ROOT IDENTIFICATION

MANUAL OF
TREES AND SHRUBS
ROOT IDENTIFICATION
MANUAL OF
TREES AND SHRUBS
A guide to the anatomy of roots of trees and shrubs
hardy in Britain and Northern Europe

D. F. CUTLER, P. J. RUDALL,
P. E. GASSON AND R. M. O. GALE
Jodrell Laboratory, Royal Botanic Gardens, Kew

London
CHAPMAN AND HALL
 First published in 1987 by
Chapman and Hall Ltd
11 New Fetter Lane, London EC4P 4EE
© 1987 The Trustees of the Royal Botanic Gardens, Kew
Softcover reprint of the hardcover 1st edition 1987

ISBN-13: 978-94-0lO-7912-9

All rights reserved. No part of this book may

be reprinted, or reproduced or utilized in any
form or by any electronic, mechanical or other
means, now known or hereafter invented,
including photocopying and recording, or in
any information storage and retrieval system,
without permission in writing from the
publisher.

BRITISH LIBRARY CATALOGUING

IN PUBLICATION DATA

Root identification manual of trees and

shrubs: a guide to the anatomy of roots
of trees and shrubs hardy in Britain and
Northern Europe.
1. Roots (Botany)
1. Cutler, D.F.
581.1 '0428 QK644
ISBN·13: 978·94·010·7912·9 e·ISBN·13: 978·94·009·3141·1
DOl: 10.1007/978·94·009·3141·1
 Contents

Foreword by Professor Keith Jones page vii

Preface ix

INTRODUCTION 1
BIBLIOGRAPHY 6
GENUS DESCRIPTIONS OF DICOTYLEDONS 7
FAMILY DESCRIPTIONS OF GYMNOSPERMS 211
APPENDIX 231
Table 1 Diagnostic characters in root anatomy of Dicotyledon trees 233
Table 2 Diagnostic characters in root anatomy of Dicotyledon shrubs
and climbers 234
Table 3 Main differences in root wood anatomy in four related
genera: Alnus, Betula, Carpinus and Corylus 236
Table 4 Diagnostic characters in root anatomy of the family
Caprifoliaceae 236
Table 5 Diagnostic characters in root anatomy of Gymnosperm trees 237
Index of scientific names 239
Index of common names 244
 Foreword

Plant anatomy is a vital part of plant descriptions and an integral component of

taxonomy. It also provides a unique means of botanical identification of plant fragments
and artefacts. The science in all of these roles has prospered for many years at the Jodrell
Laboratory which is renowned as a principal world centre for the subject. Its reputation
resides to a large extent in the substantial series of reference works on the systematic
anatomy of dicotyledons and monocotyledons written by staff of the Anatomy Section in
collaboration with researchers in other institutions. This present book, however, is the
first from Kew to describe a single organ - the root - to provide a means of identification
of a range of trees and shrubs when only the root is available. It has been inspired by the
many root samples sent to the Jodrell Laboratory over the years, often when they seem to
have been concerned with damage to the foundations of buildings. A previous book Tree
Roots and Buildings written from the Anatomy Section by Dr D. Cutler and I. Richardson
addressed itself to the frequency of damage of this sort caused by a range of tree species.
In the present book the Jodrell anatomists now expose the secrets of their diagnoses
making it possible for others to recognize the plants from which roots are derived. It is a
detailed and comprehensive work written by colleagues, all of whom are dedicated and
highly authoritative scientists. It gives me particular pleasure not only as a demonstration
of one of the important practical applications of anatomy, but also of concern for human
welfare which is the hallmark of the research carried out in all of the scientific sections of
the Jodrell Laboratory.

KEITH JONES
Keeper of the
Jodrell Laboratory
and Deputy Director
Royal Botanic Gardens
Kew

vii
 Preface

This book is the result of teamwork, not only among the authors, but also many other
people, most of whom are or were on the staff at Kew.
Over a period of about twelve years a lot of energy went into building up a large collec-
tion of reference microscope slides of shrub and tree roots. This enabled accurate identifi-
cations to be made of short lengths of root. Most of these came from inspection holes at
the foundations of buildings when it was suspected that roots might be related to
damage. The unique working collection has formed the basis for this book, but we have
added to it to extend the range of species covered and increase its value. Although the
majority of the working slides proved suitable for photography, some were not. Most of
these were replaced with new slides, but when no additional material was available,
some had to be photographed despite their quality. Roots of different diameters were
studied for many of the species examined. The descriptions include information on any
variation that occurred in these, but the most representative were selected for the illus-
trations.
Among those helping with sectioning were Timothy Lawrence and former staff
members Frederick R. Richardson and Dorothy Catling. Ian B. K. Richardson, co-author
of Tree Roots and Buildings also assisted.
Charles Erskine and his staff in the Living Collections Division have gone out of their
way to help dig up roots for us. The Photographic Section, with Tudor Harwood, Milan
Svanderlik and Andrew McRobb have worked under pressure to print the large number
of photographs.
Nigel Taylor assisted with verification of plant names, and Mary Gregory made a
critical reading of the text. Valerie Horwill typed the manuscript.
We are most grateful to all who have helped us.

ix
 Introduction

Knowledge about root structure has implications far beyond the intrinsic academic
interests of plant anatomy, physiology and ecology. It has become necessary to be able to
identify trees and shrubs from their root structure for reasons of considerable economic
importance, and for preservation of the amenity value of the plants concerned.
Roots of trees and shrubs have frequently been implicated in damage to buildings
through their effects on foundations and blockage of drains. Two recent drought periods
have emphasized the problems. When it is suspected that roots are involved in damage of
one form or another, it is most important that there should not be indiscriminate removal
of trees and shrubs. Equally, from the legal point of view, it is important that the owner of
the trees or shrubs involved should be identified. In most instances it is possible to
remove roots from the site of damage, and by an examination of their internal structure,
identify the plant from which they have come.
Root wood structure often differs from twig or branch wood structure in the same
plant. Consequently the reference books on the secondary wood of the aerial parts have
been of only limited help in making identifications.
Over a period of about twelve years, staff in the Plant Anatomy Section of the Jodrell
Laboratory at Kew have been making microscope slides of root sections from authenti-
cated sources. These have been used as reference material to assist in the identification of
roots thought to have been involved in damage to buildings or drains. This work, and a
survey of root spread that was undertaken simultaneously, led to the production of the
guide Tree Roots and Buildings by D. F. Cutler and I. B. K. Richardson.
It was decided to put the unique reference collection of microscope slides of roots to
further use, and make some of the information it contains more widely available. The
opportunity was taken to extend the collection to cover most of the trees and shrubs that
are known to be hardy out of doors in the British Isles. Consequently, many' exotics'
were studied, in addition to native plants.
These photographs have been brought together in this volume. Because of the very
large number of microscope slides available often representing several specimens with a
range of diameters, and range of variability in some species, it proved a difficult task to be
selective, and yet give a comprehensive cover in a book of manageable proportions. Illus-
trations included here show the salient points that are of diagnostic value.
In most examples a transverse section (TS), radial longitudinal section (RLS) and
tangential longitudinal section (TLS) are provided, together with a brief description of the
more important features, and notes on their diagnostic value. Both wood and bark are
described. In certain families of plants, the genera cannot be distinguished from one
another on the basis of root anatomy alone. In such instances the description of a group of

1
 INTRODUCTION

genera may extend beyond the normal two page layout, with illustrations taken from a
number of genera or species. Examples may be found in the Rosaceae, where the sub-
family Pomoideae is described rather than genera, and in the Salicaceae where Salix and
Populus are described together.
The alphabetical arrangement by scientific name has been chosen for convenience,
rather than one based on systematics. The gymnosperms (including Ginkgo) have been
placed in a section separate from the flowering trees and shrubs, and are described within
their families. There are relatively few diagnostic characters in transverse sections of
gymnosperm woods - those without resin canals tend to look very similar to one another.
Consequently, for the gymnosperms space has been given to the longitudinal planes of
section in preference to transverse sections.
It is envisaged that most users will have some background in plant anatomy, but for
those who have not, or wish to brush up earlier knowledge, this introduction gives basic
details useful in interpreting the descriptions.
All the roots illustrated in this book show secondary growth in thickness. Some are
juvenile and have very few growth rings. All have expanded to the extent that their
original epidermis (rhizodermis) has split and in many specimens, gone.

Bark
The 'bark' of the root, as defined for this book, consists of the cork (phellem), phellogen
and phelloderm, any remaining layers of cortex, and the phloem.
Cork is produced by a cork cambium (phellogen). This may be formed in the cortex, at
varying depths, or in the outer phloem, or rarely, from the rhizodermis itself. The cork
provides protection against the ingress of pathogens.
In some species cork provides few features of diagnostic value. In others there are
characteristic layers of cells with thick or thin walls. There is variation in the way the cork
layers split and are shed. The cells may lack contents, or contain dark coloured substances
frequently identified as tannins.
If the cortex is to the outside of the cork, it eventually splits and is shed; fragments may
persist in young roots. When cortical cells persist to the inner side of the cork, they
remain alive, and frequently become involved in cell division, so accommodating growth
in root diameter without tearing. A clear dilatation (expansion) zone may be present, or
the junction between cortex and the dilating ends of phloem rays may be indistinct. Since
no developmental studies were carried out during the preparation of this book, we have
decided to make no distinction between dilating ray ends and divided cortical cells in
descriptions. The cortex may consist entirely of parenchymatous cells, or it may contain
sclereids and fibres, or both. Some cells may be tannin-filled, crystals of various sorts may
be present and sometimes diagnostic features such as mucilage cells or secretory ducts
may occur.
Phloem is sufficiently variable for diagnostic characters to be identified in transverse
sections. Normally two distinct tissues are evident, the rays and the conducting cells.
Rays may remain the same width through the phloem, or they may expand or dilate to
varying extents towards their outer ends. Dilatation may be due to cell expansion, cell
division or both. The conducting tissues may all be thin-walled, consisting of sieve-tubes,
companion cells and parenchyma (or sieve cells, albuminous cells and parenchyma in

2
 INTRODUCTION

gymnosperms). However, in many species sclereids and fibres are also present. These
thick-walled cells may be scattered at random, arranged in strands or groups, or in
tangential bands, alternating with thin-walled cells.

Cambial zone
The cambial zone consists of the vascular cambium itself, and the immediate undifferen-
tiated phloem and xylem cells to either side. After the first year of secondary growth, the
cambium generally assumes a cylindrical or compressed cylindrical form. In some
species, the cambium may curve distinctly towards the root centre at each of the wide
rays.

Xylem
The primary xylem, with its archs of protoxylem is often distinct at the centre of second-
arily thickened roots. There is generally no central pith. The number of protoxylem archs
is often related to the initial width of the primary root, and has little if any diagnostic
value in the dicotyledons. Many gymnosperms are diarch.
Nearly all the characters of diagnostic value come from secondary xylem. Transverse
sections show some features clearly, and longitudinal sections show details of others. In
narrow roots it can be difficult to make accurate tangential longitudinal sections, quite
frequently there are features of both radial and tangential cell arrangement in such sec-
tions.
In transverse section, growth rings mayor may not be distinct. Some species regularly
have ring or semi-ring porous vessel distribution, whereas others are diffuse porous.
Dendritic and other distinctive types of vessel arrangement are noted when they are
present. All of these characters appear to be more variable in roots than in stems. Some
species have only solitary vessels, but it is more common to have both solitary and
grouped vessels. Characteristic vessel groupings are noted in the descriptions. Vessel
diameter is very variable in roots, and even in the same species examples may be found
with narrow vessels and others with much wider vessels. The growing conditions have a
marked effect on vessel width. In this book we have described vessel width when appro-
priate, but the dimensions that relate to our samples may represent only part of the
possible range for a species. Because of this, no actual measurements are given in descrip-
tions, and details of equivalents of width and cell wall thickness are given in the following
section which acts as a general guide.
In the longitudinal sections (LS), details of vessel to vessel (intervascular) wall pitting,
and vessel to ray pitting are noted. This is of high diagnostic value. In the gymnosperms,
tracheid wall pitting and cross-field (tracheid to ray) pitting have similar significance.
Vessel perforation plate types are also described; these are generally easy to see, and are
very valuable in diagnosis. Tyloses are described when present.
Sometimes vessels or tracheids (and fibres) may have a tertiary spiral (helical) thicken-
ing of the wall next to the lumen.
Fibres in roots frequently have thinner walls than those in stems. Mention is made in
descriptions of wall thickness and lumen width when appropriate. Often root samples
are too narrow or otherwise inadequate for detailed studies to be made of the complete

3
 INTRODUCTION

range between fibres and tracheids and their intermediate forms. In diagnosis this could
be of importance, but because of the practical difficulties we have noted only when fibres
have conspicuously bordered pits (technically making them fibre-tracheids) rather than
non-bordered or only very slightly bordered pits. For the same reasons, only imperforate
tracheary cells with very distinct bordered pits have been recognized as tracheids in this
work.
The distribution and abundance of axial parenchyma is recorded for transverse
sections. It is often more abundant in roots than stems of a given species. In LS, note is
made of important features of the axial parenchyma, for example, if it is storied, or if cells
are mainly fusiform.
Rays are described in both TS and LS, since both views are essential before the ray type
can be discerned. Ray width can only be established accurately from the TLS. Details are
given about the shape of rays, whether they are homocellular with all cells similar in
shape, or heterocellular, composed of cells with distinctly different shapes. Many narrow
roots have not developed the mature ray types. It is not wise to use the established ray
classifications for this reason. Height of rays is of importance, and is recorded.
The presence and type of crystals, cell contents (particularly starch granules), secretory
canals or ducts are recorded as appropriate.
In this book the text, captions and illustrations must be used together to gain infor-
mation useful for identification. The textual descriptions by themselves are not complete,
but emphasize details of value for diagnosis.
Tables are provided in the appendix to assist in the process of identification.
The best book can act only as a guide to the possible identity of roots. Comparison of the
unknown with accurately identified reference microscope slides is normally necessary
before the process of identification can be completed.

Measurements
There is a great deal of variation in cell width and wall thickness in roots of a given
species, much more so than in twigs or mature stem wood. Consequently accurate
measurements are not generally very useful in identification of roots. Often the root
sample is inadequate for statistically accurate measurements to be made. It was decided
not to give actual figures in the descriptions for these reasons. A general guide may be of
interest to some readers, so the descriptive terms used are defined here.
Fibre wall thickness and lumen width is given in general terms. In fibres with narrow
lumina, the double wall thickness (of adjacent fibres) is greater than the width of the
lumen. In fibres with wide lumina, the double wall thickness is narrower than the lumen.
Fibres described as thin-walled, have single walls narrower than 2 J.tm.
Vessels are described as wide when the tangential lumen diameter is greater than
100 J.tm, and narrow when less than 50 J.tm.
The size of intervascular pits may be of diagnostic value, and tends to be less variable
within a species than many other measurements in roots. Those described as coarse are
over 6.5 J.tm in diameter, and fine pits are 4 J.tm or less. Where pit size is unspecified, the
pits are between 4 and 6.5 J.tm in diameter. In Betula, the pits are minute, generally less
than 3.5J.tm.

