd, isolated records in southeast England, in England, and it seems that subtle shifts in
mostly in Kent; by 1900 just four sites were climate have a big impact on its distribution.
known in England and it was thought that The species is wintergreen, and the amount
the Lizard Orchid was almost extinct. Then, of rainfall during the growing period of
during World War I and the 1920s there was September-April has a marked influence on
a marked expansion northwards with the the flowering success, with a dry autumn and
species turning up unpredictably, often just a winter limiting flowering. Furthermore, flower
single plant but sometimes in larger numbers, buds are initialised and formed over two
north to Yorkshire and west to Devon. A seasons so the rainfall over a two-year period
remarkably large proportion of the new sites is important in determining flowering success.
were found by schoolgirls. This expansion Conversely, severe drought may kill many plants
peaked around 1927 with 36 populations, but and wipe out small populations.
the record is slightly clouded as Lizard Orchids Lizard Orchid favours areas of long but
had been dug up and transplanted to gardens sparse grass that is cut once or twice a year.
(sometimes from as far away as France) and None of the English sites are grazed but the
at least two sites in this period are thought to species is tolerant of grazing, at least by cattle,
have originated with ‘garden escapes’. Adding a although rabbits have caused severe damage at
further complication, plants were sometimes times, especially during hard winters, as have
transplanted by well-meaning naturalists, often slugs. Some sites are managed to open up the
for ‘safe-keeping’, to new sites or even new vegetation and prevent scrub invasion (with the
counties. Whatever their origin, in addition to Newmarket site, for example, being burnt every
the counties detailed in ‘Range’ there are older fifth year during the winter), but those on dune
records from north Devon, Wiltshire, Isle of grassland do not need much management.
Wight, north Essex, Hertfordshire, Berkshire,
Buckinghamshire, Norfolk, Bedfordshire,
Worcestershire, Lincolnshire and northeast
Yorkshire.
In another change of fortunes, there was a
sudden drop in the number of colonies after
1934, although the geographical spread was
maintained. From the mid-1940s to 1993 the
total number of populations remained stable at
004 systematic.indd 360 29/1/09 12:35:39

Genus ANACAMPTIS
Loose-fLowered,
PyramidaL and
Green-winGed orchids
005 systematic.indd 361 29/1/09 12:40:29
362 l GENUS ANACAMPTIS
This genus formerly included just the Pyramidal Orchid. It is now a rather heterogeneous
combination of superficially different species, three of which occur in Britain.
The genus contains about 20 species, in The flowers do not produce nectar and may
northern, central and southern Europe, North rely on deceit to attract bees as pollinators.
Africa and southwest Asia, east to Afghanistan Pyramidal Orchid, and perhaps also Green-
and south to Yemen in the Arabian Peninsula. winged Orchid, do, however, produce a sugary
sap in the walls of the spur and this may
Classification
function as a reward for insect visitors.
From 1817, when it was described and named,
until 1997, this genus contained just one Growth pattern
species, Pyramidal Orchid. However, genetic The aerial stem grows from a pair of spherical
studies then led to the transfer into Anacamptis tubers, as in the genus Orchis (see p.200).
of 19 species from the genus Orchis, including
Green-winged and Loose-flowered Orchids. Vegetative reproduction
Surprisingly, the genus Anacamptis is closer to Additional tubers may be formed at the base
the tongue orchids Serapias, bee orchids Ophrys of the aerial stem and these will go on to form
and lizard orchids Himantoglossum than to the separate plants when the connecting stem dies
genus Orchis from which those additional 19 off in the autumn.
species came.
Name
Floral structures The generic name Anacamptis derives from the
In Loose-flowered and Green-winged Orchids Greek anakampto ‘to bend back’. It is a reference
the two pollinia narrow to a caudicle (stalk) and to the structure of the flower of Pyramidal
each is attached to their own viscidium which, Orchid but there is disagreement as to which
in turn, is enclosed in a bursicle. The two-lobed particular bit of the flower is being referred to.
stigma lies on the roof of the mouth of the It may be the long spur of the flower, the ‘guide-
spur. In Pyramidal Orchid the two pollinia are plates’ at the base of the lip or the position of
attached to a single strap-shaped viscidium. the pollinia.
O 10 May, Norfolk. Sadly, although relatively common

not so long ago, Green-winged Orchid has been banished
from around half its range.
005 systematic.indd 362 29/1/09 12:40:29

LOOSE-FLOWERED ORCHID l 363
Loose-fLowered orchid
Anacamptis laxiflora
Formerly: Orchis laxiflora; Other names: Jersey Orchid

This stately orchid provides a real splash of colour to wet meadows in the spring. In Britain
it is confined to the Channel Islands, but it is widespread in the Mediterranean region,
although everywhere it is threatened by drainage and agricultural ‘improvement’.
Identification
A ‘typical’ orchid with a tall, loose spike of
purple flowers. The leaves are narrow and held
erect. The lip is purple with a long white central
zone that is unspotted or has some darker
purple markings. The central lobe is shorter
than the side-lobes and the sides of the lip are
very sharply folded downwards; the lateral
sepals are held vertically upwards. This gives the
lip a tall but ‘flattened’ profile.
Similar species
Early Purple Orchid is superficially similar but
has broader leaves that are usually spotted and
held in a rosette flat on the ground. Its flower
spike is more compact, and the centre of the lip
is spotted. The lip itself is distinctly three-lobed
with the central lobe larger and longer than the
side-lobes.
Habitat
Damp meadows and marshy fields flushed with
base-rich water.
Flowering period
Early May to mid-June but mostly in late May.
Range
Confined to Guernsey and Jersey in the
Channel Islands. There are old records from
Co. Durham, including one dating from
1872 when it was found growing north of
Hartlepool. These plants may have originated
as casual imports with wool ‘shoddy’. World
range: Essentially Mediterranean, found from
Portugal east to Greece, Cyprus and Turkey,
and including most of the Mediterranean
islands. Occurs north to north-central France
P 27 May, Sussex (Wakehurst Place).
005 systematic.indd 363 29/1/09 12:40:34

broad white stripe down the centre of the lip.

1987-99
1970-86
The sepals are oval to oblong and blunt-tipped;
pre 1970 the lateral sepals are asymmetrical and held
erect above the rest of the flower (sometimes
touching), and the upper sepal is held over the
petals, which are slightly smaller and form a
loose hood over the column. The lip is broader
than long with two large, rounded side-lobes
and a rather shorter blunt central lobe (the
central lobe is sometimes absent altogether).
The sides of the lip are so sharply folded
downwards that the side-lobes may touch at
their tips. The spur is long and slender but
broadens towards the tip, which is usually
notched, and either straight or slightly up-
curved. The centre of the lip may have small,
dark, vertical streaks within the white central
(Seine-Maritime and Marne), Switzerland and zone, extending into the purple borders.
the former Yugoslavia. It was discovered for the
first time in Holland relatively recently but is Subspecies
extinct in Belgium. None.
How to find it Variation and varieties

The tall purple flower spikes are conspicuous. Var. rosea has pale rose-pink flowers but is rare.
The best site on Jersey is Le Noir Pré and on Var. albiflora has white flowers but is rare.
Guernsey the Orchid Fields at Les Vicheries.
DESCRIPTION
Height: 15-60cm.
Stem: Green, usually strongly washed reddish-
purple towards the tip, with two to four loose,
brownish scales at the base.
Leaves: Three to eight, keeled, pointed and
strap-shaped, well-spaced along the stem and
held rather erect, with the upper leaves small
and bract-like. The leaves appear in autumn and
are wintergreen.
Spike: The six to 20 flowers are held in a very
loose spike.
Bract: Green, washed reddish-purple or
sometimes entirely reddish-purple. The bracts
are strap-shaped, narrow, pointed and roughly
the length of the ovary (which they clasp) or
longer.
Ovary: Green, washed reddish-purple or
sometimes entirely reddish-purple. The ovaries
M 27 May, Sussex (Wakehurst Place). The more or less
are long, slender, cylindrical and twisted. two-lobed, unspotted lip is sharply folded, and the spur is
Flower: Dark purple to reddish-purple with a often upturned.
005 systematic.indd 364 29/1/09 12:40:37

LOOSE-FLOWERED ORCHID l 365
P 27 May, Sussex (Wake-

hurst Place). The lateral
sepals are held upwards,
and the upper sepal and
petals form an untidy
hood.
BIOLOGY studies have shown, however, that Loose-

flowered and Green-winged Orchids should join
the Pyramidal Orchid in the genus Anacamptis,
although this change is not universally accepted.
No specific information, but in cultivation HISTORY AND
the period between germination and the CONSERVATION
production of the first green leaves is short. Loose-flowered Orchid was first found on
Guernsey around 1788 by Joshua Gosselin,
Hybrids
a local naturalist, and discovered on Jersey in
A. x alata, the hybrid with Green-winged
1837 by C.C. Babington.
Orchid, has been recorded occasionally on
There have been declines due to habitat loss,
Jersey and once on Guernsey.
but the species is still locally common or even
Name and classification sometimes abundant on Jersey and especially
The specific name laxiflora derives from the Guernsey. Most of the populations are in reserves,
Latin laxus ‘wide’ or ‘loose’ and flos ‘flower’ and although water abstractions for agriculture
means ‘loose-flowered’. and golf courses can affect even protected
Until very recently this species was areas. Loose-flowered Orchid is naturalised at
considered to be a member of the genus Orchis, Wakehurst Place in Sussex (the Royal Botanic
and its overall similarity to some of the other Gardens’ country annexe). In 1987, 350 plants
members of that genus, such as Early Purple grown from seed collected in Crete were planted
Orchid, certainly supported this. Recent genetic out and this population is still thriving.
005 systematic.indd 365 29/1/09 12:40:38

PyramidaL orchid
Anacamptis pyramidalis
This widespread and very colourful orchid of open grassland is commonest on chalk and
limestone but could be found almost anywhere that has a hint of lime in the soil. When they
first appear, the flower spikes form a pyramid or cone, hence the name, but as more flowers
open the shape of the spike changes to more of a globe or cylinder. Pyramidal Orchid is the
county flower of the Isle of Wight.
Identification
The dense, conical spikes of unmarked
bright cerise pink flowers are distinctive.
On close examination the flowers have a
deeply three-lobed lip and a very long, thin,
down-curved spur.
Similar species
The fragrant orchids flower a little earlier in the
season (apart from Marsh Fragrant Orchid) and
have a looser, taller, thinner and more tapering
flower spike. Their flowers are slightly more
purplish-pink and their lip is not so deeply cut,
with the three lobes shorter, broader and more
uneven in size and shape. If there is any doubt,
Pyramidal Orchid can always be identified by the
two prominent raised ridges or ‘guide-plates’ at
the base of the lip, unique to the species.
Habitat
Pyramidal Orchid grows on dry, well-drained
grassland on chalk, limestone or other calcium-
rich soils, such as boulder clay. It is found on
both close-cropped turf and in taller, ranker
swards. Suitable habitats also include the
grykes of limestone pavements, cliff tops,
among Marram and Lyme-grass on sand dunes
and, very locally, the coastal machair of the
Hebrides. The species is sometimes still found
in old meadows, although this habitat has
almost vanished in most areas. It also grows
among scrub; rarely in open woodland with a
broken canopy. Pyramidal Orchid takes readily
to man-made habitats, such as road verges and
roundabouts, churchyards, old quarries, disused
O 28 June, Cambridgeshire. An opportunist, it can do well
on the verges of main roads or even on roundabouts,
005 systematic.indd 366 29/1/09 12:40:39

PYRAMIDAL ORCHID l 367
railway lines and abandoned industrial sites,

1987-99
and is one of the first orchids to move into such 1970-86
newly available habitats. Recorded up to 350m pre 1970
above sea level (Brough, Cumbria).

Flowering period
Early June to mid-August, very exceptionally
from May, but mostly mid-June and early to
mid-July.
Range
Mostly found south and east of a line from
south Devon to North Yorkshire and in
southern and especially central Ireland. The
species clearly enjoys a mild maritime climate
and ranges further north and west into coastal
districts of north Cornwall, south and north
Wales, northern Ireland and northeast England
(north to Lindisfarne). It is also found on Arran, Islay, Colonsay, Tiree, Gunna and Coll
the Isle of Man, Isles of Scilly and Channel and on the Outer Hebrides on South Uist,
Islands. In Scotland it is rare and confined Funday and Barra. World range: Essentially
to a few sites in the Borders, Lothian, Fife, Europe but extends into North Africa and
Angus, Sutherland, Kintyre and Dumfries and the Middle East. Occurs north to Denmark,
Galloway, as well as the Inner Hebrides on southern Sweden and the Baltic States, east
M 5 July, Northamptonshire.Troops of Pyramidal Orchids add a splash of colour to high summer.
005 systematic.indd 367 29/1/09 12:40:41

O 22 June, Norfolk. The

spike is conical to pyramid-
shaped when the flowers
begin to open.
to Poland and the Ukraine and south to the shaped with a pointed tip. The three or four
Mediterranean, Crimea and Caucasus. Also sheathing leaves grade into five or six non-
found on the Mediterranean islands, in North sheathing leaves that decrease in size up the
Africa in Morocco, Algeria and Tunisia, and in stem and become bract-like towards the spike.
the Middle East in Israel, Turkey, northern Iraq The lower leaves may arch gracefully outwards.
and northwest Iran. The sheathing leaves emerge in the autumn and
How to find it are wintergreen, but they have often withered
Generally common in suitable habitats, indeed by flowering time. The spike and non-sheathing
often abundant. leaves do not appear until the spring.
Spike: Very dense, the 30-100 flowers are
DESCRIPTION packed so closely that they often conceal the
Height: 10-75cm but usually 20-60cm. stem. When the first flowers break bud the
Stem: Green, slightly angled towards the tip spike is distinctly conical but as more flowers
and often rather slender and flexuous (bendy); open it elongates to become domed or oval in
the stems frequently have a distinct kink shape and sometimes eventually cylindrical.
somewhere along their length. There are two or Bract: Green, sometimes flushed purple,
three brown sheaths at the extreme base. narrow, a little longer than the ovary and
Leaves: Green, tinged grey, keeled and strap- tapering to a fine point.
005 systematic.indd 368 29/1/09 12:40:43

Ovary: Cylindrical, twisted and green, often

washed reddish-purple.
Flower: Usually vivid pink, sometimes paler
pink or reddish-pink, and fading a little as the
flower ages. The sepals are oval-lanceolate with
a variably pointed tip and dished sides, and the
petals are a little shorter and blunter. The lateral
sepals are held horizontally or slightly drooping,
to about 15° below the horizontal, and the upper
sepal and petals form a hood over the column.
The lip is white towards the base, wedge-shaped
and held projecting forwards and downwards.
It is deeply divided into three virtually equal
strap-shaped lobes. The outer lobes broaden
slightly towards their blunt, shovel-like tips,
and the central lobe is more parallel-sided with
the extreme tip pinched in and upwards. On
either side of the base of the lip there are two
prominent narrow raised ridges or ‘guide-plates’
that are extensions of the column. The ridges,
although roughly parallel, converge towards the
mouth of the spur. The spur is slender, 12-14mm
long (often longer than the ovary) and down- M 22 June, Norfolk. On a close inspection, the raised
curved. The column is white, variably washed ‘guide-plates’ at the base of the lip are characteristic.
pink. The rostellum partially blocks the entrance
to the spur, and the two stigmas lie low down on
BIOLOGY
either side of the column and both pollinia are Pollination and reproduction
joined to the same strap-shaped viscidium. The Pyramidal Orchid is pollinated by moths, both
flowers are scented, the perfume most obvious in day- and night-flying, and by butterflies. Day-
the evening and variously described as sweet or flying moths and butterflies are attracted by the
as slightly unpleasant and ‘foxy’. flower’s vivid coloration, and the various burnet
moths may additionally be attracted by its
Subspecies
similarity to the bright red spots on their wings.
None.
At night the flower’s scent would assist its
Variation and varieties detection. The pollination mechanism is highly
Var. albiflora has pure white flowers but is rare evolved. The walls of the spur contain a sugary
(very pale pink flowers are rather commoner). sap but only an insect with a suitably long
Var. sanguinea has blood-red flowers. It is proboscis can reach this. The ridges or guide
scarce and known only from the Hebrides and plates on either side of the mouth of the spur
northwest Ireland. act as a guide correctly to position the insect’s
Var. emarginata has an unlobed lip that proboscis, not only to access the spur and the
resembles a scallop shell. It is very rare. sap within but also to trigger the mechanism.
Var. angustiloba has a very deeply lobed lip. The rostellum hangs over the entrance to the
Var. fundayensis has a taller and more spur and when it is touched by an insect the
cylindrical flower spike. It was described from protective, flap-like bursicle is pushed aside and
Funday in the Hebrides but is probably extinct, the strap-shaped viscidium, complete with the
if it ever existed at all. two pollinia, sticks to the insect’s proboscis.
005 systematic.indd 369 29/1/09 12:40:45

protocorm
bud
tuber root
replacement
tuber
spring summer autumn spring summer autumn

YEAR 2 YEAR 3
M Development from seed in Pyramidal Orchid (after Rasmussen 1995).
As soon as the viscidium is exposed to the air It is not known when germination occurs
it contracts like a watch-spring and coils itself but initially a protocorm is developed. Early
around the proboscis. This not only helps researchers suggested that the seedling
to attach it more firmly but also moves the remained at this stage for three years but it may
pollinia apart so that they become separated well be just a few months before the first tuber
by 90°. Then, a few seconds later in a second develops. Following this the protocorm dies off
contraction, the pollinia swing forward so as in the late summer to leave just the small tuber
to be in a perfect position to strike the sticky in a ‘resting’ state. In the autumn a bud on the
surfaces of the two stigmas on the next flower tuber produces a short rhizome and from this
visited and deposit packets of pollen. The one or two roots develop and sometimes also
mechanism is efficient, with 65-95% of flowers the first leafy shoot. The new rhizome and roots
setting seed. have to be reinfected with fungi from the soil as
Vegetative propagation is also possible via the tuber itself did not carry any ‘infection’. This
the production of additional tubers, which plant, including the leafy shoot, overwinters,
often develop at the end of short rhizomes. and in the following spring a new tuber starts
to develop from a bud on the rhizome and the
old one starts to shrivel away. Eventually the
The aerial stem grows from a pair of rounded
shoot and rhizome also wither, and the plant
or roughly elongated tubers and there are a few
will again spend the late summer ‘resting’ as a
slender fleshy roots growing almost horizontally
tuber. In this way the annual cycle of growth is
near the surface of the soil; it is the roots that
established. Again, it has been estimated that
are ‘infected’ with fungi. The basal leaves appear
several years elapse between the first appearance
in autumn and die down in summer and after
above ground and the first flowers, but this is
flowering the orchid spends the late summer
probably far too long and the period between
and autumn ‘resting’ as a tuber. The Pyramidal
germination and flowering could be as little as
Orchid is clearly adapted to a Mediterranean
three years.
climate of mild, wet winters and hot dry summers
and its overwintering leaves are probably Intergeneric hybrids
vulnerable to hard frosts. The generally southerly x Gymnanacamptis anacamptis, the hybrid with
distribution in Britain, with colonies further fragrant orchid (presumably Common Fragrant
north concentrated near the coast, emphasises its Orchid), has been recorded very rarely from
preference for a mild winter climate. Hampshire, Gloucestershire and Co. Durham.
005 systematic.indd 370 29/1/09 12:40:45


The specific name pyramidalis means simply
‘pyramidal’ or ‘conical’. Until recently the
Pyramidal Orchid was the only member of the
genus Anacamptis, but it has now been joined
by Green-winged and Loose-flowered Orchids.
HISTORY AND
CONSERVATION
The first British record dates from 1660 when
John Ray recorded it in his Catalogus Plantarum
circa Cantabrigiam nascentium (‘A catalogue of
plants found around Cambridge’): ‘In many
places, as in a chalkie close at Hinton near
where they burn lime.’
Past and present occurrence of Pyramidal Orchid in Brit-
Britain Ireland

1500-1999
current range 681 (24%*) 241 (24%*)
% lost, 1500-1969 13.5% 25%

% lost, 1970-1986 6.5% 6%
The boundaries of the range are stable. There

have been local declines, and Pyramidal Orchid
has been lost from 20% of the historical range
in Britain and 31% in Ireland. Some of the
declines are due to the loss of old meadows and
pastures and although it is tolerant of grazing,
overgrazing may also be a threat, perhaps
especially at the few sites on the machair of
the Hebrides and western Scotland. On the
other hand, there have been some marked
increases in recent years, mostly in recently
created man-made habitats such as road verges,
and the species is a pioneer colonist of such
places. Artificial habitats have their drawbacks,
however, as ‘brownfield’ sites are developed and
old pits and quarries used for landfill.
P 21 June, Suffolk. Once all the flowers have opened the

spike is cylindrical, sometimes markedly so.
005 systematic.indd 371 29/1/09 12:40:47

Green-winGed orchid Near Threatened

Anacamptis morio
Formerly: Orchis morio; Other names: Green-veined Orchid

This beautiful orchid once graced pastures and other areas of undisturbed grassland
throughout much of England, Wales and Ireland, and the sight of a field splashed with
the purples and yellows of thousands of Green-winged Orchids and Cowslips was
commonplace. Tragically, agricultural changes have led to a dramatic decline in most areas,
and the species has become rather local, although it can still be found in very large numbers
at some favoured sites. It now largely depends on the sympathetic management of the few
remaining old pastures as well those churchyards blessed by its presence. Green-winged
Orchid is the county flower of Ayrshire.
Identification
A dainty and usually petite orchid with
unspotted leaves. It has a few relatively large
and very attractive flowers that are extremely
variable in colour, from deep violet-purple to
rose-pink or whitish, although most are a shade
of purple. Whatever the colour, the flowers are
distinctive. The sepals and petals form a hood
marked with the green or bronze veins that give
the species its name. The sides of the lip turn
downwards to form a broad and often colourful
skirt, and the long straight spur projects
conspicuously back from the flower.
Similar species
Early Purple Orchid is superficially similar but
usually has spotted leaves and also tends to have
a longer flower spike and more flowers that are
a paler purple. On close examination only the
upper sepal and petals form the ‘hood’ because
the two lateral sepals are held upright as ‘wings’.
Early Purple Orchid also lacks green or purple
veins on the sepals and finally, although both
species are found in grassland, only Early
Purple Orchid is found in woodland.
Habitat
Green-winged Orchid favours unimproved
grassland on neutral or calcareous soils where
grazing, mowing or other factors keep the
grass relatively short and the sward open. It is
M 20 May, Norfolk.The flowers are usually a deep violet-
purple, a colour that is extremely hard to reproduce in very strongly associated with old species-rich
print. grassland and is slow to colonise new sites.
005 systematic.indd 372 29/1/09 12:40:48

GREEN-WINGED ORCHID l 373
The optimum habitats are damp pastures on

1987-99
clay soils, but it is also found on dry chalk or 1970-86
limestone grassland and sometimes on sands pre 1970
and gravels, including pockets of neutral

grassland on otherwise acidic heathland. The
species can tolerate both mowing for hay and
grazing. Because unimproved damp meadows
and pastures are now rare Green-winged
Orchid is almost as likely to be found on
more marginal sites such as stabilised dunes,
lime-rich eskers, old railway cuttings and
banks, village greens (especially those which
are ‘commons’), churchyards and golf courses.
Occasionally it is found on lawns or on old
industrial sites, for example, alkaline waste
produced in the manufacture of washing soda.
It is intolerant of shade, and although it can be
found among scrub it is seldom if ever found in line from Morecambe Bay to Holy Island in
woodland. Recorded up to 305m above sea level Northumberland, although absent from much
(Co. Roscommon). of Devon and Cornwall and the Pennines.
Now it is greatly reduced and absent from
Flowering period much of northern England, the Midlands, East
Mid-April to mid-June but mostly in May. It is Anglia and the Home Counties. In Wales it
earliest in the west and exceptionally has been is largely confined to coastal regions, and in
recorded flowering in mid-March. Ireland to the central limestone belt, from Co.
Range Dublin west to Co. Galway and from southern
Widespread in England, formerly north to a Co. Tipperary north to Co. Longford, with
M 10 May, Norfolk. Green-winged Orchid can occur in large numbers, especially on clay soils, on old commons and
greens, and on unimproved pastures and meadows
005 systematic.indd 373 29/1/09 12:40:50

to flowering as the leaves are wintergreen.

Nowadays most colonies are on protected sites,
such as reserves and SSSIs or on private land
with a sympathetic landowner.
DESCRIPTION
Height: 5-50cm but usually 7.5-15cm and
rather compact, and seldom more than 30cm.
Stem: Yellowish-green, washed purple
towards the tip and slightly angled, with
two or three thin whitish sheaths at the
extreme base.
Leaves: There are up to seven bluish-green
basal leaves, varying from roughly elliptical
to lanceolate, keeled and held semi-erect, and
two or three leaves higher on the stem that are
more pointed and become bract-like below the
spike. The leaves are wintergreen, emerging in
September-October and dying down by
mid-June.
Spike: Rather open and loose with four to 14
M 12 May, Norfolk. The bold, parallel, green veins on the
sepals are diagnostic. well-spaced flowers.
Bract: Green, typically washed purple, and
a few sites in Co. Cork and just one site in lanceolate in shape, two-thirds to twice the
Northern Ireland, on the coast of Co. Down.
It is very rare in Scotland, where it is found on
the coast of Ayrshire with an old record from
near Tomintoul in the Grampians. World
range: Predominantly European, extending
marginally into North Africa and the Middle
East. It ranges north to southern Norway,
southern Sweden (Gotland) and Estonia, east
to westernmost Russia and eastern Ukraine
and south to the Mediterranean, including
many of the Mediterranean islands, the Crimea
and Caucasus. Also Lebanon, Israel, Turkey,
northwest Iran and Morocco.
How to find it
In favoured localities Green-winged Orchid
can still be found in thousands or even tens
of thousands; a site in Sussex, for example,
holds up to 50,000 flower spikes. Odd plants
can, however, be inconspicuous and are easily
missed, especially if they are very short. The
number of plants producing flowers varies
greatly from year to year, with a wet winter M 20 May, Norfolk. All the sepals and petals are arrang-
and spring probably being most conducive ed together to form a tight hood over the base of the lip.
005 systematic.indd 374 29/1/09 12:40:56

length of the ovary, which they curl around and

sheathe.
Ovary: Green, variably tinged purple,
cylindrical, strongly curved (through around
90°), ribbed and twisted.
Flower: Mostly various shades of purple but
a few are rose-pink or whitish. The sepals are
oblong-oval, the upper a little narrower than
the lateral sepals. They are generally whitish
with a variable violet-purple wash, deepest
on the upper sepal and the upper margins of
the lateral sepals, which have three to seven
bold, green or bronze parallel veins that are
visible on both their upper and underside. The
petals are coloured as the upper sepal but are
shorter, narrower and more strap-shaped. The
petals and sepals form a ‘hood’ that encloses
the column. The lip is rather broader than long
and divided into three lobes. The side-lobes are
rather larger than the central lobe, rounded,
often with crinkled (crenate) edges and folded
downwards. The central lobe is roughly equal
in length to the side-lobes. The lip is typically
violet-purple, whiter in the centre and at the
mouth of the spur, and there are usually some
violet-purple blotches and spots in this paler
central area. The column is whitish, washed
purple around the pollinia. The spur is coloured
as the lip, around half the length of the ovary,
narrow but flattened towards the tip, where it
is often notched and slightly curved upwards. M 24 May, Norfolk. Var. alba. A variable but usually very
At least some flowers have a vanilla-like scent, small proportion of plants are white; the stripes on the
sepals are retained.
and the white-flowered forms may, on average,
have a stronger perfume although some people
cannot smell it at all.
BIOLOGY
Subspecies
Bees, especially bumblebees, pollinate this
None.
species. It produces no nectar and so is thought
Variation and varieties to rely upon deceit to attract insects, although as
Var. alba has white flowers, sometimes very with Early Purple and Pyramidal Orchids it has
faintly tinged pink, but still with green veins. also been suggested that potential pollinators
It is usually scarce (0.1-1% of plants) but can access sugary sap within the walls of the
sometimes makes up as much as 15% of a spur. When the queen bees first emerge from
population. hibernation in the spring they are ‘naive’. They
Var. bartlettii is small with very small flowers have yet to learn which flowers offer a genuine
and var. churchillii is tall; neither seems well reward of nectar and are easily attracted by the
defined. brightly coloured orchids. The bees usually visit
005 systematic.indd 375 29/1/09 12:40:57


It is sometimes stated that Green-winged
Orchid is monocarpic and therefore usually
dies after it has flowered once. However, in one
population in Cambridgeshire some individual
plants flowered for 17 out of 18 years and
almost all flowered several times, interspersed
with seasons in which they merely produced a
rosette of leaves or more rarely were ‘dormant’
underground. At Iron Latch Meadow in
Essex, Green-winged Orchids reappeared and
flowered after an absence of about 30 years
when closed-canopy hawthorn scrub was
removed, showing just how long-lived this
species can be.
Seeds are thought to germinate in the
late summer or autumn, and the protocorm
probably produces the first roots and even the
first small leafy shoot during its first winter or
spring. The plant goes on to develop its first
tuber at the base of the leafy shoot, and it is this
tuber alone that will persist through the late
summer ‘resting period’, establishing the annual
cycle. It is thought that plants can flower within
three years of germination, and in cultivation
they have flowered after just two.
Hybrids
A. x alata, the hybrid with Loose-flowered
Orchid, has been recorded occasionally on
Jersey and once on Guernsey.
M 16 May, Norfolk. Some are very petite, with a few,
relatively large flowers. Intergeneric hybrids
a single flower on each of several plants before ‘Orchis x morioides’, the hybrid with Early
they realise there is no reward on offer and move Purple Orchid, has been recorded rarely and
on to more profitable species. On each orchid sporadically in England and Wales. As these
visited, they start with the lower flowers of the two species are now in different genera a new
name will be needed for this hybrid.
spike and therefore these are the ones that tend
to set seed. A cold spell in May, which disrupts Name and classification
the bees’ routine, benefits the orchid because The specific name morio means ‘dull’ or ‘stupid’.
the bees have to relearn their foraging routes The species was once known as ‘fool stones’.
once the weather warms up again. Pollination Until very recently this species was
is variably efficient; in a study in Sweden only considered to be a member of the genus Orchis.
5-30% of flowers set seed but in Britain almost Its overall similarity to some of the members
all the flowers may be pollinated, with each of that genus, such as Early Purple Orchid,
capsule producing 4,000 individual seeds. The certainly seemed to support this. Appearances
species may also reproduce vegetatively via the can be deceptive, however, and recent genetic
formation of additional tubers. studies have shown that the Green-winged
005 systematic.indd 376 29/1/09 12:40:58

Orchid, together with Loose-flowered Orchid decline significantly if fertilisers are applied to
of the Channel Islands, should join the grassland, probably due to competition with
Pyramidal Orchid in the genus Anacamptis. the increasingly vigorous grass and herbs, and
the phosphorus in fertilisers may actually be
HISTORY AND toxic to the orchids. If anything, habitat losses
CONSERVATION have accelerated in the last 20 years as the last
The first British record dates from 1634 when few pristine pastures have been ‘improved’;
Thomas Johnson published his Mercurius overall the decline in the amount of suitable
Botanicus (‘Botanical Mercury’) with the first habitat must approach 99%. A typical example
localised record being published in 1660 when is Cambridgeshire, where prior to 1900 at
John Ray noted the ‘female Fool-stones’ from least 36 sites held Green-winged Orchids.
Cambridgeshire in his Catalogus Plantarum In the period 1901-85 this figure fell to 13
circa Cantabrigiam nascentium (‘A catalogue of and between 1986 and 1990 the population
plants found around Cambridge’). collapsed to just five sites.
Once common and widespread, Green-
winged Orchid has declined dramatically in the
last 50 years. It has been lost from many sites
and has gone from 49% of the historical range
in Britain and 60% in Ireland. It is classified
as Near Threatened in Britain and is specially
protected in Northern Ireland under Schedule
8 of the 1985 Wildlife Order (NI) and in Eire
under the Flora (Protection) Order.
Past and present occurrence of Green-winged Orchid in
Atlas).
Britain Ireland

1500-1999
current range 479 (17%*) 50 (5%*)
% lost, 1500-1969 38% 55%

% lost, 1970-1986 11% 5%
The decline is almost entirely due to

agricultural changes. In the 19th century its
favoured damp meadows and pastures were
ploughed and converted to arable. In the 20th
century, especially since 1945, tractors have
replaced horses, and permanent pastures
have become less and less a feature of most
farms. Those that have survived have been
drained, ploughed and reseeded. Research M 10 May, Norfolk. This delicate rose-pink is rather
has shown that Green-winged Orchid will unusual.
005 systematic.indd 377 29/1/09 12:40:59

Genus SERAPIAS
TonGue orchids
005 systematic.indd 378 29/1/09 12:41:03

GENUS SERAPIAS l 379
This genus has only a toehold in Britain and Ireland, with two species recorded in recent
years from the coast of southwest England. We consider that a natural origin, via wind-
blown seed, is the most likely explanation for their occurrence, although they have also been
treated, albeit without any evidence, as deliberate introductions.
Distribution colour of the flower may help it to absorb heat,

Serapias is basically a Mediterranean genus, and its internal temperature may be 1-3°C
distributed from the Caucasus Mountains higher than that of its surroundings. It seems
through the Mediterranean basin west to the likely, however, that there is more involved in
Canary Islands and Azores and extending north the pollination process than merely the offer
to Brittany. All the species are closely related of shelter.
and thus there is considerable disagreement The tongue orchids are closely related and
as to the exact number. Between seven and 27 genetically very similar. The various species
species are recognised in recent literature and need to be isolated reproductively from one
consequently there are considerable problems in another to have evolved in the first place and to
reaching a correct identification. continue to maintain their separate identities.
Classification A mechanism that involved their pollinators
Genetic evidence shows that the closest would be one way for this to happen but if
relations of the tongue orchids are the bee and they merely offer ‘shelter’ there would be no
spider orchids Ophrys, the Pyramidal Orchid mechanism, apart from the size of the flower,
group Anacamptis and the lizard orchids to stop insects from carrying pollen from the
Himantoglossum. flowers of one species to another. Indeed, this
The genus Serapias is often divided into might be why there are so many hybrids that
three groups: the self-pollinating Small- involve the various tongue orchids. But, so far
flowered Tongue Orchid S. parviflora and allies; Greater Tongue Orchid S. lingua is the only
the vomeracea group (the largest) and the lingua species where additional isolating mechanisms
group, which includes Greater Tongue Orchid, have been observed. Small bees, apparently
and is distinguished by having a single dark attracted to the flower by pheromones, attempt
boss at the base of the lip (the other groups pseudocopulation with the flower (see also the
having two bosses). notes on the genus Ophrys, p.388). In these
cross-pollinated species pollination is apparently
Floral structures inefficient and although their flowers are self-
The column is slender, directed forwards and compatible, self-pollination does not occur.
tapers to a long, thin point. There are two The parviflora group, which includes Small-
pollinia, each narrowing down into a caudicle flowered Tongue Orchid, is self-pollinated.
(stalk). Both pollinia are attached to the single
viscidium, which in turn is housed in a bursicle. Growth pattern
There is no spur. At flowering time there are two to five more-
or-less spherical tubers at the base of the aerial
Pollination
stem, the one to three youngest at the end of
The vomeracea and lingua groups are cross-
long, slender rhizomes.
pollinated. The flowers have no nectar and
are thought to lure insects, especially bees and Development from seed
beetles, by providing them with shelter, both Following germination, the protocorm produces
overnight and during bad weather. The dark its first root in the autumn and the first leaf
appears shortly afterwards. The following
O 27 April, Extremadura, Spain. Greater Tongue Orchid. spring the first tuber starts to grow from a bud
005 systematic.indd 379 29/1/09 12:41:04

380 l GENUS SERAPIAS
at the base of the leafy shoot, and during the Name

summer ‘resting period’ the leaf and root die The name of the genus, Serapias, is derived
off, leaving just the tuber. From now onwards from Serapis (or Osirapis). This is a composite
the usual sequence of annual tuber replacement Graeco-Egyptian god combining the attributes
takes over. In cultivation plants may flower of Apis (the scared bull) and Osiris (ruler of
three years after seed is sown, occasionally after the underworld). Serapis attracted an orgiastic
just two. cult and in about AD 64 the Greek scholar
Dioscorides, considered one of the ‘fathers’ of
botany, gave the name Serapias to an orchid that
Additional ‘daughter’ tubers are formed at the
was reputed to be an aphrodisiac. The orchid in
base of the aerial stem, usually at the end of
question was probably actually a member of the
slender rhizomes. These will go on to form
genus Orchis.
separate plants as the connecting rhizome dies
off in the summer.
upper sepal
petal
upper sepal
petal
lateral
lateral boss sepal
bosses sepal
hypochile
side-lobe
side-lobe hypochile
lip } epichile
lip
} epichile
M Small-flowered Tongue Orchid (After Delforge 2005). M Greater Tongue Orchid.
005 systematic.indd 380 29/1/09 12:41:04

SMALL-FLOWERED TONGUE ORCHID l 381
smaLL-fLowered TonGue orchid

Serapias parviflora
This species is one of the most recent additions to the British flora, having been found for
the first time in 1989. It is confined to just one site in Cornwall, where a handful of plants
flower most years, but it could turn up almost anywhere on the south coast.
Identification Islands, and including Crete, Sicily, Corsica,