4
 INTRODUCTION

Methods used in slide preparation

Freshly collected root material was washed to remove soil, fixed in formalin acetic alcohol
for 48 hours and sectioned after washing. Dried roots needed to be boiled in water for
5-10 minutes before they were suitable for sectioning.
A Reichert OME sliding microtome was used to produce sections of between 20-30 J.lm
in thickness. This method involves firm clamping of the specimen, but avoids the need
for embedding.
Transverse and longitudinal sections were prepared and stained as described in Cutler
(1978). Some were examined in temporary mounts, but permanent slides were made for
photography. Some of the sections were stained in safranin and haematoxylin, but in the
most recent, 1% aqueous aldan blue was used as a counter stain instead of haematoxylin,
and Euparal was used as a mounting medium rather than Canada balsam.

5
 Bibliography

Selected from the more recent literature.

Bean, W. J. (1970-1980) Trees and Shrubs Hardy in the British Isles, 8th Edn, vols 1-4, John
Murray, London.
Cutler, D. F. (1976) Variation in root wood anatomy, in Wood Structure in Biological and
Technological Research (eds P. Baas, A. J. Bolton and D. M. Catling), Leiden Botanical
Series No.3, 143-156, Leiden University Press, Leiden (useful for further references to
work on root wood anatomy).
Cutler, D. F. (1978) Applied Plant Anatomy, Longman, London.
Cutler, D. F. and Richardson, I. B. K. (1981) Tree Roots and Buildings, Construction Press
(Longman), London.
Fayle, D. C. F. (1968) Radial Growth in Tree Roots, Faculty of Forestry, University of
Toronto, Toronto.
Gasson, P. (1979) The identification of eight woody genera of the Caprifoliaceae by
selected features of their root anatomy, Botanical Journal of the Linnean Society 78,267-84.
Hillier's Manual of Trees and Shrubs (1981) 5th edn, David and Charles, Newton Abbot.
Miles, A. (1978) Photomicrographs of World Woods, Building Research Establishment,
HMSO, London.
Mitchell, A. (1974) A Field Guide to the Trees of Britain and Northern Europe, Collins,
London.
Schweingruber, F. H. (1978) Microscopic wood anatomy, Swiss Federal Institute of Forestry
Research, Birmensdorf, Zurcher A. G.

6
Genus descriptions
of Dicotyledons
ABELlA Caprifoliaceae

Bark heterocellular; cells of varying shape in TLS; square, up-

Peridenn consisting of radial files of thin-walled square right and only rarely procumbent in RLS .
and rectangular cells, outer layers disintegrating; two or
more periderms may develop, one arising directly adjac-
NOTE
ent to phloem. Cortex narrow. Phloem rays not dilating
towards cortex; stone cells and fibres absent. Crystals: See Appendix, Table 4.
cluster crystals sometimes present in phloem.
SPECIES EXAMINED
Xylem
A. chinensis R.Br.; A. graebneriana Rehd.; A. schumannii
TS. Diffuse porous. Growth rings distinct in some Rehd.; A. spathulata Sieb. and Zuce.; A. triflora R.Br.;
samples, indistinct or absent in others. Vessels angular, A. umbellata Rehd.
solitary, in pairs and in occasional groups of 3 or rarely up
to 5. FiYres with narrow lumina. Axial parenchyma para-
tracheal and diffuse . Rays 1-7 cells wide.

LS. Vessels: intervascular and vessel-ray pitting mainly

opposite, occasionally scalariform; perforation plates
oblique, scalariform, with 10-30 bars, but exclusively
simple in A. triflora and A. umbellata; faint spiral thicken-
ings sometimes present. Fibres usually with conspicuous
spiral thickenings. Rays 1-7 cells wide; uniseriate rays
1-12 cells high, multiseriate rays up to 65 cells high,
usually with uniseriate tails or portions; rays markedly
1. Abe/ia trif/ora, bark and xylem TS x 80.
2. Abe/ia spathu/ata, xylem TS x 80.

8
3. Abelia trif/ora, xylem TLS x 60.
4. Abelia spathulata, xylem TLS x 150.
5. Abelia spa thulata , xylem LS x 600, showing vessels
with scalariform perforation plates, abundant starch
granules and a fibre with spiral thickenings .

6. Abelia spa thulata , xylem TLS x 380, showing fine

vessel pitting and starch granules in ray and axial paren-
chyma cells .

9
ACER Aceraceae

Bark SPECIES EXAMINED

Periderm peeling in short strips, lenticels numerous. A . palmatum Thunb.; A. platanoides L., Norway Maple;
Cortex containing one or more discontinuous tangential A. pseudoplatanus L., Sycamore.
bands of thin-walled stone cells and sometimes fibres.
Phloem rays usually dilating towards their outer ends,
sometimes with large blocks of thin-walled stone cells;
conducting tissue occasionally with tangentially parallel
bands of fibres or scattered groups of fibres or stone cells,
but often without sclerenchyma. Crystals solitary, rhom-
bic, present in cortex and phloem, usually associated
with sclerenchyma, rare or absent in rays. 7. Acerplatanoides, bark RLS x 150, showing solitary
crystals, stone cells and fibres.
Xylem
8. Acer platanoides, bark and xylem TS x 80.
TS. Diffuse porous. Growth rings distinct, sometimes
discontinuous. Vessels solitary and in clusters and radial 9. Acer platanoides, xylem TS x 45, showing vessel
chains of 2-7, thin-walled. Fibres thin-walled with wide clusters.
lumina; banded appearance of xylem due to areas of 10. Acer platanoides, xylem TLS x 150, showing rays and
fibres with slightly thicker walls. Axial parenchyma in- fibres.
distinct. Rays 1-7 cells wide.
11. Acer platanoides, xylem TLS x 380, showing vessel
LS. Vessels: intervascular pitting and vessel-ray pitting elements with coarse, alternate, polygonal pitting and faint
alternate, pits coarse, polygonal; perforation plates spiral thickenings.
oblique, simple; spiral thickenings usually present. Rays
12. Acer platanoides, xylem TLS x 380, showing vessels
uniseriate and 2-7 cells wide; uniseriate rays 1-25 cells with spiral thickenings.
high, multiseriate rays 5-65 cells high; uniseriate tails
present on many rays that are up to 3 cells wide, absent
from rays 3 or more cells wide; some rays with uniseriate
portion between two multiseriate portions; rays hetero-
cellular.

10
10

11
AESCULUS Hippocastanaceae

Bark
Periderm peeling in long strips. Cortex boundary with
phloem sometimes indistinct; containing scattered blocks
of fibres and thin-walled stone cells irregular in shape and
size. Phloem rays dilating slightly towards outer ends;
conducting tissue sometimes containing small groups of
fibres or individual fibres, as in cortex. Crystals most
common in cortex, usually absent from rays; mainly
cluster crystals but a few solitary crystals usually present.

Xylem
TS. Diffuse porous. Growth rings indistinct. Vessels soli-
tary and in pairs and short radial chains, and clusters or
aggregates of up to 9; normally angular in outline, thin-
walled. Fibres thin-walled with wide lumina. Axial
parenchyma sparse. Rays uniseriate .

LS. Vessels: intervascular pitting alternate, individual pits

polygonal; vessel-ray pits similar, but lacking obvious
borders; fine spiral thickenings present; perforation
plates simple. Rays uniseriate or very occasionally biseri-
ate in part; 1-20 or occasionally up to 30 cells high;
heterocellular.

SPECIES EXAMINED
13. Aescu/us hippocastanum, bark and xylem TS x 80.
A. hippocastanum L., Horse Chestnut.

16. Aescu/us hippocastanum, xylem TLS x 380, showing

uniseriate rays and alternate, polygonal vessel pitting.
17. Aescu/us hippocastanum, xylem TLS x 380, showing
uniseriate rays and spiral thickenings on vessels.

12
14. Aesculus hippocastanum, xylem TS x 55, showing
radial chains and clusters of vessels, and uniseriate rays.
15. Aesculus hippocastanum, xylem TLS x 150, showing
uniseriate rays.

13
AILANTHUS Simaroubaceae

Bark
Periderm consisting of radial files of thin-walled square or
rectangular cells; outer layers often breaking away in
blocks giving uneven appearance. Cortex containing large
groups of thick-walled stone cells in a discontinuous ring
up to 10 cells wide; fibres rare or absent. Phloem rays dilat-
ing gradually towards cortex; conducting tissue with
scattered thin-walled fibres, often solitary, but also in
small groups of up to 15. Crystals: cluster crystals abun-
dant in cortex and phloem; solitary crystals often associ-
ated with stone cells.

Xylem
TS. Diffuse porous. Growth rings sometimes indistinct.
Vessels mainly very wide; solitary and in pairs and clusters
of up to at least 13 cells and radial chains of up to 8 cells;
wide vessels circular or oval, narrow vessels angular in
outline. Fibres thin-walled with wide lumina. Axial paren-
chyma mainly paratracheal. Rays 1-4 cells wide.

LS. Vessels: intervascular pitting coarse, vessel-ray pitting

slightly smaller, pits alternate, oval; perforation plates
simple. Rays 1-4 cells wide; uniseriate rays 1-15 cells
high, biseriate rays 4-24 cells high, wider rays up to 30
cells high; some multiseriate rays with short uniseriate
tails or portions, outer cells of ray sometimes much taller 18. Ailanthus alrissima, bark and xylem TS x 60, showing
than wide, forming an incomplete sheath in TLS; hetero- stone celis and fibres in bark.
cellular; cells in RLS square, procumbent and upright.

SPECIES EXAMINED

A. altissima Swingle, Tree of Heaven.

14
 19. Ailanthus altissima , xylem TS x 60.
20. Ailanthus altissima , xylem TLS x 60 .

21. Ailanthusaltissima, xylem TLS x 380, showing

vessels with alternate, polygonal pitting .

15
ALNUS Betulaceae

Bark
Periderm peeling in long and short strips; cells with dark
contents. Cortex with more or less continuous cylinder of
stone cells and few fibres . Phloem rays slightly dilating
towards outer ends, fibres and stone cells usually absent,
rarely occasional stone cells present. Crystals: solitary and
occasional cluster crystals present, usually associated
with cylinder of stone cells in cortex.

Xylem
TS. Diffuse porous; growth rings often indistinct. Vessels
usually in radial chains of 3-15 or exceptionally up to 25,
also solitary and occasionally in small multiples; thin-
walled, angular. Fibres thin-walled with wide lumina.
Axial parenchyma diffuse, paratracheal and apotracheal,
with a tendency to form tangential bands. Rays mainly
uniseriate, or rarely up to 3 cells wide; occasional aggre- •
gate rays present.

LS. Vessels: intervascular pitting and vessel-ray pitting

very fine, opposite; perforation plates scalariform, with 22. Alnus glutinosa, bark and xylem TS x 60, showing a
10-20 thin bars. Rays mainly uniseriate, or rarely up to 3 cylinder of stone cells in the bark .
cells wide; 2-25 cells high; homocellular; cells in RLS
mainly procumbent.

NOTE

See Appendix, Table 3.

SPECIES EXAMINED

A. glutinosa Gaertn., Alder.

25. Alnus glutinosa, xylem TLS x 150, showing thin-

walled fibres and uniseriate rays .
26. Alnus glutinosa, xylem RLS x 150, showing a vessel
with a scalariform perforation plate and fine pitting .

16
23. Alnus glutinosa, xylem TS x 80, showing radial chains
of vessels and mainly uniseriate rays .
24. Alnus glutinosa, xylem TLS x 150, showing thin-
walled fibres and axial parenchyma and a multiseriate ray.

17
ARBUTUS Ericaceae

Bark
Periderm regular, arising in outer cortex. Cortex paren-
chymatous, fibres and stone cells absent; tannin some-
times present in some cells. Phloem rays not dilating
towards outer ends; fibres and stone cells absent from
rays and conducting tissue. Crystals absent.

Xylem
TS. Diffuse porous. Growth rings indistinct. Vessels
narrow, mainly solitary, sparse. Fibres with large lumina.
Axial parenchyma sparse. Rays 1-3 cells wide.

LS. Vessels: elements narrow and long; intervascular pit-

ting and vessel-ray pitting fine, circular or elongated,
alternate; perforation plates simple; spiral thickenings
present. Fibres: many septate. Rays uniseriate and 2-3
cells wide; uniseriate rays 1-15 cells high, multiseriate
rays 6-23 cells high, often with uniseriate tails; hetero-
cellular; uniseriate rays composed predominantly of
upright cells.

27. Arbutusandrachne, bark TS x 150, showing many

SPECIES EXAMINED tannin-filled cells .
A. andrachne L.

18
28. Arbutus andrachne, xylem TS x 150, showing
narrow, solitary vessels.
29. Arbutus andrachne, xylem TLS x 150, showing
narrow vessels, hardly wider than fibres and short, narrow
rays.

30. Arbutus andrachne, xylem TLS x 600, showing a

vessel with fine alternate and elongated pitting, and a
fibre-tracheid.

19
AUCUBA Aucubaceae

Bark
Peridenn irregular, cells mainly oval, closely resembling
stone cells with wide lumina. Cortex consisting of thin-
walled parenchyma cells; sclerenchyma absent. Phloem
rays not dilating; sclerenchyma absent. Crystals: crystal
sand present in cortex.

Xylem
TS. Diffuse porous. Growth rings indistinct. Vessels angu-
lar in outline; solitary and in pairs, clusters and radial
chains of up to 5. Fibres thicker-walled and narrower than
most vessels, with wide lumina. Axial parenchyma para-
tracheal and diffuse. Rays 1-10 cells wide, cells as wide as
or wider than most vessels, mainly square.

LS. Vessels: intervascular pitting and vessel-ray pitting

alternate to opposite or scalariform, pits coarse; perfor-
ation plates very long, scalariform, usually with more
than 40 bars, some bars bifurcating; most vessels with 32. Aucubajaponica, bark TS x 150, showing cambium,
phloem and inner cortex.
faint spiral thickenings. Fibre tracheids present. Rays 1-10
cells wide; uniseriate rays 1-11 cells high; multiseriate
rays 7 to over 100 cells high; heterocellular; cells in RLS
mainly upright; starch granules often present in ray cells
and axial parenchyma cells.

SPECIES EXAMINED

A. japonica Thunb.

31. Aucuba japonica, bark TS x 150, showing outer layers

with stone cells.

20
33. Aucuba japonica, xylem TS x 45, showing narrow
vessels, wide rays, and abundant starch granules.
34. Aucuba japonica, xylem TLS x 60, showing short,
wide rays.

35. Aucuba japonica, xylem RLS x 380, showing a vessel

with a scalariform perforation plate .
36. Aucuba japonica, xylem TLS x 380, showing vessels
with opposite and scalariform pitting and starch granules
in fibres and some ray celis.

21
BERBER/ S Berberidaceae

Bark
Periderm narrow, cells usually thin-walled. Cortex: fibres
and stone cells absent, or rarely small stone cells present.
Phloem rays dilating towards outer ends; fibres and stone
cells absent or very rarely present. Crystals: solitary crys-
tals occasionally present in phloem.

Xylem
TS. Semi-ring porous. Growth rings distinct. Vessels and
tracheids solitary and in clusters and radial multiples of
2-15, sometimes with slightly dendritic arrangement.
Fibres thin-walled with wide lumina. Axial parenchyma
usually very sparse or absent. Rays 1-6 cells wide.

LS. Vessels: intervascular and vessel-ray pitting alternate,

fine; perforation plates simple; faint spiral thickenings
often present on vessels and tracheids. Rays 1-6 cells
wide, 4-100 cells high; heterocellular; cells in RLS square
and procumbent.
:no Berberis vulgaris, bark and xylem TS x 60.

NOTE 38. Berberis vulgaris, xylem TLS x 150, showing short

uniseriate and longer multiseriate rays.
Roots conspicuously yellow.