The Serapias or tongue orchids are very Sardinia and the Balearic Islands, and also
distinctive, with the sepals, petals and base of Morocco, Algeria and Tunisia in North Africa.
the lip forming a tight tube from which the In France it occurs on the Mediterranean
outer part of the lip, usually coloured a rusty- littoral with a separate centre of distribution on
brown, sticks out like a pointed little tongue. In the Atlantic coast north to Brittany.
Small-flowered Tongue Orchid the flowers are
How to find it
relatively small (indeed, they are the smallest
The British population comprises just a few
of the Serapias), with a small lip, the projecting
plants that are very vulnerable to disturbance
tip of which is 6-11mm long x 3.5-5mm wide.
and damage; indeed, flowers have already been
There are two small dark parallel raised bosses
unwittingly broken off by admirers. This is one
at the base of the lip, and the flowers are self-
orchid that is best left alone, at least until a
pollinated in bud, so that when they open the
more substantial population exists.
pollinia have already disintegrated onto the
stigma.
Similar species
Greater Tongue Orchid has larger flowers with
a single dark boss on the lip.
Habitat
In Cornwall it is found in short, rabbit-grazed
grassland among scattered gorse on a south-
facing slope near the sea. In Europe it is found
in dry and wet grassland, dune slacks, open pine
woods, scrub and olive groves, on calcareous to
mildly acidic soils.
Flowering period
Early May to early June.
Range
Known from a single site near Rame Head
in southeast Cornwall. World range: The
Mediterranean basin from Cyprus and the
Aegean Islands west through Greece, Italy,
France, Spain and Portugal to the Canary
P 7 May, Cornwall (Mike Frost). Although currently re-

stricted to one site in Britain, this species can be expected
to appear elsewhere.
005 systematic.indd 381 29/1/09 12:41:05

DESCRIPTION BIOLOGY
(based on European populations) Pollination and reproduction
Height: 10-30cm, occasionally to 40cm. The flowers are cleistogamous, that is self-
Stem: Green, blotched or streaked with red. pollinated in bud. The yellow pollinia have
Leaves: At the base of the stem there is a already disintegrated onto the stigma by the
cluster of four to seven erect, long, narrow, time the flowers open.
strap-shaped leaves with a distinct keel. Higher
on the stem there are one to three smaller Development and growth
bract-like leaves. In Europe the leaves appear No specific information. Stems usually arise
in autumn, overwinter and die down after singly, and vegetative reproduction may be
flowering. relatively unimportant.
Spike: Three to eight flowers (rarely as many Hybrids
as 12) form a narrow spike with an indistinct None.
spiral arrangement, with each flower held
horizontally or angled upwards to 45°.
The specific name parviflora means simply ‘small
Bract: Pinkish-brown washed grey and with
flower’.
distinct dark veins; the bracts are lanceolate
and held erect, clasping the ovary. They are as
long as the ovary and hood put together or just
HISTORY AND
a little shorter, although the lower bracts are CONSERVATION
sometimes much longer than the flower. First recorded in May 1989 near Rame Head in
Ovary: Green, long, narrow, and held Cornwall, when two spikes were found. Three
upright. spikes appeared in 1990, with five each in 1991
Flower: The sepals are lanceolate and a pale and 1992, although photographers squashed
dusty-pink washed grey with distinct dark two spikes in the latter year. There was then
veins. The petals are a little shorter, tapering a gap of at least four years during which no
abruptly to a long narrow point, and purplish- plants were found, but subsequently two or
brown at the base. The sepals and petals form three spikes have appeared most years up to
a tight hood over the base of the lip and the and including 2004. Seedlings germinated at
column. The lip is 14-19mm long (extremes the Royal Botanic Gardens, Kew, from Cornish
13-22mm) and reddish-brown. The inner half seed were planted at two nearby sites in 1993
(the hypochile) is heart-shaped with two small, but did not survive.
raised, parallel, dark purple bosses at the base. It has been suggested that the plants in
The sides, which are almost black, curl up and Cornwall were ‘deliberately introduced’, but
round to form a ‘gutter’ that is totally enclosed there is no evidence for this whatsoever.
by the hood. The floor of the lip has short, The plants were found hidden among gorse
sparse, whitish hairs or is sometimes almost scrub, well away from any footpaths and in
hairless. The outer half of the lip (the epichile) a locality highly unlikely to be chosen for an
is lanceolate, much narrower than the base ‘introduction’. It seems much more likely that
and bent down and under the remainder of the plants arose naturally from wind-blown
the flower. seed. Whether this tiny population persists
or the species appears elsewhere, only time
Subspecies will tell.
None.
Variation and varieties P 1 June, Cornwall (Robin Chittenden www.harle-

None. quinpictures.co.uk)
005 systematic.indd 382 29/1/09 12:41:07

005 systematic.indd 383 29/1/09 12:41:10
GreaTer TonGue orchid

Serapias lingua
This species has only the most tenuous claim to be British, having been found on Guernsey
in 1992 and in south Devon in 1998. These may be the vanguard of further discoveries,
however, if the climate becomes more ‘Mediterranean’ due to global warming.
Identification Malta, Corsica, Sardinia and the Balearics. In

The tongue orchids are distinctive, with the North Africa recorded from Morocco, Algeria
sepals, petals and base of the lip forming a tight and Tunisia. In France it occurs north to a line
tube from which the tip of the reddish lip sticks between Nantes and Lyon.
out like a tongue. The identification of the How to find it
various species is often rather difficult as they Like the Small-flowered Tongue Orchid this
are very similar, and there are also many hybrid species is confined to just one site in Britain,
populations. It often requires the dissection with just a few plants that are vulnerable to
of a flower, but the conspicuous single dark disturbance and damage. It is best left alone.
purple boss at the base of the lip, resembling a
coffee bean, easily identifies the Greater Tongue DESCRIPTION
Orchid and its allies. All the other Serapias (from European plants)
have two bosses. Within the lingua group, Height: 10-30cm, occasionally to 50cm.
identification depends on observation of the Stem: Green, blotched or streaked with red.
precise shape of the boss, the width of the lip Leaves: At the base of the stem there is a cluster
and the overall colour of the flower. of four to eight erect, long, narrow, strap-shaped
Similar species leaves that have a distinct keel. Higher on the
stem there are one to three smaller bract-like
Small-flowered Tongue Orchid has a smaller
leaves. In Europe the leaves appear in autumn,
flower with two bosses at the base of the lip and
overwinter and die down after flowering.
is self-pollinated.
Spike: The two to six flowers, rarely as many
Habitat as eight, are arranged around the stem in a
On Guernsey it was found on dry natural compact spike.
grassland on the edge of a golf course. In south Bract: Lanceolate, held erect and clasping the
Devon it was found in a meadow that had been ovary, they are shorter than the ovary and hood,
sown with a grass mixture in the 1970s and was put together. The bracts are reddish or, more
normally cut for hay in May or June. In 1998, rarely, green tinged red towards the tip, with
when the Greater Tongue Orchid was found, distinct purple veins.
it was not mown. In Europe it is found in poor Ovary: Green, long, narrow and held upright.
grassland, wet meadows, dune slacks, scrub, Flower: The sepals are lanceolate and pale lilac-
olive groves and open woodland. grey with distinct purple veins. The petals are a
Flowering period little shorter, tapering abruptly to a long narrow
May to June. point, and are more purplish-brown at the base.
The sepals and petals form a tight horizontal
Range hood over the base of the lip and the column.
Found at single sites on Guernsey and in Devon. The lip is 22-29mm long (sometimes only
World range: The Mediterranean basin, from
P 27 April, Extremadura, Spain. The origin of the English
Greece and the former Yugoslavia west to name ‘tongue orchid’ for the members of the genus
Portugal, also the Aegean Islands, Crete, Sicily, Serapias is obvious.
005 systematic.indd 384 29/1/09 12:41:10

GREATER TONGUE ORCHID l 385
005 systematic.indd 385 29/1/09 12:41:13

16mm), variable in colour from yellowish to females, with its powerful scent. In addition,
fleshy-pink or reddish and only sparsely hairy. the large shiny boss at the base of the lip also
The basal portion (hypochile) is kidney-shaped stimulates ‘pseudocopulation’, during which the
with the side-lobes dark purple, rather angular pollinia are attached to the head of the bee as it
and curling up and around to form a ‘gutter’ that attempts to copulate with the flower. This two-
is totally enclosed by the hood. There is a single pronged strategy enables the orchid to attract
shining, dark purplish-brown raised boss at the bees both at night and on sunny days.
base of the hypochile, often crescent-shaped Vegetative reproduction may be an
with the two horns of the crescent pointing important means of reproduction and clusters
backwards and elongated into two fine ridges. It of flower spikes are frequent.
is ungrooved or, at most, only slightly grooved.
The outer half of the lip (epichile) is broadly
lanceolate and slightly longer and narrower
than the hypochile. It is usually 13-18mm long Hybrids
x 7-12mm wide but can be as small as 8mm None in Britain.
x 4mm. It is held pointing downwards and is Name and classification
variably bent backwards below the rest of the The specific name lingua means ‘tongue’.
flower. The pollinia are greenish-yellow and the
flowers are strongly scented. HISTORY AND
Subspecies CONSERVATION
Disputed. More conservative taxonomic A single plant was found on the western
treatments consider that Greater Tongue side of Guernsey in May 1992 but it has not
Orchid has several subspecies, including S. l. reappeared at this site. In June 1998 three
durieui. More radical treatments, including flower spikes were found near the coast in the
Delforge (2001), treat durieui as a synonym vicinity of Kingsbridge in south Devon. They
of a different species, S. strictiflora, that is were identified as the subspecies durieui and it
found from eastern Algeria to Morocco and in is speculated that the seed could have arrived
southern Portugal and southwest Spain. This is with ‘Saharan sand’ from North Africa.
of some relevance because the plants in south As with all the Serapias orchids that have
Devon were identified as subspecies durieui on been found in Britain, some botanists are
the basis of a narrow lip and longer and more sceptical as to its origins. They presumably
elegant sepals and petals. If this identification believe that the plants originated from seed that
was correct and if the more radical classification came from an artificial source, such as a nearby
is accepted, Serapias strictiflora would be on the plant collection or agricultural seed mix, or that
British list rather than S. lingua. deliberate fraud could be involved, with seed
being sown or even mature specimens planted
Variation and varieties out. An artificial source of seed is certainly
None in Britain. possible (see Heart-flowered Tongue Orchid,
p.416) but in both Greater and Small-flowered
BIOLOGY Tongue Orchids this seems highly unlikely,
Pollination and reproduction while there is no evidence of fraud. Given the
It is thought that most Serapias are pollinated occurrence of Greater Tongue Orchid in much
by small bees and other insects which use of southern France, including the Atlantic
the flowers for shelter. However, the Greater coast south of Vendée, a natural arrival in the
Tongue Orchid may take this strategy one stage Channel Islands or southern England seems
further. The flower mimics a nest hole and eminently likely. Whether the species becomes
lures small male bees, which emerge before the established is, however, a different matter.
005 systematic.indd 386 29/1/09 12:41:14

Genus OPHRYS
Bee and sPider orchids
005 systematic.indd 387 29/1/09 12:41:18
388 l GENUS OPHRYS
This genus includes the most exotic and intriguing of British and Irish orchids, Bee and Fly
Orchids and Early and Late Spider Orchids. Their flowers have evolved to mimic insects,
predominantly bees (but definitely not spiders). In addition to their overall appearance, the
pheromones produced by the flowers and the distribution of hairs on the lip are critically
important in perpetuating the deception. Male bees are attracted to the orchids and attempt
to mate with the flowers in a process known as ‘pseudocopulation’. In doing so they carry
pollinia from one flower to the other and become unwitting pollinators.
1
Containing more than 250 species, this is All species in this genus are cross-pollinated
undoubtedly the largest genus of European with the exception of one, the Bee Orchid,
orchids. Its distribution is centred on the which is routinely self-pollinated. The Bee
Mediterranean basin, but Ophrys can be found Orchid had clearly evolved to use the same
from the Canary Islands in the west to the mechanism as the other species, with an
Caspian Sea in the east, Fly Orchid ranges elaborate pattern and texture to its lip, but
north to Scandinavia and other species extend has largely abandoned cross-pollination,
into North Africa. However, just four species presumably relatively recently.
occur in Britain and Ireland. In the genus Ophrys the flowers have no
The number of species recognised in this nectar and offer their visitors no reward.
genus has risen sharply; only around 60 were Instead, the orchid exploits the sex-drive of
listed in the 1980s. This reflects the increasing insects with a combination of visual, olfactory
4
appreciation of the importance of the highly and tactile deceits. The pollinators are usually
specialised pollination mechanism in the bees and they are initially attracted to the
evolution of the various species. flowers by chemical signals released by the
lip of the orchid that mimic the pheromones
Classification produced by virgin female bees. These
DNA evidence suggests that the closest pheromones are complex cocktails of volatile
relatives of the genus are the tongue orchids compounds that stimulate mating behaviour in
Serapias, the lizard orchids Himantoglossum and the male bees. Not only are the flower’s version
the Pyramidal Orchid group Anacamptis. of the bee’s pheromones an exact copy of the
appropriate compounds but also the orchid
Floral structures
produces such large quantities that the male
The stiff and often elaborately designed lip
bee may even prefer the flower’s deceit to the
renders the members of this genus distinctive.
real thing.
It is usually prominently hairy in places and 7
Male bees emerge from hibernation earlier
velvety in others with a smooth, hairless central
in the spring than females, and it is thought
patch, the speculum. This is sometimes shiny,
that the orchid exploits this gap. In the absence
reflecting the light and probably therefore
of females the orchid is able to lure the males to
simulating an insect’s wings. As in most orchids
the lowermost flowers open first but the spike P 1. O. levantina; 2. O. umbilicata; 3. O. flavomarginata;
continues to elongate, sometimes substantially 4. O. (f.) iricolor; 5. O. lutea galilaea; 6. O. omegaifera
so, as more flowers open. israelitica; 7. O. aesculapiiformis; 8. O. (f.) fusca; 9. O.
mammosa; 10. O. argolia elegans; 11. O. bormuelleri;
The column is long and erect. There are two 12. O. kotschyi. All Cyprus, March. The intimate relation-
pollinia, each attached by a caudicle (stalk) to a ship with a specific insect pollinator has powered the evo-
lution of numerous species.
separate sticky viscidium. These are concealed
and protected within separate bursicles. O 14 June, Kent. Late Spider Orchid
10
005 systematic.indd 388 29/1/09 12:41:19

1 2 3
4 5 6
7 8 9
10 11 12
005 systematic.indd 389 29/1/09 12:41:23

390 l GENUS OPHRYS
its flowers but once the females have emerged Ophrys has evolved many species, reflecting the
the orchids are ignored in preference to the real many species of potential pollinator. Each one is
thing. In at least some species, however, this is precisely matched.
not the case and the flowers continue to receive As a further complication, differences
visits from the males even after the female bees in the patterning and shape of the lip and
have emerged. the distribution of hairs allow two distinct
The male bees home in on the orchid’s species of Ophrys to share the same species
pheromones, and as they get closer they catch of pollinating insect. The position and
sight of the flower and land on the lip. All three direction of the hairs on the flowers are
deceits now come into play. The male bee is critical in the positioning of the insect during
still stimulated by the scent of the flower and ‘pseudocopulation’. One species of orchid may
the patterning on the lip but now also by the stimulate the bee to have its head uppermost,
texture of the various hairy, velvety and smooth facing the column and so picking up the
portions of the lip. This stimulates the bee to pollinia on its head. A second species of
orientate himself into the ‘correct’ position, orchid stimulates the bee to have its abdomen
extend his genital apparatus and attempt uppermost, and the pollinia are attached to the
copulation. Of course, the ‘correct’ position is tip of its abdomen. In the respective flowers, the
the one in which he will pick up pollinia in pollinia and stigma are positioned according to
the first flower visited and deposit the pollinia the orientation of the bee.
on the stigma of the second flower and so on. The orchid-pollinator match in Ophrys
Eventually this process of ‘pseudocopulation’ is is very close but not always perfect. All the
interrupted and the bee leaves. The male bees members of the genus are interfertile, and
may show signs of exhaustion but ejaculation there are a large variety of hybrids. It is a
of sperm has not been observed, although it reflection of how good the system is, however,
has been recorded in a similar process involving that these hybrids tend to occur rarely and in
an ichneumon wasp and the Australian orchid, very small numbers.
Cryptostylis leptochila.
Growth pattern
The orchid is totally dependent on the bee.
At flowering time all Ophrys species have two
If there is no pollination, there will be no seed
more-or-less spherical tubers side by side at
and therefore no more orchids; a powerful
the base of the aerial stem together with some
engine to drive the evolution of the orchid.
short, thick roots. The roots appear in the
Against a ‘noisy’ background of multitudes
autumn, simultaneously with the leaves, and
of chemical signals, each species of bee must
are ‘infected’ with fungi. The pattern of growth
be able to produce a unique pheromone in
and the annual replacement of tubers are very
order to attract potential mates and reproduce
similar to that of the genus Orchis (see p.200).
successfully. It may do this by having a unique
combination of relatively common chemicals Fungal partners
in its sex pheromones or by using chemicals In common with the other tuberous orchids,
that are in themselves unique. Bees use both fungal activity takes place in the roots and
systems and the various Ophrys orchids are able rhizome but not in the tuber (indeed, the tuber
to mimic both systems, too. For the orchid, contains the orchid’s food reserve and may
the better and more precisely it mimics its even be protected from fungal infection by
pollinator the better its chance of successful ‘fungicide’, see p.9). As the orchid passes the
pollination. Each species of orchid therefore summer ‘resting period’ as a tuber this presents
evolves to mimic the pheromones of a particular a problem. How can the fungal ‘infection’ be
species of bee in order to maximize its chances carried from one year to the next? In damper
of attracting pollinators. In this way the genus climates there is a good chance that the fungi
005 systematic.indd 390 29/1/09 12:41:23

GENUS OPHRYS l 391
can persist in the soil but in the hot, dry, Greek for ‘eyebrow’, perhaps an allusion to the
Mediterranean summers adult fungi cannot hairy lip (or the hairy fringe to the petals).
live in the parched soils and must survive as Alternatively, Ophrys was the name given in
spores. To guarantee continuity the members Ancient Greece by Pliny the Elder to a small,
of the genus Ophrys probably carried their two-leaved plant used to dye eyebrows and hair.
fungal ‘infection’ from one year to the next in The plant in question was probably Common
the surface layers of their tuber. Observations Twayblade, and Pliny’s name Ophrys may have
of Late Spider Orchid show that the terminal derived from a Greek word for snake due to the
bud on the tuber starts to grow at the same similarity of the flowers of Common Twayblade
time as the onset of wet weather in the autumn. to the head and tongue of a snake.
Simultaneously the fungal mycelium starts
to grow outwards from certain points on the
tuber’s surface. The orchid’s new roots grow
outwards through this fungal mycelium and
the fungus probably enters the roots through
the root hairs. As the next season’s tuber starts
to develop it too is ‘infected’ (and will carry the
fungus through the next resting period) via the
soil, through minute, hair-like projections on
its surface.
Seed probably germinates in the spring
to produce a rounded protocorm. In Late
Spider Orchid and probably other species the
protocorm produces a root in the first autumn
and then a small tuber the following spring.
In the summer ‘resting period’ the protocorm
and root die off, leaving just the tuber. In
the autumn the bud on the top of the tuber
produces a short rhizome and roots. The first
leafy shoot appears in the second spring. In
cultivation flowering plants can sometimes be
produced within three years of germination.
Additional tubers can be formed at the base
of the aerial stem, perhaps often at the end
of slender rhizomes, and these will go on to
form separate plants as the connecting stem
dies off in the autumn. It seems, however, that
vegetative reproduction is either rare (Late
Spider Orchid) or uncommon (Early Spider
Orchid) among the British species.
Name M Unlike all the other members of the genus Ophrys,
Bee Orchid is self-pollinated. Being independent of insect
The origin of the generic name Ophrys is pollinators, it has done well in Britain, and is one of the
contentious. It may derive from ‘ophrus’, the most widespread and successful species.
005 systematic.indd 391 29/1/09 12:41:25

392 l GENUS OPHRYS
fLy orchid Vulnerable, BAP

Ophrys insectifera
Formerly: Ophrys muscifera

Fly Orchid shows the most wonderful example of insect mimicry among British orchids, but
rather than flies it has evolved to lure male digger wasps to act as inadvertent pollinators as
they attempt to copulate with its flowers. For human observers the small flowers are often
hard to spot but once seen have a magnetic attraction, drawing the eye back again and again.
Fly Orchid is relatively common but inconspicuous in woodland and scrub on chalk in
southern England and occurs at scattered localities in a variety of lime-rich habitats elsewhere
in Britain north to Cumbria and Anglesey. It is also found in central and western Ireland.
Identification
Very distinctive, the individual flowers are
indeed like little flies. The purplish-brown lip
forms the ‘body’ and the lustrous slate-blue
speculum shines like folded ‘wings’. The two
glistening depressions at the base of the lip are
the ‘eyes’ and above these the dark, wire-like
petals look just like little ‘antennae’.
Similar species
None.
Habitat
Very varied, although it usually grows on
calcareous soils over chalk and limestone. It
is found in open deciduous woodland and in
southern England is particularly associated
with beechwoods. It favours the better-lit areas
in glades, rides and along the edge of woodland,
as well as shaded road banks and open scrub,
but sometimes grows in deeper shade such as
overgrown hazel coppice. In southern England
Fly Orchid is infrequently found on open
grassland in old pits, quarries and on spoil
heaps, but when it does grow in such habitats
it may occur in large numbers; it has been
recorded rarely from slumped coastal cliffs. In
northern England and Ireland it is probably
more frequent in open areas and in addition to
wooded sites is found on limestone pavements
and rocky hillsides. It grows in alkaline fens and
O 23 May, Co. Clare. In the west of Britain and Ireland

Fly Orchid will grow in the open, on limestone pavements
and in fens.
005 systematic.indd 392 29/1/09 12:41:26

FLY ORCHID l 393
on the margins of turloughs in western Ireland

and also in fens on Anglesey, often among
tussocks of Black Bog-rush. Recorded up to
390m above sea level (Helbeck Wood, Brough,
Cumbria).
Flowering period
Late April to early July but mostly in late May
and early June.
1987-99
1970-86
pre 1970
Range
It is widespread but very local. In southern
England Fly Orchid is more-or-less confined to
the North and South Downs in Kent, Sussex
and Surrey, the chalk of Hampshire, Dorset,
Wiltshire and the Chilterns, and also the
Cotswolds. There are scattered records away
from these areas in Somerset, Warwickshire,
Northamptonshire, Suffolk, Nottinghamshire,
Derbyshire and more widely in Yorkshire and
the Morecambe Bay area of north Lancashire
and Cumbria. The northernmost sites are in
the upper Eden Valley of the north Pennines
in Cumbria. In Wales it is now confined to
Anglesey and in Ireland to a central belt from
southern Co. Mayo, Co. Galway and Co.
Clare, through north Tipperary, Co. Offaly
and Westmeath to Laois and Co. Kildare.
P 27 May, Hampshire. Fly Orchid can grow in heavy
shade but visits by suitable pollinators are probably more
likely where it catches the sun for part of the day.
005 systematic.indd 393 29/1/09 12:41:28

394 l GENUS OPHRYS
005 systematic.indd 394 29/1/09 12:41:31

FLY ORCHID l 395
World range: Confined to Europe, where it has

the most northerly distribution in the genus
Ophrys, ranging north to c. 67° in Norway
and to central Sweden, southern Finland and
the Baltic States. Occurs south to Spain, the
Balearic Islands, Italy, northern Greece and
Romania and east to the Moscow region.
How to find it
The tall slender spikes of Fly Orchid, with their
small, well-spaced flowers, can be very hard to
see among other vegetation and even on a bare
woodland floor they can vanish with ease. But,
if you can get your ‘eye-in’ where one Fly Orchid
is found, there may be plenty more. Within the
range in southeast England it can be found at
many woodland and scrubby sites on chalk or
limestone. In northern England sites include
Whitbarrow Scar (Cumbria) and Castle Eden
Dene (Co. Durham) and in Ireland it is locally
common in The Burren (Co. Clare).
DESCRIPTION
Height: 15-60cm.
Stem: Pale green and slender, with one or two
M 23 May, Co. Clare. The flower has evolved to mimic
basal sheaths. Groups of up to ten plants may digger wasps.The petals form the antennae and the bluish
grow together. ‘speculum’ on the lip resembles the glint of the wasp’s
folded wings.
Leaves: There are two to five shiny, dark green
or bluish-green leaves. The lower are narrow O 21 May, Hampshire.
and strap-shaped, flaccid but keeled and usually
pointed at the tip, and the upper one or two oval-oblong with a blunt tip but look narrower
are narrower, more pointed and loosely sheathe because their edges are rolled back and the
the stem. The leaves emerge in autumn and are inner face is concave. The lateral sepals are held
wintergreen. horizontally and the upper sepal is vertical but
Spike: Although initially bunched, the one to arches forward over the column to a variable
ten flowers (exceptionally as many as 20) are extent. The petals are dark purplish-brown
well-spaced along the stem by the time the with short, fine, downy hairs and are much
uppermost has opened. smaller than the sepals (less than half as long);
Bract: Dark green or bluish-green and their edges are rolled back to give them a fine,
lanceolate, often with the edges rolled inwards; filiform appearance and they point forward.
the lower bracts are rather longer than the The lip is longer than wide and hangs down
ovary but towards the tip of the spike they are a almost vertically. It is divided into three lobes,
little shorter. with two relatively short, narrow side-lobes
Ovary: Pale green, slender, cylindrical, six- spreading outwards at the base and a broad
ribbed and held upright but curving at the tip terminal lobe that is in turn notched or forked
to hold the flower facing outwards. at the tip. The lip is velvety in texture (the
Flower: The sepals are yellowish-green and are side-lobes are hairier), rich dark reddish-brown
005 systematic.indd 395 29/1/09 12:41:32

396 l GENUS OPHRYS
P 10 May, Kent. The

male digger wasp, excited
by the pheromones pro-
duced by the flower, as
well as by its appearance
and texture, attempts
‘pseudocopulation’ with
the orchid and in the
process acts as pollinator.
or purplish-brown, becoming a little paler Variation and varieties

towards the tip and duller with age. A more-or- There is considerable variation in the colour
less square, shining, pale slate-blue band across and markings of the lip.
the centre forms the speculum, and at the base Var. ochroleuca has a lip that is pale yellowish-
of the lip there are two shining ‘pseudoeyes’. green with a white speculum. It has been
The column is short and reddish-brown with recorded from Kent, Hampshire, Wiltshire and
a small, circular stigmatic cavity at its base and Hertfordshire.
a short, blunt ‘beak’. The two yellow pollinia Var. flavescens is a little darker than var.
have their viscidia enclosed in two off-white ochroleuca, with a yellowish-brown lip, pale blue
bursicles. or whitish speculum and greenish-brown petals.
It has been recorded from Gloucestershire.
Subspecies Var. subbombifera has a very broad, rounded
None. central lobe, notched as usual at the tip. Very
005 systematic.indd 396 29/1/09 12:41:34

FLY ORCHID l 397
rare, it has been recorded from Surrey and Historie of Plantes in which he noted: ‘The
Hampshire. Bee, the Fly and the Butterfly Satyrions grow
Var. parviflora has flowers that are around half upon barren chalky hill…adjoining to a village
the normal size. It is very rare. in Kent named Greenhithe, upon Longfield
Var. luteomarginata has a broad yellow border to downs by Southfleet’…. ‘ likewise in a field…
the central lobe of the lip, often also a yellow stain half a mile from S. Albons.’
or tip to the side-lobes and yellow or green tips to There have been considerable losses
the petals. It has been recorded rarely in Surrey, since 1597 with the species now occupying
Hampshire, Gloucestershire and Anglesey. only 42% of its historical range in Britain,
and it is classified as Vulnerable. Much of
BIOLOGY the decline took place long ago, especially in
East Anglia. However, in Ireland, where Fly
Pollination and reproduction Orchid is now found in just over 50% of its
Pollination is by male digger wasps Argogorytes historical range, rather more of the losses
mystaceus and also A. fargeii, a species that is rarer have been recent. Causes include woodland
and emerges a little later in the season. The wasps clearances and ‘coniferisation’ but perhaps
are attracted by pheromones emitted by the equally important has been the maturation
orchid and by the shape and texture of the flower. of woodland and scrub due to changes in
They attempt to copulate with it and during this forest management or its abandonment
‘pseudocopulation’ the pollinia are attached to altogether. In such cases increasing shade
their heads. Pollination rates are low, with less probably means that the flowers are seldom
than 20% of flowers setting seed (Summerhayes, pollinated and the population declines and
1968 quotes rates of 2.1% and 7.5% for two eventually disappears. Fly Orchid is either
samples of approximately 1,000 plants). extinct or has not been reported recently from
Devon, the Isle of Wight, Middlesex, Essex,
Norfolk, Cambridgeshire, Huntingdonshire,
The aerial stem grows from a pair of tubers, the
Worcestershire, Herefordshire, Shropshire,
younger of which is often stalked. The first leaf
Staffordshire, Leicestershire, Lincolnshire and
appears in the winter after germination and the
Co. Durham. Similarly in Wales there have
first tuber is formed in the second year.
been no recent reports from Monmouthshire,
Hybrids Glamorganshire and Denbighshire. There is an
x Bee Orchid has been recorded at two sites old, unconfirmed record from near Killiecrankie
in the Avon Gorge in Somerset and one site in in Perthshire in Scotland.
West Sussex, but it is very rare. It is sometimes
Past and present occurrence of Fly Orchid in Britain and
called O. x pietzschii but this is an invalid name. Ireland (based on presence or absence in 10km squares
O. x hybrida, the hybrid with Early Spider of the National Grid; data from the New Atlas).
Orchid, has occurred occasionally in Kent.
Britain Ireland
Name and classification total historical 264 31
The specific name insectifera derives from the range, 1500-1999
Latin insecta meaning ‘insect’ and fero meaning current range 110 (3.9%*) 16 (1.6%*)
‘bearing’, hence ‘insect-bearing’.
% lost, 1500-1969 47.5% 26%
% lost, 1970-1986 10.5% 22.5%
HISTORY AND
% lost, total 58% 48.5%
CONSERVATION
The first British record dates from 1597, when
John Gerard published The Herball, or General
005 systematic.indd 397 29/1/09 12:41:35

398 l GENUS OPHRYS
Bee orchid
Ophrys apifera
Looking, even feeling exotic and very special, the Bee Orchid is a standard bearer for orchids
and orchid conservation. Even for those with just a passing interest in wildlife, the discovery
of a Bee Orchid can be a special event, and people take great pride in ‘their’ local Bee Orchids.
An opportunistic and adaptable species, it often turns up in unexpected places; indeed, it
has even taken to garden lawns in recent years. Well named, the flowers look very much like
a bee and have evolved to attract male bees as pollinators. The mechanism has, however,
been abandoned, and almost all plants are now self-pollinated, a process that produces
copious seed, no doubt part of the secret of its success. It is widespread in England and parts
of Wales and Ireland, occurring in a wide variety of open grassland sites. Bee Orchid is the
county flower of Bedfordshire.
Identification
Unmistakably a ‘bee’ orchid, the flower looks
like a bumblebee. It has large pink sepals and
slender, parallel-sided, greenish or pinkish-
brown petals that form the bee’s ‘antennae’. The
lip is a ‘velvety’ maroon-brown with a pattern of
creamy markings and noticeably hairy rounded
side-lobes.
Similar species
In most of Britain and Ireland it is the only
‘bee’ orchid and therefore distinctive, but in
southern England care should always be taken
to distinguish it from Early and Late Spider
Orchids.
Late Spider Orchid is superficially very
similar but usually has a broader, more
rectangular lip with a distinctive forward or
downward pointing yellowish nib at its tip. In
the Bee Orchid the tip of the lip is almost always
tucked under and out of sight. In addition, the
petals of Late Spider Orchid are always pinkish,
short and very distinctly triangular or conical
in shape. Late Spider Orchid is not usually self-
pollinating and its pollinia never dangle loose.
Early Spider Orchid has yellowish sepals
and petals and a shiny slate-blue speculum on
its lip. It lacks any creamy or yellow markings.
O 24 June, Norfolk. As successive flowers open, the spike

lengthens. The pointed tip of the lip is normally tucked
under and out of sight.
005 systematic.indd 398 29/1/09 12:41:36

BEE ORCHID l 399
Habitat glacial debris) and, especially in Ireland, fens.

Very varied and defying any easy The species is tolerant of heavy grazing and
generalisations, although Bee Orchid is trampling. Although it is usually found in open
essentially a species of open grassland. Most sunny places it has been recorded growing in
sites are on light, well-drained soils that are shady woodland with little other vegetation.
low in nutrients, but it also grows on heavy In woods, it flowers a month later than nearby
clays, in areas that may have standing water plants in the open (Summerhayes, 1968).
in the winter, or on flushed, slumped clay Bee Orchids often behave like ‘weeds’,
cliffs. It can occur on closely cropped swards, colonising areas of bare or disturbed ground.
in areas with much bare ground or in ranker They increase rapidly in numbers until a
grassland among scrub. It is usually supposed closed sward develops and then, being ‘poor
to favour calcium-rich soils overlying chalk competitors’, they disappear. On the other
and limestone as well as chalky boulder hand, they can thrive in permanent grassland
clay, but it is much more widespread and and such colonies may last for many years. It
any poor, free-draining soil may be suitable, occurs from sea level up to 335m (Parsley Hay,
although there may well be some factor Derbyshire).
that makes the soil locally more alkaline.
Flowering period
Many habitats are obviously man-made,
Early June to late July, sometimes from late May,
such as road verges, railway embankments
but at least in southern England most plants
and cuttings, old quarries, pits, spoil-heaps,
will have finished flowering by the end of June.
gravel pits, brownfield industrial sites and
garden lawns. Bee Orchids are also found in Range
more natural areas such as sand dunes, dune It is widespread in England north to Cumbria
slacks, limestone pavement, eskers (ridges of and Co. Durham, although sparse or absent in
M 16 June, Norfolk. Despite an aura of ‘rarity’, Bee Orchid is an adaptable and successful species and can be found in
large numbers, often at ‘new’ sites.
005 systematic.indd 399 29/1/09 12:41:37

400 l GENUS OPHRYS
005 systematic.indd 400 29/1/09 12:41:40

BEE ORCHID l 401
flowering plants, but the total population,

1987-99
1970-86
including vegetative plants and those ‘dormant’
pre 1970 underground, is much more stable.
DESCRIPTION
Height: 10-45cm, sometimes to 65cm.
Stem: Pale green with two scale leaves at the
base.
Leaves: There are up to six pale green basal
leaves that are clearly veined, keeled and strap-
shaped but become narrower, more pointed
and loosely clasping higher on the stem. There
are also one or two bract-like non-sheathing
leaves towards the flower spike. The leaves
appear in September-November and are
often scorched or otherwise damaged by the
summer.
north Devon and Cornwall, and the number Spike: Loose, with two to seven flowers,
and density of colonies declines steadily sometimes as many as 12.
north and west of a line from Bristol to Bract: Pale green, lanceolate, pointed at the tip
Hull. In Wales it is largely confined to areas and much longer than the ovary.
on or near the north and south coasts. It is Ovary: Green, boldly ribbed but not twisted,
also found on the Channel Islands and the held upright but slightly curved; the tip bends
Isle of Man, and occurs throughout Ireland further over to hold the flower facing outward.
but is very local and absent from large areas. Flower: The sepals are oval, tapering slightly
Very rare in Scotland, with a 1908 record towards the tip, concave and often hooded.
from Kircudbrightshire, but found on an They are various shades of pink, from pale rose
old industrial site in east Ayrshire in 2003. to a deep pink tinged lilac, and have three to
World range: Europe, the Middle East and five variably obvious green veins. Occasionally
North Africa. Found north to Holland, the sepals are white. The lateral sepals are held
northwest Germany, the Czech Republic and horizontally or a little below the horizontal
the Ukraine and south to the Mediterranean and are swept backwards, whereas the upper
Islands, Crimea, Caucasus, Turkey, Lebanon sepal is held upright but very frequently bends
and Israel. In North Africa occurs in backwards to lie almost horizontally behind
Morocco, Tunisia and Algeria. the flower. The petals are much shorter and
How to find it are strap-shaped with their margins rolled
Throughout its range Bee Orchid is rather back to make them appear even narrower; they
local and as likely to be found on man-made are greenish through pinkish-brown to pink
sites as in old, species-rich grassland or scrub. with fine white hairs. The lip is tongue-shaped
It is worth looking for in any area that has with the sides and front strongly moulded
been heavily disturbed to produce lots of bare downwards and two relatively small conical
ground, especially if it is on poor, chalky or sides lobes at the base that are conspicuously
sandy soils, which are slow to be recolonised hairy on the outer side. The tip of the lip has
by vegetation. In established colonies two lobes with a pointed nib in the shallow
there are huge variations in the number of notch between them but appears rounded
because the entire tip is curled up underneath.
O 18 June, Norfolk. At the base of the lip there is an elongated,
005 systematic.indd 401 29/1/09 12:41:41

402 l GENUS OPHRYS
semi-circular, hairless, dull orange area that is until they are released, with the viscidia at their
bordered by narrow maroon-brown and pale bases enclosed in pale yellowish-green bursicles.
yellow bands. The speculum radiates from these At the base of the column the circular stigmatic
and is a broader band of dull purple that is in cavity is yellowish with a horizontal band of
turn bounded with a pale yellow band. The orange-brown.
markings form a U or H-shape below the basal
Subspecies
area, sometimes irregular and asymmetrical.
None.
The side-lobes are also bounded by dull purple
and pale yellow bands. The remainder of the Variation and varieties
lip is velvety maroon-brown. The column is As with the other species in the genus Ophrys,
greenish and held more-or-less at 90° to the the flowers are rather variable in structure,
lip. In profile it is said to resemble the head of colour and pattern.
a duck with the anther at the tip forming an Var. trollii ‘Wasp Orchid’ has a long narrow
elongated ‘beak’. The pollinia are yellow and central lobe to the lip that tapers to a point. The
their caudicles (stalks) lie in parallel grooves side-lobes are often longer and narrower than
normal, being held away from the central lobe.
The lip is marbled asymmetrically with yellow
and rusty-brown and the speculum is either
distorted or absent. It occurs regularly at a few
sites in the West Country but is otherwise rare.
The tip of the lip in ‘normal’ plants sometimes
fails to fold under and they then have an
elongated, pointed lip but retain the normal
markings. For a time, ‘Wasp Orchid’ was
considered to be a distinct species.
Var. belgarum has an oval lip, lacking well-
defined side-lobes but with hairy ‘shoulders’. It
has symmetrical markings with a horizontal
yellow band across the middle and smaller
vertical yellow bands; there is no speculum.
First described in 1998 it has been identified
widely in southern England.
Var. bicolor has the outer half of the lip dark
brown, grading to pale, unmarked greenish or
pale brown at the base. There is no speculum. It
is very rare but has been recorded from Dorset,
Essex/Suffolk and Anglesey.
Var. chlorantha lacks anthocyanin pigments
and has whitish sepals, yellow petals and a
bright greenish-yellow lip with a ‘ghost’ pattern.
It is rare but occurs widely, especially in the
south and east. Var ‘flavescens’ has also been
described. This is a less extreme version of
var. chlorantha with a pale brown lip that has a
normal but faded pattern. It is apparently very
M 26 June, Norfolk. The column is said to resemble the
head and beak of a duck. In these flowers the pollinia rare and easily confused with the old, faded
have already become stuck to the stigmas. flowers on normal plants.
005 systematic.indd 402 29/1/09 12:41:42

BEE ORCHID l 403
of a few hours the pollinia drop downwards on

their stalks and dangle in front of the stigma. In
this position the slightest breeze will waft them
on to its sticky surface and pollination will take
place. This mechanism is efficient and a large
proportion of flowers are pollinated. Each pod is
estimated to contain 6,000-10,000 seeds. Rarely,
trollii the flowers may be cross-pollinated. ‘Pseudo-
copulation’ involving bees has been recorded and
belgarum Bee Orchid has hybridised with several other
Ophrys; hybridisation is only possible if it is
cross-pollinated at least occasionally.
Vegetative propagation may occur and
in cultivation several small ‘daughter’ tubers
were noted growing at the end of long, slender
rhizomes.
bicolor
chlorantha
friburgensis
atrofuscus
Var. friburgensis has oval petals similar in

shape to the sepals but not quite as large. They
are pink or very rarely greenish. It is very rare
but has occurred widely; Bee Orchids with
slightly enlarged petals occur more commonly.
Var. atrofuscus has an unmarked, dark,
chocolate-coloured lip. It was found in Sussex
in 2001.
BIOLOGY
Bee Orchid is usually self-pollinated. Soon
after the flower opens the anther releases the
pollinia, which have unusually long, thread-like
flexible stalks. At their base these stalks are
each in turn attached to a viscidium, and these M 26 June, Norfolk. Soon after the flower opens the pol-
linia are released and dangle like little balls on the end
remain in their bursicles (protective pouches), of their flexible stalks, waiting for a breeze to blow them
effectively anchoring the pollinia. In the space onto the stigma.
005 systematic.indd 403 29/1/09 12:41:44

404 l GENUS OPHRYS
Development and growth Name and classification

The aerial stem grows from a pair of tubers. The specific name apifera derives from the Latin
It used to be thought that Bee Orchid was apis ‘bee’ and fero ‘to bear’ or ‘to carry’ and means
monocarpic and therefore flowered just once ‘bee-bearing’.
before dying. This was given as the explanation
for the large fluctuations in the number of HISTORY AND
flowering plants. The opposite is the case, CONSERVATION
however, as it is relatively long-lived. In a ten- The first British record dates from 1597, when
year study in Cambridgeshire the ‘half-life’ of John Gerard published The Herball, or General
successive generations in a population of Bee Historie of Plantes in which he noted: ‘The
Orchids averaged 6.6 years and varied from Bee, the Fly and the Butterfly Satyrions grow
5.8-11.2 years (the ‘half-life’ is a measure of upon barren chalky hill…adjoining to a village
the life expectancy of the orchid after its first in Kent named Greenhithe, upon Longfield
appearance above ground and marks the point downs by Southfleet’ … ‘likewise in a field…
at which 50% of the population that emerged in half a mile from S. Albons.’
any given year have died). Some plants live for The Bee Orchid is specially protected in
at least ten years after their first appearance and Northern Ireland under Schedule 8 of the 1985
may flower for three years in a row. Wildlife Order (NI)
Bee Orchids can spend one or two years
Past and present occurrence of Bee Orchid in Britain and
‘dormant’ underground but do not flower again Ireland (based on presence or absence in 10km squares
when they next emerge. They seem to need at of the National Grid; data from the New Atlas).
least a year’s growth as a non-flowering rosette
Britain Ireland
to build-up enough reserves to flower again.
At any stage in their life they can only flower if 1500-1999
they have reached a minimum leaf area, and if current range 785 (27.5%*) 93 (9%*)
they do not have a big enough spread of leaves
the rosette will wither early, in May. Damage % lost, 1500-1969 10.5% 37%
to the plants by snails and slugs tends to be % lost, 1970-1986 6% 12%
slight but rabbits and deer can cause severe % lost, total 16.5% 49%
problems. * current range as a % of the total number of 10km squares
Seed probably germinates in the spring but
there is no clear picture of the period between There has been a modest decline in Britain,
germination and the first appearance above with just 16.5% of the total range lost as sites
ground. A 19th century author noted that have been destroyed by ploughing, spraying
germinating seeds, protocorms and plantlets or, in the case of quarries and old pits, are
with small leafy shoots were found around either filled with rubbish or become overgrown
adult plants in March. Young Bee Orchids with scrub. The losses have to some extent
often appear above ground in the late winter as been compensated for by the colonisation of
non-flowering plants with a single leaf but these new sites and the expansion of the range to
frequently wither and die off rather quickly. the north and west, especially in man-made
Hybrids habitats: Bee Orchid is one of the more
O. x albertiana, the hybrid with Late Spider opportunistic and adaptable British orchids.
Orchid, was recorded from Kent in 1828. There has been a much more significant decline
x Fly Orchid is very rare but has been recorded in Ireland, especially in the period before 1930.
in Somerset and West Sussex (it is sometimes The explanation for this is not clear.
called O. x pietzschii but this is an invalid name).
005 systematic.indd 404 29/1/09 12:41:44

LATE SPIDER ORCHID l 405
LaTe sPider orchid Vulnerable

Ophrys fuciflora
A close relative of the Bee Orchid, this is one of the rarest and most localised species in Britain
and is restricted to a handful of sites in east Kent. A large proportion of the population is
confined to cages as protection from grazing rabbits, and a grassy slope scattered with wire-
mesh boxes is a strange introduction to such a rare plant. The flower has evolved to trick
one particular species of bee into acting as an inadvertent pollinator, but the appropriate
bee does not occur in England. The mechanism is therefore largely defunct, few flowers
are pollinated and little seed is produced. Nevertheless, this species manages to cling on,
perhaps because the individual plants are relatively long-lived.
Identification
Unmistakably a ‘bee’ orchid, this species has
pink sepals and petals and a broad, dark, velvety
lip. The size and colour of the petals and the
colour of the sepals are variable, however, as is
the shape and pattern of markings of the lip,
but it usually has furry ‘shoulders’ and always
has a projecting nib at the tip.
Similar species
Bee Orchid is rather similar but has longer,
narrower petals that are strap-shaped rather
than triangular and more often greenish than
pink. It has a narrower lip that is distinctly
three-lobed and never square and ‘shouldered’
(on the other hand, the lip of Late Spider
Orchid can be three-lobed). Diagnostically,
the pointed tip of the lip in Bee Orchid
normally curls back and under out of sight
and it therefore lacks the projecting nib. The
column in Bee Orchid is a little longer with a
slightly more prominent projecting ‘beak’, and
the pollinia often hang loose over the stigma, a
feature never seen in Late Spider Orchid.
Early Spider Orchid always has green sepals
and long, narrow, strap-shaped petals that are
much less downy and also usually greenish
rather than pink. The speculum on its lip is
normally H-, X- or π-shaped rather than
incorporating broken rings and circles. As in
Bee Orchid, its lip lacks a forward-pointing
nib and the flowering periods of the two spider M 12 June, Kent. The petals are always short and
orchids do not normally overlap. triangular and usually pinkish, resembling little horns.
005 systematic.indd 405 29/1/09 12:41:45

406 l GENUS OPHRYS
Habitat
Well-drained grassland on infertile chalky soils,
grazed to produce a reasonably short sward
and some bare ground and also ideally facing
south. Ground disturbance of some sort may
be important for the establishment of new
populations. Current sites are on old spoil
heaps and areas which were ploughed in the
past or heavily disturbed by rabbits prior to the
outbreak of myxomatosis. The individual plants
are relatively long-lived, and colonies can persist
when conditions are no longer suitable for
seedlings to become established. Notably, the
existing colonies tend to be very discrete and do
not expand into adjacent superficially similar
grassland.
Flowering period
Late May to late July, exceptionally to August,
but mostly in early to mid-June. At Wye NNR,
colonies that face west to southwest flower three
or four weeks later than colonies facing south.
1987-99
1970-86
pre 1970
Range
Confined to the North Downs in Kent between
Folkestone and Wye. There are old records from
other counties in southern England but all of
these are probably errors. World range: Europe,
from France eastwards to Romania; it extends
M 12 June, Kent. As the flowers open the spike elongates; north to southern Belgium, central Germany,
these flowers have very well-defined side lobes to the lip. Austria, Hungary and the Czech Republic and
005 systematic.indd 406 29/1/09 12:41:47

south to Italy and the former Yugoslavia. It is

extinct in Holland. Closely related species are
found in adjacent areas.
How to find it
Around 50% of the British population is
found in six discrete colonies at Wye NNR,
many plants are caged for protection. As with
Bee Orchid, the number of flowering plants
varies widely between seasons but the overall
population is much more stable.
DESCRIPTION
Height: 5-30cm, sometimes to 37.5cm.
Stem: Grey-green.
Leaves: Grey-green and prominently veined.
The three to five lanceolate basal leaves form a
loose rosette with the lowest flat to the ground
and the remainder held up to 30° above the
horizontal. There are one to three narrower and
more pointed leaves that loosely clasp the stem.
The rosette appears above ground in October
and withers while the plant is still in flower,
usually disappearing by early July.
Spike: The two to nine flowers (rarely as many
as 14) are well-spaced along the stem.
Bract: Grey-green, lanceolate, about one to
one-and-a-half times the length of the ovary,
which they clasp at the base. M 14 June, Kent. The flowers are similar to Bee Orchid
but the pointed tip of the lip is not tucked out of sight.
Ovary: Grey-green, slim and cylindrical, boldly
six-ribbed, slightly twisted, and also curved at the base, called ‘ears’ or auricles. The lip is
through about 90° to hold the flowers facing almost square in shape but usually broadens
outwards. towards the tip. The edges are strongly moulded
Flower: The sepals are broadly oval with their downwards apart from the tip, where there is a
edges rolled back and under and a blunt tip. prominent forward or downward projecting nib
They vary from very pale pink to a rich, dark set into a notch. This nib is usually yellowish
pink and have a prominent green midrib on and sometimes three-lobed. The lip is a rich,
the outer surface and one to three green veins dark chestnut-brown or maroon-brown, usually
on the inner face. The lateral sepals are held paler around the edges and velvety in texture.
horizontally or slightly drooping at the sides On either side of the base of the lip there are
of the flower, whereas the upper sepal may be swellings or ‘shoulders’ that are variable in size;
vertical or project out horizontally over the lip in some plants these swellings are large, well-
but more often curves forwards in a graceful defined and rounded and thus more obviously
arch. The petals are much shorter, triangular, lateral lobes. The lip is particularly hairy on the
velvety-hairy and pink (often a distinctly deeper ‘shoulders’ and along the sides below them. At
pink, sometimes almost flame-coloured, at the the base of the lip below the column is a more-
base). They may have swellings on each side or-less semicircular, smooth, orange-brown
005 systematic.indd 407 29/1/09 12:41:50

408 l GENUS OPHRYS
patch that is usually bordered by a narrow,

creamy ‘necklace’. Radiating from this necklace
the speculum is a complex and extremely
variable pattern of smooth maroon or lilac-
brown markings bounded by creamy-yellow
lines that often forms an irregular star or other
geometric shape around a central dark circle.
The column is held at about 90° to the lip and
is yellowish-green, becoming distinctly greener
towards the tip. The stigmatic cavity is dark with
a small, round, black ‘pseudoeye’ on either side.
The projecting rostellum or ‘beak’ is small and
the pollinia are yellow.
Subspecies
None.
There is a great deal of variation in the shape
and colour of the lip and colour of the petals
and sepals. One named variety is worth
mentioning:
Var. flavescens lacks anthocyanin pigments
and has whitish sepals and petals and a pale
greenish lip with a very faint pattern. It is
very rare.
BIOLOGY
This species is pollinated by insects, and in
Europe bees of the genus Eucera are the main
pollinators. The appropriate bees do not occur
in England, however, and insect-pollination
has not been recorded, although it must occur;
pollen-beetles are possible alternative vectors.
Very few ripe seedpods have been found in
Kent but despite this enough viable seed is
produced to maintain the populations. Self-
pollination may occur occasionally, even in the
bud (Summerhayes, 1968).
Vegetative reproduction is either very rare
or does not occur at all. This statement is
based on the observation that very few new
plants appear within 10cm, or even 30cm, of
existing orchids.
O 12 June, Kent.The column, as in Bee Orchid, resembles

a duck’s head but is a little shorter.
005 systematic.indd 408 29/1/09 12:41:53

Development and growth with plants appearing regularly at just five

The aerial stem grows from a pair of tubers. sites although there are probably still sporadic
It had been thought that Late Spider Orchid appearances at several more. The low point
was monocarpic and flowered just once before was in the mid-1980s, and there has been
dying. The opposite is the case, as it is rather a substantial recovery since then, perhaps
long-lived. In studies in Kent the ‘half-life’ of a due in part to the weather but certainly to
population averaged 12.5 years and varied from better management. There is now a total of
7.1 to 16.8 years (the ‘half-life’ is a measure of approximately 500 plants, almost half of which
the life expectancy of the orchid after its first are at Wye NNR. The population there rose
appearance above ground and marks the point from about 50 in 1987 to around 220 in 1998.
at which 50% of the population that emerged Habitat has been lost because of agricultural
in any given year will have died). Periods of changes but also due to changes in grazing
underground ‘dormancy’ are frequent with
about 13% of plants dormant each season,
and plants may spend one or two years
underground before re-emerging.
There is no information on the length of
the period between germination and the first
appearance above ground but it is probably
three or four years. Many plants flower in their
first year above ground.
Hybrids
O. x albertiana, the hybrid with Bee Orchid,
has been recorded rarely from Kent.
O. x obscura, the hybrid with Early Spider
Orchid, has been recorded rarely in Kent. The
parent species seldom occur together and the
flowering periods do not normally overlap.
The specific name fuciflora derives from fucus
‘drone’ (i.e. a male bee but also meaning ‘paint’
or ‘dye’, especially red or purple colours) and flos
‘flower’ and thus presumably ‘bee-flowered’.
HISTORY AND
CONSERVATION
The first British record dates from 1828
when the Rev. Gerard E. Smith noted it, ‘on
the southern declivities of chalky downs near
Folkestone’ (The English Flora).
A Red Data Book species, it is classified
as Vulnerable and specially protected under
Schedule 8 of the Wildlife and Countryside
Act 1981. Late Spider Orchid has only ever
M 14 June, Kent. Ophrys x albertiana, the hybrid with Bee
been reliably recorded from Kent, with about Orchid. Under suspicion for some years, the identity of this
20 historic localities. It is now much reduced, plant as a hybrid was finally confirmed by DNA analysis.
005 systematic.indd 409 29/1/09 12:41:54

410 l GENUS OPHRYS
patterns and the reduction in rabbit numbers

following myxomatosis, with grassland
reverting to scrub. All the regular sites are now
within SSSIs and are managed with the species
in mind, but at its former localities grazing was
abandoned many years ago. The Late Spider
Orchid is probably best adapted to a ‘dynamic’
system. It needs ground disturbance and some
bare ground to provide suitable conditions
for seedlings to become established, as well
as grazing and sometimes hand-mowing to
maintain a suitable short sward for the adult
plants to thrive.
The preferred pollinator is not present
in Britain and few flowers produce seed.
To counter this, hand-pollination has been
undertaken at Wye. Fortunately, with a very low
level of mortality in adult plants (less than 5%
per annum), only small numbers of seedlings
need to survive and flower in order to maintain
the population.
Past and present occurrence of Late Spider Orchid in Brit-

Britain Ireland
1500-1999
% lost, 1500-1969 33.3%

% lost, 1970-1986 0%
% lost, total 33.3%
O 14 June, Kent. The lip shape in the small English

population is extraordinarily variable; in this plant the
sepals are largely greenish.
005 systematic.indd 410 29/1/09 12:41:55

EARLY SPIDER ORCHID l 411
earLy sPider orchid

Ophrys sphegodes
Just about the earliest orchid to flower in spring (vying with Early Purple Orchid for the
honour), this species is almost entirely restricted to the south coast between Dorset and
Kent. Although ‘Nationally Scarce’ it can be locally common and has appeared in vast
numbers at Samphire Hoe, Dover, on spoil excavated from the Channel Tunnel. Indeed, the
prospect of such a ‘rare’ plant dotted in hundreds or even thousands across the downland
turf is almost a mockery of the term itself. The flowers may look like a spider to human eyes
but have in fact evolved to trick solitary bees into acting as pollinators.
Identification
Unmistakably a ‘bee’ orchid, the circular deep
brown lip resembles the body of a large garden
spider and is marked with a lustrous bluish
speculum. The sepals are green, and the petals
are yellowish-green, narrow and strap-shaped
with wavy edges.
Similar species
Bee and Late Spider Orchids both have similar
large brownish lips but their specula are
bordered by a narrow creamy line and their
sepals and petals are pink.
Habitat
It is found on old species-rich grassland on
chalk or limestone, growing both in short
closely-cropped turf (even lawns) and in slightly
ranker swards. It has some preference for
previously disturbed areas, such as old quarries,
spoil heaps and tracks; as with Late Spider
Orchid heavy ground disturbance may aid the
successful establishment of new colonies. It has
been recorded rarely from shingle and almost all
the current sites are near the sea.
Flowering period
Late March to early June. Tends to flower
earliest in Dorset, from early or mid-April to
early or mid-May and may average a few days
later in Sussex and Kent. Flowering times are
very variable, however, both from year to year
and between colonies, even in the same area.
P 14 May, Kent. Many Early Spider Orchids are very

small.
005 systematic.indd 411 29/1/09 12:41:57

412 l GENUS OPHRYS
For example, plants at Samphire Hoe in Kent

1987-99
bloom perhaps two weeks earlier than those at 1970-86
Lydden, just 6km away. pre 1970
Range
Largely confined to the coasts of Dorset, East
Sussex and Kent; in Dorset there are good
populations between St Aldhelm’s Head and
Durlston; in East Sussex now found mostly
between Beachy Head and Seaford and at
Castle Hill near Brighton; in Kent there are
around 30 colonies, most along the coast
but including one or two inland sites such
as Queendown Warren. Away from these
strongholds there have been isolated recent
records in west Gloucestershire (from 1975),

south Wiltshire (1988), Northamptonshire
(2001, the first record for 230 years) and west
Suffolk (where a single plant appeared from
1991 at Lakenheath, the first record since
1793). Introduced and naturalised near
Bishop’s Stortford in Hertfordshire. World
Range: Mainland Europe from Spain eastwards
to Greece and north to Belgium, Luxemburg,
central Germany, Austria and the Czech
Republic.
How to find it
One of the best and most accessible sites is
Samphire Hoe below Shakespeare Cliff at
Dover. This was created using five million cubic
metres of marl dug from beneath the seabed
during the construction of the Channel Tunnel.
In 1998 there were 61 plants but by 2004 this
has increased to 9,000. In East Sussex up to
50,000 plants have been recorded at Castle Hill,
and it is also found at Cuckmere Haven and the
Seven Sisters, and in Dorset the most accessible
site is Durlston Head.
DESCRIPTION
Height: Mostly five to 15cm, sometimes to
20cm or rarely 35cm, but at Samphire Hoe
plants may be as tall as 45cm.
M 14 May, Kent. The greenish sepals and petals are dis-
tinctive. Stem: Yellowish-green, thick and fleshy.
005 systematic.indd 412 29/1/09 12:41:58

Leaves: Grey-green to green and prominently

veined. The three or four lower leaves are short
and broadly strap-shaped with a blunt tip; the
lowermost is held nearly horizontally but the
remainder are more upright. The upper leaves
are narrower, more pointed and loosely sheathe
the stem. Some plants are wintergreen but in
most the leaves appear in the early spring. All
have withered by the end of June.
Spike: Most have two to seven flowers,
exceptionally as many as ten, but at Samphire
Hoe some tall plants may have as many as 17
blooms widely spaced along the stem.
Bract: Pale green, lanceolate, blunt-tipped and
slightly longer than the ovary, which they more-
or-less sheathe at their base.
Ovary: Green and boldly six-ribbed.
Flower: The sepals are oval-oblong with blunt
tips and their margins rolled under. They are
usually pale yellowish-green in coloration but
rarely are more whitish. The petals are more M 14 May, Kent.The ‘speculum’ is usually a lustrous blue
strap-shaped, shorter and narrower, with a or slate-grey, and the flower also has a pair of ‘pseudo-
eyes’ on either side of the stigma.
squared-off tip and edges that are frequently
waved or undulate. They are often a slightly on the upper with two ‘pseudo-eyes’, one on
deeper yellowish-green and their margins each side; it has been reported to contain ‘sugar’
may be more brownish with some fine hairs. (Summerhayes, 1968). The pollinia are yellow.
Very rarely the petals are pinkish. The lateral The flowers fade rapidly, the lip becoming a dull
sepals are held horizontally and the upper sepal pale yellowish-brown or grey-brown.
vertically although the latter usually arches
Subspecies
forward over the column. The lip is almost
None.
circular in shape with the edges moulded
downwards to give it a convex profile. It is Variation and varieties
deep purplish-brown, usually paler and more The shape and coloration of the lip is very
yellowish around the lower edge and velvety in variable and rarely the speculum may be red
texture with a smooth, slightly lustrous, slate- rather than blue.
grey, lead-coloured or bluish mark in the centre Var. flavescens lacks anthocyanin pigments and
or towards the base (the speculum), often in has a greenish or golden lip, sometimes brownish
the form of an H, X or the Greek letter π. On towards the edges, with merely a ‘shadow’
either side of the base of the lip there are two speculum. It is rare but can easily be confused
roughly conical swellings or haunches that with a normal flower that has faded with age.
are variable in size, and the lip is particularly
hairy around these and along the edges below BIOLOGY
them. The column is greenish-white and held Pollination and reproduction
at about 90° to the lip with the rostellum at It is cross-pollinated by bees. The specific
its tip forming a short blunt beak. At the base pollinator is the male solitary bee Andrena
of the column the stigmatic cavity is circular, nigroaenea, and the pollinia are attached to
maroon-brown on the lower half and pale green the front of its head as it attempts ‘pseudo-
005 systematic.indd 413 29/1/09 12:41:59

414 l GENUS OPHRYS
copulation’. The mechanism can be successful is presumably being produced to sustain the
with about 25% of flowers recorded as setting current colonies despite the apparently low
seed at Samphire Hoe, but pollination rates rates of pollination.
can be very much lower with a lack of suitable
pollinators being the most likely explanation for Development and growth
the low seed-set. It has also been suggested that The aerial stem grows from a pair of tubers, the
British populations are probably mostly self- younger of which is often stalked.
pollinated. Indeed, the pollinia are occasionally Early Spider Orchid is relatively short-lived.
released to dangle in front of the stigma as in Few plants survive for more than three years
Bee Orchid, making self-pollination possible. after their first emergence above ground and
The overall low rates of pollination suggest, the majority appear just once, flower and then
however, that self-pollination is not routine. die (i.e. they are monocarpic), although a tiny
Vegetative reproduction is thought to minority can live for at least ten years. The
be uncommon or rare and as Early Spider number of plants above ground varies widely
Orchid is a short-lived orchid, dependent on from year to year, largely correlated with the
seed to maintain its numbers, sufficient seed amount of rainfall over the previous winter, and
plants may spend one or two years ‘dormant’
underground.
Early Spider Orchid has a complex life cycle.
This complicates attempts to understand its
population dynamics and life history on the
basis of an annual census. In a study at Castle
Hill, Sussex, over a winter season, plants began
to appear above ground in early September
and continued to emerge through the winter,
with peaks in November-December and more
especially in March-May. By flowering time
therefore some plants had been above ground
for just two months and some for six. Those
that appeared early in the winter included
most of the older plants that had flowered the
year before and all these early plants suffered
grazing damage. Previously unrecorded plants,
presumably seedlings emerging above ground
for the first time, appeared throughout the
season but especially from March onwards.
However, around 75% of all the plants recorded
up till March did not survive above ground
to be counted in the annual census in May,
with grazing being the likely cause of their
disappearance.
There is no information on the period
between germination and the first appearance
above ground, but it is probably in the region
of one to three years. Early Spider Orchids first
M 8 May, Dorset. The lip fades rapidly to a dull brown
and thus as successive flowers open only one or two at a
appeared on spoil from the Channel Tunnel
time will be in ‘good condition’. four years after it had been spread.
005 systematic.indd 414 29/1/09 12:42:00

Hybrids needs careful management to maintain the

O. x hybrida, the hybrid with Fly Orchid, has populations. Grazing by sheep is beneficial,
occurred occasionally in Kent. as long as they are removed during the period
O. x obscura, the hybrid with Late Spider when the orchids flower and set seed. Any
Orchid, has been recorded rarely in Kent. The future spread may depend upon the creation
parent species seldom occur together and the of suitable bare ground and the sort of drastic
flowering periods do not normally overlap. disturbance that created such ideal conditions
at Samphire Hoe.
The specific names sphegodes derives from the Past and present occurrence of Early Spider Orchid in Brit-
Greek sphex ‘wasp’ or sphekeion ‘small, wasp-like ain and Ireland (based on presence or absence in 10km
spider’ and means ‘wasp-like’.
Britain Ireland
HISTORY AND
CONSERVATION 1500-1999
The first British record dates from 1650 current range 17 (0.6%*) 0
when William How published his Phytologia
% lost, 1500-1969 71.5%
Britannica natales exhibens Indigenarum
Stirpium sponte emergentium (‘A British botany % lost, 1970-1986 1.5%
presenting the origins of native wild plants’) in % lost, total 73%
which he noted it in Northamptonshire, ‘upon *current range as a % of the total number of 10km squares
an old Stone pit ground…hard by Walcot a
mile from Barnack…Dr. Bowle’.
A Red Data Book species and although
no longer classified as Near-threatened it
remains specially protected under Schedule
8 of the Wildlife and Countryside Act 1981.
The Early Spider Orchid has vanished from
at least 73% of its historical range but the
majority of the losses occurred long ago, in
the 19th century. This was probably largely
due to the cultivation of grasslands following
the Enclosures, although the retreat towards
the coast of southeast England suggests that
climatic factors may also have been involved.
It is now extinct in the Channel Islands, west
Cornwall (where recorded at Upton Towans in
1961), the Isle of Wight (a few records, the last
in 1992), Hampshire (very few records, the last
near Lepe in 1978), West Sussex, Surrey (last
recorded from Titsey in 1942 but this record
probably originated with plants transplanted
from Queendown Warren in Kent), Essex,
Oxfordshire (1920, with an unconfirmed record
in 1975), Bedfordshire, Cambridgeshire and
Denbighshire (a 19th century record).
Many sites for the Early Spider Orchid are M 14 May, Kent. Early Spider Orchids can have just one
now protected but its grassland habitat also flower; this plant has especially brownish petals.
005 systematic.indd 415 29/1/09 12:42:01

416 l OTHER SPECIES
oTher sPecies
Several other species of orchid have been recorded from time to time in Britain and Ireland.
This is a selection:
Short-spurred Fragrant Orchid Heart-flowered Tongue Orchid

Gymnadenia odoratissima Serapias cordigera
Recorded from Black Hall Rocks in Co. Found in 1996 at Monkton on the Isle of
Durham in 1912, but doubt has been cast on Thanet in Kent, with two or three plants in
the identity of the specimen. 1997. They were on a south-facing bank at
the bottom of a cliff in an abandoned chalk
Frivald’s Fragrant Orchid quarry. This is a common and widespread
Gymnadenia frivaldii species around the Mediterranean and occurs
Found and photographed on wet heathland in north to Finistere. Despite apparently excellent
Dorset in 1972 or 1973, its normal range is the credentials, these plants almost certainly
Balkans and its occurrence here is a mystery originated from wind-blown seed coming from
(Ettlinger 1997). plants growing in pots on a patio just 300m
False Musk Orchid (Dwarf Alpine upwind of the nature reserve.
Orchid) ‘Bertoloni’s Mirror Orchid
Chamorchis alpina Ophrys bertolonii’
A single plant was reported from the New A single plant found in Dorset in 1976
Forest in Hampshire in 1976. Among the generated much controversy as to its origins,
smallest of European orchids, this is a species but it eventually emerged that a holidaymaker,
of damp, calcareous soils in the mountains returning from Italy, had planted it. ‘Ophrys
of Scandinavia and the Alps. It was probably bertolonii’ has since been divided into several
planted. species, and the precise identity of the Dorset
Pale-flowered Orchid plant cannot be confirmed.
Orchis pallens
A few specimens were recorded in the 1920s
in Hampshire but there are no further details
(Ettlinger 1997).
Scarce Tongue Orchid

Serapias neglecta
Found in 1918 in a cornfield on the Isle of
Wight, the roots were dug up and taken into
cultivation. Rather localised in coastal regions
around the central Mediterranean, this species
may have been introduced with imported
cereal seed.
005 systematic.indd 416 29/1/09 12:42:01

ORCHID FAMILY TREE l 417
Orchid Family Tree

Cypripedium (lady’s-slippers)
Subfamily
Cypripedioideae
Cephalanthera (helleborines i)
tribe Neottieae
Neottia (Bird’s-nest Orchid etc.)
Subfamily
FAMILY Epidendroideae Epipactis (helleborines ii)
tribe Gastrodieae
ORCHIDACEAE Epipogium (Ghost Orchid)
tribe Calypsoeae
Corallorhiza (Coralroot Orchid)
tribe Malaxideae Liparis (Fen Orchid)

Hammarbya (Bog Orchid)
tribe Goodyera (Creeping Lady’s-tresses)

Spiranthes (lady’s-tresses)
Cranichideae
subtribe
Herminium (Musk Orchid)
Habenariinae
Subfamily Dactylorhiza (marsh orchids)
Orchidoideae
tribe Gymnadenia (fragrant orchids)
Orchideae Platanthera (butterfly orchids)

Pseudorchis (Small White Orchid)
Orchis (’typical’ orchids)
subtribe Neotinea (Dense-flowered Orchid etc.)

Orchidinae
Himantoglossum (Lizard Orchid)
Anacamptis (Pyramidal Orchid etc.)
Serapias (tongue orchids)
Ophrys (bee orchids)
All orchids are members of the family Orchidaceae. Its most fundamental subdivisions are the
subfamilies, three of which are represented in Britain and Ireland: the Cypripedioideae, with just one
species – Lady’s-slipper; the Epidendroideae, a heterogeneous group, many of which are woodland
orchids that grow from rhizomes; and the Orchidoideae, a more homogenous group, all of which
grow from a swollen underground tuber. The subfamilies are further divided into tribes, genera and,
finally, individual species.
This family tree follows Bateman (2006) and has been constructed using the latest evidence from
genetic research. It gives a much better idea of relationships than is possible in a simple list. For
example, Bog Orchid and Creeping Lady’s-tresses lie next to each other in the field guide, but it can
be seen that they are members of separate subfamilies and not at all closely related.
006 systematic.indd 417 29/1/09 12:49:20

418 l VICE-COUNTIES
vice-cOunTies
112
111
108 109
107
110
105 106
95 94
93
104 96
92
97
91
89 90
103 88
98
87
85 84
86
102 83 82
77 78 81
101
75 79 80 68
35 100
40 72
67
34 39 74 73
36 70 66
27 28 33 37 38
32 69 65
29 62
26 25 30 71
24 31 64
16 17 22 60 61
23
15 18 52 51 59 63
19 21 54
9 14 58
20 49 50
10 57 56
8 13 48
7 11 12 39 53
2 47 40 55 28 27
5 6
1 4 38 31
46 43 37 32
3 36 29 26 25
45 44 42 30
33 23 24 20 19
41 35
34 7 21 18
22
S 6 12 17 16
5 8 15
4 13 14
9 11
2 3
10
1
006 systematic.indd 418 29/1/09 12:49:21

VICE-COUNTIES l 419
England Isle of Man 107 East Sutherland

1 West Cornwall 71 Isle of Man 108 West Sutherland
2 East Cornwall Wales 109 Caithness
3 South Devon 35 Monmouthshire 110 Outer Hebrides
4 North Devon 41 Glamorganshire 111 Orkney
5 South Somerset 42 Breconshire 112 Shetland
6 North Somerset 43 Radnorshire Channel Islands
7 North Wiltshire 44 Carmarthenshire S Channel Islands
8 South Wiltshire 45 Pembrokeshire
9 Dorset Ireland
46 Cardiganshire H1 South Kerry
10 Isle of Wight 47 Montgomeryshire
11 South Hampshire H2 North Kerry
48 Merioneth H3 West Cork
12 North Hampshire 49 Caernarvonshire
13 West Sussex H4 Mid Cork
50 Denbighshire H5 East Cork
14 East Sussex 51 Flintshire
15 East Kent H6 Co. Waterford
53 Anglesey H7 South Tipperary
16 West Kent
17 Surrey Scotland H8 Co. Limerick
18 South Essex 72 Dumfries-shire H9 Co. Clare
19 North Essex 73 Kirkcudbrightshire H10 North Tipperary
20 Hertfordshire 74 Wigtownshire H11 Co. Kilkenny
21 Middlesex 75 Ayrshire H12 Co. Wexford
22 Berkshire 76 Renfrewshire H13 Co. Carlow
23 Oxfordshire 77 Lanarkshire H14 Laois
24 Buckinghamshire 78 Peebles-shire H15 Southeast Galway
25 East Suffolk 79 Selkirkshire H16 West Galway
26 West Suffolk 80 Roxburghshire H17 Northeast Galway
27 East Norfolk 81 Berwickshire H18 Offaly
28 West Norfolk 82 East Lothian H19 Co. Kildare
29 Cambridgeshire 83 Midlothian H20 Co. Wicklow
30 Bedfordshire 84 West Lothian H21 Co. Dublin
31 Huntingdonshire 85 Fife H22 Meath
32 Northamptonshire 86 Stirlingshire H23 Westmeath
33 East Gloucestershire 87 West Perth H24 Co. Longford
34 West Gloucestershire 88 Mid Perth H25 Co. Roscommon
36 Herefordshire 89 East Perth H26 East Mayo
37 Worcestershire 90 Angus H27 West Mayo
38 Warwickshire 91 Kincardineshire H28 Co. Sligo
39 Staffordshire 92 South Aberdeen H29 Co. Leitrim
40 Shropshire 93 North Aberdeen H30 Co. Cavan
53 South Lincolnshire 94 Banffshire H31 Co. Louth
54 North Lincolnshire 95 Moray H32 Co. Monaghan
55 Leicestershire 96 Easterness (Inverness-shire) H33 Fermanagh
56 Nottinghamshire 97 Westerness H34 East Donegal
57 Derbyshire (Inverness-shire) H35 West Donegal
58 Cheshire 98 Main Argyll H36 Tyrone
59 South Lancashire 99 Dunbarton H37 Co. Armagh
60 West Lancashire 100 Clyde Islands (Buteshire) H38 Co. Down
61 Southeast Yorkshire 101 Kintyre H39 Co. Antrim
62 Northeast Yorkshire 102 South Ebudes H40 Co. Londonderry
63 Southwest Yorkshire (Inner Hebrides)
64 Mid-west Yorkshire 103 Mid Ebudes (Some alternative names are add-
65 Northwest Yorkshire (Inner Hebrides) ed in parentheses to allow easier
66 Co. Durham 104 North Ebudes reference to modern or historical
67 South Northumberland (Inner Hebrides) counties.)
68 Cheviot 105 West Ross
(north Northumberland) (Ross & Cromarty)
69 Westmorland 106 East Ross
70 Cumberland (Ross & Cromarty)
006 systematic.indd 419 29/1/09 12:49:21