SPECIES EXAMINED

B. x stenophylla (B. danoinii Hook. x B. empetrifolia Lam.);

B. vulgaris L.; Barberry.

39. Berberis vulgaris, xylem TLS x 150.

40. Berberis vulgaris, xylem RLS x 380, showing a hetero-
cellular ray.

22
 . ·..
.);....t; ,.; , I

•
·

23
BETULA Betulaceae

Bark
Periderm not peeling; some cells containing tannin. Cortex
with more or less continuous cylinder of stone cells and
fibres (fibres relatively few). Phloem rays dilating slightly
towards cortex; occasionally with stone cells; conducting
tissue in triangular blocks, often including stone cells,
sometimes in large groups, and also with occasional
fibres. Crystals: solitary and cluster crystals present in
cortex associated with sclerenchyma cylinder, and rarely
present in phloem rays.

Xylem
TS. Diffuse porous. Growth rings distinct. Vessels nor-
mally in radial chains of 3-9, and exceptionally up to 15,
also solitary and in pairs and multiples of up to 6; thin-
walled, often angular. Fibres thin-walled with wide
lumina. Axial parenchyma usually sparse, paratracheal and
scattered. Rays 1-3 cells wide.

LS. Vessels: intervascular pitting and vessel-ray pitting

minute, alternate; perforation plates scalariform, circular
or oval, with 5-15 thin, widely spaced bars. Rays uniseri-
ate and 2-3 cells wide; uniseriate rays 1-15 cells high;
multiseriate rays 6-20 cells high or occasionally up to 45
cells high, often with uniseriate tails; heterocellular; cells
in RLS procumbent or square, marginal cells occasionally
upright.
41. Betula sp., bark TS x 80, showing a continuous ring
of stone cells and fibres and dilating rays in phloem.

NOTE

See Appendix, Table 3.

SPECIES EXAMINED

B. pendula Roth.; Betula sp.; Birch.

24
42. Betula sp ., xylem TS x 80, showing a growth ring
boundary, vessels solitary and in clusters.
43. Betula sp., xylem TLS x 80, showing vessels with
minute pitting, uniseriate and multiseriate rays.

44. Betula sp., xylem RLS x 380, showing two vessels

with scalariform perforation plates and minute vessel
pitting .

25
BIGNONIA Bignoniaceae

Bark
Peridenn very wide, irregular on outside; stratified, with
alternating bands of 2-3 layers of thin-walled, square or
rectangular cells and 1-3 layers of thicker-walled cells.
Cortex absent or very narrow . Phloem rays dilating slightly
towards outer ends; conducting tissue with tangentially
elongated blocks of fibres and stone cells several cells
wide, sometimes fibres absent in young roots. Crystals:
cluster crystals present in phloem rays .

Xylem
IS. Diffuse porous to semi-ring porous. Growth rings
distinct. Vessels solitary and occasionally in pairs. Fibres
with wide lumina. Axial parenchyma scattered, with some
cells directly adjacent to vessels. Rays wider than wedges
of axial xylem (20 cells wide in first year), widening con-
siderably from the centre of the root outwards. Crystals:
cluster crystals present in some ray cells .

LS. Vessels storied; intervascular pitting and vessel-ray

pitting alternate, pits coarse, polygonal; perforation
plates simple. Rays up to at least 20 cells wide and very
tall, frequently wider than radiating blocks of axial xylem;
rays heterocellular; cells in RLS mainly square and
45. Bignonia capreo!ata, bark and xylem TS x 50, show-
upright. ing stratified periderm and very wide rays .

NOTE

The radiating blocks of axial xylem are easily distin-

guished by the naked eye and give a star-like appearance
to the root.

SPECIES EXAMINED

B. capreolata L., Cross Vine.

26
46. Bignonia capreo!ata, xylem TS x 50, showing distinct
growth rings, solitary vessels and very wide rays .
47. Bignonia capreo!ata, xylem TLS x 50, showing ends
of two wide rays.

48. Bignonia capreo!ata, xylem RLS x 150, showing

vessels with simple perforation plates and cluster crystals
in some ray cells.

27
BUDDLEJA Buddlejaceae

Bark
Periderm ragged, fairly wide, with thin-walled cells.
Cortex absent. Phloem extending to periderm, fibres and
stone cells absent; rays dilating gradually towards outer
ends. Crystals absent.

Xylem
TS. Diffuse porous. Growth rings indistinct. Vessels soli-
tary, and in radial chains of up to 14 cells, and in tangen-
tial or obliquely arranged clusters of up to 7 cells. Fibres
thin-walled with wide lumina. Axial parenchyma sparse,
scattered. Rays 1-6 cells wide.

LS. Vessels: intervascular pitting alternate, polygonal;

vessel-ray pitting alternate or sometimes elongated; per-
foration plates simple; spiral thickenings present on
narrower vessels and tracheids. Rays 1-6 cells wide, 1-50
or more cells high; heterocellular; in TLS groups of small
cells surrounded by larger cells; cells in RLS square and
upright.

SPECIES EXAMINED

B. davidii Franch., Buddleja.

49. Budd/eja davldii, bark TS x 80.

28
50. Budd/eja davidii, xylem TS x 30.
51. Budd/eja davidii, xylem TLS x 60, showing tall rays .

52. Budd/eja davidii, xylem TLS x 150, showing alternate

vessel pitti ng.
53. Budd/eja davldli, xylem TLS x 380, showing spiral
thickenings on vessels, and ray cells of varying size and
shape.

52

29
BUXUS Buxaceae

Bark
Periderm ragged, several layers thick; cells irregular, thin-
walled or occasionally with thickened periclinal walls.
Cortex narrow or absent; cells thin-walled, parenchyma-
tous. Phloem rays dilating only at outer ends; fibres and
stone cells absent. Crystals: groups of solitary and double
crystals with associated smaller crystalline masses pres-
ent in phloem.

Xylem
IS. Diffuse porous. Growth rings often indistinct. Vessels
narrow, crowded, mainly solitary and with occasional
tangential pairs and radial chains of up to 4. Fibres thick-
walled. Axial parenchyma fairly frequent, scattered. Rays •
1-3 cells wide, heterocellular.

LS. Vessels: intervascular pitting and vessel-ray pitting 54. Buxus sempervirens, bark TS x 80.
fine, opposite. Perforation plates scalariform, circular to
oval, with 8-11 bars, occasionally with trabeculae. Rays
uniseriate and biseriate or rarely up to 3 cells wide; uni-
seriate rays 1-22 cells high; biseriate rays 8-25 cells high
with uniseriate tails and sometimes uniseriate portions;
heterocellular; cells in RLS procumbent, square and
upright.

SPECIES EXAMINED

B. semperoirens L., Box.

57. Buxus sempervirens, xylem TLS x 380, showing alter-

nate vessel pitting, fibre and tracheid pitting, and rays.
58. Buxus sempervirens, xylem RLS x 380, showing a
vessel with a scalariform perforation plate, and a
heterocellular ray.

30
 ,M(:'"
~

~ t~ W
:.rilll.tI' •
8 ~~~lrl~~~
I JUItf !