Orchid FlOwering periOds
Apr May Jun Jul Aug Sept
Lady’s-slipper
Red Helleborine
Sword-leaved Helleborine
White Helleborine
Lesser Twayblade
Common Twayblade
Bird’s-nest Orchid
Marsh Helleborine
Dark-red Helleborine
Broad-leaved Helleborine
Violet Helleborine
Narrow-lipped Helleborine
Dune Helleborine
Lindisfarne Helleborine
Green-flowered Helleborine
Ghost Orchid
Fen Orchid
Bog Orchid
Coralroot Orchid
Creeping Lady’s-tresses
Irish Lady’s-tresses
Autumn Lady’s-tresses
Musk Orchid
Man Orchid
Monkey Orchid
Military Orchid
Lady Orchid
Early Purple Orchid
Small White Orchid
Lesser Butterfly Orchid
Greater Butterfly Orchid
Common Fragrant Orchid
Heath Fragrant Orchid
Marsh Fragrant Orchid
Early Marsh Orchid
Frog Orchid
Common Spotted Orchid
Heath Spotted Orchid
Southern Marsh Orchid
Pugsley’s Marsh Orchid
Hebridean Marsh Orchid
Northern Marsh Orchid
Irish Marsh Orchid
Dense-flowered Orchid
Burnt Orchid
Lizard Orchid
Loose-flowered Orchid
Pyramidal Orchid
Green-winged Orchid
Fly Orchid
Bee Orchid
Late Spider Orchid
Early Spider Orchid
Apr May Jun Jul Aug Sept
006 systematic.indd 420 29/1/09 12:49:21

SITE GUIDE l 421
Site Guide
INTRODUCTION
The following is a selection of sites in Britain and Ireland where it is possible to find wild orchids.
Of course, orchids can be found in many places, but sadly more and more of the countryside is bereft
of their beauty and it is increasingly necessary to search out special places in order to experience the
magic of wild orchids.
We see this selection as a starting point and would encourage everyone to explore and discover
new places, to record the orchids that they find and to pass this information on to the local botanical
recorder. But, in the meantime, you may be in an unfamiliar area, on holiday, or just keen to find
somewhere that offers a good chance of success.
For each site we have included brief details of access (including a grid reference), habitat and
the species that may be found. Many orchid sites are accessed down a maze of lanes, and it is very
difficult to give concise and understandable written directions. To locate many of these sites you
will need a large-scale road atlas, especially one that marks the National Grid and therefore allows
you to pinpoint grid references or, even better, an OS map at 1:50,000 or 1:25,000 scale. If the site
is a reserve (and most are) further information can often be found on the internet by visiting the
managing organisation’s website. It is also worth trawling the ‘net’ for information on nearby reserves
or Roadside Nature Reserves which are often also good for orchids.
This selection includes sites for some of the rarer orchids but these are all accessible and welcome
visitors. On many reserves, guided walks are organised from time to time by the managing body and
these can be a good introduction to the site. We have not tried to include localities for every species;
one or two are so rare and vulnerable that visitors are not welcome, and some species, for example
Green-flowered Helleborine, occur in small, scattered, hard-to-find colonies in places that have no
other orchid interest. Rather, we have tried to select areas that hold a good selection of species, or have
large and spectacular displays of one or two orchids. We have also endeavoured to choose sites that
are interesting and attractive, and almost all will have a great deal of other wildlife interest, be it other
plants, butterflies, dragonflies or whatever. All have public access throughout the orchid season, with
the exception of one or two where entry is restricted to a limited number of open days. Unfortunately,
we have not been able to give details of disabled access, but it is generally rather limited.
Orchid hunting requires very little equipment. As well as a good map, a hand-lens is useful, as is
a small plastic ruler and, of course, a field guide to all the other wonderful plants that will be seen.
Finally, it should be remembered that many orchids are scarce, and some are so rare that they are
protected by law. All orchids are valuable, as indeed are the other plants around them. Please look
where you are going, stay on paths where possible, and leave the site as you would wish to find it.
Inevitably there are errors in the accounts, for which we take full responsibility. We would be
delighted to hear of any corrections or updates, or indeed any additional sites that could be included
in future editions of the book (the authors can be contacted via their website: www.norfolknature.
co.uk or via the publishers).
006 systematic.indd 421 29/1/09 12:49:21

422 l SITE GUIDE
useFul addresses
An Taisce (National Trust for Ireland), Tailor’s Hall, Back Lane, Dublin 8, Ireland
Tel: +353 (0)454 1786. Email: [email protected] Website: www.antaisce.org
CEH Monkswood, Abbots Ripton, Huntingdon, Cambridgeshire PE28 2LS
Tel: 01487 772400. Email: [email protected] Website: www.ceh.ac.uk
The Countryside Agency, Head Office: John Dower House, Crescent Place, Cheltenham,
Gloucestershire GL50 3RA
Tel: 01242 521381. Email: [email protected] Website: www.countryside.gov.uk
Countryside Council for Wales, Maes-y-Ffynnon, Penrhosgarnedd, Bangor, Gwynedd LL57 2LQ
Tel: 01248 385500. Email: [email protected] Website: www.ccw.gov.uk
Environment and Heritage Service, Commonwealth House, 35 Castle Street, Belfast BT1 1GU
Tel: 028 90 251477. Website: www.ehsni.gov.uk
Irish Wildlife Trust, 107 Lower Baggot Street, Dublin 2, Ireland
Tel: +353 (0)1 676 8588. Email: [email protected] Website: www.iwt.ie
Joint Nature Conservation Committee, Monkstone House, City Road, Peterborough PE1 1JY
Tel: 01733 562626. Email: [email protected] Website: www.jncc.gov.uk
National Parks and Wildlife Service, Department of the Environment, Heritage & Local
Government, 7 Ely Place, Dublin 2, Ireland
Tel: +353 (0)1 888 2000. LoCall: 1890 202021 (within Republic of Ireland only).
Email: [email protected] Website: www.environ.ie or www.npws.ie
National Trust, 36 Queen Anne’s Gate, London SW1H 9AS
Tel: 020 72229251. Website: www.nationaltrust.org.uk
National Trust (Northern Ireland), Rowallane, Saintfield, Ballynahinch, Co. Down BT24 7LH
Tel: 028 97 510721. Website: www.ntni.org.uk
National Trust for Scotland, Wemyss House, 28 Charlotte Square, Edinburgh EH2 4ET
Tel: 0131 2439300. Website: www.nts.org.uk
Natural England, 1 East Parade, Sheffield, S1 2ET.
Tel: 0845 600 3078. Email: [email protected] Website: www.naturalengland.
org.uk
Royal Society for the Protection of Birds, The Lodge, Sandy, Bedfordshire SG19 2DL
Tel: 01767 680551. Website: www.rspb.org.uk
Scottish Natural Heritage, 12 Hope Terrace, Edinburgh EH9 2AS
Tel: 0131 447 4784. Email: [email protected] Website: www.snh.org.uk
Scottish Wildlife Trust, Cramond House, Off Cramond Glebe Road, Edinburgh EH4 6NS
Tel: 0131 312 7765. Email: [email protected] Website: www.swt.org.uk
The Wildlife Trusts, The Kiln, Waterside, Mather Road, Newark NG24 1WT
Tel: 0870 036 7711. Email: [email protected] Website: www.wildlifetrusts.org
006 systematic.indd 422 29/1/09 12:49:22

BOTANICAL ORGANISATIONS l 423
BOTanical OrganisaTiOns
Botanical society of the British isles
Hon. General Secretary, c/o Department of Botany, The Natural History Museum, Cromwell Road,
London SW7 5BD. Website: www.bsbi.org.uk
The BSBI is a learned society of professional and amateur botanists dedicated to the study of and
interest in the British and Irish flora. It publishes a journal, Watsonia, as well as the less formal (and
eminently readable) BSBI News. In addition, the BSBI organises the system of county plant recorders,
runs field meetings and has a panel of referees to advise on plant identifications (including, of course,
orchids). BSBI members provided the vast majority of the data used to produce the New Atlas. In
short, the BSBI is the society for anyone with a keen interest in wild plants.
Hardy Orchid Society
Website: www.hardyorchidsociety.org.uk
Dedicated to the study of European orchids, including field study, photography and conservation,
with an emphasis on cultivation and propagation. Publishes a quarterly Journal with articles on all
these subjects and organises field meetings and exhibitions of cultivated orchids.
Plantlife International
14 Rollestone Street, Salisbury, Wiltshire SP1 1DX
Tel: 01722 342730. Email: [email protected] Website: www.plantlife.org.uk
The wild plant conservation charity. Plantlife International (formerly ‘Plantlife’) has an increasingly
international focus but still does much useful work in Britain. It has a small number of reserves,
runs ‘back from the brink’ projects for many declining species (including Sword-leaved Helleborine),
organised the ‘county flowers’ poll and promotes various surveys to raise awareness of wild flowers, as
well as gathering much useful information. Publishes a quarterly magazine, Plantlife, and a newsletter.
Wild Flower Society
c/o Mike Hooper, 24 Muirfield Drive, Astley, Manchester, M29 7QJ.
E-mail: [email protected] Website: www.thewildflowersociety.com
Established in 1886 for amateur botanists and wild flower lovers in the UK. Organises meetings
throughout the summer months to see and photograph British wild plants in their natural habitats.
aBBreviaTiOns
BBOWT: Berkshire, Buckinghamshire and LRWT: Leicestershire and Rutland Wildlife
Oxfordshire Wildlife Trust Trust
BC: Butterfly Conservation NE: Natural England
BCNPWT: Bedfordshire, Cambridgeshire, NNR: National Nature Reserve
Northamptonshire and Peterborough NPWS: National Parks and Wildlife Service
Wildlife Trust NT: National Trust
BSBI: Botanical Society of the British Isles RSPB: Royal Society for the Protection of
CCW: Countryside Council for Wales Birds
CEH: Centre for Ecology and Hydrology SNH: Scottish Natural Heritage
FC: Forestry Commission SSSI: Site of Special Scientific Interest
JNCC: Joint Nature Conservation Committee WT: Wildlife Trust (with prefix, e.g. Sussex WT)
LNR: Local Nature Reserve WT: Woodland Trust (without prefix)
006 systematic.indd 423 29/1/09 12:49:22

424 l SITE GUIDE
Southeast England
56 59
54 55 57
58
53
52 60
51
44 50
46 61
43 49 62
45
48
42 47 63
38
41
40
39 64 1 2
66 3
37 65
36
35 8 7 5
33 10 9 4
34 6
24
28 26 21
29 27 25 19 18 12
11
23 22 20 16
30 17 15
13
14
31 32
Kent 18 Ditchling Beacon 36 Ranmore Common, 51 Bernwood Meadows

1 Trosley Country Park West Sussex Denbies Hillside and 52 The Ashridge Estate and
2 Queendown Warren 19 Devil’s Dyke White Downs Ivinghoe Beacon
3 Sandwich and Pegwell 20 Cissbury Ring 37 Ashtead Common Bedfordshire
Bays 21 Furnace Meadow and Berkshire 53 Dunstable and
4 Langdon Cliffs Brick Kiln Rough 38 Hurley Chalk Pit Whipsnade Downs
5 Lydden/Temple Ewell 22 Levin Down 39 Greenham and 54 Totternhoe Knolls
Down 23 Kingley Vale Crookham Commons 55 Sharpenhoe Clappers,
6 Samphire Hoe Hampshire 40 Basildon Park Smithcombe and
7 Park Gate Down 24 Noar Hill 41 Lardon Chase, the Holies Sundon Hills
8 Bonsai Bank 25 Butser Hill and Lough Down 56 Barton Hills
9 Wye 26 Chapetts Copse/Hen Oxfordshire Hertfordshire
10 Marden Meadows Wood 42 Hartslock 57 Hoo Bit
East Sussex 27 Old Winchester Hill 43 Dry Sandfort Pit 58 Hexton Chalk Pit
11 Rye Harbour 28 St Catherine’s Hill 44 Parsonage Moor 59 Therfield Heath and Fox
12 Fore Wood 29 Martin Down 45 Aston Rowant Covert
13 Wild Woods at Friston 30 New Forest 46 Watlington Hill 60 The Meads
Forest The Isle of Wight 47 Warburg Reserve 61 Cheshunt Lock
14 Cuckmere Haven and 31 Tennyson Down Buckinghamshire Greater London
the Seven Sisters 32 Compton Down 48 Homefield Wood. 62 Rammey Marsh
Country Park Surrey 49 Grangelands and Pulpit 63 The Ripple
15 Mount Caburn 33 Thursley Common Hill 64 West Kent Golf Course
16 Malling Down 34 Wallis Wood 50 Dancersend and the 65 Downe Bank
17 Castle Hill 35 Box Hill Crong Meadow 66 Hutchinson’s Bank
006 systematic.indd 424 29/1/09 12:49:23

SOUTHEAST ENGLAND l 425
Close to Dover on the minor road between Lydden

KEnT and Temple Ewell; access is from the south side of the
railway bridge. Park behind the ‘George and Dragon
1. Trosley Country Park TQ 642614 Centre’ in Temple Ewell, from where the reserve is sign-
An area of woodland and chalk downland border- posted. (NNR, Kent WT)
ing the North Downs Way. Part of a much larger
conservation area, there is an interesting chalkland 6. Samphire Hoe TR 298395
flora that includes Bee, Man, Common Fragrant and Lying just west of Dover, this area of grassland
Pyramidal Orchids. bounded by a sea wall was created with waste
The car park is off the A227 between Meopham and material left over from the digging of the Channel
Wrotham, south of Gravesend. (Kent County Council) Tunnel. The spoil was deposited at Lower Shake-
speare Cliff then landscaped and sown with wild-
2. Queendown Warren TQ 827629
flower seeds collected locally. A wide variety of
Excellent species-rich chalk grassland on the site of
plants now grows here, including one of the largest
an old rabbit warren. Specialities include Early Spi-
colonies of Early Spider Orchid in Britain (10,000
der Orchid and Man Orchid (although the former
spikes in 2008).
is declining here); other species include Common
Access by car is via a tunnel through the cliffs. You
Fragrant, Common Spotted, Bee, Pyramidal, Early
can reach the tunnel from the A20; driving towards
Purple, Fly, Lady Orchids and Autumn Lady’s-tresses;
Folkestone from Dover, follow the signs for Samphire
also White and Broad-leaved Helleborines in the
Hoe. (Samphire Hoe is owned and managed by
fringing woodland.
several bodies in partnership with the White Cliffs
Near Hartlip off the A2, southeast of Gillingham. Park
Countryside Project.)
in Warren Lane, off Mount Lane, accessed from the
Lower Hartlip Road. (Kent WT, Plantlife, Swale Bor- 7. Park Gate Down TR 168457
ough Council) A chalk grassland in a dry valley fringed by wood-
3. Sandwich and Pegwell Bays TR 341632 land, this reserve has an exceptionally rich orchid
A large reserve of mixed coastal habitats including flora. It is best known for its colony of Monkey
shingle, rich dune grassland, and coastal scrub. Good Orchids (introduced here from seeds collected
for Southern Marsh Orchid, Pyramidal and Bee Or- elsewhere in Kent). These have increased in recent
chids, Marsh Helleborine and the biggest colony of years and put on a spectacular show. Other spe-
Lizard Orchids in Britain (more than 4,000 were cies include masses of Common Fragrant Orchids
counted in 2001). as well as Early Purple, Bee and Pyramidal Orchids,
Access from Sandwich Town via the Sandwich Bay Es- Common Spotted Orchid, Common Twayblade,
tate, where a substantial toll operates on the private Greater Butterfly Orchid, a few hard-to-find Musk
road.There are many footpaths along the beach, dunes Orchids and, in some seasons, a few Late Spider
and golf courses. Park on sea-front near Prince’s Old Orchids. Fly, Lady and Man Orchids also occur in
Clubhouse at Sandwich Bay. (Kent WT) small numbers.
From Sloane Street (B2068) go through Stelling Min-
4. Langdon Cliffs TR 335422 nis and turn left (i.e. straight on) at the right-angle
Lying to the east of Dover, this is part of the famous bend, then take the next three right turns and carry on
white cliffs. With fantastic views, the chalk downland through the wood; the reserve lies to the left after c.
on top of the cliffs holds many interesting plants 1.5km. (Kent WT)
with Bee, Common Fragrant, Pyramidal and Com-
mon Spotted Orchids, as well as some Early Spider 8. Bonsai Bank TR 105510
Orchids and Autumn Lady’s-tresses. Part of the Denge Wood complex, with Beech and
There are footpaths to the cliffs from Dover town cen- Sweet Chestnut mixed with conifer plantations, this
tre and the port. Signposted from the M20 and Dover wood is named after its rather stunted crop of coni-
town centre. Park past the Castle beside the Visitor fers and is an excellent site for Lady Orchid, as well
Centre. (NT) as Early Purple Orchids.
From the cross roads in Sole Street, follow Penny
5. Lydden/Temple Ewell Down TR 277453
Pot Lane north towards Thruxted. On entering the
One of Kent’s best examples of chalk downland,
woods the car park is signed to the right and the re-
which is of international importance. There is also
serve lies 1/2 mile along the forest track. (Forestry
some woodland and scrub. The speciality of the
Commission)
steep slopes is Early Spider Orchid, and there are
also Common Fragrant, Bee and Pyramidal Orchids, 9. Wye TR 074469
Common Twayblade, Autumn Lady’s-tresses, and a The habitats on this steep chalk downland include
very few Burnt Orchids. grassland, scrub and woodland, and an exceptional
006 systematic.indd 425 29/1/09 12:49:23

426 l SITE GUIDE
19 species of orchid occur here. The speciality is 14. Cuckmere Haven and Seven Sisters
Late Spider Orchid with around six distinct colonies, Country Park TV 519995
but they are hard to find and visitors should con- Lying just east of Seaford, the cliff-top chalk downs
tact the warden for advice. Early Spider Orchid and have Common Fragrant, Common Spotted, Early
Burnt Orchid also occur but are similarly elusive. Purple, Pyramidal and Bee Orchids, Autumn Lady’s-
Not so Common Fragrant Orchid, Common Spot- tresses and, in a few places, Early Spider, Frog and
ted Orchid, Common Twayblade and Pyramidal, Bee Burnt Orchids (including the late-flowering variety
and Man Orchids (the last favouring the western of Burnt).
flank of the escarpment around Dentons Field). The Park in the Seven Sisters Country Park ‘Riverside’ car
woods may yield Bird’s-nest, Lady, Fly and Greater park on the south side of the A259. Paths lead east
Butterfly Orchids. along the cliff tops all the way to Beachy Head. (Sus-
Northeast of Ashford on the minor road between Wye sex Downs Conservation Board)
and Hastingleigh; there are several car parks and
many footpaths. (NNR, NE) 15. Mount Caburn TQ 445090
Just southeast of Lewes, Mount Caburn is an impor-
10. Marden Meadows TQ 763445 tant Bronze Age hill fort; its south-facing chalk slopes
These three unimproved hay meadows have several hold Common Fragrant and Pyramidal Orchids and
pools and are surrounded by ancient hedgerows. a large population of Burnt Orchids (over 6,000
Traditionally managed, there is an amazing display of plants in some years; look in the dry valley to the
Green-winged Orchids in May, when up to 500,000 northwest of the hill fort).
have been counted. Access is from the village of Glynde, following the steep
Just west of Staplehurst off the A229. The meadows public footpath to the fort from roughly opposite the
are sandwiched between the minor road to Marden post office in Ranscombe Lane. (NNR)
and the Tonbridge-Ashford railway line. Park on the
roadside. (Kent WT)
16. Malling Down TQ 423107
Old chalk pits on a steep north-facing slope, with
a rich flora including a selection of orchids. Musk
EaST SuSSEx and Frog Orchids can be found in small numbers,
11. Rye Harbour TQ 925185 with good displays of Pyramidal and Common Fra-
A large coastal site with managed grassland, ditches, grant Orchids and stunning numbers of Common
marsh, scrub and woodland. Botanical interest is fo- Spotted Orchids. There are also Bee Orchids and
cused mainly on the ditches and sandier areas, with Common Twayblades.
orchids such as Common Spotted, Pyramidal and Easily reached by walking from Lewes town centre
Bee. Common Twayblade, Autumn Lady’s-tresses (access in Wheatsheaf Gardens); alternatively, park in
and Green-winged Orchid can also be found. the Ringmer Road lay-by on the B2192. (Sussex WT)
There is parking at Castle Water and a Visitor Centre 17. Castle Hill TQ 371070
southeast of Rye along Harbour Road. (East Sussex East of Brighton, on the South Downs between
County Council) Woodingdean and Kingston, this traditionally manag-
12. Fore Wood TQ 756126 ed chalk grassland has Britain’s largest colony of Early
A hornbeam and hazel coppice with some oak and Spider Orchids, with up to 50,000 plants recorded.
alder. There are wide grassy rides and a good spring Accessible along an un-signed track off the B2123 just
flora. Orchids include colonies of Early Purple and north of Woodingdean (park at TQ 356063 and walk
Common Spotted Orchids, Broad-leaved Hellebo- northeast for 1/2 mile). Access without a permit is
rine, Common Twayblade and Bird’s-nest Orchid. restricted to rights of way. (NNR)
Beside the village of Crowhurst, off the A2100 Battle to 18. Ditchling Beacon TQ 330130
Hastings road. (RSPB) Lying to the north of Brighton on the South Downs
13. Wild Woods at Friston Forest TV 555995 Way long-distance footpath, this Iron Age hill
This area of forestry includes extensive Beech plan- fort has chalk grassland and scrub with Common
tations with many wide grassy rides. A rich flora is Fragrant Orchid, Common Spotted Orchid and
developing, and includes a colony of several thou- Common Twayblade (and also Marsh Fragrant
sand White Helleborines. Orchid on north-facing slopes towards Plumpton).
Park in the Seven Sisters Country Park ‘Forest’ car park The site lies a little off the South Downs Way to the
north of the A259 or off minor roads to the north; northwest of the National Trust car park (accessed
there is a Visitor Centre at the car park, and way- on minor roads south from the B2116 at Ditchling).
marked walks. (FC) (Sussex WT)
006 systematic.indd 426 29/1/09 12:49:23

colony of Musk Orchid. Also Early Purple, Frog,

WEST SuSSEx Bee, Pyramidal, Common Spotted and Common
Fragrant Orchids, plus Common Twayblade and
19. Devil’s Dyke TQ 260110 Autumn Lady’s-tresses.
On the South Downs to the north of Brighton, the Leave Selborne south on the B3006 and immediately
Devil’s Dyke is the largest dry chalk combe in Britain. turn west towards Newton Valence. Then take the first
With some woodland and scrub, the Dyke has a rich left, signposted Noar Hill. There is limited parking by
flora. The list of orchids includes Common Spotted, the footpath sign at SU 738323; follow this track to
Bee and Frog Orchids and Common Twayblade. the reserve. (Hampshire WT)
Well signposted on roads north of Brighton; park in the
large car park on Devil’s Dyke Road. (NT) 25. Butser Hill Su 714198
This is one of the largest areas of chalk grassland in
20. Cissbury Ring TQ 130078
Hampshire. It contains a variety of habitats, includ-
A large and impressive Iron Age hill fort on the
ing Yew woodland and chalk heath. There is a good
South Downs with Pyramidal, Bee and Frog Orchids
orchid flora with Early Purple, Bee and Common
among its varied chalk flora.
Spotted Orchids, and Broad-leaved Helleborine.
Three miles north of Worthing, off the A24 at Findon.
Part of the larger Queen Elizabeth Country Park, south
Footpaths lead from Worthing and Findon. (NT).
of Petersfield and north of Portsmouth on the A3.
21. Furnace Meadow and Brick Kiln Rough There is a Visitor Centre and car parks. (Hampshire
Su 977277 County Council, FC)
A flower-rich meadow on clay soils, with an adjacent
ancient semi-natural woodland. The varied habitats 26. Chapetts Copse/Hen Wood Su 654230
here hold a good orchid flora, with Early Purple, A beechwood, probably a plantation that was
Common Spotted, Greater Butterfly and Bird’s- established on the site of an ancient woodland,
nest Orchids, and Broad-leaved Helleborine among with a very rich orchid flora. The star species is
others. Sword-leaved Helleborine; the largest colony in
Three miles from Petworth on the minor road to Eb- Britain by far is found here, with more than 3,400
ernoe. Park beside the church. The meadow is next to flowering plants. It even grows along the road bank
the Sussex WT reserve of Ebernoe Common. (Sussex outside the wood. Other species include White
WT, Plantlife) Helleborine, and Bird’s-nest, Fly and Common
Spotted Orchids.
22. Levin Down Su 884138 Off the A32 between West Meon and East Meon; turn
A very steep chalk downland with Juniper and south at the sign for Meon Springs Fly Fishery into
patches of acidic chalk heath. Orchids include Pyra- Coombes Lane, and park in the ride on the left after
midal Orchid and Autumn Lady’s-tresses. about 450 yards. (Hampshire WT)
Access on footpaths leading north from the minor road
between Singleton and Charlton, which is reached via 27. Old Winchester Hill Su 645209
the A286. Park in the lay-by at the crossroads in Charl- A spectacular hill fort at the west end of the South
ton. (Sussex WT) Downs, comprising chalk grassland with mixed scrub,
Yew and broad-leaved woodland with 14 species of
23. Kingley Vale Su 824088 orchid. On the south-facing slopes, large colonies of
‘The finest Yew forest in the world’ according to Greater Butterfly, Frog, Common Fragrant Orchids
English Nature. There is much of archaeological in- and Autumn Lady’s-tresses can be found, with Man,
terest too, as well as some scrub and chalk grassland. Bee and Fly Orchids, and Common Twayblade in
Species include Fly, Bee, Frog, Common Fragrant and the scrub. White Helleborine, and Early Purple and
Pyramidal Orchids and Autumn Lady’s-tresses. Bird’s-nest Orchids occur in the woodland.
On the South Downs, 41/2 miles northwest of Chich- From the A32 at Warnford, take the minor road south-
ester. There is a car park and waymarked walks. east towards Clanfield. The car park is on the right
(NNR) after 2 miles. (NNR)
HaMPSHIRE 28. St Catherine’s Hill Su 485275

An ancient hill fort near Winchester city centre, with
24. noar Hill SY 740320 a rich chalk flora including Musk and Frog Orchids,
Medieval chalk pits that now form one of the best and Autumn Lady’s-tresses.
examples of chalk grassland and scrub in Britain. Access off the B3335 south of Winchester centre onto
Rich in flowers and invertebrate life, 11 species of Garnier Road. There is a large car park. (Hampshire
orchid can be found, including a nationally important WT)
006 systematic.indd 427 29/1/09 12:49:24

428 l SITE GUIDE
29. Martin Down Su 061192 32. Compton Down SZ 375853

A vast area of chalk grassland with wonderful dis- The most southerly area of chalk in the country, this
plays of wildflowers, as well as other habitats such faces the sea. In spring there are Early Purple and
as chalk heath and scrub. Various earthworks criss- Green-winged Orchids, and later Pyramidal and Bee
cross the relatively flat downland. Specialities in- Orchids amongst a rich chalk flora.
clude Burnt Orchid, which can be found around the On the south coast of the Isle of Wight, east of Fresh-
prominent Bokerley Dyke. Other species include water Bay and north of the A3055. (NT)
Common Fragrant, Pyramidal, Bee, Early Purple,
Greater Butterfly and Common Spotted Orchids, SuRREY
hard-to-spot Frog Orchids, Southern Marsh Orchid,
and a very few Man Orchids. 33. Thursley Common Su 900417
There is open access on foot from the village of Martin, Thursley is the largest remnant of the Surrey heaths,
south of Salisbury off the A354. (NNR, NE) with bog, dry and wet heath, and woodland. South-
ern Marsh Orchid and the purple-flowered subspe-
30. new Forest Su 298081
cies of Early Marsh Orchid (pulchella) occur on the
The New Forest is one of the greatest biological trea-
wet heath (the creamy-flowered variety ochrantha
sures in Britain. An ancient system of grazing rights,
can also be found at Thursley—not to be confused
still widely exercised, has produced a rich and varied
with the very rare creamy-flowered subspecies
landscape. Much of the forest is heath or wood-
ochroleuca!).
land, with relatively few orchids, but the acid bogs
Access is from south of Godalming alongside the A3;
and valley mires are of exceptional interest. Heath
park between Elstead and Thursley. (NNR)
Spotted Orchid is very common, and the purple-
flowered subspecies pulchella of Early Marsh Orchid 34. Wallis Wood TQ 121388
is fairly frequent. Bog Orchid occurs in the more Oak-hazel coppice with a stream, ponds and pasture.
acidic bogs, such as Holmsley and Wilverley Bogs There is a rich ground flora with Broad-leaved and
(on the northern side of the Avon Water around Violet Helleborines and Common Spotted Orchid.
SU 230000 and SU 245005), the western fringe of On the minor road north of Walliswood village, just
Long Beech Inclosure (north off the A31 at Stoney east of the A29 south of Ockley. (Surrey WT)
Cross), Matley Bog (north of the B3056 21/2 miles
east of Lyndhurst), the area south and southwest of
35. Box Hill TQ 180510
An outstanding area of woodland and chalk down-
Hatchet Pond (on the B3054 11/2 miles southwest
land, Box Hill has long been famous as a destination
of Beaulieu), and at Acres Down. Bog Orchid can
for day-trippers from London. Surprisingly extensive,
be very hard to find, requiring patience and great
there are many walks and views towards the South
care, because it is easy to tread on them; some of
Downs. On the summit there is an Information
these sites are very wet with deep peat – beware!
Centre and a nineteenth-century fort. There is an
Another local speciality is Heath Fragrant Orchid,
impressive array of orchids, with Broad-leaved and
found in a few of the more calcareous mires, such as
White Helleborines, Common Twayblade, Bee, Py-
Stony Moors (north of Holmsley Camp Site), Dib-
ramidal, Common Fragrant and Common Spotted
den Bottom (west of Dibden) and Boundway Hill
Orchids, and Autumn Lady’s-tresses in reasonable
(near Sway). Lesser Butterfly Orchids are relatively
numbers. Harder to find are Green-winged, Man,
frequent on the higher, drier ground in and around
Musk, Early Purple, Fly, Frog and Greater Butterfly
the bogs and on adjacent wet heathland. Finally, the
Orchids, and Violet Helleborine.
New Forest was once home to Summer Lady's-
Box Hill is east of the A24 between Dorking and
tresses, which, although probably extinct in the wild
Leatherhead. There are several car parks around the
since 1952, is reported to have been occasionally
perimeter and also along Zig Zag Road. (NT)
reintroduced to its old sites, secretly.
36. Ranmore Common, Denbies Hillside and
THE ISLE OF WIGHT White Downs TQ 145510
A cluster of National Trust sites on the North
31. Tennyson Down SZ 327854 Downs comprising a wooded common and chalk
On land that drops steeply to the sea to end in downland. White Helleborine, and Bird’s-nest, Fly, Py-
the famous Needles, this site is a combination of ramidal and Bee Orchids occur among the scrub and
coast and downland. The chalk grassland has large trees.There are also some Man Orchids and Autumn
colonies of Early Purple, Bee, Pyramidal and Green- Lady’s-tresses.
winged Orchids. The North Downs Way links Denbies Hillside and the
Take the B3322 out of Freshwater towards the Nee- White Downs, with car parking two miles northwest of
dles, where there is car parking. (NT) Dorking off the A25. (NT)
006 systematic.indd 428 29/1/09 12:49:24

37. ashtead Common TQ 175592 hybrid Monkey x Lady Orchid, as well as Common
A relict woodland pasture with more than 2,300 Spotted, Bee, and Pyramidal Orchids, Common
pollarded oaks. Southern Marsh and Common Twayblade and White Helleborine.
Spotted Orchids, Common Twayblade and Broad- One mile south along the minor road from Goring Station;
leaved and Violet Helleborines can be found here. park by the cart track or in Goring village. (BBOWT)
Near Junction 9 of the M25; access is from Woodfield
43. Dry Sandford Pit Su 467997
Road in Ashtead. Park at Ashtead Station. (NNR)
Based around an old quarry next to Sandford Brook,
with Common Spotted and Early Marsh Orchids,
BERKSHIRE Marsh Helleborine and Common Twayblade.
In Cothill village, 2 miles from Abingdon, off the B4017
38. Hurley Chalk Pit Su 813822
Wootton Road. There is a small car park and marked
A tiny reserve in a disused chalk pit with a surpris- footpaths. (BBOWT)
ingly rich flora amidst its grassland and beechwood.
Common Spotted, Common Fragrant, Pyramidal 44. Parsonage Moor Su 462998
and Bee Orchids are all abundant. There are also Part of the Cothill Fen NNR and one of the best
White Helleborines under the Beeches. fenland sites in the county. Excellent woodland, fen,
South of the B4130 Henley to Maidenhead road. Ac- carr and mire communities with Southern Marsh,
cess from the bridleway south of the Black Boy Public Pugsley’s Marsh and Marsh Fragrant Orchids. Keep
House. (BBOWT) to the footpaths because the fen is deep in places.
Use the same car park as for Dry Sandford Pit (site
39. Greenham and Crookham Commons 43). The footpath to the reserve is opposite the Merry
Su 520643 Miller Restaurant. (BBOWT)
Some Early Purple, Bee, Green-winged, Pyramidal,
and Heath Spotted Orchids, and Autumn Lady’s- 45. aston Rowant Su 740968
tresses can be found at these sites. Also visit nearby Chalk grassland, scrub and beechwoods, now bisect-
New Greenham Park, where some very poor soils ed by the M40, where White, Narrow-lipped and
and the removal of runways from the former airfield Violet Helleborines occur amongst other species.
have created good habitats for plants. Between Stokenchurch and Lewknor. There is a car
Around 3 miles southeast of Newbury; access to the park for the northern section of the reserve signposted
commons is off the A339 through New Greenham off the A40, with a Forestry Commission car park at
Cowleaze Wood serving the southern section. (NNR)
Business Park, or from the minor road between Green-
ham and Thatcham. 46. Watlington Hill Su 702935
Fine chalk grassland and scrub with some stands of
40. Basildon Park Su 611782
Beech and Yew. There are Frog, Bee and Pyramidal
Woodland with Yew and Box, and open grassland. Bee
Orchids at the site.
Orchid, and Narrow-lipped, Broad-leaved and Violet One mile southeast of Watlington, east of the B480.
Helleborines are among the species to search for. There is a small car park. (NT).
Between Pangbourne and Streatley, northwest of Read-
ing, on the west side of the A329. (NT) 47. Warburg Reserve Su 720879
This incredibly rich site contains woodland, scrub and
41. Lardon Chase, the Holies and Lough Down grassland in the dry valley at Bix Bottom. A network
This area of National Trust-owned downland has of footpaths provides access to Broad-leaved, White,
great views over the Thames Valley, where the Violet and Narrow-lipped Helleborines, Early Purple,
river divides the Chilterns from the North Wessex Bird’s-nest and Fly Orchids. Both Greater Butterfly and
Downs.The slopes form one of the largest remaining Lesser Butterfly Orchids are present, along with Com-
areas of chalk grassland in the county and support mon Twayblade, and Bee, Common Spotted and Pyra-
a wide range of invertebrates and flowers, including midal Orchids. BBOWT’s largest reserve is a real gem.
Bee, Common Fragrant and Pyramidal Orchids. Northwest of Henley-on-Thames. Follow signposts for
Just north of Streatley, west of the A417. (NT) Bix Bottom off the A4130 at Bix. The reserve is at the
end of a narrow two-mile road; there is a car park and
OxFORDSHIRE Visitor Centre. (BBOWT)
42. Hartslock Su 616796 BuCKInGHaMSHIRE

An area of rich chalk downland and scrub over-
looking the Thames. The grassland is the sole Ox- 48. Homefield Wood Su 814867
fordshire site for Monkey Orchid, while a handful Although some of the original beechwood has been
of Lady Orchids are also present, together with the cleared and planted with conifers, there is still much
006 systematic.indd 429 29/1/09 12:49:24

430 l SITE GUIDE
mixed beechwood and open chalk grassland at this Just north of Berkhamsted, on both sides of the B4506
site. It is traditionally managed and has a rich flora, between the A41 and the B489. The main car park
with 11 species of orchid including Bird’s-nest, Com- is by the Visitor Centre, just off the B4506 between
mon Spotted, Fly and Bee Orchids and Broad-leaved Berkhamsted and Northchurch. (NT)
and White Helleborines, and it is one of only three
British sites for Military Orchid. BEDFORDSHIRE
Off the A4155 west of Marlow. Turn right before the
Dog and Badger Public House.The reserve is on the left 53. Dunstable and Whipsnade Downs
between Bockmer and Lower Woodend. (BBOWT) TL 000190
The extensive chalk grasslands here have fine views.
49. Grangelands and Pulpit Hill SP 827049 The grazed slopes are plant-rich, and Bee, Pyramidal
These two sites hold a range of habitats, including and Frog Orchids can be found.
chalk grassland, scrub and beechwood, with Bee, Southwest of Dunstable, 4 miles northwest of Ashridge
Common Fragrant, Musk, Frog, and Bird’s-nest Or- between the B4540 and B4541. There are car parks
chids and Violet, White and Narrow-lipped Helle- on the B4541. (NT)
borines.
Northeast of Princes Risborough on the A4010, on 54. Totternhoe Knolls SP 979220
the minor road from Askett to Great Missenden. Park Part old chalk quarry (the Little Hills), part earth-
in the lay-by on the left-hand side of the road. Access works with grass and scrub, and part beech planta-
by the footpath going north. (Grangelands: BBOWT; tion, these varied habitats hold Common Spotted
Pulpit Hill: NT) and Common Fragrant Orchids, Common Tway-
blade and a few Musk Orchids.
50. Dancersend and the Crong Meadow Between Dunstable and Tring on the B489. Park in the
Su 616796 picnic site car park on Castle Hill Road. (BCNPWT)
Woodland and meadows in the Chiltern Hills. The
chalk grassland has been cleared of scrub and has 55. Sharpenhoe Clappers, Smithcombe and
a rich flora, with Common Spotted, Bee, Common Sundon Hills TL 065295
Fragrant and Pyramidal Orchids; Greater Butterfly Sharpenhoe Clappers is the most prominent section
and Fly Orchids can also be found. of chalk escarpment in Bedfordshire.This whole area
Off the B4009 Wendover to Tring road. Turn towards is interesting; there are beechwoods on the hill tops
St Leonards just before the junction with the A41; park with unimproved chalk grassland on both east- and
opposite the pond. (BBOWT) west-facing slopes. White Helleborine and Pyramidal,
Common Fragrant, Bee and Fly Orchids occur.
51. Bernwood Meadows SP 606111 Reached via minor roads 1 mile southwest of Barton-
This traditionally managed old hay meadow has a le-clay. Park in the National Trust car park in Sharpen-
large colony of Green-winged Orchids. hoe village. (NT)
Northeast of Oxford, off the minor road between
56. Barton Hills TL 093295
Stanton St John and Oakley. Car parking is avail-
A large area of chalk grassland, scrub, beechwoods
able beside the reserve on the south side of the road.
and ash-maple woodland in the Chilterns. Many spe-
(BBOWT)
cies of orchid can be found including Bee, Common
52. The ashridge Estate and Ivinghoe Beacon Spotted and Common Fragrant Orchids.
SP 970131 On minor roads immediately south of Barton-le-clay.
Stretching along the northeastern edge of the Footpaths cross the area; park in Barton-le-clay. (NNR,
Chiltern escarpment on the Hertfordshire- NE)
Buckinghamshire border from Berkhamstead to
Ivinghoe Hills, this area is rich in archaeological re- HERTFORDSHIRE
mains. Bronze Age barrows survive around Iving- 57. Hoo Bit TL 117290
hoe Beacon and there is an impressive Bronze Age On the Icknield Way, this reserve has mixed broad-
hill fort at the top. The flora is excellent. There are leaved woods with a good shrub layer and some
woodlands on the chalk scarp with Fly and Bird’s- chalk grassland. Adjacent to both Pegston Hills and
nest Orchids, and Violet and White Helleborines, the Herts and Middlesex WT reserve of Telegraph
wooded commons with Early Purple Orchids, Hill, the whole area has a rich and interesting flora
Green-winged Orchids, and hard-to-find Narrow- which includes Fly, Bee and Common Spotted Or-
lipped and Green-flowered Helleborines, and chalk chids and White Helleborine.
grassland with Bee, Pyramidal, Common Fragrant, Between Luton and Hitchin. Park at Treasure’s Grove
Frog and Common Spotted Orchids. on the minor road off the A505 between Lilley and
006 systematic.indd 430 29/1/09 12:49:25

Hexton, and walk northeast along the Icknield Way.

(BCNPWT)
GREaTER LOnDOn
58. Hexton Chalk Pit TL 107299 62. Lee Valley Park: Rammey Marsh
An old grassed-over chalk quarry with a good chalk TQ 370987
flora, and five species of orchid. The whole area is This area, along the outflow from King George’s
rich in wildflowers, walks and good views. reservoir, has a magnificent display of Bee Orchids
Between Hitchin and Barton-le-Clay at Hexton. Park in in most years – up to 15,000 have been counted.
Hexton off the minor road to Lilley, and walk a short There are also a few Pyramidal Orchids.
distance south to the reserve. (Herts and Middlesex Near Enfield Lock. Park by the Swan and Pike Pool and
WT) walk north towards the M25 along the canal. (Lee
Valley Regional Park Authority)
59. Therfield Heath and Fox Covert
TL 335400 63. The Ripple TQ 467824
Therfield Heath is not a heath but a fine hillside Once a dumping ground for pulverised fuel ash,
of unimproved grassland, chalk downland and this reclaimed industrial wasteland at Barking Reach
hawthorn scrub. Alongside the grassland is a small, now holds a stunning display of hundreds of South-
mature beechwood called Fox Covert. The flora ern Marsh and Common Spotted Orchids. May and
includes Bee and Common Fragrant Orchids, with June are the best months to go.
White Helleborine in the woodland. Entrance to the site is on Thames Road/Renwick Road
At the roundabout on the A505 just west of Royston turn in Barking. (London WT)
south past the ‘Little Chef ’ and then right onto the minor
road towards Therfield. Access is from the small lay-by at 64. West Kent Golf Course TQ 425612
the top of the hill. (Herts and Middlesex WT) This small reserve is close to Down House, which
was once the home of Charles Darwin. The chalk
60. The Meads TQ 348141 grassland, scrub and woodland has a wonderful flora
One of the largest wetlands remaining in Hertford- and fauna with swathes of orchids.
shire, with flood meadows, pools and some chalk Access is via a footpath from the village of Downe,
grassland where Pyramidal Orchid can be found. along West Hill and Milking Lane. (London WT)
Between Hertford and Ware on the A119. Park in
Ware and walk west along the canal towpath to the 65. Downe Bank TQ 438609
reserve. (Herts and Middlesex WT) Lying on the North Downs, this reserve, famous for
its association with Charles Darwin as his ‘Orchis
61. Lee Valley Park Bank’, has mixed woodland, coppice and chalk grass-
This is the best site in the region to see spectacu- land. Many species of orchid occur such as Man, Bee,
lar numbers of Early Marsh Orchids. They thrive on Fly, Common Spotted and Pyramidal Orchids and
waste ash from coal-fired power stations; covering White and Broad-leaved Helleborines.
an area the size of a football pitch, they are well Downe Bank is situated midway between the villages
worth seeking out in June. There are also Southern of Downe and Cudham, just south of Downe village,
Marsh and Common Spotted Orchids. The main ar- where it is best to park. Formerly a closed reserve, the
eas of interest are at Cheshunt Lock (TL 367023) southern half is now open for visitors. Contact the Kent
(an excellent boardwalk through a large colony of WT for access to the northern half. (Kent WT)
Early and Southern Marsh and Common Spotted
Orchids); Bowyers Water (TL 367020) (near Ches- 66. Hutchinson’s Bank
hunt station) with Common Spotted, Early Marsh, TQ 381616 or TQ 386607
Southern Marsh Orchids, and Common Twayblade; One of the largest areas of chalk grassland remain-
and Fishers Green (TL 367032) (close to Lee Valley ing in the Greater London area. The grassy slopes
Information Centre), where small numbers of Bee, hold Pyramidal, Common Spotted and Man Or-
Southern Marsh and Early Marsh Orchids surround chids. Common Twayblade and White Helleborine
a large artificial pond, with Pyramidal Orchids close occur on Chapel Bank, an area of ancient woodland,
to the nearby electricity substation. scrub and grassland.
The main access from the west is from the car park Southwest of New Addington, Croydon. Access is from
close to Cheshunt station; from the east, head to the Featherbed Lane, with Hutchinson’s Bank to the east,
Lee Valley Information Centre at Hooks Marsh (off the via footpaths off Farleigh Dean Crescent (TQ 381616)
B194 at Holyfield). The Centre can provide detailed and Chapel Bank to the south and west (TQ 386607).
maps and directions to all these sites and informa- (London WT)
tion about orchid flowering times. (Lee Valley Regional
Park Authority)
006 systematic.indd 431 29/1/09 12:49:25

432 l SITE GUIDE
Southwest England and Channel Islands
48
51
47 49 50
46
45
44
43 26
38 40
39
36 35 41
37 42
30 31 34 25 22
32 21
6 33 23
29 24
27 19
28 18
17 20
5
10 16
15
4
12 14
9
13
3
11
8
7
2
53
52
1
Cornwall 20 Pepperbox Hill 41 Brown’s Folly

1 The Lizard Peninsula 21 Ham Hill 42 Folly Farm
2 Rame Head and Penlee Point 22 Jones’s Mill Gloucestershire
3 Sylvia’s Meadow (St Ann’s Chapel) 23 Pewsey Downs 43 Lower Woods
4 Creddacott Meadows 24 Parsonage Down 44 Minchinhampton and
Devon 25 Marlborough Downs Rodborough Commons
5 Dunsdon 26 Clattinger Farm Reserve 45 Strawberry Banks
6 Braunton Burrows Somerset 46 Elliot (Swift’s Hill)
7 Andrew’s Wood 27 Long Sutton Plantations, 47 Plump Hill Dolomite Quarry
8 Berry Head - Sharkham Point 28 Barrington Hill Meadows 48 Betty Daw’s Wood
9 Dawlish Warren 29 West Sedgemoor 49 Painswick Hill
Dorset 30 Porlock Bay 50 Cotswold commons and
10 Kingcombe Meadows 31 Berrow Dunes beechwoods
11 Isle of Portland 32 Ash Priors Common 51 Greystones Farm and
12 Tadnoll and Winfrith Heath 33 Great Breach Wood Salmonsbury Meadows
13 Durlston Country Park 34 Cheddar Gorge and Black Rock Channel Islands
14 Ballard Down and Nine Nature Reserve 52 Le Noir Pré, Jersey
Barrow Down ‘avon’ 53 The ‘Orchid Fields’ at Les Vicheries,
15 Corfe Mullen Meadow 35 Blagdon Lake Pumping Station Guernsey
16 Badbury Rings 36 Hellenge Hill
17 Hambledon Hill 37 Walborough
18 Fontmell and Melbury Downs 38 Netcott’s Meadow
Wiltshire 39 Ashton Court Meadow
19 West Wiltshire Downs 40 Avon Gorge/Leigh Woods
006 systematic.indd 432 29/1/09 12:49:25

SOUTHWEST ENGLAND l 433
Devon. There are good displays of Lesser Butterfly,

CORnWaLL Heath Spotted and Southern Marsh Orchids.
1. The Lizard Peninsula Between Holsworthy and Bude, off the A3072 to Pan-
SW 687133 (Kynance) crasweek. The entrance to the reserve and car park is
SW 667163 (Predannack) just before Gains Cross. (Devon WT)
The Lizard is the most southerly point of the Brit- 6. Braunton Burrows SS 464326
ish Isles and has many rare native plants. Orchids, The largest sand dune system in the UK, with a very
however, are of interest more from a local view- diverse plant community. There are vast numbers
point than a national one. Around 10 species occur, of Pyramidal Orchids, and also Southern Marsh
including Green-winged and Early Marsh Orchids, Orchids and large colonies of Marsh Helleborines
which occur virtually nowhere else in Cornwall. in the dune slacks. Recently named the UK’s first
Heath Fragrant Orchid can be locally common on Biosphere Reserve.
the Lizard’s heaths; others present include Common On the north bank of the Taw/Torridge estuary,
Twayblade and Autumn Lady’s-tresses. west of Barnstaple. There are two car parks with
Many orchids can be seen from the coastal footpath information.
between the car parks at Kynance and Predannack
Wollas. (NNR, NT) 7. andrew’s Wood Sx 707515
A fascinating reserve of ancient meadows and
2. Rame Head and Penlee Battery Sx 436492 woods, over an old field system. The flora benefits
An area of coastal grassland with extensive gorse from traditional management; there are Southern
scrub. An interesting area, it holds Cornwall’s main Marsh and Heath Spotted Orchids, with Broad-
population of Bee Orchid and is the only known site leaved Helleborine and Common Twayblade in the
in southeast Cornwall for Pyramidal Orchid. Early woodland.
Purple and Southern Marsh Orchids can be found Close to Loddiswell north of Kingsbridge, there is a
along the footpaths, as well as a few Autumn Lady’s- track to the reserve from the car park just beyond
tresses. Coldharbour Cross. (Devon WT)
Parking is available at Penlee Battery Cornwall WT re-
serve and at Rame Head, from which coastal paths 8. Berry Head - Sharkham Point Sx 945567
are easily accessed. A large coastal area comprising two headlands
and a smaller promontory, Durl Head. There is an
3. Sylvia’s Meadow (St ann’s Chapel) important flora on the cliff-tops, and in the grassland
Sx 413707 and scrub areas, with eight species of orchid. These
This small reserve of unimproved herb-rich grass- include Early Purple and Green-winged Orchids,
land is one of Cornwall’s best single sites for orchids, Common Twayblade, Bee, Common Spotted and
with eight species recorded. Heath Spotted, Lesser Pyramidal Orchids, and Autumn Lady’s-tresses.
Butterfly and Early Purple Orchids can be abundant, The site lies at the southern end of Torbay past Brix-
and there may be a few spikes of Greater Butterfly ham. (Berry Head Country Park, NNR)
Orchid. Southern Marsh Orchids and their hybrids
with Heath Spotted Orchid also occur. Common 9. Dawlish Warren Sx 983787
Spotted Orchid is a rarity in Cornwall but a few can A range of coastal habitats around a double sand
spit that extends across the mouth of the River Exe.
be found here, along with Autumn Lady’s-tresses
The dune grassland is of great botanical interest and
and Broad-leaved Helleborine.
has Autumn Lady’s-tresses in good numbers.
South of Kit Hill and west of Gunnislake, with parking
Off the A379; the car park is through the entrance
available at Honicombe. A public footpath runs south
tunnel under the railway in Dawlish Warren village.
from the A390 alongside the reserve. (Cornwall WT).
(Devon WT, NNR)
4. Creddacott Meadows Sx 234963
One of the last remnants of species-rich damp culm DORSET
grassland in Cornwall. Lesser Butterfly Orchid and
Heath Spotted Orchid occur, among others. 10. Kingcombe Meadows SY 555992
Park off the B3254 Launceston to Bude road, at Week A large area of species-rich old grassland, hedge-
St Mary. (Cornwall WT, Plantlife) rows, woodland and common land – a real ancient
landscape. Heath Spotted, Early Purple and Bee Or-
DEVOn chids are abundant.
The site is at Lower Kingcombe near Toller Porcorum.
5. Dunsdon SS 302080 There are small car parks, footpaths and a Visitor Cen-
One of the best examples of culm grassland in tre at Pound Cottage, off the B356. (Dorset WT)
006 systematic.indd 433 29/1/09 12:49:26

434 l SITE GUIDE
11. Isle of Portland SY 685700 Access is from the car park on the north side of the
Famous for its limestone quarries, there is still much B3082 Wimborne to Blandford road. (NT)
of interest on the cliff-tops and in the abandoned
quarry workings. Two are worthy of visits to search 17. Hambledon Hill ST 845125
for orchids:The Broadcroft Quarry Reserve and Tout Dramatic chalk grassland with an impressive Iron
Quarry. There is also a large car park at Portland Age earthwork. A rich chalk flora includes Early
Bill, and a variety of orchids to be found in the sur- Purple and Pyramidal Orchids.
rounding grasslands. Bee and Pyramidal Orchids and Northwest of Blandford Forum off the A350, there is
Autumn Lady’s-tresses are particularly abundant. limited parking off Duck Street and Shaftsbury Road
Access to the Isle of Portland is on the A354 from near Child Okeford. Plenty of footpaths give good ac-
Weymouth. Carry on to Southwell and the large car cess to the site. (NNR)
park at the lighthouse for the Bill. (BC)
18. Fontmell and Melbury Downs
12. Tadnoll and Winfrith Heath SY 795876 ST 884176, ST 885187
A large example of Dorset heath, bogs and mead- An outstanding area of chalk downland, scrub and
ows, part of Thomas Hardy’s ‘Egdon Heath’, with woodland, which includes Melbury Beacon. Autumn
typical flora and fauna. Heath Spotted and Southern Lady’s-tresses and Green-winged, Common Fra-
Marsh Orchids occur, along with the hard-to-find grant, Greater Butterfly, Pyramidal, Bee, Early Purple
Bog Orchid, and Autumn Lady’s-tresses. and a few Frog Orchids can all be found there
From the A352, the Tadnoll entrance is along Red- Access is via footpaths from the National Trust view-
bridge Road, while the Winfrith Heath entrance is on point car park at the top of Spread Eagle Hill. (NT,
Gatemore Road (with limited parking). (Dorset WT). Dorset WT)
13. Durlston Country Park SY 032773
Limestone grassland occurs along the south coast WILTSHIRE
of Dorset and its speciality, Early Spider Orchid, can
be found anywhere from Durlston Head west to 19. West Wiltshire Downs
Worth Matravers. Other species in the area include A wonderful landscape of natural and archaeologi-
Green-winged, Early Purple and Bee Orchids. cal features, southwest of Salisbury. There are many
Durlston Country Park lies on a minor road south from places to visit – here is a selection.
Swanage. Coombe Bisset Down (Su 111256) Rich chalk grass-
land with scrub and beechwoods. There are many
14. Ballard Down and nine Barrow Down
Bee Orchids, with the very local Burnt Orchid.
SY 050820
Clearbury Rings (Su 152245), with over 1,000
On the Isle of Purbeck chalk ridge, these downs
Burnt Orchids on the gentle, south-facing slopes of
contain steep slopes and cliff-tops, with Common
Clearbury Down.
Fragrant and Early Purple Orchids present.
Middleton Down (Su 043252), with Early Purple
Between Ballard Head and Old Harry Rocks, the
Orchids and Autumn Lady’s-tresses, among others.
downs are easily accessible on foot from car parks at
Studland (NT) and Swanage. 20. Pepperbox Hill Su 212248
15. Corfe Mullen Meadow SY 980967 Rough chalk grassland and mixed scrub with a good
A small but flower-rich meadow with a stunning selection of orchid species. Common Spotted, Frog,
colony of Green-winged Orchids. Common Fragrant and Pyramidal Orchids, White
Northwest of Poole on the B3074 at Corfe Mullen, Helleborine, Common Twayblade and Autumn
with roadside parking. (Dorset WT) Lady’s-tresses all occur.
Off the A36 southeast of Salisbury, with parking on the
16. Badbury Rings ST 964029 minor road between West Grinstead and West Dean.
An Iron Age hill fort on the Kingston Lacy Estate. (NT)
A wonderful site with a rich downland flora and a
great variety of orchid species. On the hill fort it- 21. Ham Hill Su 334616
self Frog Orchid is a speciality; other species include A small, very steep chalk downland reserve with
Common Fragrant and Common Spotted Orchids, abundant wildlife and good views. The flora includes
Common Twayblade, Early Purple, Green-winged, Common Twayblade and Common Fragrant Orchid,
Pyramidal and Bee Orchids, Greater Butterfly Or- and also a few Musk Orchids.
chid and Autumn Lady’s-tresses, with White Helle- Off the A336 Burbage to Hungerford road, between
borine and Bird’s-nest Orchid under the roadside Shalbourne and Ham on the road to Buttermere, with
Beech trees. parking on the left-hand verge. (Wiltshire WT)
006 systematic.indd 434 29/1/09 12:49:26

22. Jones’s Mill Su 168613

Wet woodland and fen meadows with Southern
SOMERSET
Marsh and Common Spotted Orchids.
Just north of Pewsey off the B3087 to Burbage; the 27. Long Sutton Plantations, Burnt House
reserve entrance is close to a small lay-by, just over the
and Monday’s Court Lane ST 461269
This is a wonderful mix of lanes, plantations and beech
railway bridge. Alternatively, walk from Pewsey along
copses, with White Helleborine, Greater Butterfly
the Kennet & Avon towpath. (Wiltshire WT)
Orchid and Common Twayblade in the woods and
23. Pewsey Downs Su 120640 Bee and Pyramidal Orchids in the open areas.
Perhaps the finest site on the Marlborough Downs, Northwest of Yeovil, off the B3165 between Long Sut-
comprising Milk Hill, White Horse Hill, Walkers Hill ton and Mortock. Park on the road verge before the
and Knapp Hill, with some of the best habitats the railway bridge.
region can offer. Common Spotted, Burnt, Bee, Frog, 28. Barrington Hill Meadows ST 290169
Common Fragrant, Lesser Butterfly, Green-winged Several species-rich unimproved meadows on the
and Pyramidal Orchids and Autumn Lady’s-tresses southern edge of the Blackdown Hills, with old
all occur here. hedges and ponds.There is a large colony of Green-
In the Vale of Pewsey between Devizes and Pewsey, winged Orchids.
southwest of Marlborough. Access is via the unclassi- Northwest of Ilminster on the minor road off the A358
fied road between the A361 and A345. Car parking is between Bickenhall and Broadway. Park beside the
at Knapp Hill. (NNR) road along Folly Drive. (NNR)
24. Parsonage Down Su 055415 29. West Sedgemoor ST 365255
Some wonderful downland and pastures with a Woodland and traditionally managed grazing mead-
history of traditional grazing management. There ows, often flooded in winter. The wet meadows
are many interesting species here; orchids include contain a wide variety of flowers; one notable field,
Green-winged, Frog and perhaps as many as 30,000 slightly higher than the rest, holds cowslips and
Burnt Orchids spread over 95 hectares, forming the Green-winged Orchids.
largest colony in Northwest Europe. East of Taunton on the A378 Langport road. Car park-
The site is at Cherry Lodge Farm, Shrewton, on the ing is east of Fivehead. (RSPB)
south edge of Salisbury Plain, between Shrewton and
Winterbourne Stoke on the B2083. (NNR)
30. Porlock Bay SS 885482
A fine area of species-rich dunes and foreshore that
25. Marlborough Downs is particularly good for orchids, with a large colony
An impressive area surrounding Marlborough, most- of Autumn Lady’s-tresses.
ly consisting of gently undulating chalk grassland.Tra- Porlock is west of Minehead on the A39. Minor roads
ditional grazing has ensured a rich flora and fauna, lead to the coast and the coastal footpath.
with many areas now designated nature reserves. 31. Berrow Dunes ST 299515
Here are two great places to visit. An area of reedbeds and dunes beside Berrow Golf
Morgan’s Hill (Su 025672) and Kingsplay Hill (Su Course. The dune slacks hold Southern Marsh Or-
006658) On the western edge of Marlborough chids and Marsh Helleborines in good numbers, and
downs with chalk grassland and scrub; both have a Heath Spotted and Early Marsh Orchids also occur,
good orchid flora. with Lizard and Bee Orchids (including var. belgarum
Cherhill Down and Oldbury Castle (Su 046694) of Bee) on the drier areas..
The chalk grassland on the southern slopes has a Just north of Burnham-on-Sea. Access is on waymarked
rich flora, including Lesser Butterfly Orchid. (NT) trails from the car park on Coast Road, Berrow (op-
posite Sandy Glade Caravan Park). (LNR, Sedgemoor
26. Clattinger Farm Reserve Su 017937
District Council)
This site consists almost wholly of unimproved
species-rich alluvial grasslands, with spectacular dis- 32. ash Priors Common ST 151289
plays of hay-meadow flowers. Green-winged Orchid, Scrub, grassland, woodland and heath habitats, with
Common Twayblade, Common and Heath Spotted a good variety of orchids: Heath Spotted, Common
Orchids, Early and Southern Marsh Orchids and Spotted, Southern Marsh and Early Purple Orchids,
Burnt Orchid (outside its normal downland habitat) and Common Twayblade all occur.
are present. Northwest of Taunton off the A358 at Ash Priors. Car
Off the B4040 between Malmesbury and Cirencester, parking is available beside the common. (Taunton and
with parking on the roadside. (Wiltshire WT) District Borough Council)
006 systematic.indd 435 29/1/09 12:49:26

436 l SITE GUIDE
33. Great Breach Wood ST 506325 limestone only 2 miles from the centre of Bristol.
One of Somerset’s largest reserves, with mixed Clifton and Durdham Downs are large areas of
woodland and grassland. In some areas orchids are limestone grassland with abundant Bee and Com-
plentiful; there are Early Purple, Greater Butterfly mon Spotted Orchids. There are smaller numbers
and Pyramidal Orchids on the steep south- and of Pyramidal and Green-winged Orchids. On the
west-facing slopes, for example. There are many west side of the gorge Leigh Woods (ST 555730)
rides and paths through the site. hold Bird’s-nest and Fly Orchids, and Broad-leaved
South of Glastonbury, between Street and Somerton. Helleborines.
Park near Butleigh Park on the minor road from the The area is easily accessed from Bristol on the A4018
B3153. (Somerset WT) and the A4. Leigh Woods are on the A369 Portishead
to Bristol road.
34. Cheddar Gorge and Black Rock
ST 482545 41. Brown’s Folly ST 795665
Limestone grassland, plantation and woodland at Calcareous grassland and mature secondary wood-
the head of Cheddar Gorge. Part of the Cheddar land with some ancient semi-natural areas. Broad-
complex, the area is botanically rich; orchids to be leaved Helleborine together with Early Purple, Bee,
found include Broad-leaved and Narrow-lipped Pyramidal and Common Spotted Orchids can be
Helleborines. found on and around the grassland. Elsewhere,
Park at Black Rock Gate, on the B3135 just northwest Bird’s-nest Orchid and White Helleborine occur
of Cheddar. (Somerset WT) within the wood.
East of Bath, off the A36 at Bathampton opposite the
aVOn golf course. (Avon WT)
42. Folly Farm ST 607605
35. Blagdon Lake Pumping Station ST 503598 Species-rich neutral grassland and ancient semi-
An excellent if unusual site for Green-winged natural woodland. There are Common Spotted and
Orchids. Heath Spotted Orchids, as well as hybrids between
Bristol Water have many open days for public access. the two. Common Twayblades are also present
Contact them for details of orchid flowering times. within the wood, with Early Purple Orchids and Vio-
(Bristol Water. Tel: 0117 966 5881) let Helleborines.
36. Hellenge Hill ST 345572 East of Bishop Sutton, south of Bristol; the site is sign-
Calcareous grassland and scrub on the south side posted off the A368, to the east of its junction with the
of the Mendips with great views. Autumn Lady’s- A37. (Avon WT)
tresses are abundant in some areas.
At Bleadon, south of Weston-Super-Mare. (Avon WT) GLOuCESTERSHIRE
37. Walborough ST 316579 43. Lower Woods ST 743876
An area of limestone grassland beside the sea wall One of England’s largest stands of ancient oak-ash
at Uphill. There are Green-winged and Early Purple woodland. It contains a diversity of habitats, includ-
Orchids and Autumn Lady’s-tresses. ing species-rich grassland, coppiced woodland and
South of Uphill village. Park by the sluice gates near the wooded common. Early Purple, Greater Butterfly
beach and walk along the sea wall. (Avon WT) and Common Spotted Orchids and Violet Hellebo-
38. netcott’s Meadow ST 476696 rine can all be found.
An area of semi-improved neutral, damp grassland Between Hawksbury and Wickwar. Park by the
with Green-winged, Early Purple, Southern Marsh Lodge on the minor road off the B4509 at Wickwar.
and Bee Orchids present in large numbers. (Gloucestershire WT)
Close to Nailsea, Bristol, with parking after the railway 44. Minchinhampton Common SO 850010
bridge. (Avon WT) and Rodborough Common SO 850038
Some of the best areas of limestone grassland on
39. ashton Court Meadow ST 545720
the edge of the Cotswolds. There are great views,
Part of the Ashton Court Estate and close to the
and Minchinhampton has much of archaeological
centre of Bristol. In May the meadow is full of
interest. Many species of orchid have been re-
Green-winged Orchids.
corded among the varied flora.There are Common
Off the A369 Bristol to Portishead road. Park in the lay-
Spotted, Pyramidal, Early Purple, Green-winged,
by on Beggar Bush Lane, the B3129. (Avon WT)
Common Fragrant, Bee, Frog and Greater But-
40. avon Gorge ST 553731 terfly Orchids, as well as Autumn Lady’s-tresses,
A fantastic area, the Avon gorge cuts through the Common Twayblade and White, Broad-leaved and
006 systematic.indd 436 29/1/09 12:49:27

Narrow-lipped Helleborines. Both commons are Broad-leaved, White, Narrow-lipped, and Green-
worth exploring. flowered Helleborines.
Off the A46 between Stroud and Nailsworth. (NT) Access to Buckholt Wood at various points from minor
roads east off the A46, 3 miles northeast of Painswick.
45. Strawberry Banks SO 910033 (NNR)
This limestone grassland on a west-facing slope has
a rich flora. There is also some mixed scrub and a 51. Greystones Farm and Salmonsbury
small stream. Greater Butterfly, Bee, Common Spot- Meadows SP 173209
ted, Green-winged, Early Purple and Pyramidal Or- With traditional hay meadows, grazing land and
chids all occur. It is reached by walking through Three streams, this is an attractive place to visit. Early
Groves Wood Nature Reserve, where Broad-leaved Marsh and Southern Marsh Orchids are abundant
Helleborine can be found. in the meadows.
East of Chalford between France Lynch and Oakridge vil- Close to Bourton-on-the-Water; park and follow the
lages.There is a small car park. (Gloucestershire WT) Oxfordshire Way, off Moor Lane. (Gloucestershire
WT)
46. Elliot (Swift’s Hill) SO 877067
This area of limestone grassland and old quarry THE CHannEL ISLanDS
workings is carefully managed by grazing, and 11
species of orchid regularly occur. These include 52. Le noir Pré, Jersey
Pyramidal, Common Fragrant, Early Purple and Fly Two adjoining wet meadows, the speciality here is
Orchids, and Autumn Lady’s-tresses. Loose-flowered Orchid. There are also Southern
Just northeast of Stroud, on the minor road to Elcombe. Marsh, Heath Spotted and Common Spotted Or-
(Gloucestershire WT) chids.
The entrance is at Le Chemin de L’Ouziere, a minor
47. Plump Hill Dolomite Quarry SO 661171
road off Le Grand Route de Mielle (Five Mile Road).
An interesting site with steep exposed rock faces
There is an open day in late May, and unrestricted
and a limestone flora on the quarry floor. There is a
access for several weeks in the summer. (National
good colony of Autumn Lady’s-tresses.
Trust for Jersey)
South of Mitcheldean beside the A4136. Park in the
lay-by. (Gloucestershire WT) 53. The ‘Orchid Fields’ at Les Vicheries,
Guernsey
48. Betty Daw’s Wood SO 696284 An area of low-lying wet meadows in southwest
An ancient Sessile Oak woodland, part of a larger Guernsey. The rich flora includes Loose-flowered
complex, with a good ground flora that includes Orchid as well as Heath Spotted, Common Spotted
Bird’s-nest Orchid and White Helleborine. and Southern Marsh Orchids.
West of Gloucester, northwest of Newent, at Four In the St Peter in the Wood region. The reserve lies
Oaks. Park and walk from the village. (Gloucester- close to the main coast road along the Rue du Douit
shire WT) du Moulin and the Rue des Vicheries, where there is a
49. Painswick Hill SO 869120 La Société Guernesiaise information board.
Excellent chalk grassland on a steep-sided hill fort,
where Bee, Fly, Musk, Frog and Greater Butterfly
Orchids, and Autumn Lady’s-tresses can be found.
South of Gloucester on the B4073 to Painswick. There
are several footpaths and two long-distance paths, the
Wysis Way and the Cotswold Way, meet here.
50. Cotswold commons and beechwoods
The finest Cotswold beechwoods occur on the
scarp and dip slopes between Gloucester and
Stroud around Cranham, Sheepscombe and Pains-
wick. The whole area contains important examples
of ancient woods, grassland and streams and there
are many designated nature reserves. Buckholt
Wood (SO 894131) is one of the finest with an
interesting shrub layer and good ground flora. Or-
chids typical of beech woodland occur here, such
as Bird’s-nest and Greater Butterfly Orchids and
006 systematic.indd 437 29/1/09 12:49:27

438 l SITE GUIDE
East anglia
12 13
10 14
11
15 16
17
8
18
23 19
7
22 21 20
6
5 25
3 4 24
1 26
2 27
28
29
30
31
32
Cambridgeshire Suffolk
1 Hayley Wood 19 Reydon Wood
2 Thriplow Meadows 20 Wink’s Meadow
3 Beechwood 21 Redgrave and Lopham Fens
4 Fulbourn Educational Reserve 22 Market Weston Fen
5 Devil’s Dyke 23 Rex Graham Reserve
6 Wicken Fen 24 Bradfield Woods
7 Soham Meadows 25 Chippenhall Green
8 Upwood Meadows 26 Wolves Wood
norfolk 27 Groton Wood
9 Narborough Railway Line Essex
10 Holme Dunes 28 West Wood
11 Holkham 29 Hatfield Forest
12 Wells Woods 30 Roding Valley Meadows
13 Beeston Regis and Sheringham Commons 31 Langdon
14 Overstrand 32 Grays Chalk Quarry
15 Foxley Wood
16 Buxton Heath
17 Thompson Common
18 New Buckenham Common
006 systematic.indd 438 29/1/09 12:49:27

EAST ANGLIA l 439
in plant life with Southern Marsh and Early Marsh

CaMBRIDGESHIRE Orchids (in a variety of colour forms).
1. Hayley Wood TL 294534 Signposted from Wicken village, on the A113 northeast
An ancient woodland with a long recorded history of Cambridge. With nature trails and a Visitor Centre.
of management.There are fine displays of woodland (NNR, NT)
flowers in spring, with Early Purple, Common Spot-
ted and Bird’s-nest Orchids.
7. Soham Meadows TL 608722
An area of traditionally managed unimproved
On the B1046 between Cambridge and Sandy, about
grassland with Bee and Common Spotted Orchids,
3 miles from Gamlingay. Park on the road verge.
(BCNPWT) Common Twayblade and a small number of typically
elusive Frog Orchids.
2. Thriplow Meadows TL 445470 Alongside the A142 Soham bypass, with parking in the
A small area of traditionally managed unimproved lay-by. (BCNPWT)
grassland. Bee Orchids are among the rich flora on
the dry ground, with good populations of Early and 8. upwood Meadows TL 251825
Southern Marsh Orchids in the wetter areas. A permanent pasture on calcareous boulder clay.
Parking is by the Village Hall in Thriplow, south of Cam- The ridge and furrow landscape indicates an an-
bridge off the B1368. A footpath leads to the reserve cient arable history, but the meadows have been
from the road junction. (BCNPWT) grassed and traditionally managed for at least 300
years. They have a rich flora with large numbers of
3. Beechwood LnR TL 486548 Green-winged Orchids in May.
A beech plantation on thin chalky soils, with a fine Two and a half miles southwest of Ramsey, which is
colony of White Helleborines (3,800 were counted on the B1040. Go down Meadow Road and park in
in 1998) and a few Common Twayblades. Green- Upwood. (BCNPWT)
flowered Helleborine has been recorded here.
From the roundabout on the A1307 southeast of Cam- nORFOLK
bridge turn northeast onto the minor road towards
Cherry Hinton; then turn right towards Fulbourn. Park 9. narborough Railway Line TF 750118
in the lay-by opposite the wood. (BCNPWT) This sunny railway embankment was built with chalk
4. Fulbourn Educational Reserve TL 528560 ballast, and now supports a rich chalk flora. Com-
Ancient wet meadows with alder woods and scrub; mon Twayblade, and Southern Marsh and Common
Common Twayblade, Early Marsh and Southern Spotted Orchids occur along the path, with Marsh
Marsh, Common Spotted and Bee Orchids can be Helleborines in the pits on the sides.
found in the rough pasture. Take the A47 from Narborough towards Swaffham;
On the minor road southeast out of Fulbourn leading there is a small car park beside the railway line. (Nor-
to Balsham. Park on the roadside. (BCNPWT). folk WT)
5. Devil’s Dyke TL 616616 10. Holme Dunes TF 697438

This massive 8-mile long earthwork, a ditch and This site includes a range of coastal habitats, with
rampart about 60 feet high, is thought to have been good dune and saltmarsh communities. Marsh Hel-
built in the sixth century. It bisects Newmarket’s July leborine, Early Marsh Orchid (including the red-
race course and continues towards Reach. It sup- flowered coccinea), and Southern Marsh Orchid are
ports important areas of chalk grassland notable for common.
colonies of Lizard Orchids (especially to the south- Access is from the A149 east of Hunstanton, through
east of the A45). Other species include Pyramidal Holme village. There are car parks at the beach and
and Common Spotted Orchids. at the reserve centre one mile further east. (NNR,
A footpath runs along the dyke from the roundabout at Norfolk WT)
the junction of the A1303 and A1304, 2 miles south-
11. Holkham TF 892441
west of Newmarket. The Dyke can also be accessed
The westward continuation of Wells Woods (site
from the B1102 between Swaffham Prior and Bur-
12), but with more extensive dune slacks on the
well.
seaward side of the pines. Marsh Helleborine and
6. Wicken Fen TL 554702 Southern Marsh Orchid can be abundant, with scat-
One of Britain’s oldest nature reserves, Wicken Fen tered Common Spotted, Pyramidal and Bee Or-
lies nine miles northeast of Cambridge, and contains chids. Creeping Lady’s-tresses occurs very locally in
reedbeds, sedge-fen, ditches, open water, peat dig- the pinewoods.
gings, and birch and sallow carr. The open fen is rich Access is from Lady Anne’s Drive (north off the A149,
006 systematic.indd 439 29/1/09 12:49:28