,; ,
.CL Ii ~ III

~~ ~ ~ :s [) M ~
~
~
~
t' ~ II II'" JtI
J1. Ihl '1
~ l: ~ tJ. r

,
~ It:
1'1'
~ 1:
~

"'
~ .,..~
or ~
~ ,~~ ~

lArS ~ ~:1 ~
I~ ;U p
•
I
IIr:
l- IIR' l1li9 ill. ;r:
~~~' A
~

..~ 7
~

~
~ >'
]I

55 rlAl.~J Yo
.-
t
m
• . ~~
'-Il ~
- -- ,v ~

56. Buxus sempervirens, xylem TS x so.

56. Buxus sempervirens, xylem TLS x 150, showing
uniseriate rays of tall cells and biseriate rays with uniseriate
tails of tall cells.

31
CAMELLIA Theaceae

Bark
Periderm: outer layers irregular, often with dark contents.
Cortex narrow, parenchymatous; fibres and stone cells
absent. Phloem rays dilating slightly towards outer ends;
conducting tissue in small wedge-shaped areas; fibres
and stone cells absent or rare. Crystals absent or rarely
solitary crystals present in cortex.

Xylem
TS. Diffuse porous. Growth rings indistinct. Vessels nar-
row, angular, usually solitary and occasionally in pairs or
groups of up to 5. Fibres thick-walled. Axial parenchyma
abundant, scattered. Rays crowded, 1-4 cells wide.

LS. Vessels: intervascular and vessel-ray pitting oval or

elongated, sometimes scalariform; perforation plates
oblique, scalariform, with 15-25 fine, closely spaced bars.
Rays 1-4 cells wide, 1-100 or more cells high; markedly
heterocellular, with uniseriate tails and portions; cells in
RLS usually square or upright in uniseriate portions, pro-
cumbent in multiseriate portions.

SPECIES EXAMINED 60. Camellia cusp/data, xylem TS x 150, showing narrow

C. cuspidata Veitch.; C. japonica L.; Camellia. angular vessels.
61. Camellia cuspidata, xylem TLS x 150, showing
uniseriate rays and heterocellular multiseriate rays with
uniseriate tails.
62. Camelliajaponica, xylem TLS x 150, showing
uniseriate rays and heterocellular multiseriate rays with
uniseriate portions and tails.

63. Camellia cuspidata, xylem RLS x 150, showing

heterocellular rays, scalariform vessel and vessel-ray pits
and scalariform perforation plates.
64. Camellia cuspidata, xylem RLS x 380, showing
scalariform vessel-ray pitting, a scalariform perforation
59. Camellia cuspidata, bark and xylem TS x 60 . plate, and tracheids.

32
33
CARPINUS Carpinaceae

Bark
Periderm peeling in thin strips; individual cells rectangular
or square, thin-walled, occasionally with dark contents.
Cortex with more or less continuous cylinder of stone cells
and fibres . Phloem rays dilating towards outer ends in all
but the youngest roots, occasionally with stone cells;
conducting tissue in radially elongated or triangular
blocks, without sclerenchyma. Crystals: solitary and
cluster crystals associated with sclerenchyma cylinder in
cortex, also in ray parenchyma, rare or absent in conduct-
ing tissue.

Xylem
TS. Diffuse porous. Growth rings fairly distinct. Vessels
solitary, in radial chains of 2-10 and occasionally in
clusters. Fibres thin-walled with large lumina. Axial paren-
chyma sparse, para tracheal and scattered. Rays 1-5 cells
wide, or rarely aggregate in larger roots; individual ray
cells rectangular in the radial direction. Broader rays
sometimes noded at growth rings.

LS. Vessels: intervascular pitting and vessel-ray pitting

alternate, coarse, polygonal; vessel-ray pits narrowly
bordered; perforation plates simple; tracheids present,
with similar pits to those of vessels and sometimes with
faint spiral thickenings. Rays 1-4 or 5 cells wide; uni-
seriate rays 1-21 cells high, multiseriate rays 7-92 cells 65. Carpinus betu/us, bark and xylem TS x 80, showing
cylinder of stone cells and fibres in cortex, and radial
high, sometimes with uniseriate portions and tails;
chains of vessels in xylem.
heterocellular. Crystals: solitary crystals occasionally
present in ray cells.

NOTE

See Appendix, Table 3.

SPECIES EXAMINED

C. betulus L., Hornbeam.

34
66. Carpinus betulus, xylem TS x 80.
67. Carpinus betulus, xylem TLS x 150, showing
uniseriate rays and multi seriate rays with uniseriate tails,
narrow vessels and tracheids, and fibres.

68. Carpinus betulus, xylem TLS x 380, showing a vessel

with coarse, alternate, polygonal pitting, and uniseriate
rays.
69. Carpinus betulus, xylem RLS x 380, showing a
heterocellular ray with alternate vessel-ray pitting.

35
CASTANEA Fagaceae

Bark
Peridenn consisting of few layers of rectangular cells,
often with dark contents. Cortex with groups of 7-50
thick-walled fibres; stone cells rarely present. Phloem with
groups of fibres as in cortex; rays not dilating towards
outer ends. Crystals: solitary crystals present, associated
with fibres in cortex and phloem.

Xylem
IS . Diffuse porous. Growth rings usually indistinct.
Vessels sometimes in oblique lines, solitary, varying in
size from very wide to narrow . Fibres sometimes thin-
walled, with wide lumina. Axial parenchyma abundant,
paratracheal and apotracheal, scattered, or sometimes in
short uniseriate tangential bands. Rays uniseriate.

LS. Vessels : intervascular pitting alternate; vessel-ray pit- 71 . Castanea sativa, bark TS x 150, showing periderm
ting circular or elongated with narrow borders, often and outer cortex with groups of thick-walled fibres .
orientated in several directions; perforation plates
simple. Rays uniseriate, or rarely biseriate; 1-50 cells
high; heterocellular; cells in RLS mainly square ana pro-
cumbent, also upright.

NOTE

Distinguishable from Oak, Quercus spp. by absence of

multiseriate rays.

SPECIES EXAMINED

C. sativa Mill., Sweet Chestnut.

70. Castanea sativa, bark TS x 150, showing periderm

and cortex .

36
72. Castanea sativa, xylem TS x 40, showing wide vessels
and uniseriate, occasionally biseriate rays .
73. Castanea sativa, xylem TS x 80. 74. Castanea sativa, xylem TLS x 150, showing fibres,
axial parenchyma strands, uniseriate rays and one ray with
a biseriate portion.

75. Castanea sativa, xylem RLS x 380, showing fibre-

tracheids and a ray with vessel-ray pitting orientated in
several directions.

37
CATALPA Bignoniaceae

Bark
Periderm often peeling off in ragged strips; sometimes
including a layer of thick-walled cells. Cortex paren-
chymatous, with occasional small groups of 1-15 thick-
walled fibres . Phloem rays not dilating widely towards
cortex; conducting tissue consisting of regularly arranged
tangential rows of cells, sometimes with air spaces
between them, and with occasional small groups of 1-15
fibres interspersed. Crystals absent.

Xylem
TS . Diffuse to slightly semi-ring porous. Growth rings
more or less distinct. Vessels solitary and in clusters and
radial chains of up to 10. Fibres with wide lumina. Axial
parenchyma paratracheal and apotracheal, more abundant
in early wood. Rays 1-2 or rarely 3 cells wide.

LS. Vessels: intervascular pitting alternate; vessel-ray

pitting similar with narrow borders, rarely slightly
elongated; perforation plates simple. Fibres occasionally
septate. Rays 1-3 cells wide, 1-25 cells high, heterocellu-
lar; cells in RLS mainly procumbent, occasionally square
and upright.

SPECIES EXAMINED

C. speciosa Engelm., Western Catalpa.

76. Catalpa speciosa, bark TS x 60.

38
78. Catalpaspeciosa , xylem TLS x 150, showing vessels
with alternate pitting and simple perforation plates, fibres
and short narrow rays .

79. Catalpa speciosa, xylem RLS x 150, showing hetero-

cellular rays and alternate vessel-ray pitting.

39
CEANOTHUS Rhamnaceae

Bark LS. Vessels: intervascular pitting and vessel-ray pitting

Periderm consisting of irregularly-shaped cells with dark alternate, pits oval or polygonal; vessel elements long,
contents, often crushed; outer outline very irregular. with oblique simple perforation plates. Rays 1-3 cells
Cortex poorly distinguished from periderm to outside and wide; uniseriate rays 1-20 cells high, consisting of tall
phloem to inside, containing irregular groups of thick- cells (as seen in TLS); multiseriate rays 5-30 cells high,
walled fibres and stone cells with wide lumina. Phloem consisting of cells with square or more rounded outlines
rays not dilating outwards towards the cortex; conduct- (in TLS); heterocellular; cells in RLS mainly square and
ing tissue capped with large blocks of fibres and stone upright. Crystals: solitary crystals occasionally present in
cells at the boundary of phloem and cortex. Crystals: ray cells.
cluster crystals present in cortex and phloem, solitary
crystals associated with fibres.
SPECIES EXAMINED

Xylem C. american us L.; C. arboreus Greene; C. griseus McMinn.;

TS. Diffuse porous. Growth rings sometimes distinct. C. impressus TreI.; C. masonii McMinn.; Californian Lilac.
Vessels solitary, in radial chains of up to 6 cells and
occasionally in small clusters. Vessels of variable diam-
eter, those in C. impressus being widest, C. americanus the
narrowest. Fibres with wide lumina, often with gelatinous
inner walls, reducing the width of the lumina. Axial paren-
chyma paratracheal and scattered. Rays 1-3 cells wide; SO. Ceanothus impressus, bark and xylem TS x 60,
individual ray cells square or rectangular, usually larger showing blocks of fibres and stone cells in phloem and
in area than fibres. cortex, and wide vessels in xylem.
81. Ceanothusamericanus, xylem TS x 60, showing
narrow vessels and conspicuous rays.

40
82. Ceanothus arboreus, xylem TS x 60, showing one
growth ring .
83. Ceanothusarboreus, xylem TLS x 60, showing
vessels with simple perforation plates, and short, narrow
rays.

84. Ceanothus americanus. xylem TLS x 150, showing a

vessel with fine pitting and simple perforation plates,
fibres, and short, narrow rays .
85. Ceanothus impressus, xylem RLS x 150, showing a
heterocellular ray and alternate vessel-ray pitting .

41
CELASTRUS Celastaceae

Bark
Peridenn wide, consisting of alternating tapering bands,
several cells wide, of thick-walled rectangular cells, some
with dark contents, and thin-walled cells; outer layers
peeling away in wide strips. Cortex parenchymatous.
Phloem rays dilating towards cortex; sclerenchyma
absent, except for very rare stone cells in conducting
tissue. Crystals: cluster crystals present in cortex and
phloem.

Xylem
TS . Diffuse porous. Growth rings indistinct. Vessels soli-
tary and in pairs and clusters of up to 5; vessels variable in
diameter; unlignified tyloses sometimes present in
vessels. Fibres with wide lumina. Axial parenchyma rare or
absent. Rays 1-11 cells wide.

LS. Vessels: intervascular pitting alternate, pits fine, oval;

vessel-ray pitting similar, but some pits horizontally
elongated; faint spiral thickenings present in narrow
vessels and tracheids; perforation plates simple. Fibres
septate. Rays 1-11 cells wide and up to at least 70 cells
high; markedly heterocellular; ray cells usually taller than
broad in ILS, mainly square and upright in RLS.
86. Celastrus monospermus, bark and xylem TS x 60.
SPECIES EXAMINED

C. hookeri Prain; C. monospennus Roxb.

42
~. Celastrus hookeri, bark and xylem TS x 150, showing
thick- and thin-walled cells in periderm.
88. Celastrus hookeri, xylem TLS x 380, showing a vessel
with alternate pitting and spiral thickenings, septate fibres,
and a heterocellular ray.

89. Celastrus hookeri, xylem RLS x 380, showing a

heterocellular ray containing abundant starch granules.

43
CLEMATIS Ranunculaceae

Bark
Periderm consisting of few layers of thin-walled rectangu-
lar cells. Periderm forming between each year's growth of
phloem tissue, and tissue tending to break off at peri-
dermal layer. Cortex absent. Phloem rays dilating towards
outer ends, consisting of thin-walled parenchymatous
cells; conducting tissue with semi-circular areas of thin-
walled cells surrounded on outer edge by crescent shaped
blocks of fibres. Crystals absent.

Xylem
TS. Semi-ring porous. Growth rings fairly distinct. Vessels
solitary and in clusters of up to 7. Fibres thin- or thick-
walled. Axial parenchyma abundant in areas around
groups of vessels. Rays 1-15 or more cells wide, width
variable.

LS. Vessels: intervascular pitting and vessel-ray pitting

alternate, fine; perforation plates simple; vessels storied
in places. Fibres short, storied. Axial parenchyma storied.
Rays up to 15 or more cells wide in TLS; up to 120 cells
high; heterocellular; cells square and upright in RLS.

SPECIES EXAMINED 90. Clematis montana var. rubens, bark and xylem TS x
C. x jackmanii Moore (c. lanuginosa x C. viticella); C. mon- 150, showing a band of fibres in outer phloem .
tana DC. var. rubens; C. patens Morr. and Dcne.; Clematis
sp.; Clematis.

44
 91 . Clematis montana var. rubens, xylem TS x 60, show-
ing wide rays .

92. Clematis montana var. rubens, xylem TLS x 50,

showing narrow vessel elements, short fibres, and tall,
wide rays .

93. Clematis x jackmanii, xylem LS x 380, showing

storied tissues and vessel elements with polygonal,
alternate pitting and horizontal, simple perforation plates.

45
CORNUS Cornaceae

Bark
Periderm consisting of radial files of square or radially
elongated cells, usually with dark contents; outer layers
sometimes peeling in broad strips. Cortex parenchyma-
tous or consisting of a discontinuous band of thick-walled
stone cells without associated fibres. Phloem rays dilating
towards outer ends; sc1erenchyma often absent, but large
blocks of stone cells present in C. controversa. Crystals
solitary, present in phloem and cortex, often associated
with stone cells.

Xylem
TS. Diffuse porous. Growth rings well defined. Vessels
solitary and in pairs, occasionally in radial chains of up to
6, angular, thin-walled. Fibres with wide lumina. Axial
parenchyma paratrachealand scattered, sometimes in
broken uniseriate tar:entiallines. Rays 1-4 cells wide.

LS. Vessels: intervascular and vessel-ray pitting opposite,

some pits hOrizontally elongated; perforation plates very
long and oblique, scalariform, with 40 or more bars. Rays
uniseriate and 2-4 cells wide; uniseriate rays 1-27 cells
high, multiseriate rays 7-45 cells high, often with uni-
seriate portions and sometimes very long uniseriate tails;
heterocellular, cells in RLS mainly square and upright.

SPECIES EXAMINED 94. Comus amomum 'Grandiflorum', bark and xylem

TS x 80.
C. amomum Mill. 'Grandiflorum'; C. australis CA. Mey;
C. chinensis Wanger.; C controversa Hems!.; C. macro-
phylla Wall.; C. nuttallii Audub.; C. officinalis Sieb. and
Zucc.; C. sanguinea L.; C. walleri Wanger.; Comus sp.;
Dogwood.

46
96. Comus controversa, xylem TLS x 60 .
96. Comus controversa, xylem TLS x 150, showing a
vessel with fine opposite pitting and scalariform
perforation plates, and multiseriate heterocellular rays with
uniseriate portions and tails .

97. Comus controversa, xylem TLS x 380, showing a

vessel element with a scalariform perforation plate.

47
CORYLUS Corylaceae

Bark
Periderm: cells in regular radial files, outer layers often
with dark contents. Cortex with more or less continuous
cylinder of fibres and stone cells near border with phloem,
often extending into phloem rays. Phloem rays dilating
slightly towards cortex. Crystals: cluster crystals and
solitary crystals present in cortex and phloem, mainly
associated with sclerenchyma cylinder.

Xylem
TS . Diffuse porous. Growth rings distinct. Vessels nar-
row, often angular, usually in radial chains of 1-12 and
occasionally solitary and in clusters of up to 10. Fibres with
wide lumina. Axial parenchyma sparse, diffuse. Rays 1-4
cells wide, and occasional aggregate rays present.

LS. Vessels: intervascular and vessel-ray pitting usually

alternate, sometimes opposite, rarely oval; perforation
plates scalariform, with 3-10 widely spaced, often thick .•
bars; faint spiral thickenings sometimes present. Rays 1-4
cells wide, 2-100 or more cells high, multiseriate rays ,
sometimes with uniseriate portions and tails; slightly
heterocellular; cells in RLS procumbent, square and
upright.

NOTE

See Appendix, Table 3.

98. Cory/us a vel/ana , bark and xylem TS x 80, showing a
cylinder of fibres and stone cells at phloem-cortex border
SPECIES EXAMINED and vessels in radial chains .
C. avellana L., Hazel.

48