440 l SITE GUIDE
opposite the entrance to Holkham Hall); the areas to Just east of the B1149 Norwich to Holt Road north-
the west of the car park, including the dunes past the west of Heavingham.There is a small car park.
west end of the pines, are best. (NNR)
17. Thompson Common TL 934967
12. Wells Woods TF 913454 A fascinating place to visit, with open water (Thomp-
These coastal dunes are planted with Corsican son Water), carr, grassland, scrub and woodland as
Pines, but contain some relict areas of dune slacks, well as a number of pingos (shallow ponds formed
especially at ‘The Dell’, where Southern Marsh and by glacial activity). Early Marsh (subspecies incarna-
Common Spotted Orchids and Marsh Helleborine ta) and Southern Marsh Orchids can be found, and
are abundant, with a few Common Twayblades and possibly Pugsley’s Marsh Orchid.
Bee Orchids as well. Creeping Lady’s-tresses occurs Accessed from Watton on the A1075 to Thetford. Car
very locally in the pinewoods. parking is available just before Stow Bedon at the
Access is from Wells beach car park (signposted from Great Eastern Pingo Trail car park. (Norfolk WT)
the town centre);The Dell lies to the north of the path
450 yards west of the kissing gate.
18. new Buckenham Common TM 090906
This traditionally managed common has never been
13. Beeston Regis and Sheringham Commons ploughed. The site’s speciality is a large colony of
TG 166425 Green-winged Orchids.
A small but very diverse site with rough grass- Four miles south of Attleborough on the B1113 at New
land, alkaline mires, acid heath and a little wood- Buckenham, with parking available beside the com-
land. Marsh Helleborine and Common Spotted mon. (Norfolk WT)
and Marsh Fragrant Orchids are abundant, as are
a confusing mixture of marsh orchids (many Pug- SuFFOLK
sley’s Marsh Orchids and intermediates, plus a few
‘pure’ Southern Marsh). Common Twayblade and There are many well-managed Roadside Nature Re-
Bee, Pyramidal and Heath Spotted Orchids occur serves in Suffolk that hold some interesting and local
in small numbers; Early Purple and Lesser Butterfly orchids; look out for the wooden marker posts. In par-
Orchids are also present but hard to find. ticular, the Claydon and A140 roundabouts along the
Access is from the lay-by on the south side of the A149, A14 hold huge colonies of Pyramidal Orchids.
1 mile east of Sheringham.
19. Reydon Wood TM 480788
14. Overstrand TG 246411 This small ancient wood is managed as a ‘Commu-
The slumped clay cliffs at Overstrand have many nity Wood’ by local people. There is hazel coppice,
flushes and seepages, and hold large numbers of the ancient Ash stools, and many rides. Several orchid
delightful red coccinea form of Early Marsh Orchid, species occur here, including Early Purple, Com-
as well as Southern Marsh, Common Spotted and mon Spotted and Bird’s-nest Orchids and Common
Bee Orchids and Marsh Helleborines. Twayblade.
Access is from the cliff-top car park at Overstrand, on On the B1126 Southwold to Wangford road. Park at
the coast east of Cromer. Take the path to the beach the end of Wood Lane. (Suffolk WT)
and walk north, scrambling up the slopes from time to
20. Wink’s Meadow TM 303799
time (it can be very slippery).
This is a traditionally managed meadow with an excel-
15. Foxley Wood TF 049229 lent flora. Species to be found include Common Tway-
The largest area of ancient woodland in Norfolk, blade, Common Spotted, Green-winged, Early Purple,
with wide, grassy, flower-rich rides. Early Purple, Pyramidal and Bee Orchids, and the only colony of
Common Spotted and a very few Greater Butterfly Frog Orchids in Suffolk (they are hard to find).
Orchids can be seen here. Accessed via Metfield on the B1123. Take the road
Fifteen miles northwest of Norwich at Foxley village, off opposite the garage, parking on the concrete pad near
the A1067 Norwich to Fakenham Road. There is a car the meadow entrance. (Suffolk WT, Plantlife)
park at the wood. (Norfolk WT)
21. Redgrave and Lopham Fens TM 046797
16. Buxton Heath TG 172216 This large reserve lies on the border between
An area of heath and mire with a small alkaline Suffolk and Norfolk in the Waveney Valley. The inter-
stream. The mix of vegetation is very interesting nationally important valley fen is flanked by wood-
with Marsh Helleborine, Marsh Fragrant Orchid, land, open water, reed and sedge beds and wet
Common and Heath Spotted Orchids and South- heath. Southern and Early Marsh Orchids, Common
ern and Pugsley’s Marsh Orchids (and some confus- Twayblade, and a few Marsh Fragrant Orchids are
ing intermediates). present.
006 systematic.indd 440 29/1/09 12:49:28

EAST ANGLIA l 441
Signposted between Redgrave and South Lopham on

the B1113; there are car parks and a Visitor Centre.
ESSEx
(NNR, Suffolk WT)
28. West Wood TL 624332
22. Market Weston Fen TL 981789 An ancient woodland on chalky boulder clay, with
A fine remnant of a valley fen dominated by Great a rich ground flora that includes Early Purple and
Fen Sedge, with adjacent heath and ponds. Spe- Greater Butterfly Orchids.
cies include Marsh Fragrant, Common Spotted and Midway between Thaxted and Great Sampford on the
Southern Marsh Orchids. An array of hybrids can B1051. Park beside the road. (Essex WT)
usually be found too. 29. Hatfield Forest TL 547202
West of Diss on the B1111, off the minor road between One of the last medieval royal forests to retain
Hopton and Coney Beeston Road. (Suffolk WT) much of its original character and composition. The
site contains mixed habitats of ancient woodland,
23. Rex Graham Reserve pasture, glades and marshy ground, separated by
This Suffolk WT reserve protects one of the few ditches and banks. The ground flora includes Violet
remaining British sites for Military Orchid. The Trust Helleborine and Bird’s-nest, Bee and Pyramidal Or-
opens the reserve to the public on 1-2 days each chids in the woods and glades, and Early Marsh and
year in late May or early June, to allow viewing of Common Spotted Orchids in the fens.
this very special orchid. Contact the Trust for details. Park off the A120 at Takely near the M11 Junction 8.
(Suffolk WT) (NNR, NT)
24. Bradfield Woods TL 935581 30. Roding Valley Meadows TO 430943
An ancient wood, one of the richest in the country, A traditionally managed river valley with hay mead-
comprising coppice-with-standards. Parts are known ows, scrub and woodland beside the River Roding.
to have been traditionally managed since 1252. The wet meadows hold many Southern Marsh
There is a diverse ground flora, and Early Purple Orchids.
Orchid can be found in spring, with Green-flowered Park next to the David Lloyd Centre, Roding Lane,
Helleborine elusive in mid-summer. Chigwell. There are other footpaths on the Loughton
On the road between Bradfield St George and Felsham, side of the River. (Essex WT)
12 miles southeast of Bury St Edmunds. (NNR, Suffolk
WT) 31. Langdon TQ 659874
An unusual site with much social history. The aban-
25. Chippenhall Green TM 285757 doned ‘plotlands’ are patchworks of old gardens and
A very impressive colony of Green-winged Orchids grassland bordered by hedgerows and orchards.
grows on this area of common land ‘enclosed’ by Green-winged Orchids can be seen in their thou-
cattle grids. sands in the Lincewood area, and Common Spot-
Southeast of Fressingfield on the B1116. Take the mi- ted, Green-winged, Pyramidal and Bee Orchids can
nor road to Chippenhall Green at Rackham’s Corner. be found in the area of Marks Hill.
West of Basildon off the B1036. There is a car park
26. Wolves Wood TM 054440 and Visitor Centre. (Essex WT)
An ancient woodland with hazel and hawthorn cop-
pice and scrub. The wide rides have a good ground
32. Grays Chalk Quarry TQ 611787
A long-disused quarry now with woodland, a small
flora with several species of orchid, including Violet
area of chalk grassland and a pond. There are many
Helleborine under the coppice.
orchid species present, with good numbers of Man,
On the A1071, 2 miles east of Hadleigh and 8 miles
Pyramidal and Bee Orchids.
west of Ipswich. (RSPB)
Park in the public car park on London Road near Grays
27. Groton Wood TL 976428 town centre. The entrance to the site is opposite, via a
A stand of ancient woodland with old lime coppice, short path. (Essex WT)
traditionally managed. The rides have an interesting
flora with Early Purple Orchid and Violet Hellebo-
rine, with a particularly good colony of helleborines
under the old hazel coppice on the west side of
the wood.
North of the A1071; along a maze of lanes 3 miles
west of Hadleigh, between Castling’s Heath and Kersey
Tye. (Suffolk WT)
006 systematic.indd 441 29/1/09 12:49:28

442 l SITE GUIDE
Central England
12
13 14
8
6 11
1
7 15
3 16
5 9
2 17
10 18
19 20 4 28
21 29
27
46
26 31 44
45
30 43
22 42
41
25
36
35 40
23 24
39
34
38
32 37
33
Cheshire Lincolnshire 24 Ufton Fields 35 Monkwood

1 Witton Lime Beds 12 Messingham Sand Quarry 25 Draycote Meadow 36 Wyre Forest
13 Kingerby Beck Meadows Leicestershire 37 Tiddesley Wood
Staffordshire
14 Donna Nook and 26 Lea Meadows 38 Windmill Hill
2 Rod Wood
Saltfleetby Dunes 27 Cloud Wood 39 Trench Wood
3 Coombes Valley
15 Candlesby Hill Quarry 28 Muston Meadows 40 Eades Meadow
Derbyshire 16 Heath’s Meadows 29 Cribbs Meadow northamptonshire
4 Hilton Nature Reserve 17 Ancaster Valley
30 Great Merrible Wood 41 Stoke Wood End Quarter
5 Rose End Meadows 18 Moulton Marsh
Rutland 42 Glapthorn Cow Pastures
6 Derbyshire Dales Shropshire 43 Short Wood
7 Deep Dale 31 Prior’s Coppice
19 Wem Moss 44 Collyweston Quarries
8 Miller’s Dale Quarry 20 Granville Country Park Herefordshire 45 Bedford Purlieus
nottinghamshire 21 Llynclys Common 32 Davies Meadows 46 Barnack Hills and Holes
9 Bentinck Banks 22 Wenlock Edge 33 The Doward Reserves
10 Wilwell Farm Cutting Warwickshire Worcestershire
11 Eaton Wood 23 Snitterfield Bushes 34 The Knapp and Papermill
Northeast of Northwich on the far side of the Trent &

CHESHIRE Mersey Canal. Access is from the minor road west of the
Flash. Park just before the village of Marston on the B5075.
1. Witton Lime Beds SJ 660746
These lime beds were produced by the chemical STaFFORDSHIRE
and salt industries near Northwich. The Aston’s
Flash area is notable for the magnificent show of 2. Rod Wood SJ 997531
Marsh Fragrant Orchids. More than 10,000 spikes With some fine meadows, woodland and marsh, the
have been recorded on the site in some years. interesting flora here includes Common Twayblade
Marsh Helleborine also occurs. and Early Purple Orchid.
006 systematic.indd 442 29/1/09 12:49:29

CENTRAL ENGLAND l 443
Near Leek. Accessed from the A523 Leek to Ashbourne Turn off the A6 towards Sheldon between Taddington
road; go past the RSPB Coombes Valley reserve (Site 3 and Ashford-in-the-water. Access is along public foot-
below) to the top of the hill. Park past the cattle grid. paths. (Plantlife)
(Staffordshire WT)
8. Miller’s Dale Quarry SK 140731
3. Coombes Valley SK 005525 One of the Wye Valley reserves, which also include
A steep valley among old woodland with the Chee Dale (SK 120727), and Priestcliffe Lees (SK
Coombes Brook running the length of the reserve. 147730). They have some excellent habitats, and a
There is some good flower-rich grassland here, with range of orchids can be found. These include Early
Common Spotted and Greater Butterfly Orchids. Purple, Common Fragrant and Common Spotted
Southeast of Leek; take the A523 Leek to Ashbourne Orchids, and Common Twayblade.
road and go along the minor road signposted to East of Buxton, with parking at Miller’s Dale off the
Apesford. (RSPB, Staffordshire WT) B6049 or at Monsal Dale Station. (Derbyshire WT)
DERBYSHIRE nOTTInGHaMSHIRE
4. Hilton nature Reserve SK 249315 9. Bentinck Banks SK 493550
With lakes, pools, woodland and grassland, there are A triangle of disused railway lines; the embankments
stunning displays of Southern Marsh Orchids here. have some good limestone grassland, with Common
Also several thousand Common Twayblades. Fragrant Orchid, Frog Orchid, Common Twayblade
West of Derby, off the A516 to Willowpit Lane, north of and Common Spotted Orchid.
Hilton. (Derbyshire WT) Just south of Mansfield, off the B6018 between Kirkby
in Ashfield and the minor road to Nuncargate. Park on
5. Rose End Meadows SK 293567 the verge. (Nottinghamshire WT)
A series of traditionally managed meadows, wetland
and woodland with a great variety of wildflowers. Spe- 10. Wilwell Farm Cutting SK 568352
cies include Pyramidal, Common Fragrant and Bee Or- One of the best wildflower sites in Nottinghamshire,
chids, and there are many Common Spotted Orchids. with grassland, acid fen and scrub on an old indus-
Above Cromford between the A5012 and the B5036 trial site. There is a good display of Green-winged
Cromford to Wirksworth road. Park in Cromford village. Orchids, as well as Bee, Common Spotted, and
(Derbyshire WT) Southern Marsh Orchids and Common Twayblade.
Vast hybrid ‘swarms’ of orchids also occur.
6. Derbyshire Dales Just off the B680 between Ruddington and Wilford.
Five valleys make up the Derbyshire Dales NNR. Parking is available beside the reserve. (Nottingham-
These are two of the best for orchids. shire WT)
Lathkill Dale (SK 190658) is one of the finest,
with stunning scenery and woodland, grassland
11. Eaton Wood SK 727772
Ancient pasture woodland with a visible ridge and
and evidence of the lead-mining industry. Common
furrow system that holds several species, including
Twayblade, Common Spotted Orchid and Com-
Greater Butterfly, Early Purple and Common Spotted
mon Fragrant Orchid can all be found among a very
Orchids, and perhaps Broad-leaved Helleborine.
rich flora, with thousands of Early Purple Orchids on
Off the minor road from Upton to Eaton at East Retford,
the grassy slopes.
opposite Gamston Wood. (Nottinghamshire WT)
Southwest of Bakewell, with numerous footpaths
across the dale. There is parking at Monyash on the
B5055 and at Over Haddon on minor roads from
LInCOLnSHIRE
Bakewell. 12. Messingham Sand Quarry SE 908032
Cressbrook Dale (SK 170731) is a beautiful place, Old flooded lagoons with fringing vegetation and
with Early Purple, Common Spotted and Common remnant heath, with woodland, grassland and marsh.
Fragrant Orchids, and a few Dark-red Helleborines Several orchid species occur, including Pyramidal,
and Common Twayblades. Common Spotted, Early Purple and Bee Orchids.
Northwest of Bakewell between Cressbrook and Litton. South of Scunthorpe at Messingham, on the A159. Ac-
Park at Monsal Dale off the B6465 or at Tideswell cess is off the B1400 opposite Scallow Grove Farm. A
Dale off the B6049. track leads to a small car park. (Lincolnshire WT)
7. Deep Dale SK 165698 13. Kingerby Beck Meadows TF 051941
Fantastic upland limestone grassland, with stunning A series of herb-rich meadows with mixed bound-
displays of Early Purple Orchids in spring. Other ary hedges. Orchids present include a colony of the
orchids to be found include Common Twayblade. diminutive Frog Orchid.
006 systematic.indd 443 29/1/09 12:49:29

444 l SITE GUIDE
Northwest of Lincoln near Market Rasen.The meadows carr, woodland and a fen community that includes
lie between Kingerby Beck and North Owersby; access Early Marsh and Lesser Butterfly Orchids.
from North Owersby on the minor road to Glentham. Between Wem and Ellesmere, near Northwood on the
Park near the entrance. (Lincolnshire WT) B5063. Park on the minor road to Dobson’s Bridge just
west of Northwood. (Shropshire WT, NNR)
14. Donna Nook and Saltfleetby-
Theddlethorpe Dunes TF 467917 20. Granville Country Park SJ 719124
A large area of coastal saltmarsh and sand dunes, with Based around reclaimed mine workings near Tel-
Bee and Pyramidal Orchids and Marsh Helleborine ford, this site contains woodland and old grassy pit
in the dune slacks. The area of freshwater marsh can mounds; the meadows have vast numbers of South-
have fantastic displays of Early and Southern Marsh ern Marsh Orchids and hybrids.
Orchids and their hybrids.The Saltfleetby reserve oc- Just outside Telford, on Granville Road, Oakengates,
cupies the coast between Mablethorpe North End in Wellington. (Shropshire WT, Telford & The Wrekin
the south and Saltfleet Haven in the north. Council)
Access to Donna Nook is from North Somercotes,
which is on the A1031 coast road; park at Stonebridge. 21. Llynclys Common SJ 273238
Do not enter the danger area when the ranges are A mixture of habitats: woods, meadows, old quar-
in use. Saltfleetby is easily accessed from several car ries and scrub, where up to 12 orchid species have
parks off the A1031. (NNR, NE, Lincolnshire WT) been recorded. These include Common Spotted,
Early Purple, Pyramidal, Greater Butterfly and Bee
15. Candlesby Hill Quarry TF 460682 Orchids, and Common Twayblade.
An old chalk and lime pit on the eastern edge of the South of Oswestry on the A495. Park in the lay-by west
Lincolnshire Wolds. There is wood and scrub, and of Llynclys crossroads opposite Dolgoch. (Shropshire
newly colonised chalk grassland with Bee and Com- WT).
mon Spotted Orchids, among others.
Northwest of Skegness off the A1028 Skegness to 22. Wenlock Edge SO 595988
Louth road, just north of Gunby Corner roundabout. A magnificent limestone escarpment running from
Park on the verge. (Lincolnshire WT) Much Wenlock to Craven Arms, with habitats rang-
ing from scrub woodland and grassland to marsh.
16. Heath’s Meadows TF 484640 The pockets of limestone grassland have good dis-
Old meadows divided into small fields surrounded by plays of Pyramidal and Greater Butterfly Orchids,
large hedgerows.The meadows are traditionally man- along with other limestone species.
aged and support a good grassland flora, including Harton Hollow Woods (SO 481878), on minor
Green-winged and Common Spotted Orchids. roads off the B4368, is an excellent place to start a
West of Skegness on the A158, just southwest of walk along the escarpment. The National Trust has a
Burgh-le-Marsh at Bratoft Ings. Access is from Ings car park here. (NT, Shropshire WT)
Lane. Park on the roadside. (Lincolnshire WT)
17. ancaster Valley SK 984434 WaRWICKSHIRE
A narrow, steep-sided valley with limestone grassland,
scrub and woodland. Bee and Common Fragrant There are several interesting Roadside Nature Re-
Orchids can be found here. Early Purple Orchid oc- serves in the county, notably on the A429 Ettington
cur in the beech woodland. bypass, which has a spectacular display of Bee and
On the south side of the A153 Sleaford to Grantham Pyramidal Orchids.
road near Ancaster. Access is via the bridleway sign-
23. Snitterfield Bushes SP 200603
posted east of the Ancaster crossroads. Park in the vil-
One of the best ancient woodlands in the county,
lage. (Lincolnshire WT)
the ash and birch woods have a good shrub layer
18. Moulton Marsh SK 344336 and ground flora, with Early Purple, Greater Butter-
The bank on the south side of the River Welland has fly and Common Spotted Orchids, Common Tway-
a fantastic display of Pyramidal Orchids, with over blade, and Broad-leaved Helleborine.
4,000 spikes. Near Stratford-on-Avon, on the Bearley Road out of
Off the A17 south of Fosdyke Bank, on a minor road Snitterfield; the reserve is on both sides of the road,
that leads on to the river. There is a car park. (Lin- with a car park. (Warwickshire WT)
colnshire WT)
24. ufton Fields SP 378615
SHROPSHIRE Old limestone workings with scrub, grassland and
ponds and a good range of orchid species, with Com-
19. Wem Moss SJ 472342 mon Twayblade, Greater Butterfly, Common Spotted
One of Shropshire’s lowland mires, with open water, and Bee Orchids.
006 systematic.indd 444 29/1/09 12:49:29

CENTRAL ENGLAND l 445
Between Royal Leamington Spa and Southam, off the ash-maple and ash-wych elm coppice is a rich flora,
A425 at Ufton. There is a car park available. (War- with a large colony of Early Purple Orchids and
wickshire WT) Broad-leaved and Violet Helleborines; the often-wet
25. Draycote Meadow SP 448706 rides contain Common Spotted Orchids.
Warwickshire’s best meadow, with thousands of Southwest of Oakham, on minor roads connecting
Green-winged Orchids among other specialities. Braunston to Leighfield. Park on the road to Leighfield
Southwest from Rugby, with access off the B4453 at Lodge. (LRWT).
Draycote village. A public bridleway gives access to the
meadow. (Warwickshire WT) HEREFORDSHIRE
32. Davies Meadows SO 375485
LEICESTERSHIRE With three unimproved meadows, an ancient or-
26. Lea Meadows SK 506115 chard, marshland and old hedgerows, this is an
Unimproved meadows beside a clear stream. Signs enchanting place to visit. The meadows contain
of ridge and furrow agriculture betray medieval us- Green-winged and Common Spotted Orchids.
age, but the fields are now traditionally managed. On the A480 from Hereford to Lyonshall. Entrance
They contain a rich flora that includes great displays is just past the Three Horseshoes pub at Eccles
of Common Spotted and Heath Spotted Orchids. Green; park on the road verge. (Herefordshire WT,
Northwest of Leicester; take the minor road to Newton Plantlife)
Linford from the A46. The site is on Ulverscroft Lane.
33. The Doward Reserves SO 549161
(LRWT)
Several ancient oak and beech woodland reserves
27. Cloud Wood SK 899188 with areas of unimproved limestone grassland. They
A traditionally coppiced ancient woodland with good skirt the Doward, a rock outcrop in a bend of the
rides and glades. Several orchids can be found includ- River Wye. At Woodside (SO 555147), the lime-
ing Bee, Bird’s-nest, Common Spotted and Greater stone grassland has a good flora with Early Purple
Butterfly Orchids and Broad-leaved Helleborine. and Greater Butterfly Orchids. Leeping Stocks (SO
Northeast of Ashby de la Zouch on the minor road 548162) has White and Broad-leaved Helleborines
linking Griffydam on the B5324 and Tonge on the and Greater Butterfly Orchid, while White Rocks
A453. Park in the lay-by. (LRWT) (SO 550158) has Bee Orchid.
28. Muston Meadows SK 824367 South of Ross-on-Wye on the A40 to Whitchurch, from
One of the best lowland meadows in England with where the reserves are signposted. (Herefordshire WT)
unimproved ridge and furrow grassland. Among its
rich flora is a colony of more than 10,000 Green- WORCESTERSHIRE
winged Orchids.
West of Grantham off the A52 at Muston. Park off the 34. The Knapp and Papermill SO 751522
minor road south to Woolsthorpe. (LRWT, NNR) One of the finest nature reserves in the Midlands, this
is a mixture of old meadows, woods and orchards be-
29. Cribbs Meadow SK 899188 side the Leigh Brook. Species include Green-winged,
Ancient grassland, ponds and a disused railway line.The Early Purple, Greater Butterfly and Common Spotted
fields lie on boulder clay and support a rich flora, includ- Orchids, and Common Twayblade.
ing Green-winged and Common Spotted Orchids. Southwest of Worcester on the A4103. Follow minor
Between Wymondham and South Witham on the mi- roads from Bransford along the Bransford–Smith End
nor road between Sewstern and Thistleton. Park on the
Green–Alfrick Pound road. The reserve entrance is on
road verge. (LRWT, NNR)
the left after about 3 miles, where the road crosses the
30. Great Merrible Wood SP 834962 Leigh Brook. There is limited parking near the bridge.
A mixed woodland with a very ancient history. With (Worcestershire WT)
ash, oak, and a varied shrub layer, the ground flora
includes Broad-leaved and Violet Helleborines. 35. Monkwood SO 804606
Southwest of Uppingham off the Horninghold to Great A coppiced woodland important for butterflies
Easton road, southeast of the crossroads with the and moths, but also supporting many Early Purple
B664.There is roadside parking. (LRWT) Orchids, some Common Spotted and Greater But-
terfly Orchids, and a few, hard-to-find Broad-leaved
RuTLanD and Violet Helleborines.
Northwest of Worcester; near Grimley, on the minor
31. Prior’s Coppice SK 834052 road to Monkwood Green from the A443. (Worces-
An ancient woodland on a steep slope. Under the tershire WT, Butterfly Conservation)
006 systematic.indd 445 29/1/09 12:49:30

446 l SITE GUIDE
36. Wyre Forest SO 743740 Access is only possible on open days – contact the
The Wyre forest covers over 6,000 acres and con- Worcestershire WT. (Worcestershire WT)
tains some of the best stands of ancient woodland
in Britain (it is the largest continuous area of ancient nORTHaMPTOnSHIRE
woodland in England). Although predominantly Ses-
sile Oak, just over half of the forest consists of conifer
41. Stoke Wood End Quarter SP 800861
This ancient oak and ash woodland has an excellent
plantations. The Dowles Brook runs through one of
flora on the mainly calcareous clay soils. Orchids
the richest and most diverse areas of the forest, includ-
include Early Purple Orchid, Broad-leaved Helle-
ing meadows and abandoned orchards. Eleven species
borine, Common Twayblade and Greater Butterfly
of orchid have been recorded, including Sword-leaved
Orchid.
Helleborine, although there are just a few scattered
North of Desborough on the B669 to Stoke Albany.
populations of the latter that are hard to find. Com-
Park in the lay-by. (BCNPWT, WT)
mon Spotted Orchid is abundant in some rides, and
other species include Common Fragrant Orchid and 42. Glapthorn Cow Pastures TL 005903
Green-winged Orchid in some of the meadows (and A scrubbed-over pasture, the resulting thickets are a
perhaps also Heath Fragrant Orchid). haven for wildlife. The area has a rich flora, including
West of Bewdley on the A456 Ludlow road, with park- Early Purple and Common Spotted Orchids.
ing and a Visitor Centre at the FC’s Callow Hill, which Access from the minor road halfway between Glap-
gives access to the North Worcestershire path; the thorn and Benefield, north of Oundle. Park on the road
New Parks Beech Wood area is a good place to start. verge. (BCNPWT)
37. Tiddesley Wood SO 929462 43. Short Wood TL 015913
An ancient mixed woodland near Pershore with a Rich deciduous woodland and coppice on boulder-
good shrub layer. Common Spotted Orchid is wide- clay (said to be the finest Bluebell wood in the coun-
spread; there are also Early Purple and Greater But- ty), with Early Purple, Bird’s-nest, Greater Butterfly
terfly Orchids and Common Twayblade, with Bee and Common Spotted Orchids, Common Twayblade,
Orchids in the open grassland; Bird’s-nest Orchid and Broad-leaved and Violet Helleborines.
and Violet Helleborine are also present but can be Northwest from Oundle to Southwick on a minor road.
hard to find. Access is along a signposted footpath on this road.
Next to the minor road signposted to Besford and (BCNPWT)
Croome from Pershore on the A44. Park beside the 44. Collyweston Quarries TF 004038
wood. (Worcestershire WT) A series of old grassed-over quarries, the open-cast
38. Windmill Hill SP 072477 pits and mines have created a ‘hills and holes’ appear-
Windmill Hill has a small area of limestone grassland ance, and a rich flora has developed. Common Tway-
and scrub. There is a spectacular display of Pyrami- blade, Man, Bee, and Common Fragrant Orchids are
dal Orchids as well as Common Spotted Orchid, present, while Pyramidal Orchid is abundant.
Common Twayblade and small numbers of Bee and South of Stamford, on the A43 between Easton-on-the-
Greater Butterfly Orchids. hill and Collyweston. (BCNPWT)
East of Evesham on the south side of the B4510, close 45. Bedford Purlieus TL 034997
to the Fish and Anchor Inn and just before the brow A large, remarkably diverse woodland reserve,
of the hill; park on the roadside opposite the entrance. which many botanists consider to be the richest of
(Worcestershire WT) all British woods – more than 400 species of vascu-
39. Trench Wood SO 930589 lar plants have been recorded on the site. Fly and
An ancient woodland, with the wood managed Greater Butterfly Orchids can be found here, as can
partly as high forest and partly as scrub or coppice. large numbers of Common Spotted Orchids.
Orchids present include Common Spotted, with North of Oundle on the A47; park on the farm track be-
smaller numbers of Pyramidal and Greater Butterfly tween Wansford and the King’s Cliffe road. (FC, NNR)
Orchids. 46. Barnack Hills and Holes TF 075045
Southeast of Droitwich on the Shernal Green to Sale Green A superb area of calcareous grassland on long-
minor road, just west of the M5. (Worcestershire WT) abandoned medieval limestone workings. There is
40. Eades Meadow an important Man Orchid colony (more than 1,000
A traditionally managed meadow on heavy clay soils, spikes) as well as Early Purple, Pyramidal, Bee, Com-
Eades Meadow has a spectacular display of Green- mon Fragrant and Common Spotted Orchids.
winged Orchids in the spring. Common Spotted Orchid, East of Stamford; the reserve is off the B1443 on mi-
Common Twayblade, Greater Butterfly Orchid and a nor roads at the western end of Barnack Village, with
few Common Fragrant Orchids are also present. several car parks around the perimeter. (NNR)
006 systematic.indd 446 29/1/09 12:49:30

NORTHERN ENGLAND l 447
northern England and Isle of Man
2
4
3
28
29 27
30
32 31
5 26
8
6
34 33 7
12
35
11 9
36 13 10 14 25
17
18
19 20 24
23
21
16
15 22
northumberland 18 Brae Pasture

1 Holy Island 19 Malham Tarn Estate
2 Beltingham River Gravels 20 Grass Wood
3 Williamston Reserve 21 Sherburn Willows
Cumbria 22 Brockadale
4 Geltsdale 23 Skipwith Common
5 Augill Pasture 24 Wharram Quarry
6 Little Asby Inrakes and Outrakes 25 Flamborough Cliffs
7 Smardale Gill County Durham
8 Waitby Greenriggs 26 Coatham Marsh
9 Hutton Roof Crags 27 Castle Eden Dene
10 Arnside Knott 28 Hawthorn Dene
11 Latterbarrow 29 Wingate Quarry
12 Whitbarrow 30 Raisby Hill Grassland
13 Sandscale Hawes 31 Bishop Middleham Quarry
Lancashire 32 Upper Teesdale
14 Gait Barrows Isle of Man
15 Ainsdale Sand Dunes 33 Ayres Visitor Centre
16 Nob End 34 Moaney and Crawyn’s Meadows
Yorkshire 35 Close Sartfield
17 Southerscales 36 Dalby Mountain
006 systematic.indd 447 29/1/09 12:49:30

448 l SITE GUIDE
East of Carlisle, south of the A69 off the B6413 at

nORTHuMBERLanD Castle Carrock. Park in the RSPB car park. (RSPB)
1. Holy Island nu 100432 5. augill Pasture nY 817147
The island is around three miles long, and sand Species-rich grassland and some woodland with
dunes extend along much of the north shore. The Common Fragrant and Greater Butterfly Orchids,
dunes and dune slacks form an excellent orchid among others.
habitat, especially at the Snook at the western end Just east of Brough off the A66 to Augill House Farm.
of the island. The great speciality is Lindisfarne Hel- Park at the end of the road. (Cumbria WT)
leborine, formerly treated as a subspecies of Dune
Helleborine (which it greatly resembles). Following
6. Little asby Inrakes and Outrakes
recent genetic research, the Lindisfarne Helleborine nY 699101
is now treated as a distinct species endemic to the A small area of rough grazing notable for a colony of
island. The population is small (150-300 plants) and Small White Orchids, as well as Northern and Early
scattered, and it is advisable to contact the warden Marsh Orchids, Common Spotted Orchid, Early
for advice; tel. 01289 381470. Other notable species Purple Orchid and Heath Fragrant Orchid.
include Coralroot Orchid in very small numbers in West of Kirkby Stephen, which lies on the A685. The
the dune slacks, Marsh Helleborine, Northern and area lies north of the minor road immediately north
Early Marsh Orchids and Common Spotted Orchid. of Little Asby.
The road to the island is signposted off the A1 at West 7. Smardale Gill nY 727070
Mains Inn. It is impossible to cross the causeway for A disused railway line with a rich orchid flora; Fly and
at least 2 hours either side of high water, and the Frog Orchids can be found in small numbers, along
strength of the wind and the height of the tide must with Greater Butterfly, Common Spotted, Northern
also be taken into account.Tide tables are displayed on Marsh and Common Fragrant Orchids, Common
the causeway, which is a tarmac road. Twayblade, and Marsh Helleborine. A few Lesser
2. Beltingham River Gravels nY 785640 Butterfly Orchids are also here but hard to find.
Grassland polluted with heavy metals, woodland West of Kirkby Stephen, off the A685. Access is from
and scrub where ‘Tyne Helleborine’, the inland form the south side of the minor road running from Kirkby
of Dune Helleborine, can be found. Stephen to Crosby Garrett. (Cumbria WT, NNR)
Off the A69 south of Bardon Mill, on the minor road 8. Waitby Greenriggs nY 760085
from Beltingham to Willimontswick. There is roadside An old railway line running down a beautiful valley,
parking and a walk along the river to the reserve. with woodland and herb-rich meadows. The rich
(Northumberland WT) flora includes Marsh Helleborine, all three fragrant
3. Williamston Reserve nY 681521 orchids, Fly, Frog, Northern Marsh and Common
This small reserve on the eastern bank of the River Spotted Orchids, and Common Twayblade.
South Tyne comprises birch and willow woodland; East of Ravenstonedale at Waitby, off the A685. Park
this has grown on gravel and silt contaminated by close to the railway bridge. (Cumbria WT)
‘tailings’ from lead and zinc mines. The reserve is no- 9. Hutton Roof Crags SD 543783
table for its large colony of ‘Tyne Helleborine’, the A large area of woodland, scrub, heath and fantas-
inland form of Dune Helleborine. Common Spot- tic limestone pavement, with great views. Dark-red
ted Orchid is also present. Helleborine can be found amongst the grikes, and
Off the A689 Alston to Brampton road, south of Slaggy- Fly Orchids grow in the grassland.
ford village, signposted to Barhough Hall. Park by the Off the A6070, between Burton and Hutton Roof. Park
road; the entrance to the reserve is via a stile immedi- on the roadside or walk from Hutton Roof, Burton or
ately over the river bridge. (Northumberland WT)
Holme villages. (Cumbria WT)
CuMBRIa 10. arnside Knott SD 456775
Woodland, scrub and limestone pavement sloping
4. Geltsdale nY 561557 down to saltmarsh, with great views. Several spe-
Twelve-thousand acres of upland heather moorland, cies of orchid are present, including Early Purple, Fly
working farms, old coal and limestone workings and and Lesser Butterfly Orchids and Broad-leaved and
woodland, along the River Gelt. There are some Dark-red Helleborines.
very rich habitats where several species of orchid On the south side of the Kent estuary. Park in the
can be found, including Greater Butterfly Orchid National Trust car park signposted from Arnside village,
and Common Fragrant Orchid. which is at the end of the B5282. (NT)
006 systematic.indd 448 29/1/09 12:49:31

11. Latterbarrow SD 441827 16. nob End SD 743070

Excellent limestone grassland, scrub and woodland An unusual site made up entirely of alkaline indus-
with Early Purple and Greater Butterfly Orchids. trial waste, generated through soda crystal manu-
Near Grange-over-Sands just off the A590 at Long- facture in the 19th century. Its weathered surface
howe End. Park beside the road just past the Derby now has a unique flora, with eight species of orchids.
Arms. (Cumbria WT) Common Spotted Orchid and Early Marsh Orchid
12. Whitbarrow SD 436859 (subspecies coccinea) are common. Northern and
Largely limestone, Whitbarrow Scar is just over 650 Southern Marsh Orchids occur, and Common
feet high. The reserve sits on top; it consists of lime- Fragrant Orchid and Common Twayblade are also
stone pavement, grassland, juniper scrub and wood- common. Harder to find are Marsh Helleborine and
land. It is an interesting place to visit, and Dark-red Bee Orchid. The marsh orchid hybridisation at this
Helleborine and Fly Orchid can be found here. site is stunning.
Southwest of Kendal off the A5074. There is a small Just off the A6053 west of Little Lever, near Bolton.
car park at the Kennels at Witherslack. Access is via Access is on foot, with parking near Rock Hall Visitor
steep and rocky paths. (Cumbria WT, FC, NNR) Centre in Moses Gate Country Park.
13. Sandscale Hawes SD 200756

This extensive dune system holds 10 species of or-
YORKSHIRE
chid, with specialities including the largest popula-
tion of Coralroot Orchid in England (3,000 were 17. Southerscales SD 742769
counted across five slacks in 1991), Dune and Part of the much larger Ingleborough NNR, this re-
Green-flowered Helleborines (often more than serve has good limestone pavement with its clints
1,000 spikes of each), as well as Marsh Helleborine, and grikes, and acidic and limestone grassland. Early
Northern Marsh Orchid and Bee and Pyramidal Purple Orchid, Dark-red Helleborine and Common
Orchids. Twayblade are present.
North off the A590 to the northwest of Dalton-in- Off the B6255 Ingleton to Hawes road in North York-
Furness, on the minor road to Roanhead. Park at the shire. Park at the Old Hill Inn or on the roadside at
beach car park and walk west along the beach and Chapel-le-Dale. (Yorkshire WT, NNR)
then into the dunes (the first ‘blow-out’ after 550 yards
holds a few Coralroot Orchids, but the farther slacks 18. Brae Pasture SD 790741
are better). (NT, NNR) A small area of limestone pasture and damaged
limestone pavement that has a very rich flora; Early
LanCaSHIRE Purple and Frog Orchids can be found here.
On the B6479 between Horton-in-Ribblesdale and
14. Gait Barrows SD 483775 Selside. Park on this road. A Public Footpath sign points
Some of the finest limestone pavement and Yew to the reserve.
woodland in the country. Dark-red and Broad-
leaved Helleborines, and Common Fragrant, Com- 19. Malham Tarn Estate SD 890660
mon Spotted and Northern Marsh Orchids are Upland hay meadows and limestone grassland, lime-
important parts of a fascinating flora. stone pavements and the NNR at Malham Tarn make
Gait Barrows is just in Lancashire, east of Silverdale and up this wonderful landscape. The variety of habitats
close to Arnside Knott. Off the minor road linking Carn- have Dark-red Helleborine and, in the wetter areas,
forth to Arnside, with roadside parking. (NNR) Early, Northern and Pugsley’s Marsh Orchids.
15. ainsdale Sand Dunes SD 397129 Northwest of Skipton on minor roads from the A65;
Part of the huge Sefton Coast wildlife area, the spe- the estate is north from Malham village, where there is
ciality here is Dune Helleborine, which is common, a Visitor Centre, parking and footpaths. (NT, NNR)
as well as Green-flowered Helleborine (both these
helleborines can also be found in the pine planta- 20. Grass Wood SD 983652
tions to the rear of the dunes). Other orchids include A wooded limestone hillside that slopes down to
Marsh Helleborine, Bee and Pyramidal Orchids and the River Wharfe. The wood has formed on a series
Early Marsh Orchid (both pink-flowered plants and of limestone terraces, and has rocky outcrops and
the red-flowered subspecies coccinea). Northern open areas. Early Purple and Bird’s-nest Orchids can
and Southern Marsh Orchids also occur here. be found, among others.
There is open access to the dunes, which stretch from In Wharfedale, north of Grassington alongside the
Southport to Formby, with plenty of car parks. Ainsdale Grassington to Coniston minor road. There is a small
beach car park is a useful start. (NNR) car park. (Yorkshire WT)
006 systematic.indd 449 29/1/09 12:49:31

450 l SITE GUIDE
21. Sherburn Willows SE 487326 Off the A1085 Redcar to Middlesbrough road. Park off
An interesting mix of habitats, with magnesian lime- Tod Point Road or on the sea front on Manjuba Road.
stone grassland, scrub, and fen to the side of a small (Tees Valley WT)
stream. Common Spotted and Bee Orchids and
27. Castle Eden Dene nZ 435397
Common Twayblade can be seen.
This spectacular wooded gorge is on magnesian
West of Selby; access along Mill Dyke, off the B1222 limestone and boulder clay. Fly and Bird’s-nest Or-
between Sherburn-in-Elmet and South Milford. Park on chids can be found in the woods, and Common
New Lane or walk from South Milford Station. (York- Spotted and Common Fragrant Orchids occur in
shire WT) the open grassland.
22. Brockadale and Thompson Meadow On the south side of Peterlee, east of the A19. Castle
SE 513174 Eden Dene is signposted from the A19 and from Peter-
Brockadale is a wooded valley with some old quarry lee town centre. Car parking is at the main entrance,
workings and traditionally managed meadows. One and a small car park is available at Oakerside Dene
of these, Thompson Meadow, is on a sloping hillside Lodge. (NNR)
beside the River Went. The rich limestone flora in- 28. Hawthorn Dene and Meadow nZ 433457
cludes Common Spotted, Common Fragrant and and Beacon Hill nZ 440455
Early Purple Orchids. A coastal site with extensive, steep-sided mixed
Between Wentbridge and Kirk Smeaton, off the B6474 woodland on limestone and some excellent
south of Pontefract, just east of the A1.The reserve car grasslands with a great flora. Early Purple, Lesser
park is on an unmarked track out of Little Smeaton. Butterfly and Bird’s-nest Orchids can be found
(Yorkshire WT, Plantlife) among the ground flora in the woods, with Bee,
23. Skipwith Common SE 669378 Common Fragrant, Common Spotted and North-
A large area of heath, marsh and woodland with ern Marsh Orchids in the pastures.
Common Twayblade, Common Spotted Orchid and At Hawthorn off the B1432, between Easington and
Broad-leaved Helleborine. Seaham on the Durham coast. The site is at the end
Northeast of Selby, Skipwith is north off the A163 on of a minor road signposted ‘Quarry Traffic’. Park in the
minor roads after North Duffield. Park and walk from lay-by opposite the cottage. Access to Beacon Hill is
the village. along the Coastal Footpath or through the southern
end of Hawthorn Dene. (Durham WT, NT).
24. Wharram Quarry SE 858653
A disused chalk quarry, with steep chalk slopes and 29. Wingate Quarry nZ 373375
grassland. A typical chalk flora has developed, and An important example of magnesian limestone
Pyramidal Orchid, Common Spotted Orchid and grassland – one of Britain’s rarest habitats. There are
abundant Bee Orchids can be seen. large numbers of Common Fragrant Orchids here.
Southeast of Malton close to Wharram-le-Street, on the Northwest of Hartlepool, off the A181 west of Wing-
B1248 Malton to Wetwang road. Park on the roadside ate.There is a car park. (Durham WT)
close to the old railway line. (Yorkshire WT) 30. Raisby Hill Grassland nZ 337375
25. Flamborough Cliffs Ta 240722 Magnesian limestone grassland with fen and pools.
The spectacular chalk cliffs attract thousands of sea- There is a small disused quarry which has an impor-
birds but also support a varied flora on the cliff-tops. tant population of Dark-red Helleborines.
Off the A177 from Middlesbrough at Coxhoe Quarry.
Pyramidal and Common Spotted Orchids occur
(Durham WT)
here and Northern Marsh Orchids can be found
in wet flushes. 31. Bishop Middleham Quarry nZ 330324
East of Bridlington on the B1255 to Flamborough. This large disused quarry on the Durham limestone
There is a car park at North Landing and access to is superb. It holds the largest colony of Dark-red
the clifftop footpaths. (Yorkshire WT) Helleborines in Britain, with around 2,000 spikes
(perhaps more than all the other populations put
COunTY DuRHaM together). Other species include Pyramidal and Bee
Orchids, Common Spotted Orchid, Marsh Fragrant
26. Coatham Marsh nZ 586248 Orchid and Common Twayblade.
A wetland reserve of lakes, marsh and wet meadows. Northwest of Middlesbrough, off the A177 west
The wide variety of wildflowers include Common through Bishop Middleham. The reserve is on a minor
Fragrant Orchid, and good displays of Northern road north of the village. Park in the lay-by opposite the
Marsh and Bee Orchids. entrance. (Durham WT)
006 systematic.indd 450 29/1/09 12:49:32

32. upper Teesdale nY 907282

UpperTeesdale is a botanist’s paradise, with the famous
reserves of Cronkley and Widdybank Fell above Cow
Green Reservoir on the ‘sugar’ limestone. At lower
altitudes, wonderful flower meadows still exist along
the River Tees from Langdon Beck to Middleton-in-
Teesdale.There are many orchids in the meadows and
around the paths. Heath Spotted, Common Spotted
and Early Purple Orchids and Common Twayblade
can be found. As well as Early Marsh, Northern Marsh
and Lesser and Greater Butterfly Orchids, there can
be spectacular displays of hybrids.
North of Middleton-in-Teesdale on the B6277 there is
an excellent Durham WT Visitor Centre at Bowlees. In-
formation on good walks in the area is available from
there. (NNR)
ISLE OF Man
33. ayres Visitor Centre nx 435038
The shingle near the northern tip of the island sup-
ports an unusual lichen-heath community. Dense-
flowered Orchid grew here from 1966-1986 at its
only British site. The lichen heath lies behind the
dunes, and Pyramidal Orchid can be found here. In
the dune slacks there are Early Marsh and Northern
Marsh Orchids.
On the A10 Ballaghennie Road, west of Bride. There is
a Visitor Centre and car park. (Manx WT)
34. Moaney and Crawyn’s Meadows
SC 375957
Traditional hay meadows with a rich flora.The mead-
ows are on the edge of the Ballaugh Curragh wet-
land. Heath Spotted Orchids occur in abundance.
Off the A14 from Sulby to Ballaugh Curragh. Park in
the lay-by. (Manx WT, Plantlife)
35. Close Sartfield SC 358956
Part of the Ballaugh Curragh, this is the largest and
most important wetland on the Isle of Man. It is a
mixed area with bog, willow and birch woodland,
and rich hay meadows. There is an amazing display
of many thousands of orchids flowering from late
May, including Heath Spotted, Common Spotted,
Early Marsh and Northern Marsh Orchids, and
Common Twayblade.
On minor roads off the B9 between Ballaugh village
and Sulby Glen.The entrance and car park are located
about 25 yards along a track. (Manx WT).
36. Dalby Mountain SC 233769
A traditional moorland with good displays of Heath
Spotted Orchids.
The A27 Dalby to Round Table road passes through
the reserve; there is parking on this road close to the
track to Eary Cushlin. (Manx WT)
006 systematic.indd 451 29/1/09 12:49:32

452 l SITE GUIDE
Wales
anglesey Pembrokeshire
1 Newborough 19 Dowrog Common
3 9
Warren 20 St David’s Head
2 4 2 South Stack Cliffs Carmarthenshire
3 Cemlyn 21 Allt Rhyd y Groes
1 4 Cors Goch 22 Pembrey Burrows
10 Gwynedd
5 Glamorganshire
5 Caeau Tan y Bwlch 23 South Gower Cliffs
6 8 6 Morfa Harlech
7 12 24 Oxwich Bay
7 Morfa Dyffryn 25 Crymlyn Bog
14 8 Roman Steps
13 26 Kenfig
Conwy 27 Merthyr Mawr
11 9 Great Orme Monmouthshire
Wrexham 28 Cwm Clydach
10 Marford Quarry 29 Pentwyn Farm
15 17 30 Springdale Farm
Cardiganshire
16 11 Ynyslas 31 Newport Wetlands
21 Powys
20 18 12 Lake Vyrnwy
19
13 Dyfnant Meadows
14 Llanymynech
28
29 Rocks
22 15 Elan Estate
25 30
16 Nant Irfon
23 24 26 17 Burfa Bog
27 31
18 Pwll y Wrach
Feilw Common. Heath Spotted Orchid is common

anGLESEY on Holyhead Mountain and four species of orchid,
including Northern Marsh, can be found at Penrhos
1. newborough Warren SH 415635 Feilw Common.
Lying at the southern tip of Anglesey, this is one of West of Holyhead on minor roads via Llaingoch and
the largest sand dune systems in Britain, covering Twr to South Stack. For Penrhos Feilw Common (The
more than 3,000 acres. Although two-thirds of this Range) follow minor roads south from South Stack, or
area has been planted with Corsican Pine, there the B4545 from Dyffryn to Trearddur, and then take
are still some fantastic dunes and dune slacks in the the minor coast road through Penrhosfeilw, parking at
treeless south-eastern section of the Warren. Dune Gors-goch. (RSPB)
Helleborine (once again a British endemic) is a spe-
ciality, as are Northern and Early Marsh Orchids, 3. Cemlyn SH 331932
Common Twayblade and Marsh Helleborine. On the north coast of Anglesey, a shingle storm
Access is from the Menai Bridge along the A4080 to beach has sealed off the entrance to Cemlyn Bay
Aberffraw. There are several car parks along this road to form a saline lagoon. There are Autumn Lady’s-
around Newborough, and footpaths crossing the dunes tresses on the more stable parts of the beach.
to the foreshore. (NNR, CCW) Off the A5025 between Dyffryn and Cemaes, on the
minor road from Tregele. There are car parks at both
2. South Stack Cliffs SH 207823, SH 218805 the eastern (Traeth Cemlyn) and western (near Bryn
Situated three miles west of Holyhead on Holy Aber) ends of the shingle ridge. (NT, North Wales
Island at the western extremity of Anglesey, this WT)
area has some of the finest coastal cliff scenery in
Wales. Inland, there are areas of maritime heath- 4. Cors Goch SH 504817
land rising to 853 feet at Holyhead Mountain, with A large and diverse reserve with areas of fen, heath
a separate area of heath to the south at Penrhos and grassland. The orchids here include Early Purple,
006 systematic.indd 452 29/1/09 12:49:32

WALES l 453
Green-winged, Early Marsh and Northern Marsh limestone pavement, with Autumn Lady’s-tresses on
Orchids, Common Fragrant and Lesser Butterfly the short turf capping.
Orchids, and Marsh Helleborine. There are several From Llandudno, Marine Drive encircles the head and
splendid fens on Anglesey, but this is the only one a minor road bisects the plateau, while a tramway also
with open access and with footpaths and a board- climbs to the top.The whole area is a Country Park.
walk. For information and permits for the others
contact the CCW. WRExHaM
Cors Goch is off the A5025 turning to Llanbedrgoch.
Parking is available in the lay-by. (North Wales WT, 10. Marford Quarry SJ 357560
CCW) An old sand and gravel quarry, now grassed over
and developing an interesting flora, which includes
GWYnEDD Common Spotted, Pyramidal and Bee Orchids.
North of Wrexham on the B5445, between Rossett
5. Caeau Tan y Bwlch SH 431488 and Gresford west of the village of Marford. The re-
A sloping calcareous grassland site with wet flushes; serve is on Springfield Lane; park beside the railway
more than 2,000 Greater Butterfly Orchids can be bridge. (North Wales WT)
seen in some years. Heath Spotted and Common
Spotted Orchids also occur. CaRDIGanSHIRE
Off the A499 Caernarfon to Pwllheli road, near Clyn-
nog-fawr on the Lleyn Peninsula. (North Wales WT, 11.Ynyslas Sn 610941
Plantlife) Lying on the southern side of the Dyfi estuary, the
dunes here are excellent, with Marsh Helleborine,
6. Morfa Harlech SH 555350 Early Marsh Orchid (including the red-flowered
One of two major sand dune systems near Harlech, subspecies coccinea), Northern Marsh, Southern
Morfa Harlech has a rich flora. There are various Marsh, Pyramidal and Bee Orchids.
orchids present, including Early Marsh Orchid of the Leave the A487 from Machynlleth onto the B4353,
red-flowered subspecies coccinea, and also Marsh and turn north at Ynyslas onto the minor road to the
Helleborine. dunes. There is a car park on the sand beside the In-
Morfa Harlech can be reached by turning off the A496 formation Centre. (NNR, CCW)
to the public car park at Harlech and walking along
the beach. (NNR, CCW) POWYS
7. Morfa Dyffryn SH 560250
The second of the dune systems between Harlech 12. Lake Vyrnwy SJ 015191
and Barmouth, Morfa Dyffryn has extensive areas The largest artificial lake in Wales, fringed by scrub
of dune slack with Early Marsh Orchid of the sub- woodland and meadows with heather and grass
species coccinea, Marsh Helleborine and Green- moorland on the slopes. There are large areas of
flowered Helleborine. conifer plantation and deciduous woodland too.
Morfa Dyffyn can be reached from public car parks Heath Spotted Orchid is locally common and Less-
at each end of the reserve, south of Llanbedr on the er Twayblade occurs, although it is hard to find; look
A496. (NNR, CCW) on damp north-facing slopes under heather.
Northwest of Welshpool, off the B4393 from Llanfyllin
8. Roman Steps SH 500775
to Llanwddyn, from where the road continues on to
This area of hill pasture and moorland is well known
circumnavigate the lake. The Information Centre is on
for its population of Lesser Twayblades. Look for
the minor road 100 yards south of the west end of the
these orchids in places where the footpath passes
through stands of heather. dam. (RSPB)
From the B4573 at Harlech follow the minor roads to 13. Dyfnant Meadows SH 998155
Cwm Bychan (SH 647314). Then follow the Roman A typical traditionally managed upland pasture, com-
Steps footpath for around 1 mile. prising eight fields, with acidic bogs and flushes con-
taining large numbers of Heath Spotted Orchids.
COnWY Near Dyfnant (in Dyfnant Forest), off the B4395 at
Hendre, then on minor roads and forest tracks. Park by
9. Great Orme SH 780832 the reserve gates. (Montgomeryshire WT)
Lying immediately north of Llandudno, this massive
limestone headland covers around two square miles 14. Llanymynech Rocks SJ 267218
and rises to 675 feet, forming the eastern boundary An outcrop of limestone, part of an old quarry with
of Conwy Bay. Dark-red Helleborine grows on the some woodland and a rich grassland flora; several
006 systematic.indd 453 29/1/09 12:49:33

454 l SITE GUIDE
orchids occur including Greater Butterfly, Early Pur- Just outside St David’s on the A487 to Fishguard, with
ple, Bee and Pyramidal Orchids. footpaths leading from the road. There is a small car
Near Llanymynech, between Oswestry and Welshpool, park on the western boundary at SM 772275. (NT)
off the A483, with a footpath from the small village
20. St David’s Head SM 734272
of Pant. Park at the end of the cul-de-sac called Pant
The walks along the Pembrokeshire Coast Path
Underhill Lane. The reserve is on the border between
close to St David’s encompass cliff grassland and
England and Wales. (Shropshire WT)
coastal heathland, with good displays of Common
15. Elan Estate Sn 930652 Spotted, Heath Spotted and Pyramidal Orchids.
A vast mosaic of moorland, blanket bog, Sessile Common Twayblade and Lesser Butterfly Orchid
Oak woodland, conifer plantations, rivers and res- can also be found here.
ervoirs around the Rivers Elan and Claerwen. In the Access is from St David’s on the B4583 to Whitesand
river valleys the meadows have Common Fragrant, Bay.
Heath Spotted and Greater Butterfly Orchids; very
much harder to locate is Bog Orchid in the upland CaRMaRTHEnSHIRE
plateau bogs.
West of Rhayader on the B4518; the reservoirs are 21. allt Rhyd y Groes Sn 760480
well signposted. There is a Visitor Centre with full de- Ancient woodland on the slopes of a rocky river
tails of the 80 miles of leafleted walks and nature valley; Greater Butterfly Orchids can be found in the
trails, and a Countryside Ranger service. adjacent meadows.
The site is in the Cambrian Mountains west of Llanwr-
16. nant Irfon Sn 840550 tyd Wells, on minor roads from Rhandirmwyn (off the
A mixture of upland habitats in a spectacular rocky A483) to the A482 Lampeter road. There are various
valley with some beautiful Sessile Oak woodland. paths from the minor roads. (NNR, CCW)
There are also extensive areas of pasture, meadows,
flushes and upland streams with Early Purple and 22. Pembrey Burrows SS 415007
Common Fragrant Orchids, among others. Dunes and a forest of Corsican Pines on former
On the minor road from Abergwesyn to Terrain. Nant dunes; the wide, grassy and sheltered rides are an
Irfon is part of a large protected area, which includes excellent habitat for Bee Orchid and Marsh Hel-
the National Trust reserve of Abergwesyn Common leborine.
and another NNR at Claerwen.There is general access Northwest of Swansea, off the B4311 at Burry Port.
along paths from either end of the valley. (NNR) Car parking is at Pembrey Country Park.
17. Burfa Bog SO 275613
The wet woodland, grasslands and mire have large GLaMORGanSHIRE
colonies of Heath Spotted Orchids that produce a
marvellous display in summer. 23. South Gower Cliffs SS 470844
Off the B4362 Walton to Ditchyeld Bridge road, near Large areas of the Gower Peninsula are protected
Presteigne. Park on the road verge. (Radnorshire as nature reserves or are in National Trust owner-
WT) ship. There are a wide variety of habitats; the South
Gower Cliffs Reserve contains interesting cliff-top
18. Pwll y Wrach SO 165326 and old dune grassland, with Green-winged and
Deciduous woodland beside the River Enig with Early Purple Orchids.
Early Purple and Bird’s-nest Orchids among the in- Access to the Gower is via Swansea or via Junction 47
teresting flora. of the M4. There is car parking at the beach car park
Pwll y Wrach is in the Brecon Beacons National Park at Port Eynon. (WT)
off the A479 Talgarth to Abergavenny road, southeast
of Talgarth. Park at the reserve. (Brecknock WT) 24. Oxwich Bay SS 506870
Oxwich Bay, on the south shore of the Gower Pen-
PEMBROKESHIRE insula, contains large areas of dunes and dune slacks,
woodland on limestone and freshwater marsh.There
19. Dowrog Common SM 775273 are a good number of orchid species with Pyramidal
Some fantastic heathland and bog, grassland, willow and Southern Marsh Orchids, the coccinea subspe-
carr and pools. With the neighbouring commons of cies of Early Marsh Orchid, Marsh Helleborine and,
Tretio and Waun Fawr, this is a large and botanically more rarely, Bee Orchid and Green-flowered Hel-
rich area. There are good colonies of Lesser Butter- leborine in the dune slacks, and Broad-leaved Hel-
fly and Heath Spotted Orchids, and Southern Marsh leborine in Nicholaston and Oxwich Woods.
Orchids in the wetter parts. Oxwich village is on a minor road south off the A4118,
006 systematic.indd 454 29/1/09 12:49:33

WALES l 455
with open access to the dunes and beach and marked limit) on a steep slope beside a river gorge. Little
paths in Nicholaston and Oxwich Woods. (NNR, grows on the ground, but Bird’s-nest Orchid can be
CCW) found in some numbers.
West of Abergavenny on the A465, accessed via foot-
25. Crymlyn Bog SS 694947
paths from minor roads which run from Clydach to
Just north of Swansea docks, this is the largest area Llanelly Hill. (NNR, CCW)
of lowland fen in Wales and is of international im-
portance for its valley mire communities. The veg- 29. Pentwyn Farm ST 523095
etation includes rich fen, reedbed and old pasture. A small hill farm reserve of unimproved hay mead-
There are Southern Marsh, Common Spotted and ows with great views towards the Forest of Dean.
Heath Spotted Orchids and hybrids. Several orchid species can be found, including
Access is via minor roads leading off the A4217 and Common Twayblade, Early Purple, Greater Butter-
A483 around Kelvey Hill. There is a Visitor Centre on fly, Green-winged and Common Spotted Orchids
the west side. (NNR, CCW) among the meadows and lanes.
Located at Penallt near Monmouth, the reserve is adja-
26. Kenfig SS 780820 cent to the Bush Inn with parking. (Gwent WT)
North of Porthcawl, this extensive area of calcare-
ous dunes in various stages of development is one 30. Springdale Farm ST 401992
of the richest orchid sites in Wales. The speciality is Woodland and grassland on limestone. The woods
Fen Orchid (of the broad-leaved variety ovata). This contain Early Purple Orchid, Common Twayblade
grows in the younger dune slacks and has been in and Broad-leaved Helleborine, and there are also
decline in recent years due to a lack of suitable new many Common Spotted Orchids.
slacks. It is hard to find; the best way is to contact Park beside the almshouse on the minor road south
the Information Centre in advance to check flower- between Usk and Llantrisant. (Gwent WT)
ing dates, which can vary a lot (any time from early 31. newport Wetlands ST 334834
June onwards), and to make sure someone will be A new reserve on the edge of Newport with reed-
on hand to give precise directions. Other notable beds and wet grassland. Heath Spotted and South-
species include a few Green-flowered Helleborines, ern Marsh Orchids can be seen from the many
and also Broad-leaved Helleborines of the unusual paths.
subspecies neerlandica (known as ‘Dutch Hellebo- The car park is on West Nash Road between Nash
rine’), although this is erratic in its appearances and village and Uskmouth Power Station. (NNR, CCW)
hard to find. Other orchids include Marsh Hellebo-
rine, Early Marsh Orchid (including the subspecies
coccinea), Southern Marsh, Pyramidal and Green-
winged Orchids, and Common Twayblade.
Kenfig can be reached from Junction 37 of the M4
and is signposted from North Cornelly, Pyle and Porth-
cawl. (Bridgend County Borough Council, Tel: 01656
743386; NNR)
27. Merthyr Mawr SS 870771
The newest NNR in Wales is adjacent to Kenfig. It
has sand, perched sand dunes on top of limestone,
and limestone outcrops, pools and scrub – a vast
and interesting area. The dune slacks hold the usual
complement of orchids, including large numbers of
Marsh Helleborines, and Autumn Lady’s-tresses.
The reserve has open access, with parking at Can-
dleston Castle at the end of a minor road through Mer-
thyr Mawr village, located off the B4265 southeast of
Bridgend. It can also be accessed from the beach car
park at Newton, Porthcawl.
MOnMOuTHSHIRE
28. Cwm Clydach Woodlands SO 207123
Important beechwoods (close to their northern
006 systematic.indd 455 29/1/09 12:49:33

456 l SITE GUIDE
Scotland
1
10
3
7
13
11
12
6
14
20 15
18 19 21
24
16
17
25
4
28
27
22 26
23
Shetland Highland 15 St Cyrus 23 Feoch Meadows

1 Lumbister 7 Beinn Eighe and Torridon argyll and Bute Fife
8 Inchnadamph 16 Ballachuan Hazelwood 24 Tentsmuir Point
Orkney
9 Invernaver 17 Knapdale
2 Birsay Moors and 10 Talich East Lothian
Cottascarth Perth and Kinross
11 Glen Affric 25 Aberlady Bay
18 Black Wood of Rannoch
Outer Hebrides 12 Abernethy Forest and 19 Keltneyburn Dumfries and Galloway
3 Balranald Loch Garten 20 Killiecrankie 26 Grey Mare’s Tail
Inner Hebrides Moray angus Borders
4 Killinallan Dunes, Islay 13 Culbin Forest 21 Glen Cova 27 Gordon Moss
5 Mull aberdeenshire ayrshire 28 St Abb’s Head
6 Tokavaig Wood, Skye 14 Glen Tanar Estate 22 Auchalton Meadow
006 systematic.indd 456 29/1/09 12:49:34

SCOTLAND l 457
Early Marsh and Northern Marsh Orchids, Com-

SHETLanD ISLES mon Spotted and Common Fragrant Orchids and
1. Lumbister Hu 485967 Common Twayblade. The heaths on either side of
A series of lochs among heather moorland and sea the lane contain Lesser Butterfly Orchid and Heath
cliffs. This reserve covers over 4,000 acres; Heath Spotted Orchid. A small colony of Marsh Hellebo-
Spotted Orchids are easy to spot, but Lesser Tway- rines is a speciality here.
blade is very inconspicuous here. On the northern coast of Islay beside Loch Gruinart.
Located west of the A968 and northeast of Whale There is limited parking beside the kissing gate.
Firth; access to the site is from the lay-by just north-
west of Mid Yell on the Island of Yell, north of Mainland. 5. Mull
The Daal of Lumbister, a steep narrow gorge, makes a Mull is a diverse island, with mountains, sea cliffs
good walk. (RSPB) and dune habitats – and great wildlife. Thirteen
species of orchid have been recorded here. Broad-
ORKnEY ISLES leaved Helleborine and Early Purple Orchid can be
found in woodland near Tobermory Lighthouse,
2. Birsay Moors and Cottascarth HY 368197 while pulchella Early Marsh Orchids occur around
Heather moorland with dry and wet heath. This Loch Spelve in the south. Other species to look
large area includes the raised mire of Glim’s Moss for include Small White, Greater Butterfly, Lesser
and a calcareous valley mire known as the Dee of Butterfly, Common Fragrant, Common Spotted and
Durkadale. Orchids found here include Lesser Tway- Northern Marsh Orchids.
blade, and Heath Spotted, Northern Marsh and Mull can be accessed by ferry from the mainland at
Early Marsh Orchids. Oban.
On Mainland, on both sides of the B9057.The entrance
to Cottascarth is north of Finstown off the A966, sign- 6. Tokavaig Wood, Skye nG 620120
posted ‘RSPB’. Park in the farmyard. (RSPB) This wood is on the southern shore of Loch
Eishort on a wide variety of rock types and sup-
OuTER HEBRIDES port a varied flora. Lesser Twayblade grows in
the sandstone and quartzite areas; limestone ar-
The well-protected machair of the north coast of eas hold Frog Orchid, Dark-red Helleborine and
North Uist is home to the endemic Hebridean Marsh Common Twayblade.
Orchid. Other species typical of the machair include The Isle of Skye can be reached from the A87, crossing
‘Hebridean’ Spotted Orchid (Common Spotted Orchid at the Kyle of Lochalsh. The woods are off the A851
of the hebridensis subspecies), Northern Marsh Or- towards Ord and Tokavaig. (NNR)
chid, Early Marsh Orchid of both the pink-flowered
incarnata and red-flowered coccinea subspecies, HIGHLanD
Lesser Butterfly Orchid, Frog Orchid and Common
Twayblade. 7. Beinn Eighe and Torridon nG 950621
3. Balranald nF 714699 Some of the finest mountain scenery in Scotland,
An area of machair and dunes interspersed with with mountain habitats, grassland, moorland, cliffs
lochs and marshes, on the west coast of North and woodland. There is a rich flora in places with
Uist. Predominantly grazing land, the large stretches Creeping Lady’s-tresses in the Caledonian pine-
of machair are herb-rich, with Frog Orchid; the woods.
marshes have Northern Marsh and Early Marsh Between Loch Torridon and Loch Maree; enclosed by
Orchids. the A896 and A832, northwest of Kinlochewe.There is
West of the A865 between Tigharry and Paiblesgarry, a Visitor Centre with information on walks. (NNR)
northwest of Bayhead; follow signposts for Hougharry.
The cottage at Goular has information and maps. 8. Inchnadamph nC 250220
(RSPB) This is Scotland’s largest area of limestone, with
limestone pavements and outcrops at the head of
InnER HEBRIDES Loch Assynt. Species of interest include Dark-red
Helleborine, Common Fragrant, Frog and Small
4. Killinallan Dunes, Islay nR 3071 White Orchids.
These low dunes, which stretch for a few hundred To the east of the A837 at the eastern end of Loch
yards behind the beach, are good for Pyramidal and Assynt, east of Lochinver and north of Ullapool.
Frog Orchids, with small numbers of Early Purple, (NNR)
006 systematic.indd 457 29/1/09 12:49:34

458 l SITE GUIDE
9. Invernaver nC 681612 Davan and Kinord. Among several species of orchid

Seashore, dunes, machair and moorland by the are Lesser Twayblade, Creeping Lady’s-tresses and
mouth of the River Naver, this area has some of the Heath Fragrant Orchid.
most interesting plant communities in the whole of Off the B976 just south of Aboyne. There is a Visi-
Scotland. Small White, fragrant (probably Common tor Centre across the Water of Tanar from Braeloine.
Fragrant) and Greater Butterfly Orchids and Dark- (NNR)
red Helleborine can be found.
15. St Cyrus nO 743635
On the north coast of Sutherland around the village of
A wonderful part of Montrose Bay, there are dunes
Bettyhill. Access from Invernaver village, off the A836.
and slacks, and coastal grasslands and cliffs with a
Slightly to the west at Durness, the limestone cliffs
rich flora, including Northern Marsh, Heath Fragrant,
have several orchid species too.
Heath Spotted and Lesser Butterfly Orchids.
10. Talich nH 850786 North of Montrose off the A92.There is a car park and
An area of woodland, meadow and marsh on the Visitor Centre just south of St Cyrus. (NNR)
bed of a former loch. The wet pastures contain
Heath Fragrant, Heath Spotted, Early Purple and aRGYLL anD BuTE
Lesser Butterfly Orchids.
North of Invergordon on the A9, at Rhynie off the 16. Ballachuan Hazelwood nM 763146
B9165. (Scottish WT) An abandoned hazel coppice with a good ground
flora. Heath Fragrant and Northern Marsh Orchids,
11. Glen affric nH 235240 and Sword-leaved Helleborine can be found.
One of the most attractive valleys in Scotland, there At the southern tip of Seil Island, west of the A816. Ac-
are remnants of Caledonian pine forest, grassland cess from Ballachuan Farm on the B8003 near Cuan.
and lochs, with Northern Marsh and Lesser Butter- (Scottish WT)
fly Orchids.
Southwest of Inverness, on a minor road into the Glen 17. Knapdale nR 720835
from Cannich on the A831.There are several car parks An area of ancient Atlantic oak woodland and conif-
and footpath information. (NNR). erous plantation beside Loch Sween, Knapdale is a
good site for Sword-leaved Helleborine.
12. abernethy Forest and Loch Garten At the head of Loch Sween, at the north end of the Kin-
nH 980188 tyre Peninsula. Off the B8025 to Tayvallich. (Scottish
Pine forest, lochs and wet moors – part of a rem- WT)
nant of Caledonian pine forest. A large area with
an interesting flora, there are several orchid species, PERTH anD KInROSS
including Creeping Lady’s-tresses and Lesser Tway-
blade. 18. Black Wood of Rannoch nn 570536
Off a minor road between Boat of Garten and Nethy- The most extensive patch of original Caledonian
bridge. Park at the RSPB centre. (RSPB) pine forest in Perthshire, with Lesser Twayblade and
Coralroot Orchid present.
MORaY West of Pitlochry on the B8019 and B846, to the
south side of Loch Rannoch; there are two car parks
13. Culbin Forest nH 997614 with information boards.
This vast area stretches from Nairn in the west to
19. Keltneyburn nn 771496
Findhorn in the east. There are sand dunes which
Keltneyburn is a steep wooded gorge with an adja-
have been extensively planted with Corsican and
cent herb-rich meadow, Balchroich Meadow. There
Scots Pines, which hold Creeping Lady’s-tresses,
are eight species to see here, including Bird’s-nest,
Lesser Twayblade and Coralroot Orchid.
Greater Butterfly, Heath Fragrant and Small White
Off minor roads north of the A96.There are several car Orchids.
parks in the area. Turn west off the B846 c. 5 miles west of Aberfeldy
towards Fortingall, then north off this minor road, just
aBERDEEnSHIRE west of Keltneyburn bridge, on a rough track to the
reserve. (Scottish WT)
14. Glen Tanar Estate nO 480964
The Dee’s wooded valley has areas of relict Scots 20. Killiecrankie nn 910620
Pine and birch woodland. The surrounding slopes This site includes oak woodland, farmland, birch-
contain bog and moorland, and extend to the peak clad crags and moorland with wet lime-rich areas.
of Beinn a’Bhuird. In the valley bottom are Lochs The boggy areas have Northern Marsh Orchid,
006 systematic.indd 458 29/1/09 12:49:35

SCOTLAND l 459
and Lesser Twayblade can be found amongst the A large bay to explore, with many footpaths.
heather. Bird’s-nest Orchid occurs in the wood- Northeast of Edinburgh. Access via the A198 with a
land. car park just outside the village of Aberlady. (NTS)
Overlooking the famous Pass of Killiecrankie, west
of the A9 to the north of Pitlochry. The reserve is DuMFRIES anD GaLLOWaY
signposted from the centre of Killiecrankie village.
(RSPB) 26. Grey Mare’s Tail nature Reserve
nT 185145
Famous for its waterfall, the botanical richness of this
anGuS mountainous landscape should be explored. There
21. Glen Clova nO 330730 are upland heaths, blanket bog, rocky slopes, a loch
and wonderful scenery. It is a good place to search
One of the five Glens of Angus, Glen Clova is
for Lesser Twayblade under the heather.
popular with walkers. Within it grow Heath Fra-
In the Moffat Valley, northeast of Moffat on the A708
grant, Small White, Frog and Common Spotted
Moffat to Selkirk road. There is a Visitor Centre and
Orchids.
car park. (NTS)
Access from the B955, to the north of the village of
Kirriemuir. Parking is available at Clova.
BORDERS
aYRSHIRE 27. Gordon Moss nT 635425
The largest area of semi-natural woodland in the
22. auchalton Meadow nS 335036 Borders, this site consists of dense birchwoods on
These meadows on the site of old limestone work- peat moss. Rich in plants, there are Coralroot Or-
ings contain a good orchid flora, with Frog and chids, Lesser Butterfly Orchids and several other
Northern Marsh Orchids, Lesser and Greater But- species to be found.
terfly Orchids, and Common Twayblade. Just outside Gordon, south of the A6105 Gordon to
Park off the B741 near Broomland Cottage, between Earlston road. Park at the reserve entrance on this
Girvan and Maybole. (Scottish WT) road. (Scottish WT)
23. Feoch Meadows nx 263822 28. St abb’s Head nT 914692
Traditionally managed herb-rich meadows which As well as great coastal scenery and seabird colo-
have retained a good flora, including Greater and nies, the grasslands along the top of the cliffs have
Lesser Butterfly, Heath Fragrant, Frog and Small many Early Purple Orchids.
White Orchids. From the A1107 take the B6438. This leads to the
Off the A714 Newton Stewart to Barrhill Road at car park at Northfield Farm. (Scottish WT, NTS,
Killantringan. (Scottish WT) NNR)
FIFE
24. Tentsmuir Point nO 500270
Tentsmuir Forest lies on the southern bank of the
Tay Estuary across the river from Dundee. There is
an extensive dune system, one of the fastest grow-
ing in Britain, with a rich flora that includes a very
few Coralroot Orchids in the dune slacks.
Access from Tayport near to the B945.Walk from here
along the beach, or park at the Forestry Commission
car park at Kinshaldy Beach. (FC, NNR)
EaST LOTHIan
25. aberlady Bay nT 465801
The heathland and sand dunes around Aberlady
Bay hold a rich flora with several orchid species,
including Pyramidal, Common Spotted, Frog and
Heath Spotted Orchids, and Common Twayblade.
006 systematic.indd 459 29/1/09 12:49:35

460 l SITE GUIDE
Ireland (Republic of Ireland and northern Ireland)
28 29
30
26 31
27
32
22
33
34
21 25
24
20 23
36 35
19
18
17
16
14 12
15
11 10
2 13
9 7
8 6
1 4
3
Co. Kerry 13 Coole/Kilcolgan/Ardrahan 27 Lough Beg

1 Killarney National Park 14 Lough Corrib Co. antrim
Co. Clare 15 Inishmore Island 28 The Giant’s Causeway
2 The Burren National Park 16 Slyne Head Peninsula and 29 Carrick-a-rede and Whitepark Bay
Aillebrack 30 Fairhead and Murlough Bay
Co. Waterford
17 Connemara National Park 31 Slieveanorra
3 Tramore Dunes
Co. Mayo 32 Shane’s Castle
Co. Wexford
18 Lough Carra 33 Lough Neagh
4 Bannow Bay
19 Urlaur Lakes Co. Down
5 Cahore Polders and Dunes
20 Lough Conn and Lough Cullin 34 Belfast Harbour Reserve
Co. Wicklow 21 Blacksod Bay 35 Killard Point
6 Wicklow Mountains National
Co. Sligo 36 Murlough NR
Park
22 Mullaghmore
7 Bray Head
Co. Laois Co.Monaghan
8 Ballyprior Grassland 23 Kilroosky Lough Cluster
9 Clonaslee Eskers and Derry Bog Co. Fermanagh
Co. Galway 24 Crom Estate
10 River Shannon Callows 25 Castle Coole
11 Glenloughaun Esker Co. Londonderry
12 Rahasane Turlough 26 Umbra
006 systematic.indd 460 29/1/09 12:49:35