 99. Cory/us ave//ana, xylem TLS x 150, showing vessels
with fine pitting, axial parenchyma and rays.

100. Cory/us ave//ana, xylem TLS x 380, showing a vessel

element with fine, opposite pitting, and several uniseriate
rays.

101. Cory/us ave//ana, xylem RLS x 380, showing vessels

with alternate and opposite pitting and scalariform
perforation plates .

49
COT/NUS Anacardiaceae

Bark
Periderm narrow, cells often with slightly thickened walls.
Cortex narrow, interspersed with secretory canals. Phloem
rays dilating slightly towards outer ends; conducting
tissue with large sieve cells; fibres and stone cells absent;
interspersed with abundant secretory canals. Crystals:
both cluster crystals and solitary crystals abundant in
phloem and cortex.

Xylem
TS. Diffuse porous. Growth rings sometimes distinct
owing to tangential ring of narrow vessels in early wood
at growth ring boundary. Vessels solitary and in radial
chains and clusters of up to 10. Fibres often gelatinous.
Axial parenchyma mainly paratracheal. Rays 1-2 or some-
times 3 cells wide.

LS. Vessels: intervascular and vessel-ray pitting alternate,

polygonal; perforation plates simple. Rays 1-2 or rarely
up to 3 cells wide; 1-30 or more cells high; cells in RLS
mainly procumbent. Crystals rarely present in ray cells.

SPECIES EXAMINED

C. obovatus Raf. x coggyria Scop., Smoke Tree, Venetian

Sumach.
102. Cotinus obovatus x coggvria, bark and xylem
TS x 150, showing secretory canals in cortex and phloem .

50
103. Cotinus obovatus x coggyria, xylem TS x 60.
104. Cotinus obovatus x coggyria, xylem TLS x 380,
showing vessel pitting, and solitary crystals in some ray
celis.

105. Cotinus obovatus x coggyria. xylem RLS x 150,

showing vessels with alternate pitting and simple
perforation plates, vessel-ray pitting and procumbent ray
celis. some containing abundant starch granules .

51
CYTISUS Leguminosae

Bark
Peridenn consisting of several layers of thin-walled
rectangular cells; outer layers often in wedge-shaped
blocks, sometimes peeling in wide strips. Cortex narrow,
parenchymatous, sometimes with scattered groups of
1-20 gelatinous fibres. Phloem rays dilating towards
cortex; conducting tissue more or less alternating with
groups of fibres. Crystals absent.

Xylem
TS. Diffuse to semi-ring porous. Growth rings generally
indistinct. Vessels solitary, in pairs and clusters of up to 8
cells; sometimes in dendritic arrangement. Fibres thin-
walled with wide lumina, often gelatinous. Axial
parenchyma para tracheal; individual cells wider and
thinner-walled than fibres. Rays 1-11 cells wide.

LS. Vessels: intervascular pitting and vessel-ray pitting

alternate, pits oval, vestured. Spiral thickenings some-
times present, particularly on narrow vessels; perforation
plates simple; vessels storied. Axial parenchyma storied;
cells mainly fusiform. Rays 1-11 cells wide; uniseriate
rays 1-8 cells high, multiseriate rays 5 to about 100 cells
high; heterocellular; cells in RLS procumbent, square and
upright.

SPECIES EXAMINED

C. hirsutus L.; C. multiflorus Sweet 'Incarnatus'; C. pur-

gans Spach.; C. scoparius Link.; Cytisus sp.; Broom.

52
108. CVtisus hirsutus, xylem TS x 150.
109. CVtisus sp ., xylem TLS x 150, showing storied,
narrow vessels with simple perforation plates, and axial
parenchyma, non-storied fibres, and rays .
110. Cvtisus multiflorus 'Incarnatus', xylem TLS x 150,
showing storied axial parenchyma, fibres and rays
111. CVtisus hirsutus, xylem RLS x 150, showing a
heterocellular ray and vessel-ray pitting.

106. Cvtisus multiflorus 'Incarnatus' , bark and xylem

TS x 80.
107. CVtisus sp., xylem TS x 60, showing narrow vessels
in oblique multiples .

53
DAPHNE Thymelaeaceae

Bark
Periderm narrow, up to about 5 cells wide; outer layers
consisting of collapsed and broken cells; individual cells
thin-walled. Cortex with a more or less continuous ring of
fibres and stone cells. Phloem rays usually dilating
towards outer ends; conducting tissue broken up by
irregularly distributed groups of thick-walled gelatinous
fibres. Crystals: solitary crystals abundant in association
with stone cells in cortex, cluster crystals rare in ray cells.

Xylem
TS. Diffuse porous. Growth rings indistinct. Vessels
solitary and in pairs, and radial chains and clusters of up
to 5. Fibres in blocks, often gelatinous; sometimes thin-
walled and difficult to distinguish from axial parenchyma
cells. Axial parenchyma paratracheal and in some areas
forming most of axial tissue. Rays 1-4 cells wide, some-
times containing solitary crystals.

LS. Vessels: intervascular pitting coarse, pits oval or 112. Daphne mezereum f. alba, bark and xylem TS x SO,
polygonal, alternate; vessel-ray pitting slightly smaller, showing a young root.
circular; perforation plates simple; spiral thickenings 113. Daphne mezereum f. alba, xylem TLS x 150, show-
present on some narrow vessels and tracheids. Rays 1-4 ing crystals in some ray cells.
cells wide; uniseriate rays 1-16 cells high, multiseriate
rays 5-48 cells high; heterocellular; cells in RLS square
and procumbent.

SPECIES EXAMINED

D. mezereum L. forma alba, Mezereon.

54
114. Daphne mezereum f. alba, xylem RLS x 380, show-
ing a vessel with alternate, polygonal pitting and a hetero-
cellular ray.

55
DEUTZIA Philadelphaceae

Bark 1-17 cells high, consisting mainly of upright cells, multi-

seriate rays 19-90 cells high; heterocellular; cells in RLS
Periderm stratified, composed of layers of thin-walled
mainly square and upright, very few procumbent.
rectangular cells alternating with bands of wider, irregu-
larly-shaped parenchyma cells together with blocks of
thick-walled stone cells. Cortex very narrow or absent,
SPECIES EXAMINED
with periderm arising almost directly adjacent to phloem.
Phloem rays dilating towards outer ends; conducting D. x maliflora Rehd. !D. x lemoinii (D. gracilis x parviflora)
tissue capped at outer edge by blocks of up to 15 stone x D. purpurascensJ; D. scabra Thunb.; Deutzia sp.
cells similar to those in periderm. Crystals absent.

Xylem
TS. Diffuse porous. Growth rings indistinct. Vessels
narrow, mainly solitary, and also in pairs; thin-walled, 115. Deutzia x ma/if/ora, bark and xylem T3 x 150.
angular; tracheids frequent. Fibres narrow with narrow
lumina. Axial parenchyma sparse, diffuse. Rays 1-6 cells 116. Deutzia sp., xylem T3 x 150.
wide; ray cells several times wider than fibres, often
exceeding vessels in area.

LS. Vessels: intervascular and vessel-ray pitting fine, oval

to horizontally elongated, opposite; perforation plates
scalariform, long and oblique, with up to 35 closely
spaced bars; tracheids frequent, narrower and much
longer than vessel elements, with spiral thickenings.
Fibres: most septate. Rays 1-6 cells wide; uniseriate rays

56
117. Deutzia x rna/if/ora, xylem TLS x 60, showing tall,
multiseriate and shorter, narrower rays.
118. Deutzia x rna/if/ora, xylem TLS x 150, showing a
vessel with fine alternate pitting, and with spiral thicken - 119. Deutzia scabra, xylem LS x 600, showing fibre-
ings. tracheids with spiral thickenings.
120. Deutzia x malif/ora, xylem RLS x 380, showing a
scalariform perforation plate and a heterocellular ray.

57
DIERVILLA and WEIGELA Caprifoliaceae

Bark LS. Vessels: intervascular pitting and vessel-ray pitting

Periderm consisting of radial files of thin-walled rectangu- mainly opposite, scalariform in places and sometimes
lar cells, the outer layers often disintegrating; two or alternate; perforation plates very oblique, scalariform
more periderms may develop, one arising adjacent to with up to 40 or more bars. Fibres very occasionally with
phloem. Cortex parenchymatous or collenchymatous, spiral thickenings. Rays 1-4, occasionally up to 7 cells
sometimes containing small groups of 1-5 thick-walled wide; uniseriate rays 1-20 cells high, multiseriate rays up
angular fibres with narrow lumina. Phloem rays dilating to 150 cells high, many with uniseriate tails; markedly
towards cortex, conducting tissue sometimes containing heterocellular; cells of varying shape in TLS, square,
small groups of angular thick-walled fibres, or less upright and less often procumbent in RLS.
frequently blocks of up to about 20 small, thick-walled
stone cells near the phloem-cortex boundary. Crystals:
cluster crystals often in phloem, sometimes entirely NOTE
absent; solitary crystals in D. rivularis. See Appendix, Table 4.

Xylem SPECIES EXAMINED

TS. Diffuse porous. Growth rings usually indistinct. D. lonicera Mill.; D. rivularis Gatt.; D. sessilifolia Buckl.;
Vessels usually angular, solitary, in pairs and very D. x splendens Kirchn. (D. lonicera x sessilifolia); W. decora
occasional radial chains and clusters of up to 5. Fibres with Nakai; W. floribunda C. A. Mey.; W. florida A.DC.; W. prae-
wide lumina. Axial parenchyma paratracheal and diffuse. cox Bailey.
Rays 1-4 and occasionally up to 7 cells wide.

58
121. Weige/a praecox, bark and xylem TS x 60, showing 126. Weige/a praecox, xylem RLS x 380, showing a
a multiple periderm . scalariform perforation plate and fibre-tracheids.
122. Diervi//a rivu/aris, bark and xylem TS x 60 , showing
small stone cells in outer phloem and cortex.
123. Weige/a decora, bark and xylem TS x 80.
124. Weige/a praecox, xylem TLS x 60, showing narrow
rays with uniseriate tails.
125. Weige/a praecox, xylem TLS x 380, showing a vessel
with opposite and elongated pitting, a scalariform
perforation plate, and several uniseriate portions of rays.

59
ELAEAGNUS Elaeagnaceae

Bark
Periderm consisting of regular radial files of rectangular
cells, some with dark contents; outer layers crushed and
peeling in strips. Cortex narrow, parenchymatous. Phloem
rays dilating towards cortex; conducting tissue capped
with triangular blocks of many thin-walled fibres. Crys-
tals: cluster crystals present in variable numbers in rays,
phloem and cortex, sometimes abundant.

Xylem
TS. Diffuse porous. Growth rings indistinct. Vessels
solitary and in pairs, occasionally in radial chains of up to
4. Fibres with wide lumina. Axial parenchyma sparse. Rays
1-5 cells wide.

LS. Vessels: intervascular and vessel-ray pitting fine,

alternate; perforation plates simple, horizontal. Rays 1-5
cells wide; uniseriate rays 1-21 cells high; multiseriate 127. E/aeagnus umbel/ata, bark and xylem TS x 60.
rays 7-82 cells high, often with uniseriate tails; hetero- 128. E/aeagnus pungens 'Variegata', bark and xylem
cellular, cells in RLS procumbent, square and upright. TS x 60.

SPECIES EXAMINED

E. pungens Thunb. 'Variegata'; E. umbellata Thunb.;

Oleaster.

60
129. E/aeagnus pungens 'Variegata' , bark and xylem
TS x 150.

130. Elaeagnus umbellata, xylem TLS x 150, showing

narrow vessel elements with fine alternate pitting, and tall
rays .

131. E/aeagnus umbellata, xylem TLS x 150, showing

wide vessel elements with fine alternate pitting, and short ,
narrow rays .

61
ESCALLONIA Escalloniaceae

Bark
Peridenn very irregular in appearance; cells thin-walled.
Cortex narrow or absent, parenchymatous. Phloem
usually extending to periderm; rays not dilating widely
towards outer ends; conducting tissue interspersed with
thick-walled fibres, either solitary or in small groups.
Crystals absent.

Xylem
TS. Diffuse porous. Growth rings usually indistinct.
Vessels mainly solitary, and occasionally in tangential
pairs or clusters of up to 4. Fibres moderately thick-
walled. Axial parenchyma paratracheal and apotracheal.
Rays 1-3 cells wide; cells rounded, and similar in appear-
ance to axial parenchyma cells.

LS. Vessels: intervascular and vessel-ray pitting opposite

and elongated; perforation plates oblique, scalariform,
with 10-20 or more narrow bars. Rays 1-3 cells wide;
usually up to 40 cells high with uniseriate tails; hetero-
cellular; cells in RLS mainly square and upright.

SPECIES EXAMINED

E. viscosa Forbes.

132. Escallonia viscosa, bark TS x 150, showing

scattered, thick-walled fibres in phloem and cortex .

62
133. Escallonia viscosa, bark and xylem TS x 150, show-
ing irregular outline of xylem.
134. Escallonia viscosa, xylem TS x 150, showing several
scalariform perforation plates.

135. Escallonia viscosa, xylem LS x 150, showing hetero-

cellular rays, and vessels with scalariform perforation
plates.

63
fUGAL YPTUS Myrtaceae

Bark
Periderm consisting of layers of thin-walled square or
rectangular cells, usually alternating with layers of cells
with thicker, often lignified walls; periderm often also
forming deep in the pericycle, cells outside the new layer
of periderm being worn away. Cortex narrow or some-
times absent; when present, with small blocks of 1-20
fibres interspersed throughout; secretory canals often
present. Phloem rays dilating slightly, sometimes widely
at outer ends, consisting of parenchymatous tissue,
sometimes with secretory canals; conducting tissue
consisting of thin-walled cells alternating with irregularly
shaped blocks of 4-25 fibres. Crystals: solitary crystals
often present in phloem.

Xylem
T5. Diffuse porous. Growth rings often indistinct. Vessels
solitary, rarely in pairs, occasionally forming loose
oblique lines in late wood; circular or oval. Fibres often
with wide lumina. Axial parenchyma vasicentric and
diffuse. Rays uniseriate, or rarely 2 cells wide.

L5. Vessels: intervascular pitting alternate or occasionally

elongated, vestured; vessel-ray pitting larger than inter-
136. Eucalyptus coccifera, bark and xylem TS x 50, show-
vascular pitting, often elongated; perforation plates
ing blocks of fibres in phloem and cortex.
simple; tyloses often present. Rays usually uniseriate, or
occasionally 2 cells wide in places; 1-25 cells high; cells
square and procumbent in RLS.

SPECIES EXAMINED

E. coccifera Hook. f.; E. gunnii Hook. f.; E. paroifolia Cam-

bage; Gum Tree.

139. Eucalyptus coccifera, xylem TLS x 150, showing

uniseriate rays and elongated vessel-ray pitting.
140. Eucalyptus coccifera, xylem TLS x 380, showing
vestured vessel pitting.

64
137. Eucalyptus coccifera, bark TS x 150, showing outer
bark with two periderms of alternating thick- and thin -
walled cells.
138. Eucalyptus coccifera, xylem TS x 55, showing
solitary vessels and uniseriate rays .

65
EUONYMUS Celastraceae

Bark
Periderm often peeling in short strips; cells rectangular,
often with dark contents. Cortex narrow, parenchyma-
tous; fibres and stone cells absent. Phloem rays dilating
towards outer ends; fibres and stone cells absent, or stone
cells rarely present in rays. Crystals: cluster crystals
present in phloem and cortex.

Xylem
TS. Diffuse to semi-ring porous; growth rings distinct.
Vessels narrow, solitary, occasionally in groups of 2-4,
and in tangential groups of 2-4 at beginning of growth
rings. Fibres with moderately thickened walls. Axial
parenchyma apotracheal, scattered. Rays 1-2 cells wide.

LS. Vessels: intervascular pitting and vessel-ray pitting

fine, opposite; marked spiral thickenings often present;
perforation plates simple, oblique. Rays uniseriate or
often 2 cells wide in places, 2-55 cells high; cells in RLS
usually square, occasionally procumbent or upright.

141. Euonvmusjaponicus, bark TS x 80.

SPECIES EXAMINED

E. europaeus L.; E. japonicus Thunb.; Spindle Tree.

144. Euonvmus europaeus, xylem RLS x 380, showing

vessels with spiral thickenings, fibre-tracheids, and
uniseriate rays.
145. Euonvmusjaponicus, xylem RLS x 380, showing
vessels with simple perforation plates, and vessel-ray
pitting .

66
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142. Euonymusjaponicus, xylem TS x 80.

143. Euonymusjaponicus, xylem TLS x 150, showing tall
narrow vessel elements, and uniseriate rays with
occasional biseriate portions .

67
FAGUS Fagaceae

Bark
Periderm often peeling in short or long strips; cells
rectangular, sometimes with tannin. Cortex with more or
less continuous cylinder of thick-walled stone cells and
occasionally groups of fibres. Phloem rays dilating
towards outer ends, large rays often with group of stone