IRELAND l 461
CO. KERRY CO. WaTERFORD

1. Killarney national Park R 970860 3. Tramore Dunes S 580005
The mountainous park to the south and west of A series of fixed dunes and dune slacks with Bee
Killarney around Lough Leane has good bog and Orchids present.
moorland vegetation, and there are extensive South of Waterford on the R675. Park in the town of
woodlands around the famous Lakes of Killarney. Tramore.
There are many interesting orchids and Irish Lady’s-
tresses has been recorded here. CO. WExFORD
The National Park Visitor Centre is at Muckross
House, at Muckross on the N71, where information 4. Bannow Bay S 804064
on various walks is available. A mosaic of sand dune habitats on both sides of the
estuary leading into Bannow Bay. There are damp
slacks where both Bee and Pyramidal Orchids grow.
CO. CLaRE On the south coast between Wexford and Waterford;
the R734 leads to Fethard on the west side, and the
2. The Burren R 324954 R736 leads to Newtown on the east side.
One of Europe’s finest areas of limestone pave-
ment, and of outstanding botanical interest. There 5. Cahore Polders and Dunes T 220462
are turloughs (seasonal loughs), fens and some An important wildlife site on the southeast coast.
small stands of woodland, with dunes covering a A 2/2 mile-long sand dune system extends south
large area on the west coast of Co. Clare around from Cahore Point; behind the dunes there are
Lisdoonvarna and Kilfenora. The Burren is bisected areas of grassland, wetland and drainage channels.
by the N67, and many other minor roads allow Pyramidal Orchid occurs on the fixed dunes at the
easy exploration. Of the 30 species of orchid found northern end. The newer dunes to the south also
in Ireland, 23 have been recorded in The Burren. have a rich flora, and Marsh Helleborine has been
Dense-flowered Orchid is the real speciality. This recorded.
favours short, closely grazed turf; it is widespread Between Wexford and Arklow, south of Courtown.Take
but hard to spot. Another speciality is Early Marsh minor roads off the R742 to reach the site.
Orchid of the spotted-leaved subspecies cruenta
(‘Flecked Marsh Orchid’), which is found around CO. WICKLOW
some of the loughs in the east of the region. Early
Purple Orchids are abundant and spectacular, and 6. Wicklow Mountains national Park
Heath and Common Spotted Orchids are wide- T 099980
spread (with white-flowered Common Spotted This National Park covers much of upland Wicklow,
Orchids of the variety okellyi common here). Irish extending to Blessington Reservoir in the west, Var-
Marsh Orchid is scattered and sometimes found in try Reservoir in the east, Cruagh in the north and
dry as well as wet grassland habitats. Fly Orchids Lybagh in the south; the Park also includes Glen-
occur in open habitats, on rocky hillsides and in fens dalough Wood Nature Reserve. There is heathland,
around the turloughs; Common Twayblade and, lat- blanket bog and upland grassland, small rivers and
er in the season, Dark-red Helleborine and Marsh scattered lakes, with a rich flora; the Liffey Head
Fragrant Orchid can also be found. blanket bog is one of the best of its kind in eastern
The Burren National Park covers a fairly small area Ireland. Small White Orchid and Bog Orchid occur
in the east; the green road at R 304945 gives good here but are difficult to find.
access. Other productive areas include Loch Gealain, The Visitor Centre is in Glendalough Valley off the R756
with numerous ‘Flecked Marsh Orchids’, Lough Bun- Green Road close to the Upper Lough (where there is
ny, and the pastures near the coast at Poullsallagh, a car park). It is popular walking country with many
which have some Dense-flowered Orchid colonies. footpaths taking in the major habitat types.
Keelhilla at Slieve Carron on the northeastern edge 7. Bray Head O 284174
of The Burren is also a good area; Fly and Dense- This coastal site is a plateau of high ground with
flowered Orchids occur there (roadside parking for rocky knolls and sea cliffs. Heath is the principal
a few cars). The grid reference above is for Mul- habitat but in some areas there is calcareous grass-
laghmore (National Trust for Ireland) in the centre land with several species of orchids, including Bee,
of the region, but the whole of the Burren is worth Fragrant, Pyramidal and Common Spotted Orchids
exploring. and Common Twayblade.
006 systematic.indd 461 29/1/09 12:49:35

462 l SITE GUIDE
Off the R761 out of Bray; walk from Bray or Greystones. Pyramidal, Heath Spotted, Common Spotted, Great-
The site is signposted from the road. er Butterfly, Fragrant and Fly Orchids occur here.
Similar habitats occur just slightly north around Ca-
CO. LaOIS ranavoodaun turlough south-east of Kilcolgan. The
dry, calcareous grassland among the limestone pave-
8. Ballyprior Grassland S 572924 ment here holds Early Marsh, Lesser Butterfly, Fra-
Located at the north end of the Castlecomer Plateau grant and Dense-flowered Orchids, Broad-leaved
and largely underlain by limestone, this site contains Helleborine and Autumn Lady’s-tresses. On the
much orchid-rich calcareous grassland, with Early Ardrahan limestones, the Ballinderreen turlough has
Purple, Fragrant and Common Spotted Orchids. similar species with the addition of Fly Orchid.
Green-winged Orchid has also been recorded here. Coole Lough (and Visitor Centre) is off the N18, north-
The site is 2 1/2 miles south of the village of Stradbally, west of Gort. (M 430040). Caranavoodaun turlough is
which is on the N80 between Portlaoise and Carlow. off the N18 SE of Kilcolgan (M 4317). Ballinderreen
turlough is S of Ballinderreen on the N67 (M 3914).
9. Clonaslee Eskers and Derry Bog S 265110
A series of glacial moraines and eskers, including 14. Lough Corrib M 121429
Derry Bog. The wide ranging habitats support an The second largest lake in Ireland, this is surround-
interesting flora including Fragrant and Fly Orchids. ed by some excellent habitats, including spring-fed
Three miles west of Clonaslee, northwest of Portlaoise fen, raised bog, limestone pavement, woodland and
on the R422. orchid-rich grassland. Pyramidal, Common Spotted,
Early Purple, Frog, Fragrant and Greater Butterfly
Orchids and Marsh Helleborine can all be found.
CO. GaLWaY There are good populations of Irish Lady’s-tresses
10. River Shannon Callows n 014304 around the lough and on the Doorus Peninsula.
Callows are seasonally flooded wet grasslands. To the north of Galway City, with information points at
These form the largest area of lowland semi-natural Oughterard on the N59 and Cong on the R345; there
grassland in Ireland. They have a good range of char- are various picnic spots around the Lough.
acteristic species, varying according to their toler- 15. Inishmore Island L 830090
ance to flooding. There are some areas of limestone The largest of the three Aran Islands, Inishmore is
pavement and calcareous grassland where Green- geologically an extension of The Burren in Co. Clare.
winged Orchid occurs, among other species. There are some fantastic cliffs, sand dunes, machair,
Along the River Shannon between the towns of Athlone orchid-rich grassland and limestone pavement, as
and Portumna;The NationalTrust for Ireland has a reserve well as ancient meadows surrounded by stone walls.
at Mongan Bog (N 032318) and an information point The rich flora includes Common Twayblade, Early
with parking at Clonmacnoise National Monument. Purple, Common Spotted and Heath Spotted Or-
11. Glenloughaun Esker M 8226 chids. Dense-flowered Orchid is a speciality, and Bee
A fine example of dry, mostly unimproved, orchid- Orchid occurs in the coastal machair.
rich grassland on an esker. The flora includes large The Aran Islands can be reached by ferry from Ros-
populations of Green-winged and Early Purple saveal or Doolin in Co. Clare and explored by foot.
Orchids. 16. Slyne Head Peninsula and aillebrack
Three miles southwest of Ballinasloe, off the N6, along L 620445
the minor roads to Kilnahown. In the far west of Galway, this low-lying peninsula
12. Rahasane Turlough M 472238 has some extensive areas of machair (particularly
On the course of the River Dunkellin, this turlough good at Mannin Bay and Aillebrack) and sand dunes
is surrounded by damp grassland, limestone out- backed by a mosaic of tiny fields, grassland, heath,
crops and scrubby woodland. A flora similar to that lakes, marshes and fens. The specialities are Dense-
of The Burren occurs here. flowered Orchid and, very locally in spring-fed fens,
Just west of Craughwell, southeast of Galway City on Pugsley’s Marsh Orchid. The dry grassland supports
the N6. Access from the N6, north on a minor road Early Purple and Green-winged Orchids, and both
towards Athenry. Greater and Lesser Butterfly Orchids.
The area lies west of Ballyconneely on minor roads off
13. Coole Lough M 430040 the R341.
An area of low-lying limestone karst with deciduous
woodland, pasture and limestone heath. Espe- 17. Connemara national Park L 670520
cially interesting are the turloughs, with limestone Connemara has mountains, bogs, heaths, grass-
pavement extending to their edges in some places. lands and woodlands. The predominant habitats are
006 systematic.indd 462 29/1/09 12:49:36

IRELAND l 463
blanket bog and heathland. Lesser Twayblade is pres-

ent but, as usual, is hard to find.
CO. SLIGO
Entrance to the park is on the Clifden side of Letter-
22. Mullaghmore G 710580
frack village on the N59.There is a car park and Visitor
This tiny site measures just 100 x 200 yards, but it is
Centre, numerous trails and helpful staff. (NPWS)
extraordinarily orchid-rich; Pyramidal, Fragrant, Frog,
Lesser Butterfly and Northern Marsh Orchids all
CO. MaYO occur, as do Common Twayblade and Marsh Helle-
borine. Common Spotted Orchid, the red-flowered
18. Lough Carra M 190740
coccinea subspecies of Early Marsh Orchid, and
A large, shallow, marl lake surrounded by limestone,
Heath Spotted Orchid are found in their hundreds.
with fens, woodland, grassland and limestone pave-
On a headland off the N15 between Sligo and Bun-
ment. Early Purple and Dense-flowered Orchid
occur. Nearby Lough Mask has Irish Lady’s-tresses doran, on the R279 to Mullaghmore.
along its eastern shore.
Alongside the N84, between Castlebar and Ballinrobe.
CO. MOnaGHan
Access from minor roads around the lough, and park 23. Kilroosky Lough Cluster H 490276
on the east side south of Carrownacon.There are many Lough Kilroosky, Lough Burdautien, Lough Summer-
picnic areas on the east side of Lough Mask on the hill and Dummy’s Lough are a series of marl lakes,
‘scenic road’. low in nutrients and surrounded by fen and species-
19. urlaur Lakes M 520899 rich freshwater marsh. Nearby lakes are surrounded
Three small calcareous lakes, Lough Nanoge, Lough by reed swamp and wet woodland. Many species
Roe and Urlaur Lough, lie in the upper catchment of orchid can be found here, including Marsh Hel-
of the River Lung, a major tributary of the Boyle. leborine.
Swamps and reedbeds fringe the lakes, and in some Northwest of Clones, straddling the border with North-
areas there is flushed, species-rich heath with Lesser ern Ireland.
Butterfly, Pyramidal and Heath Spotted Orchids.
Common Twayblade can also be found. CO. FERManaGH
North of Ballyhaunis, on either side of the N83 near
Urlaur. Access via minor roads.
24. Crom Estate H 455655 - 361245
On the shores of Upper Lough Erne, this is one
20. Lough Conn and Lough Cullin G 216048 of Ireland’s most important conservation areas. Or-
There are fine oak woodlands around the south- chids include Common and Heath Spotted, Early
ern shores of both these lakes, each with a mixed Purple, Fragrant, Bird’s-nest, and Greater and Lesser
shrub layer and good ground flora. Sword-leaved Butterfly Orchids, plus Broad-leaved Helleborine
Helleborine has been recorded here. Lesser Tway- and Common Twayblade.
blade occurs further up the slopes on the blanket The Visitor Centre and car park lie west of Newtown-
bog, while there are important populations of Irish butler on the minor road to Crom, with walks through
Lady’s-tresses around the lake margins. the wetlands and woodlands. (NT)
There is information at the nearby town of Ballina. Pic-
nic sites are on Lough Conn at Crossmolina on the 25. Castle Coole H 378788
N59, and on Lough Cullin beside the R318. The landscaped grounds of Castle Coole have walks
in woodland to Lough Coole, where Common
21. Blacksod Bay F 700330 Twayblade, Bird’s-nest Orchid and Broad-leaved
The Mullet Peninsula and Blacksod Bay form an Helleborine can be found.
important wildlife habitat. There are fixed dunes, Just east of Enniskillen on the A4. (NT)
machair and dune heath, as well as alkaline fens
and coastal habitats. The dunes and machair can be CO. LOnDOnDERRY
found in the north of the peninsula to the west of
Termoncarragh Lough, Tonamace and Cross Lough; 26. umbra C 726358
such habitat also occurs on the eastern shore of A dune system on the north coast, with woodland
the bay, around Doolough, Srah and Dooyork. Or- and scrub behind the beach of Magilligan Strand.The
chids to be found include Lesser Butterfly, Pyramidal, rich flora in the dune slacks includes Early Purple,
Common Spotted and Early Marsh, and Pugsley’s Common Spotted, Northern Marsh, Early Marsh
Marsh Orchid has also been recorded here. (two subspecies, incarnata and coccinea), Pyramidal
Situated in northwest Mayo, Belmullet is on the R313, and Bee Orchids, Common Twayblade and Marsh
from where the peninsula can be explored. Helleborine.
006 systematic.indd 463 29/1/09 12:49:36

464 l SITE GUIDE
Access from the A2 Coleraine to Limavady road; the 32. Shane’s Castle J 111881
site is adjacent to the Umbra level crossing on the A2, Lying on the north shore of Lough Neagh, this
and has roadside parking. The whole Magilligan dune area includes woods, parkland and farmland. Castle
system is worthy of exploration. (Ulster WT) Meadow holds carpets of Common Spotted Or-
chids; there are Broad-leaved Helleborines in the
27 Lough Beg H 9895
woods, and Irish Lady’s-tresses have been recorded
Just north of Lough Neagh, this area holds Irish La-
along the shores of the lough.
dy’s-tresses which can be found in damp grassland
Enter via the park gate 1 mile west of Antrim on the
on the western shore. Early Marsh Orchid (subspe-
A6 Randalstown road. (RSPB)
cies pulchella) also occurs.
A good area to look is between the Ballydermott Road 33. Lough neagh J 050620
and Church Island (H 966953). Ireland’s largest freshwater lough, bounded by
meadows, reedbeds and woodland. There are many
CO. anTRIM nature reserves including the Montiaghs NNR and
Lough Neagh NNR. Irish Lady’s-tresses (albeit much
28. The Giant’s Causeway C 954453 reduced), Lesser Butterfly Orchid and Early Marsh
These famous polygonal columns of layered basalt Orchid (subspecies pulchella) can be found.
are a World Heritage Site. Several species of orchid Situated to the west of Belfast, many minor roads lead
can be found along and near to the North Antrim to the shore with picnic areas.The grid reference is for
Coastal Path and beside paths to the Causeway, the Discovery Centre on Oxford Island where informa-
including Frog, Common Spotted, Heath Spotted, tion can be obtained regarding access and flowering
Northern Marsh and Early Purple Orchids, and Irish times from the staff and the Environment & Heritage
Lady’s-tresses has recently been found here. Service wardens. (Craigavon Borough Council)
On the B146 2 miles from Bushmills. There are car
parks and a Visitor Centre. (NT) CO. DOWn
29. Carrick-a-rede and Whitepark Bay 34. Belfast Harbour Reserve J 398795
D 062450 The reserve and surrounding area hold Bee Orchid,
On the North Antrim Coast east of the Giant’s Common Spotted and Northern Marsh Orchids
Causeway, this rocky outcrop is accessible only and good displays of Dactylorhiza hybrids. Pyramidal
via a rope bridge. The grasslands in this area have Orchid and Common Twayblade may also be found.
Common Twayblade, Early Purple, Greater Butterfly, 3 miles north of Belfast city centre, off the A2 on the
Fragrant, Common Spotted, Northern Marsh and south side of Belfast Lough. Speak to the warden for
Pyramidal Orchids. At nearby Whitepark Bay, the access arrangements. (RSPB)
dunes, grassland and scrub, with limestone cliffs to
the east, support similar species. 35. Killard Point J 610433
On the North Antrim coast road; there are picnic areas, Killard Point marks the southern limit of Strangford
footpaths and a nature trail to Whitepark Bay. (NT) Lough, and has dunes, lime-rich boulder clay grass-
land, scrub, heath and low cliffs. One of the best
30. Fairhead and Murlough Bay D185430 orchid sites in Northern Ireland: Common Tway-
On the north coast 4 miles east of Ballycastle, this blade, and Common Spotted, Heath Spotted, Early
area of rugged cliffs has limestone outcrops along- Marsh, Early Purple and Green-winged Orchids oc-
side a wooded area behind Murlough Bay. Common cur in good numbers and there are also Bee, Frog,
Twayblade, Bird’s-nest, Bee, Pyramidal, Northern Pyramidal and Northern Marsh Orchids in smaller
Marsh and Common and Heath Spotted Orchids
numbers.
all occur.
Approximately 7 miles east of Downpatrick. Take the
Access from the A2, parking at Murlough Bay. There
minor road between Kilclief and Ballyhornan from the
are several paths and viewpoints. (NT)
A2. Park in one of the lay-bys. (NNR)
31. Slieveanorra Forest D 132265
Mixed woodland and moorland; there are forest 36. Murlough nature Reserve J 414351
tracks to the summit of Slieveanorra where there This large area of dunes forms a peninsula that proj-
are interesting plots of peat bog at various stages ects into Dundrum Bay.The rich flora includes Com-
of formation. The bogs contain a range of species, mon Spotted, Heath Spotted, Pyramidal, Bee and
including Lesser Twayblade and Heath Spotted Or- Northern Marsh Orchids, and Common Twayblade.
chid. On the coast, about 3 miles north of Newcastle. Access
Slieveanorra is on the Altarichard road off the Bally- from the A2; park at Dundrum and walk over Down-
money to Cushendun Scenic Route. shire Bridge to the reserve. (NT, NNR)
006 systematic.indd 464 29/1/09 12:49:37

GLOSSARY l 465
Glossary
achlorophyllose lacking the green pigment chlorophyll and therefore unable to photosynthsise.
actinomorphic radially symmetrical; with more than one plane of symmetry.
adventitious buds and roots that appear in abnormal places on the stem.
ancient woodland woodland that has maintained a more or less continuous cover of trees, probably
for thousands of years.
annular ring-shaped.
anther the pollen-bearing, male reproductive organ. In most orchids, the pollen is grouped into two
pollinia.
anther cap in some orchids, such as the helleborines Epipactis, the anther lies on top of the column
and is hinged or stalked. It may be contrastingly coloured.
anthocyanins group of pigments that produce purple or reddish colours.
apomixis reproduction by seed, originating from unfertilised egg cells.
asymbiotic when a symbiotic fungus is absent.
auricles ear-like structures.
autogamy self-pollination with pollen from the same flower (see self-pollinate).
back-cross cross between a hybrid and one of its parent species.
base-rich soil with a high concentration of calcium or magnesium and a pH above 7.0.
bog plant community on wet, acidic peat.
bosses irregular swellings.
bract structure at the base of a flower stalk, varying in size and shape, but often leaf-like.
bulbils tiny, round growths, e.g. along the rim of the leaf of Bog Orchid, which can separate and are
capable of developing into a new plant.
bursicle the pouch-like structure on the column of some orchids that contains and protects the
viscidium (q.v.).
calcareous rich in calcium carbonate, e.g. chalk, limestone or sea shells.
Caledonian woodland ancient pine woodland, a relict of the ‘Forest of Caledon’ that supposedly
once covered Scotland.
capsule the dry seed pod of an orchid.
carapace hardened shell.
caudicle the stalk present in some orchids that attaches the pollinium to the viscidium (qq.v.).
cilia minute, thickened or fleshy hair-like structures.
ciliate with cilia projecting from the margin.
chlorophyll a green pigment, important in photosynthesis, found in discrete organelles (chloroplasts)
in the cells of plants, usually in the leaves.
cleistogamy self-pollination in bud; after which the bud may remain closed or may open.
clinandrium depression on the top of the column, below the anther and behind the stigmatic zone,
in which the pollinia lie.
clone individual of identical genetic make-up to its ‘parent’ that results from asexual, vegetative reproduction.
column specialist structure characteristic of orchid flowers in which the stamens and stigmas are
fused together.
crenate with scalloped margins.
cross-pollinate pollination in which pollen from one flower fertilises another; usually taken to mean
a flower on a different plant.
006 systematic.indd 465 29/1/09 12:49:37

466 l GLOSSARY
decurved curved downwards.

deflexed bent sharply downwards.
deltoid shaped like the Greek letter ‘delta’, i.e. triangular.
diploid having two matching sets of chromosomes. This is the normal state for plant cells.
ectomycorrhiza association of a fungus with the roots of a plant where the fungus forms a layer on
the outside of the roots.
endomycorrhizal association of a fungus with the roots of a plant in which the fungus penetrates
the tissue of the root.
epichile outer portion of the lip, often heart-shaped, in those orchid genera where the lip is divided
into two (e.g. Epipactis, Cephalanthera, Serapias).
epidermis ‘skin’ or surface layers.
epiphyte plant growing on the surface of another plant without receiving nutrition from it.
esker glacial debris, often sands and gravels.
fen plant community on alkaline, neutral or very slightly acidic soil.
filiform thread-like.
flexuous wavy.
eutrophication enriched with plant nutrients, often leading to a luxuriant and eventually stifling
growth of vegetation.
geitonogamy fertilised by pollen from another flower on the same plant.
glandular hair short hair tipped with a small spherical gland containing oil or resin.
hanger a wood on a steep hillside.
herbarium (plural: herbaria) a collection of dried and pressed plant material.
hooded formed into a concave shape resembling a monk’s cowl.
hybrid plant originating from the fertilisation of one species by another.
hybrid swarm population in which the barriers between two species have largely or completely
broken down. Hybridisation is commonplace and at random, producing a population that forms
a continuous range of intermediates between the two parent species.
hybrid vigour when the first generation of hybrids between two species are exceptionally large and
robust.
hyperchromic intensely coloured, with an excessive amount of pigmentation.
hyperresupinate when the ovary and/or pedicel twist through 360° to position the lip at the top of
the flower, e.g. Bog Orchid.
hypha (plural: hyphae) fine, thread-like structures that make up the body of a fungus.
hypochile inner portion of the lip, often cup-shaped, in those orchid genera where the lip is divided
into two (e.g. Epipactis, Cephalanthera, Serapias).
intergeneric hybrid a hybrid whose parents are in two different genera.
internode section of stem between two nodes.
lanceolate narrowly oval, tapering to a more or less pointed tip.
lax loose, not dense.
lignify become hardened and woody.
Leblanc chemical process used to produce washing soda.
lip highly modified third petal of an orchid; also known as the labellum.
lough Irish term for loch.
machair sandy, lime-rich soil with a species-rich sward of short grasses and herbs. The machair is
confined to the coasts of western Ireland and western Scotland.
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GLOSSARY l 467
meadow grassy field from which stock are excluded for at least part of the year so that it can be cut
for hay.
monocarpic flowering once and then dying.
mutualism an intimate relationship between two or more organisms from which all derive benefits.
mycorhizome early stages in the development of the underground rhizome in which the seedling is
nourished entirely by fungi.
mycorrhiza association of a fungus with the roots of a plant in which the fungus may form a layer on
the outside of the roots (ectomycorrhizal, q.v.) or penetrate the tissue of the root (endomycorrhizal,
q.v.).
mucro very short, bristle-like tip.
mycelium the mass of branching filaments that make up the body of a fungus.
mycotrophic acquiring nutrition from fungi.
native growing in an area where it was not introduced, either accidentally or deliberately, by
humans.
node point on a stem from which leaves, flowers or lateral stems grow.
non-sheathing leaf a leaf with its base clasping the stem but not completely encircling it.
ovary female reproductive organ that contains the ovules.
ovule organ inside the ovary that contains the embryo sac, which in turn contains the egg.
pH measure of acidity.
parasitic organism that lives on or at the expense of other organisms.
pasture grassland that is grazed for some or all of the year but not cut.
petals inner row of ‘perianth segments’, one of which is modified to form the lip.
patent projecting more or less at right-angles.
papilla (plural: papillae) small, nipple-like projection.
pedicel stalk of the flower; very short in many orchids, with the cylindrical and sometimes slender
ovary forming the apparent ‘stalk’.
pendant hanging downwards.
peloton a coil-like structure formed by fungi inside the cells of an orchid.
pheromone chemical secreted by an animal, especially an insect, that influences the behaviour or
development of others of the same species.
photosynthesis production of food by green plants. In the presence of chlorophyll and light energy
from the sun, carbon dioxide and water are converted into carbohydrates and oxygen.
phototrophic acquiring nutrition through the process of photosynthesis.
pollen single-celled spores containing the male gametes.
pollinium (plural: pollinia) regularly-shaped mass of individual pollen grains which is transported as
a single unit during pollination; the pollinia are often divided into two.
propagules various vegetative portions of a plant such as a bud or other offshoots that aid in dispersal
and from which a new individual may develop.
protocorm initial stage of development for every orchid formed by a cluster of cells.
pseudobulb swollen or thickened portion of stem, covered in the leaf bases. It fulfils the same storage
function as a bulb or tuber; found in Fen and Bog Orchids and common in tropical species.
pseudocopulation attempts by an insect to copulate with an insect-mimicking flower.
pseudopollen structures in a flower that imitate pollen in order to attract insects, e.g. in the
Cephalanhera helleborines.
reflexed bent back or down.
resupinate when the ovary and/or pedicel twist through 180° to position the lip at the bottom of
the flower.
006 systematic.indd 467 29/1/09 12:49:37

468 l GLOSSARY
reticulation marked with a network of veins.

rhizoid hair-like structure on the surface of a protocorm or mycorhizome that facilitates the entrance
of fungi (i.e. a root hair).
rhizome underground stem that lasts for more than one year from which roots and growth buds
emerge; also a horizontal stem, either growing along the surface or underground.
rostellum a projection from the column that often functions to separate the pollinia from the stigma
and thus prevents self-pollination (the rostellum is actually a modified sterile third stigma). The
rostellum may exude a viscidium (q.v.).
runner horizontal stem that grows along or just below the surface of the soil.
saprophytic plants, fungi, etc. that feed on dead organic matter.
scale leaf a leaf that is reduced to a small scale.
secondary woodland woodland in which the continuity of tree cover has been broken for a substantial
period of time.
secund all facing in the same direction.
self-pollinate pollination of a flower by pollen taken from the same plant; usually used in the context
of autogamy, where pollination is by pollen from the same flower, but sometimes also geitonogamy,
where the pollen comes from another flower on the same plant.
sepal outer row of ‘perianth segments’ that form the protective covering of the bud. In orchids they
are either green or brightly coloured and form a conspicuous part of the flower.
sessile stalkless.
symbiosis an intimate relationship between two or more organisms; formerly used for a
relationship where all participants derive benefits (q.v. mutualism) but now used in a broader
sense to include parasitism
sheathing leaf main leaf on an orchid with a base that completely encircles the stem.
sinus indentation between two lobes on the lip, used especially when describing the flowers of marsh
orchids and spotted orchids in the genus Dactylorhiza.
speculum pattern on the lip, often with a metallic lustre or shine, in the bee and spider orchids, genus
Ophrys.
spur sack-like extension of the base of the lip which contains nectar in some species of orchid.
stamens male reproductive organs of a flowering plant.
staminode sterile stamen that forms a prominent shield-shaped structure within the flower of
Lady’s-slipper.
stigma receptive surface of the female reproductive organs to which the pollen grains adhere.
subsecund almost second, i.e. almost all facing to one side.
sympodial pattern of growth in which the tip of the stem or rhizome either terminates in a flower
spike or dies each year. Growth continues from buds formed at the base of the old stem.
synonym former scientific name.
synsepal structure formed when the two lateral sepals are joined for almost their entire length, found
in Lady’s-slipper.
tetrads group of four pollen grains originating from a single ‘mother’ cell.
tetraploid having four sets of chromosomes.
tubers swollen underground roots or stems, functioning as storage organs.
turbary an area where peat or turf is cut for fuel.
turlough Irish term, used for a seasonal lake on limestone in which the water level may fall dramatically
in summer.
viscidium (plural viscidia) detachable sticky exudation from the rostellum that attaches the pollinia
(sometimes via a short stalk, the caudicle) to a visiting insect (qq.v.).
zygomorphic having only one plane of symmetry.
006 systematic.indd 468 29/1/09 12:49:38

BIBLIOGRAPHY l 469
Sources of Information
and Bibliography
In researching this book we have consulted many books and articles, and a complete list of consulted
works appears below. We would, however, like to acknowledge several works in particular that have
been especially valuable. A great debt is owed to the Wild Orchids of Britain (Summerhayes 1968),
which beautifully summarises knowledge on the orchid flora up to that point. More recent advances,
and especially the matter of subspecies, varieties and hybrids, were expertly presented by D.M.
Turner Ettlinger in Notes on British and Irish Orchids (1997) and Illustrations of British and Irish
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Hill (Sankey 2000), Dorset ( Jenkinson 1991), Hampshire and the Isle of Wight ( Jenkinson 1995),
Scotland (Allan & Woods 1993), Suffolk (Sanford 1991) and Sussex (Lang 2001). Two other works
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139.
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INDEX l 479
Index of Species
aestivalis, Spiranthes 188 dunensis, Epipactis 113
albida, Gymnadenia 236 Dune Helleborine 113
albida, Leucorchis 236 Dwarf Alpine Orchid 416
albida, Pseudorchis 236 Early Coralroot 160
alpina, Chamorchis 416 Early Marsh Orchid 276
AnACAmPtiS 361 Early Purple Orchid 228
anthropophora, Orchis 202 Early Spider Orchid 411
anthropophorum, Aceras 202 ebudensis, Dactylorhiza 325
aphyllum, Epipogium 136 EPiPACtiS 73
apifera, Ophrys 398 EPiPOGiUm 135
atrorubens, Epipactis 85 ericetorum, Dactylorchis 303
Autumn Lady’s-tresses 181 ericetorum, Orchis 303
‘Bertoloni’s Mirror Orchid’ 416 False Musk Orchid 416
‘bertolonii, Ophrys’ 416 Fen Orchid 142
Bee Orchid 398 Fly Orchid 392
bifolia, Platanthera 243 Frivald’s Fragrant Orchid 416
Bird’s-nest Orchid 67 frivaldii, Gymnadenia 416
Bog Adder’s-mouth 152 Frog Orchid 288
Bog Orchid 152 fuchsii, Dactylorhiza 294
borealis, Gymnadenia 263 fuciflora, Ophrys 405
Bracted Green Orchis 288 fuschii, Dactylorchis 294
Broad-leaved Helleborine 92 fuschii, Orchis 294
Burnt-tip Orchid 345 Ghost Orchid 136
Burnt Orchid 345 GOODYERA 165
calceolus, Cypripedium 25 Greater Butterfly Orchid 249
cambrensis, Dactylorhiza purpurella 332 Greater Tongue Orchid 384
CEPHALAntHERA 32 Green-flowered Helleborine 126
Chalk Fragrant Orchid 256 Green-veined Orchid 372
chlorantha, Platanthera 249 Green-winged Orchid 372
coccinea, Dactylorhiza incarnata 276 GYmnADEniA 254
Common Fragrant Orchid 256 HAmmARBYA 151
Common Spotted Orchid 294 Heart-flowered Tongue Orchid 416
Common Twayblade 62 Heart-leaved Twayblade 56
conopsea, Gymnadenia 256 Heath Fragrant Orchid 263
CORALLORHiZA 159 Heath Spotted Orchid 303
Coralroot Orchid 160 Hebridean Marsh Orchid 325
cordata, Listera 56 helleborine, Epipactis 92
cordata, neottia 56 HERminiUm 192
cordigera, Serapias 416 HimAntOGLOSSUm 353
Creeping Lady’s-tresses 166 hircinum, Himantoglossum 354
cruenta, Dactylorhiza incarnata 276 Hooded Ladies’-tresses 174
CYPRiPEDiUm 23 incarnata, Dactylorhiza 276
‘Dutch Helleborine’ 97 incarnata, Dactylorhiza incarnata 276
DACtYLORHiZA 272 insectifera, Ophrys 392
damasonium, Cephalanthera 48 Irish Lady’s-tresses 174
Dark-red Helleborine 85 Irish Marsh Orchid 334
Dense-flowered Orchid 341 Jersey Orchid 363
densiflora, Gymnadenia 267 ‘Lapland Marsh Orchid’ 321
densiflora, Gymnadenia borealis 263 Lady’s-slipper 25
densiflora, Gymnadenia conopsea 267 Lady Orchid 222
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480 l INDEX
lapponica, Dactylorhiza traunsteineri 321 ovata, neottia 62

Late Spider Orchid 405 Pale-flowered Orchid 416
latifolia, Dactylorchis 276 pallens, Orchis 416
latifolia, Orchis 276 paludosa, Hammarbya 152
laxiflora, Anacamptis 363 paludosa, malaxis 152
laxiflora, Orchis 363 palustris, Epipactis 78
leptochila, Epipactis 107 parviflora, Serapias 381
Lesser Butterfly Orchid 243 pendula, Epipactis 126
Lesser Rattlesnake Orchid 166 Pendulous-flowered Helleborine 126
Lesser Rattlesnake Plantain 166 phyllanthes, Epipactis 126
Lesser Twayblade 56 PLAtAntHERA 241
Lindisfarne Helleborine 122 praetermissa, Dactylorchis 309
lingua, Serapias 384 praetermissa, Dactylorhiza 309
LiPARiS 141 praetermissa, Orchis 309
Lizard Orchid 354 PSEUDORCHiS 235
Loesel’s Twayblade 142 Pugsley’s Marsh Orchid 316
loeselii, Liparis 142 pulchella, Dactylorhiza incarnata 276
longifolia, Cephalanthera 41 purpurata, Epipactis 101
Loose-flowered Orchid 363 purpurea, Orchis 222
maculata, Dactylorhiza 303 purpurella, Dactylorchis 328
maculata, neotinea 341 purpurella, Dactylorhiza 328
majaliformis, Dactylorhiza purpurella 332 purpurella, Orchis 328
Man Orchid 202 pyramidalis, Anacamptis 366
Marsh Fragrant Orchid 267 Pyramidal Orchid 366
Marsh Helleborine 78 Red Helleborine 34
mascula, Orchis 228 repens, Goodyera 166
militaris, Orchis 214 romanzoffiana, Spiranthes 174
Military Orchid 214 rubra, Cephalanthera 34
Monkey Orchid 207 sancta, Epipactis 122
monorchis, Herminium 193 Scarce Tongue Orchid 416
morio, Anacamptis 372 SERAPiAS 378
morio, Orchis 372 Short-spurred Fragrant Orchid 416
muelleri, Epipactis 107, 113 simia, Orchis 207
muscifera, Ophrys 392 Small-flowered Tongue Orchid 381
Musk Orchid 193 Small White Orchid 236
Narrow-leaved Helleborine 41 Southern Marsh Orchid 309
Narrow-leaved Marsh Orchid 316 sphegodes, Ophrys 411
Narrow-lipped Helleborine 107 spiralis, Spiranthes 181
neerlandica, Epipactis helleborine 97 SPiRAntHES 172
neglecta, Serapias 416 straminea, Pseudorchis 236
nEOtinEA 340 Summer Lady’s-tresses 188
nEOttiA 54 Sword-leaved Helleborine 41
Newfoundland Orchid 236 ‘Tyne Helleborine’ 118
nidus-avis, neottia 67 traunsteineri, Dactylorhiza 316
Northern Marsh Orchid 328 traunsteinerioides, Dactylorhiza 316
occidentalis, Dactylorchis 334 trifida, Corallorhiza 160
occidentalis, Dactylorhiza 334 ustulata, neotinea 345
occidentalis, Dactylorhiza comosa 334 ustulata, Orchis 345
occidentalis, Dactylorhiza majalis 325, 334 vectensis, Epipactis 126
occidentalis, Orchis 334 Violet Helleborine 101
ochroleuca, Dactylorhiza incarnata 276 viride, Coeloglossum 288
odoratissima, Gymnadenia 416 viridis, Dactylorhiza 288
OPHRYS 387 White Helleborine 48
ORCHiS 199 ‘Young’s Helleborine’ 97
ovata, Listera 62 ‘youngiana, Epipactis’ 97
006 systematic.indd 480 29/1/09 12:49:42

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What is the book about?

What is the book about?

What are some of the species mentioned in the acknowledgements section?

What are some of the species mentioned in the acknowledgements section?

Orchids

of Britain and Ireland

A Field and Site Guide

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A Field and Site Guide

Anne and Simon Harrap

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Published in 2005 by A&C Black Publishers Ltd,

Printed in China by Lion Productions Ltd.

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SOURCES OF INFORmATION AND BIBLIOGRApHY 469

INDEx OF SpECIES 479

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Fungi form several kinds of mycorrhiza:

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fungi and rare orchids

germinAtion And underground growth

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cross-pollination versus self-pollination

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marshes and fens

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heaths and moors

Pits, quarries and railway lines

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road verges, green lanes, churchyards and lawns

dunes and dune slacks

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Variation and varieties

Name and classification

history and conservation

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Distribution produces two buds at its tip: the larger will

O 31 May, Khabarovsk, eastern Sibera. Lady’s-slipper

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Lady’s-sLipper Critically Endangered, BAP

Identification north-facing slope. They nestle in a sheltered

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