cells and fibres projecting towards xylem rays; conduct-
ing tissue thin-walled with few fibres and stone cells
round periphery. Crystals: solitary and cluster crystals
present in cortex and sometimes also in phloem.

Xylem
TS. Semi-ring porous. Growth rings distinct. Vessels
solitary, in tangential bands of 2-5, and in clusters of up
to 6; sometimes very wide. Fibres thin- or thick-walled.
Axial parenchyma abundant, apotracheal and paratracheal.
Rays: uniseriate rays with tortuous path round large
vessels, and multiseriate rays 2-20 cells wide, slightly
noded at growth rings. Crystals rarely present in ray cells,
solitary.

LS. Vessels: intervascular pitting opposite or scalariform;

vessel-ray pitting scalariform, often orientated in several 146. Fagus svlvatica, bark and xylem TS x 150, showing a
directions; perforation plates both simple and less com- cylinder of stone cells and fibres at cortex-phloem
monly scalariform, with 3-15 narrow bars. Rays uni- boundary, solitary crystals in bark, and a wide ray in xylem
and phloem.
seriate and 2-20 cells wide; uniseriate rays 1-15 cells
high, multiseriate rays 8-150 cells high, sometimes with
characteristic 'shoulder' where end of ray closely adjac-
ent to end of ray above or below, the two rays separated
by very few axial elements; cells in RLS mainly square
and procumbent.

SPECIES EXAMINED

F. sylvatica L., Beech.

68
147. Fagus sylvacica, xylem TS x 80, showing rays of
varying width.

148. Fagussylvacica , xylem TLS x 150, showing fibres,

axial parenchyma, and wide and narrow rays.

149. Fagus sylvatica, xylem RLS x 380, showing a vessel

with a scalariform perforation plate, fibres, fibre-tracheids ,
and ray cells, some with a solitary crystal.

69
FICUS Moraceae

Bark SPECIES EXAMINED

Periderm consisting of several layers of rectangular cells F. carica L., Fig.

with thickened periclinal walls, sometimes with dark
contents. Cortex narrow, parenchymatous; latex canals
often present. Phloem rays dilating towards outer ends;
conducting tissue in long wedge-shaped areas with scat-
tered thick-walled, sometimes gelatinous, fibres; latex
canals often present in phloem rays. Crystals: solitary and
double crystals and occasional cluster crystals present in
periderm, cortex and phloem.

Xylem
TS . Diffuse porous. Growth rings indistinct. Vessels
solitary and in radial chains of 2-7 cells and occasionally
in clusters of up to 6 cells; vessel walls often thickened
where adjacent to other vessels; tyloses often present.
Fibres thick-walled, often gelatinous, in small blocks in
parenchymatous areas. Axial parenchyma forming bulk of 150. Ficus carica, bark TS x 150, showing periderm,
ground tissue. Rays 1-5 cells wide; cells large and thin- cortex with one latex canal, and scattered solitary crystals.
walled.
151. Ficus carica, xylem TS x 50, showing many vessels
in short, radial chains .
LS. Vessels: intervascular pitting alternate, fine, oval to
polygonal or occasionally elongated; vessel-ray pitting
oval, elongated, sometimes scalariform; perforation
plates simple, often transverse; vessel elements some-
times storied. Axial parenchyma storied. Rays 1-5 cells
wide; uniseriate rays 2-12 cells high, multiseriate rays up
to 70 or more cells high, often with uniseriate tails;
heterocellular; cells in RLS procumbent, square and
upright.

70
152. Ficus carica, xylem TS x 150, showing vessels , 154. Ficus carica, xylem TLS x 380, showing a vessel with
gelatinous fibres and axial parenchyma. fine, alternate pitting.

153. Ficus carica, xylem TLS x 150, showing storied axial 155. Ficus carica, xylem RLS x 380, showing vessel-ray
parenchyma, and wide rays. pitting.

71
FOREST/ERA Oleaceae

Bark
Periderm consisting of thin-walled cells, sometimes with
outer layers thicker-walled, lignified; often splitting into
large triangular or rectangular blocks. Cortex consisting of
parenchymatous cells. Phloem rays dilating at outer ends;
large groups of stone cells present at border between
phloem and cortex; fibres absent. Crystals absent.

Xylem
TS. Diffuse porous to semi-ring porous. Growth rings
fairly distinct. Vessels narrow, solitary and in clusters of
up to 6, sometimes with dendritic arrangement. Fibres
with wide lumina. Axial parenchyma very sparse, para-
tracheal. Rays 1-2 (or rarely 3) cells wide; ray cells some-
times as large as vessels.

LS. Vessels: intervascular pitting and vessel-ray pitting

fine, alternate; perforation plates simple, oblique. Rays
1-2 (or rarely 3) cells wide, usually uniseriate with biseri-
ate portions, cells in uniseriate portions often with
oblique walls in TLS; heterocellular; cells in RLS pro-
cumbent, square and upright.

SPECIES EXAMINED

Forestiera sp.

156. Forestiera sp., bark and xylem TS x 60, showing

blocks of stone celis in phloem .

72
157. Forestiera sp ., bark RLS x 150, showing a block of
stone cells .

158. Forestiera sp., xylem TLS x 150, showing narrow

vessels, fibres, and narrow heterocellular rays .

159. Forestiera sp., xylem TLS x 380, showing a vessel

with fine alternate pitting, fibres, and rays .

73
FORSYTHIA Oleaceae

Bark
Peridenn: cells thin-walled, often square or rectangular,
cells in innermost layers occasionally narrow, thick-
walled and lignified; groups of cells on outside often
splitting to form triangular or rectangular blocks. Cortex
with groups of thick-walled stone cells, or in some roots
stone cells scarce or absent; in some cases a distinct
cylinder of parenchymatous cells with numerous cross
walls (pericycle) present. Phloem rays dilating slightly
towards outer ends; fibres absent; stone cells usually
absent. Crystals usually absent.

Xylem
TS. Diffuse or semi-rIng porous. Growth rings fairly
distinct. Vessels crowded, mainly solitary and sometimes
in clusters of up to 4. Fibres with narrow or wide lumina.
Axial parenchyma sparse, paratracheal. Rays 1-3 cells
wide.

LS. Vessels: intervascular and vessel-ray pitting alternate,

fine; perforation plates slightly oblique, simple, or rarely
scalariform or reticulate with thin bars; spiral thickenings
sometimes present; fibre tracheids present, often with
spiral thickenings. Rays 1-3 cells wide; uniseriate rays
1-30 cells high, multiseriate rays 5-40 cells high, often
with uniseriate tails; heterocellular; cells in RLS mainly
square and upright and sometimes procumbent.
160. Forsvthia suspensa, bark and xylem TS x 150.

SPECIES EXAMINED

F. x intennedia Zab. (F. suspensa x viridissima) var. spec-

tabilis; F. suspensa Yah!.

74
161. Forsythia suspensa, bark TS x 150, showing a block
of thick-walled stone cells in phloem.
162. Forsythia x intermedia var. spectabilis, xylem
TS x SO.

163. Forsythiasuspensa, xylem TLS x 150, showing

vessels with fine alternate pitting, fibre-tracheids, and
narrow heterocellular rays.

75
FRAXINUS Oleaceae

Bark
Periderm: cells thin-walled; groups of cells often splitting
into wedge-shaped blocks. Cortex interspersed with
groups of thick-walled fibres and stone cells, sometimes
forming a continuous sclerenchyma cylinder. Phloem rays
dilating slightly near cambium then becoming wider at
outer ends; conducting tissue with groups of fibres and
stone cells, as in cortex. Crystals: crystal sand present in
cortex and phloem.

Xylem
IS . Semi-ring porous. Growth rings usually distinct.
Vessels solitary and in radial pairs with adjoining walls
thickened at corners, also occasionally in clusters and
radial chains of up to 4 cells; vessel width variable. Fibres
with wide lumina. Axial parenchyma paratracheal and
sparsely scattered. Rays 1-2 cells wide.

LS. Vessels: intervascular pitting and vessel-ray pitting

alternate, fine; perforation plates simple. Rays 1-2 cells
wide; 1-21 cells high; heterocellular; cells often thick-
walled; cells in RLS procumbent and square, with
marginal cells sometimes upright.

SPECIES EXAMINED

F. excelsior L., Ash.

164. Fraxinus excelsior, bark and xylem TS x 80, showing
groups of fibres and stone cells in cortex and phloem.

167. Fraxinus excelsior, xylem TLS x 80.

168. Fraxinus excelsior, xylem TLS x 150, showing vessel
elements with fine, alternate pitting and short, narrow
rays .

76
166. Fraxinus excelsior, xylem TS x SO, showing
unusually narrow vessels .
166. Fraxinus excelsior, xylem TS x SO, showing a
complete narrow growth ring and both wide and narrow
vessels .

77
FUCHSIA Onagraceae

Bark
Periderm consisting of several layers of thin-walled, regu-
larly arranged square and rectangular cells; sometimes
peeling in long strips round outside edge. Cortex narrow,
parenchymatous; fibres and stone cells absent. Phloem
rays dilating slightly towards outer ends, cells of rays and
conducting tissue thin-walled, fibres and stone cells
absent. Crystals: raphides present in cells in cortex and
phloem.

Xylem
TS. Diffuse porous. Growth rings indistinct. Vessels thin-
walled, angular, solitary or more usually in clusters, and
radial chains of 2-7 or more. Fibres thin-walled with wide
lumina. Axial parenchyma diffuse. Rays 1-4 cells wide.

LS. Vessels sometimes storied; intervascular and vessel-

ray pitting alternate, polygonal or elongated, sometimes
with indistinct borders. Perforation plates simple; tyloses
often present. Fibres septate, often short and storied. Rays
1-4 cells wide, 3-20 cells high, or sometimes rays fused
and appearing taller; markedly heterocellular; cells thin-
walled; cells in RLS procumbent, square and upright.

SPECIES EXAMINED

F. magellanica Lam. 'Riccartonii'; Fuchsia sp.

169. Fuchsia magellanica 'Riccartonii', bark and xylem

TS x 60 .

172. Fuchsia magel/anica 'Riccartonii', xylem RLS x 150,

showing a heterocellular ray and alternate vessel-ray
pitting .

173. Fuchsia magellanica 'Riccartonii', xylem LS x 380,

showing vessels with fine, alternate pitting, and starch
granules in some cells .

78
170. Fuchsia mage/lanica 'Riccartonii', bark LS x 380,
showing cortical parenchyma cells containing raphides .
171. Fuchsia mage/lanica 'Riccartonii' , xylem TLS x 150,
showing vessels with alternate pitting, septate fibres, and
narrow heterocellular rays.

79
GARRYA Garryaceae

Bark
Periderm consisting of radial files of square and rectangu-
lar cells, most thin-walled, or occasionally thick-walled;
cells in outer layers disintegrating, giving uneven sur-
face. Cortex mainly parenchymatous with occasional
groups of 1-10 thick-walled fibres. Phloem rays dilating
slightly, then widely towards cortex, sometimes includ-
ing thin-walled stone cells with wide lumina; conducting
tissue in blocks with scattered fibres, capped with large,
triangular, more or less continuous blocks of thin-walled
fibres with a few stone cells. Crystals absent.

Xylem
TS. Diffuse porous. Growth rings indistinct. Vessels
narrow, solitary and occasionally in pairs. Fibres narrow,
thick-walled. Axial parenchyma paratracheal and diffuse.
Rays usually 1-6 cells wide, occasionally up to 15 cells
wide.

LS. Vessel elements long and narrow with long tails; inter-
vascular and vessel-ray pitting fine, alternate; perforation
plates oblique, scalariform with 2-4 widely spaced bars;
fine spiral thickenings present. Fibre-tracheids present. 174. Garrya e/liptica, bark and xylem TS x 60, showing
Rays 1-6 cells wide, occasionally up to 15 cells wide; fibre blocks in phloem.
uniseriate rays usually less than 10 cells high, multiseriate
rays up to more than 100 cells high; heterocellular; cells
generally taller than broad in TLS, in RLS mainly square
and upright, few procumbent.

SPECIES EXAMINED

G. elliptica Lind!.

80
175. Garrya el/iptica, xylem TS x 60, showing centre of
root, narrow vessels , and wide and narrow rays .
176. Garrya el/iptica, xylem TLS x 150, showing narrow
vessels with fine, alternate pitting, and tall , heterocellular
rays .

177. Garrya el/iptica , xylem RLS x 600, showing a vessel

with fine spiral thickenings, a scalariform perforation plate ,
and fibre-tracheids .

81
HAMAMELIS Hamamelidaceae

Bark
Peridenn with regular radial files of narrow, radially
elongated cells, sometimes thick-walled and occasionally
with dark contents. Cortex parenchymatous, cells often
with dark contents, and rarely thick-walled. Phloem ' •
narrow with rays not dilating strongly; conducting tissue
interspersed with groups of 1-15 thick-walled fibres.
Crystals: solitary and occasional cluster crystals present in
phloem and cortex.

Xylem
TS . Diffuse porous. Growth rings indistinct. Vessels
narrow, often angular, solitary and in radial chains and
clusters of 2-6. Fibres moderately thick-walled. Axial
parenchyma diffuse . Rays 1-3 cells wide.

LS. Vessels: intervascular and vessel-ray pitting opposite

and scalariform; perforation plates oblique, scalariform,
with 10-20 narrow bars. Rays 1-3 cells wide, 2-50 or more
cells high; heterocellular; cells in RLS procumbent,
square, and upright.
178. Hamamelis virginiana, bark TS x 150, showing some
cortex and ray cells with starch granules and small groups
of fibres and solitary crystals in phloem.
SPECIES EXAMINED

H. virginiana L., Witch Hazel.

181. Hamamelis virginiana, xylem RLS x 150, showing

vessels with opposite and scalariform intervascular and
vessel-ray pitting and scalariform perforation plates.
182. Hamamelis virginiana, xylem RLS x 380, showing
vessels with opposite and scalariform pitting and a scalari-
form perforation plate .

82
 179
179. Hamamelis virginiana, bark and xylem TS x 60.
180. Hamamelis virginiana, xylem TLS x 150, showing
narrow vessels, fibres and narrow, heterocellular rays.

83
HEBE Scrophulariaceae

Bark
Peridenn often irregular, cells thick-walled and occasion-
ally with dark contents. Cortex parenchymatous, inner
part interspersed with groups of fibres and occasional
stone cells near boundary with phloem. Sc1erenchyma
varying from sparse to abundant, in interrupted ring
around phloem. Phloem: conducting tissue in continuous
cylinder, without rays . Crystals absent.

Xylem
TS. Diffuse porous. Growth rings indistinct. Vessels
narrow, solitary and in radial chains of up to 10. Fibres
with wide lumina. Axial parenchyma very sparse or
absent. Rays absent.

LS. Vessels often storied; intervascular pitting alternate or

often slightly elongated; perforation plates simple. Fibres
often storied. Rays absent.

SPECIES EXAMINED

H. speciosa R. Cunn. 'Cranleighensis'; H. speciosa

R. Cunn. 183. Hebe speciosa 'Cranleighensis', bark and xylem
TS x 150, showing groups of wide fibres in phloem and
cortex.

84
184. Hebe speciosa 'Cranleighensis', xylem TS x 80; note
the absence of rays.
185. Hebe speciosa 'Cranleighensis' , xylem TLS x 380,
showing a vessel with alternate and elongated pitting .

186. Hebe speciosa 'Cranleighensis' , xylem LS x 600,

showing vessel pitting and fibres .

85
HEDERA Araliaceae

Bark LS. Vessels: intervascular pitting opposite, alternate or

sometimes elongated; vessel-ray pitting circular or
Periderm: outer cells with thickened, lignified outer peri-
usually elongated; perforation plates simple. Fibres
clinal walls; inner cells usually thin-walled. Cortex paren-
usually septate. Rays 1-10 cells wide; uniseriate rays 3-20
chymatous, sometimes without sclerenchyma, sometimes
cells ~igh; multiseriate rays 6-200 or more cells high,
with groups of fibres with wide lumina, and rarely stone
sometimes with uniseriate tails; heterocellular; cells in
cells; vertical secretory canals present in cortex and
RLS procumbent, square and upright.
p~loem. Phloem rays dilating slightly, conducting tissue
wIth groups of fibres with wide lumina, and rarely stone
cells. Crystals: cluster crystals present in cortex and
SPECIES EXAMINED
phloem, sometimes very abundant.
H. helix L., Ivy.
Xylem
TS. Semi-ring to diffuse porous. Growth rings fairly in-
distinct. Vessels thin-walled, solitary and in radial chains
and tangential pairs or clusters of up to 5. Axial paren-
chyma frequent, paratracheal. Fibres with wide lumina.
Rays 1-10 cells wide.

188. Hedera helix, bark TS x 380, showing phloem sieve

cells, parenchyma cells containing cluster crystals, and a
secretory canal.

187. Hedera helix, bark TS x 150, showing abundant

cluster crystals and secretory canals in cortex and phloem.

86
189. Hedera he/ix, bark TS x 80, showing scattered
groups of thin-walled fibres in phloem. 191. Hedera he/ix, xylem TLS x 80, showing narrow
190. Hedera he/ix, xylem TS x 80, showing angular vessels, septate fibres and rays of two sizes .
vessels and paratracheal parenchyma. 192. Hedera helix, xylem RLS x 380, showing a vessel
with opposite and elongated pitting and a simple perfor-
ation plate.

87
HIBISCUS Malvaceae

Bark
Peridenn tending to split into wedge-shaped blocks.
Cortex narrow, parenchymatous. Phloem with well
defined triangular regions consisting of alternate layers of
conducting tissue and fibres with wide lumina; phloem
rays widely dilating towards outer ends, parenchyma-
tous. Crystals: cluster crystals present in cortex and
phloem.

Xylem
TS. Diffuse to seml-nng porous. Growth rings fairly
distinct. Vessels usually angular, solitary, in clusters and
radial chains of up to about 9 cells, or occasionally in small
tangential chains. Fibres thin-walled. Axial parenchyma
abundant; both paratracheal and diffuse. Rays 1-4 cells
wide.

LS. Vessels: intervascular and vessel-ray pitting alternate,

fine; perforation plates simple. Fibres often storied. Fibre-
tracheids present, storied, with spiral thickenings. Axial 193. Hibiscus syriacus, bark TS x 60, showing blocks of
parenchyma storied. Rays 1-4 cells wide; 3-36 cells high; fibres in phloem .
heterocellular; cells in RLS procumbent, square and
upright.

SPECIES EXAMINED

H. syriacus L.

196. Hibiscus syriacus, xylem TLS x 150, showing vessels

with fine alternate pitting, fibres, and heterocellular rays
containing starch granules.
197. Hibiscus syriacus, xylem RLS x 150, showing a
heterocellular ray.

88
194. Hibiscus syriacus, xylem TS x 60, showing vessel
multiples and conspicuous axial parenchyma.

195. Hibiscus syriacus, xylem TS x 150.

89
HYDRANGEA Hydrangeaceae

Bark
Peridenn: cells thin-walled or occasionally with thickened
periclinal walls. Cortex absent or very narrow, paren-
chymatous. Phloem cells thin-walled; fibres and stone
cells absent; rays dilating towards outer ends. Crystals:
raphides present in phloem.

Xylem
TS. Diffuse porous. Growth rings usually indistinct.
Vessels solitary and in tangential and radial pairs, and in
clusters and radial chains of up to 6; narrow and often
angular. Fibres with wide lumina. Axial parenchyma very
sparse. Rays 1-6 cells wide; cells often as wide as vessels.

LS. Vessels: intervascular and vessel-ray pitting scalari-

form and opposite or rarely alternate; perforation plates
scalariform and long, with 20-60 very fine bars; tyloses
often present. Fibres septate. Rays 1-6 cells wide, 8-50
cells high, often with uniseriate tails; markedly hetero-
cellular; cells in RLS mainly square and upright.

SPECIES EXAMINED

H. paniculata Sieb., Hydrangea sp. 198. Hydrangea paniculata, bark and xylem TS x 60,
showing very narrow vessels .

201. Hydrangeapaniculata, xylem RLS x 150, showing

scalariform, intervascular and vessel-ray pitting and
upright ray cells.
202. Hydrangea paniculata, xylem RLS x 380, showing a
scalariform perforation plate and vessel-ray pitting .

90
199. Hydrangea sp., xylem TLS x 150, showing narrow
vessels with scalariform pitting, septate fibres, and
narrow, heterocellular rays .
200. Hydrangea sp., xylem TLS x 150.

91
HYPERICUM Guttiferae

Bark
Periderm: outer layers often with dark contents and peel-
ing in short strips; inner region often in alternating layers
of narrow cells with dark contents and radially wider cells
without contents. Cortex parenchymatous; sclerenchyma
usually absent although walls of parenchyma cells some-
times slightly thickened; secretory canals present, inter-
spersed throughout cortex and sometimes in phloem,
consisting of central canal surrounded by thin-walled
epithelial cells. Phloem rays dilating very little towards
cortex; sclerenchyma usually absent; secretory canals
present. Crystals absent.

Xylem
TS. Diffuse porous. Growth rings sometimes indistinct.
Vessels narrow, angular, solitary and in pairs and radial
chains and clusters of up to 7. Fibres with wide lumina.
Axial parenchyma mainly paratracheal, sometimes abun-
dant. Rays 1-4 cells wide.

LS. Vessels: intervascular and vessel-ray pitting alternate, 204. Hvpericum x inodorum, bark TS x 380, showing
polygonal or sometimes elongated and variously orien- cortex with secretory canals.
tated; perforation plates simple, oblique. Fibres septate.
Rays 1-4 cells wide, 2-40 cells high; multiseriate rays
often with uniseriate tails; heterocellular; cells in RLS
procumbent, square and upright.

SPECIES EXAMINED

H. androsaemum L., Tutsan; H. x inodorum Mill. (H. andro-

saemum x H. hircinum); Hypericum sp.

203. Hvpericum androsaemum, bark TS x 150, showing

periderm and one secretory canal in cortex .

92
206. Hypericum x inodorum, bark and xylem TS x 80.
207. Hypericum sp ., xylem TLS x 150, showing rays, fusi-
206. Hypericum x inodorum, xylem TS x 60. form axial parenchyma and narrow vessels .
208. Hypericum sp ., xylem RLS x 380, showing vessel
pitting, simple perforation plates, and abundant starch
granules in axial parenchyma.

93
fLEX Aquifoliaceae

Bark tyloses occasionally present. Rays abundant, in two dis-

tinct sizes; uniseriate and 2-7 cells wide; uniseriate rays
Periderm slightly peeling; cells rectangular, regularly
1-15 cells high, multiseriate rays 8-35 cells high, often
divided, sometimes with thickened periclinal walls.
with uniseriate tails; heterocellular; cells in RLS procum-
Cortex narrow, parenchymatous, fibres absent, stone cells
bent, square and upright. Crystals sometimes present in
rarely present. Phloem: fibres and stone cells usually
ray cells and parenchyma, as described above.
absent; rays dilating slightly towards outer ends. Crystals:
solitary, double and cluster crystals and crystal sand
present in cortex and phloem.
SPECIES EXAMINED

I. aquifolium L., Ilex sp., Holly.

Xylem
TS. Diffuse or semi-ring porous. Growth rings usually
distinct. Vessels usually solitary, rarely in pairs and radial
chains of up to 5; often angular; frequency and size vari-
able . Fibres thin-walled with wide lumina. Axial paren-
chyma abundant, scattered. Rays 1-7 cells wide, hetero-
209. /lex aquifolium, bark and xylem TS x 80.
cellular, cells sometimes wider than vessels. Crystals
occaSionally present ~;-. ray cells and rarely in axial paren- 210. /lex aquifolium, xylem TS x 80 .
chyma; solitary, double and cluster crystals and crystal
sand. 211. /lex aquifolium, xylem TLS x 150, showing fine,
opposite vessel pitting, and rays.
LS. Vessels: intervascular pitting opposite; vessel-ray pit- 212. /lex aquifolium, xylem TLS x 380, showing crystals in
ting opposite or scalariform; spirals present; perforation two ray cells.
plates scalariform, oval, with up to 35 bars; thin-walled
213. /lex aquifolium, xylem LS x 380, showing fibre-
tracheids and vessels with spiral thickenings.
214. /lex aquifolium, xylem RLS x 380, showing vessel
and fibre pitting and a scalariform perforation plate.

94
95
JASMINUM Oleaceae

Bark SPECIES EXAMINED

Periderm: cells usually fairly thin-walled and often in 1. humile L.; 1. nudiflorum Lind!.; 1. officinale L.; 1. x steph-
irregular files; occasionally groups of cells splitting to anense Lemoine (j. beesianum x 1. officinale); Jasminum sp.;
form triangular blocks. Cortex with groups of stone cells Jasmine.
and rarely occasional fibres; sderenchyma often in con-
centric discontinuous, or rarely continuous, rings. Phloem
rays dilating slightly, often widely at outer ends, con-
ducting tissue with rings of sderenchyma as in cortex.
Crystals absent.

Xylem
TS. Diffuse or semi-ring porous. Growth rings usually
distinct. Vessels mainly solitary, and also in dusters of up
to 6. Fibres with wide lumina. Axial parenchyma fairly
abundant, mainly para trachea!. Rays 1-3 cells wide; cells
often square.

LS. Vessels: intervascular and vessel-ray pitting alternate,

fine; perforation plates simple; spiral thickenings often
present. Fibre tracheids with spiral thickenings often
present. Rays 1-3 cells wide, 2-40 cells high; multiseriate
rays often with uniseriate tails; markedly heterocellular;
cells in RLS mainly square and upright and sometimes
procumbent.

215. Jasminum humi/e, bark and xylem TS x 80 .

216. Jasminum humile, xylem TS x 55.

96
217. Jasminum sp., xylem TS x 60.
218. Jasminum humile, xylem TLS x 150, showing fine
vessel pitting and narrow rays.
219. Jasminum officina/e, xylem RLS x 150, showing a
heterocellular ray and fine vessel-ray pitting.
220. Jasminum humile, xylem RLS x 380, showing inter-
vascular and vessel-ray pitting .

97
JUGLANS Juglandaceae

Bark
Peridenn consisting of radial files of thin-walled rectangu-
lar cells, very few with dark contents; outer layers peeling
away in wide strips. Cortex narrow, sometimes with small
scattered groups of gelatinous fibres. Phloem rays dilating
towards cortex; conducting tissue between rays triangu-
lar, containing alternating blocks of gelatinous fibres;
groups of up to 10 large, thin-walled stone cells scattered
throughout conducting tissue. Crystals: cluster crystals
abundant in phloem and cortex.

Xylem
TS. Diffuse porous. Growth rings usually distinct. Vessels
mainly very wide, solitary and in pairs and radial chains
of up to 6 and small clusters of up to 5. Tyloses sometimes •
present. Fibres thin-walled with wide lumina. Axial paren- ~ut:e:;:~-'
chyma scattered and in incomplete tangential uniseriate
bands, some cells directly adjacent to vessels. Rays 1-5
cells wide.

LS. Vessels: intervascular pitting and vessel-ray pitting

coarse, alternate, pits polygonal or oval; perforation
plates simple. Rays 1-5 cells wide; uniseriate rays 1-22 ;
cells high, multiseriate rays 8-51 cells high; heterocellu-
lar; cells in RLS square, upright and procumbent.

SPECIES EXAMINED

J. nigra L.; J. regia L.; Walnut.

221. Jug/ans regia, bark TS x 60, showing periderm,
blocks of phloem fibres and scattered cluster crystals.

98
 223

222. Jug/ans regia, xylem TS x 50, showing very wide

vessels, some with tyloses.
223. Jug/ans regia, xylem TLS x 150.

224. Jug/ans regia, xylem TLS x 380, showing coarse ,

polygonal vessel pitting.

99
KERRIA Rosaceae

Bark
Peridenn: outer layers irregular, often with dark contents.
Cortex narrow, parenchymatous, with small groups of
fibres near border with phloem. Phloem rays dilating
slightly towards cortex; conducting tissue with groups of
1-20 phloem fibres at outer edge. Crystals: cluster crystals
present in cortex and phloem.

Xylem
TS. Diffuse porous. Growth rings distinct. Vessels solitary
and in clusters of up to 4. Fibres moderately thick-walled.
Axial parenchyma mainly paratracheal, also occasionally
apotracheal. Rays uniseriate and 2-6 cells wide, or some-
times up to 10 cells wide. Crystals: solitary crystals rarely
present in ray cells.

LS. Vessels: intervascular and vessel-ray pitting alternate

and elongated, sometimes scalariform; perforation plates
simple. Rays uniseriate and 2-6 cells wide; uniseriate rays
1-10 cells high, multiseriate rays usually several hundred
cells high; heterocellular; cells in RLS mainly upright in
uniseriate rays, square and upright in multiseriate rays.

SPECIES EXAMINED

K. japonica DC. 225. Kerria japonica, bark and xylem TS x 80.

100
226. Kerriajaponica, xylem TS x 80.
m. Kerria japonica, xylem TLS x 150, showing very tall,
multiseriate rays and shorter, uniseriate rays.

228. Kerria japonica, xylem RLS x 380, showing inter-

vascular and vessel-ray pitting.

101
KOLKWITZIA Caprifoliaceae

Bark
Peridenn consisting of radial files of thin-walled square
and rectangular cells, outer layers disintegrating; two or
more periderms may develop, one arising directly
adjacent to phloem. Cortex narrow. Phloem rays dilating
very slightly to outer ends; stone cells and fibres absent.
Crystals absent.

Xylem
TS. Diffuse porous. Growth rings usually indistinct.
Vessels angular, solitary, in pairs and occasional groups of
three. Fibres with narrow lumina. Axial parenchyma para-
tracheal and diffuse. Rays 1-5 cells wide.

LS. Vessels: intervascular and vessel-ray pitting alternate,

opposite and scalariform in places; perforation plates
scalariform, with 10-25 bars. Fibres with faint spiral
thickenings. Rays 1-5 cells wide, 1-100 or more cells high,
markedly heterocellular; in TLS ray cells of varying
shape, often much taller than broad, especially in uni-
seriate tails and portions and along ray edges; in RLS cells
mainly square and upright.

NOTE

See Appendix, Table 4. 229. Kolkwitzia amabilis, bark and xylem T8 x 60,
showing two periderms in bark .
SPECIES EXAMINED

K. amabilis Graebn., Beauty Bush.

102
230. Kolkwitzia amabilis, xylem TS x 150.
231. Kolkwitzia amabilis, xylem TLS x 60, showing
markedly heterocellular rays .

232. Kolkwitzia amabilis, xylem LS x 380, showing a

vessel with a scalariform perforation plate , and fibre-
tracheids with spiral thickenings.

103
LABURNUM Leguminosae

Bark present, less frequent than narrow vessels; narrow

vessels and tracheids often storied. Axial parenchyma
Periderm consisting of layers of rectangular cells, some- storied, each fusiform cell usually with one transverse
times with dark contents, and sometimes with periclinal
dividing wall. Rays 1-12 cells wide; uniseriate rays 1-8
walls thicker than anticlinal walls; outer layers peeling in
cells high, multiseriate rays 5-55 or more cells high, in
wide strips. Cortex narrow, with scattered groups of
some samples exceeding 300 cells high; heterocellular;
gelatinous fibres and sometimes groups of up to 10 thin-
cells in RLS mainly square and procumbent.
walled stone cells. Phloem rays dilating slightly towards
cortex; conducting tissue interspersed with groups of
fibres, with occasional stone cells. Crystals: solitary
SPECIES EXAMINED
crystals sometimes present in cortex and phloem, mainly
associated with fibres. L. alpinum Bercht. and Pres!; L. anagyroides Med.; L. x
watereri Dipp. (= L. alpinum x L. anagyroides); Laburnum.
Xylem
TS. Diffuse porous. Growth rings indistinct. Vessels soli-
233. Laburnum a/pinum, bark and xylem T8 x 60.
tary, in pairs and clusters of up to 10 cells; both wide and
narrow. Fibres in tangential bands, mainly gelatinous 234. Laburnuma/pinum, xylem T8 x 60.
with narrow lumina. Axial parenchyma paratracheal and
235. Laburnum x watereri, xylem T8 x 150.
often abundant; cells often approximately same width as
narrow vessels, but with thinner walls. Rays 1-12 cells 236. Laburnum x watereri, xylem TLS x 150, showing
wide. storied, narrow vessel elements and axial parenchyma.

LS. Vessels: intervascular and vessel-ray pitting alternate;

m. Laburnum x watereri, xylem TLS x 600, showing
spiral thickenings in storied, narrow vessels and tracheids.
pits oval or polygonal; vesturing visible on some pits;
perforation plates simple; spiral thickenings present on 238. Laburnum x watereri, xylem RLS x 380, showing
many narrow vessels and tracheids; tracheids often intervascular and vessel-ray pitting .

104
105
LAURUS Lauraceae

Bark
Periderm: outer layers in large broken blocks, inner layers
consisting of rectangular thin-walled cells; cells often
with dark contents. Cortex with a more or less continuous
ring of small stone cells, usually with wide lumina; fibres
infrequent or absent. Phloem rays dilating to outer ends,
ending at ring of stone cells; conducting tissue in more or
less continuous blocks with solitary, scattered fibres .
Crystals absent.

Xylem
IS. Diffuse porous. Growth rings indistinct. Vessels soli-
tary and in pairs and radial multiples of up to about 8;
often angular. Fibres usually with wide lumina. Axial
parenchyma paratracheal and diffuse. Rays 1-5 or more
cells wide.

LS. Vessels: intervascular pitting alternate, pits circular or

oval; vessel-ray pits circular or elongated, orientated in
several directions; vessel elements long, with oblique,
simple perforation plates. Rays 1-5 or more cells wide;
uniseriate rays 1-11 cells high, multiseriate rays 4-55 cells
high; heterocellular; cells in TLS irregular in shape,
usually with tall cells in uniseriate portions; large pro-
portion of square cells in RLS.

SPECIES EXAMINED
240. Laurus nobi/is, xylem TS x 80.
L. nobilis L., Bay Laurel.

239. Laurus nobilis, bark and xylem TS x 60.

106
 241
241. Laurus nobi/is, xylem TLS x 60.
242. Laurus nobi/is, xylem TLS x 380, showing alternate
vessel pitting and fibres and ray cells containing starch
granules .

243. Laurus nobi/is, xylem RLS x 150, showing hetero-

cellular rays and vessel-ray pitting .
244. Laurus nob/lis, xylem RLS x 380, showing vessel-ray
pitting and starch granules in ray cells, fibres and axial
parenchyma .

107
LAVANDULA Labiatae

Bark
Periderm: cells often with dark contents, sometimes thick-
walled. Cortex parenchymatous; cells often with dark
contents; interspersed with large groups of fibres. Phloem
rays not dilating widely towards cortex; conducting
tissue in outer region with large groups of fibres. Crystals:
abundant solitary crystals associated with sclerenchyma
in cortex and phloem; cluster crystals also present,
mainly in phloem.

Xylem
IS . Diffuse porous. Growth rings often indistinct. Vessels
mainly solitary or occasionally in clusters of up to 3. Fibres
fairly thick-walled. Axial parenchyma abundant, mainly
apotracheal and often in short tangential rows. Rays 1-3
cells wide.

LS. Vessels: intervascular and vessel-ray pitting alternate,

fine . Perforation plates simple; faint spiral thickenings
present. Fibres often with bordered pits. Rays 1-2 or
sometimes up to 3 cells wide; 1-20 or more cells high,
occasionally with uniseriate tails and portions; slightly
heterocellular; cells in RLS mainly square and procum- 246. Lavandula sp ., xylem TS x 60.
bent.

SPECIES EXAMINED

Lavandula sp., Lavender.

246. Lavandula sp ., bark TS x 60 .

108
247. Lavandula sp., xylem TLS x 150.
248. Lavandula sp., xylem LS x 600 , showing fibre-
tracheids with spiral thickenings.

249. Lavandula sp., xylem LS x 380, showing vessels with

alternate pitting and faint spiral thickenings .

250. Lavandula sp ., xylem RLS x 150, showing a vessel

element with a simple perforation plate and a hetero-
cellular ray .

109
LIGUSTRUM Oleaceae

Bark
Peridenn: cells often lignified; sometimes groups of cells
splitting to form triangular blocks. Cortex sometimes with
small groups of fibres interspersed throughout, though
these may be sparse or absent; stone cells rarely present.
Phloem rays dilating slightly towards outer ends; con-
ducting tissue sometimes with blocks of fibres and rarely
stone cells, at or near junction with cortex. Crystals
usually absent or rarely small solitary crystals present.

Xylem
T5. Diffuse or seml-nng porous. Growth rings fairly
distinct. Vessels mainly solitary, and in clusters of up to 6.
Fibres with wide lumina. Axial parenchyma absent or very
sparse, usually paratracheal. Rays 1-3 cells wide.

L5. Vessels: intervascular and vessel-ray pitting alternate,

rarely slightly elongated; perforation plates simple; fibre- 2~5;':'1~~~11J~~#~~~tQ~~R-~~J
tracheids also sometimes present; faint spiral thickenings
occasionally present on vessels and fibre-tracheids. Fibres
occasionally septate. Rays 1-3 cells wide, often with both 251. Ligustrum vulgare, bark and xylem TS x 60.
multiseriate and uniseriate portions or uniseriate tails;
2-30 cells high; markedly heterocellular in TLS; cells in
RLS procumbent and square and occasionally upright.
Crystals: small solitary crystals sometimes present in ray
cells.

SPECIES EXAMINED

L. vulgare L., Privet.

254. Ligustrum vulgare, xylem TLS x 380, showing vessel

wall pitting .
256. Ligustrum vulgare, xylem RLS x 380, showing soli-
tary crystals in some ray cells.

110
252. Ligustrum vulgare, xylem TS x 80.
253. Ligustrum vulgare, xylem TLS x 150.

111
LIQUIDAMBAR Altingiaceae

Bark
Peridenn: outer layers irregular, sometimes peeling in
narrow strips; consisting of thin-walled cells, occasion-
ally with dark contents. Cortex narrow; cells often with
slightly thickened lignified walls; occasionally small
groups of 2-10 thick-walled fibres present; large sec-
retory canals occasionally present. Phloem narrow; rays
not dilating markedly towards cortex; small groups of
1-10 thick-walled fibres present, occasionally with associ-
ated stone cells. Crystals: abundant cluster crystals pres-
ent in cortex and phloem.

Xylem
TS. Diffuse porous. Growth rings sometimes indistinct.
Vessels narrow, thin-walled, angular, solitary and
occasionally in radial chains and clusters of 2-6. Fibres
thick-walled. Axial parenchyma paratracheal and apo-
tracheal, abundant. Rays 1-3 cells wide.

LS. Vessels: intervascular and vessel-ray pitting alternate

or scalariform; perforation plates scalariform, with 10-20
bars. Fibres and fibre-tracheids present. Rays uniseriate and
1-3 cells wide; 2-40 cells high; multiseriate rays usually
with uniseriate tails and markedly heterocellular; in RLS
cells in multiseriate portions mainly procumbent and cells
in uniseriate portions mainly square and upright.

SPECIES EXAMINED
256. Liquidambar styraciflua, bark TS x 150.
L. styraciflua L., Sweet Gum.

112
2E1. Liquidambar styraciflua, bark and xylem TS x 60,
showing a large secretory canal in the bark.
258. Liquidambarstyraciflua, xylem TS x 150.

259. Liquidambar styraciflua, xylem TLS x 150.

260. Liquidambar styraciflua, xylem RLS x 380, showing
a vessel with a scalariform perforation plate.

113
LIRIODENDRON Magnoliaceae

Bark obliquely; perforation plates scalariform, usually oval,

Peridenn often with ragged appearance; several layers with 3-15 thin, widely spaced bars. Rays 1-5 cells wide,
2-18, or occasionally up to 80 cells high; heterocellular;
wide. Cortex wide, consisting of thin-walled parenchyma-
cells in RLS mainly procumbent and square, rarely up-
tous cells, with occasional very thin-walled stone cells;
right.
fibres absent. Phloem rays dilating towards outer ends;
small groups of 1-15 fibres with wide lumina inter-
spersed in conducting tissue. Crystals absent.
SPECIES EXAMINED

L. tulipifera L., Tulip Tree.

Xylem
IS. Diffuse or semi-ring porous. Growth rings usually
fairly distinct. Vessels solitary, in clusters of 2-5 and in
radial chains of 2-15; often crowded and relatively thin-
walled. Fibres often with fairly wide lumina. Axial paren-
chyma scattered, sometimes abundant. Rays 1-5 cells
wide; often noded at growth rings.

LS. Vessels: intervascular pitting opposite or scalariform; 261. Liriodendron tu/ipifera, bark T8 x 60.
vessel-ray pitting scalariform, sometimes orientated 262. Liriodendron tu/ipifera, xylem T8 x 45.

114
 284. Liriodendron tulipifera, xylem TLS x 150, showing
opposite and scalariform vessel pitting and scalariform
perforation plates .

266. Liriodendron tulipifera, xylem RLS x 150, showing a

heterocellular ray and scalariform vessel-ray pitting.
266. Liriodendron tulipifera, xylem RLS x 380, showing
opposite and elongated vessel pitting.

115
LON/CERA Caprifoliaceae

Bark
Periderm consisting of uniseriate layers of thin-walled,
square or rectangular cells, sometimes alternating with
multiseriate bands of thicker-walled cells or fibres. Cortex
narrow. Phloem rays not dilating; conducting tissue
usually containing blocks of 1-20 thick-walled angular
fibres . Crystals: cluster crystals usually abundant.

Xylem
TS. Diffuse porous or semi-ring porous. Growth rings
usually distinct. Vessels frequently angular, solitary, in
pairs and clusters of up to 5. Fibres with narrow lumina.
Axial parenchyma paratracheal and diffuse. Rays 1-6 cells 267. Lonicera orienta/is, bark and xylem TS x 80, showing
wide. semi-ring porous xylem.

LS. Vessels: intervascular and vessel-ray pitting fine, alter-

nate; perforation plates simple, occasionally scalariform
with 1-30 bars; faint spiral thickenings sometimes
present. Rays 1-6 cells wide, 7-200 or more cells high;
multiseriate rays usually with uniseriate tails; markedly
heterocellular, tails composed of predominantly upright
cells; in RLS, cells mainly square and upright, few cells
procumbent.

NOTE

See Appendix, Table 4.

SPECIES EXAMINED

L. x americana K. Koch (L. caprifolium x L. etrusca);

L. caprifolium L.; L. discolor Lind!.; L. etrusca Santi; L. fra-
grantissima Lind!. and Paxt.; L. nitida Wils.; L. orientalis
Bieb.; L. prolifera Rehd.; Honeysuckle.

116
268. Lonicera caprifo/ium, xylem TS x 60.
269. Lonicera disc%r, xylem TS x 60.

270. Lonicera disc%r, xylem TLS x 60.

271. Lonicera nitida, xylem RLS x 380, showing a hetero-
cellular ray.

117
MAGNOLIA Magnoliaceae

Bark ~
1. U
~
1
.... ~
)l ~ .4 [(1)g ..... UlUt.. "-.4
~"'" p
Periderm narrow, irregular, with thin-walled cells. Cortex ~
,A'):~~II:f~~~~~-111(~';#-~-+...cAII!~~ ~ .,I;

parenchymatous, rarely with thin-walled stone cells or rD' ~ P'<....:

groups of up to 10 fibres. Phloem rays dilating towards E: 'r:I J ~...., ~U~'.:lIAI"'"
cortex; conducting tissue interspersed with few groups of "" l ~ ~ ~ rot Wi IIJ
2-15 fibres. Crystals absent. .All
~ F""~ ~
", ~ ~'W'oC1~r-..~~1-.- ~
Xylem
TS. Diffuse porous. Growth rings fairly indistinct. Vessels
~ ~ ;~ }\"~~~~U~d~'Nl.
solitary and in radial chains and clusters of 2-8. Fibres
~~I ~ ~ ~ ~~
.It"
with wide lumina. Axial parenchyma sparse. Rays 1-3 cells
wide. I¥l .lIZ t.l ~ B-"
~ ~ kf '~~~~~~~~~~r~~~~~~~

..
LS. Vessels: intervascular and vessel-ray pitting scalari-
form; perforation plates oblique, scalariform, with 5-20 .. ~ ~a.rt.W4f&.r.J~ ~
~ m~,ftt:fJ[],K K:.l ~
bars. Rays 1-3 cells wide, 1-82 cells high; in some species
mainly uniseriate with occasional biseriate portions; in
~ ~ r~~~~~~m~~~~
M. x soulangiana mainly 2-3 cells wide; slightly hetero- )t; ~.-,~Mff IBI! ~
~ ~ ~~ ~~ ~
cellular; cells in RLS mainly procumbent and square.

273 .JI[ AnI "t:

_.
'ft.~
.-..ov.
rHl #IIJ
SPECIES EXAMINED ~ 1N~

M. campbellii Hook. f. and Thoms.; M. denudata Des- m. Magnolia sieboldii, xylem TS x 60.
rouss.; M. sieboldii K. Koch; M. x soulangiana Soulange-
Bodin (M. denudata x M. liliiflora) .

Z12. Magnolia sieboldii, bark TS x 55.

118
274. Magnolia campbellii, xylem TLS x 150, showing
scalariform vessel pitting and short rays .
275. Magnoliadenudata, xylem TLS x 150, showing
scalariform vessel pitting and tall rays .
276. Magnolia campbellii, xylem RLS x 150, showing a
heterocellular ray with ray-axial parenchyma pitting and
scalariform vessel-ray pitting .
2n. Magnolia sieboldii, xylem RLS x 380, showing a
vessel with a scalariform perforation plate .

119
MAHONIA Berberidaceae

Bark
Periderm: outer layers very irregular, individual cells vary-
ing in shape, usually approximately square, rarely in
rectangular files. Cortex mainly parenchymatous with a
few scattered fibres. Phloem rays dilating slightly towards
cortex; conducting tissue with fibres in scattered groups
of 1-20 or more. Crystals: solitary crystals very rare or
absent in cortex and phloem.

Xylem
TS. Diffuse porous. Growth rings indistinct. Vessels soli-
tary, in pairs and small groups of 3 or 4; arrangement
sometimes weakly dendritic; vessels narrow, angular in
outline; diameter of narrowest vessels similar to that of
fibres. Fibres wide with wide lumina. Axial parenchyma
mainly paratracheal, sparse. Rays 1-5 cells wide, cells
square or rectangular, sometimes wider than narrowest
vessels.

LS. Vessels: intervascular and vessel-ray pitting alternate;

pits fine, oval in outline; spiral thickenings present on
most vessels; perforation plates simple. Fibres septate.
Rays 1-5 cells wide; uniseriate rays 1-10 cells high, multi-
seriate rays 5-80 cells high; heterocellular; cells in RLS
mainly square or procumbent. 278. Mahonia aquifo/ium, bark TS x 150, showing
irregular periderm.

SPECIES EXAMINED

M. aquifolium Nutt., Oregon Grape; M. japonica DC.

281. Mahoniaaquifo/ium, xylem TLS x 150.

282. Mahonia aquifo/ium, xylem RLS x 380, showing
alternate vessel pitting and simple perforation plates.

120
279. Mahonia aquifolium. bark and xylem TS x 60.
280. Mahonia aquifolium. xylem T S x150.

121
MORUS Moraceae

Bark right and often with dark contents. Crystals: solitary,

Peridenn narrow, outer layers sometimes peeling in short rarely present in axial parenchyma cells.
strips. Cortex fairly narrow, cells mainly parenchymatous
and often with dark contents, with occasional small
groups of up to 10 thick-walled fibres . Phloem very wide, SPECIES EXAMINED
rays dilating widely to cortex, occasionally with groups of M. alba L.; M. nigra L.; Morus sp.; Mulberry.
fibres as in cortex; conducting tissue interspersed with
small groups of gelatinous, thick-walled fibres. Crystals:
solitary and cluster crystals present in cortex and phloem,
sometimes extremely abundant.

Xylem
TS. Diffuse porous. Growth rings often indistinct. Vessels
both narrow and wide, crowded, solitary and in clusters
and radial chains of up to 20. Fibres often gelatinous,
thick-walled and in conspicuous blocks. Axial parenchyma
abundant, sometimes forming much of ground tissue;
cells sometimes with dark contents . Rays 1-6 cells wide.

LS. Vessels: intervascular pitting mainly alternate, poly-

gonal and sometimes elongated; vessel-ray pitting often
elongated, sometimes alternate; perforation plates
simple. Rays 1-6 cells wide, up to 100 or more cells high;
heterocellular; cells in RLS procumbent, square and up-

283. Morus alba, bark TS x 60, showing periderm with

disrupted outer layers and ray and cortex cells with dark
contents.
284. Morus nigra, bark TS x 150, showing abundant
solitary crystals .

122
286. Morus nigra, xylem T8 x 60.
286. Morus nigra, xylem T8 x 150.

2If1. Morus nigra, xylem TL8 x 150.

288. Morus sp., xylem TL8 x 150, showing alternate
vessel pitting .

...
287

123
NO THO FA GUS Fagaceae

Bark
Periderm: cells in regular, radial files, radially elongated,
with dark contents. Cortex with more or less continuous
cylinder of fibres and stone cells. Phloem rays not dilating
widely; conducting tissues interspersed with blocks of
thick-walled fibres sometimes continuous with cortical
sclerenchyma cylinder. Crystals: solitary, mainly associ-
ated with sclerenchyma.

Xylem
TS. Diffuse porous. Growth rings indistinct. Vessels soli-
tary and in groups of up to 5 or rarely more. Fibres moder-
ately thick-walled. Axial parenchyma sparse, mainly apo-
tracheal. Rays 1-2 or occasionally up to 3 cells wide.

LS. Vessels: intervascular pitting opposite or sometimes 289. Nothofagus obliqua, bark and xylem T8 x 60.
elongated, coarse; vessel-ray pitting mainly scalariform,
often apparently vestured (pseudovestured); perforation 290. Nothofagus obliqua, xylem TL8 x 150, showing
plates both simple and scalariform, with 1-10 or more narrow vessels and rays.
bars. Fibres septate. Rays 1-2 or sometimes up to 3 cells
wide, 1-30 or more cells high, often with uniseriate tails;
markedly heterocellular; cells in RLS procumbent, square
and upright. Crystals: chains of crystals sometimes pres-
ent in axial parenchyma cells.

SPECIES EXAMINED

N. obliqua Bl., Southern Beech.

124
291. Nothofagus obliqua, xylem RLS x 150, showing
opposite and scalariform vessel-ray pitting and ray celis
containing starch granules .

125
OLEARIA Compositae

Bark
Peridenn irregular; cells thin-walled and occasionally
slightly thick-walled. Cortex parenchymatous, sometimes
with large, more or less rectangular, blocks of fibres as in
phloem. Phloem with discontinuous concentric rings of
sclerenchyma consisting of more or less rectangular
blocks of 10-100 or more fibres; rays dilating towards
outer ends. Crystals absent.

Xylem
TS. Diffuse porous. Growth rings sometimes indistinct.
Vessels in clusters and tangential and radial chains of up to
10. Fibres often with wide lumina. Axial parenchyma abun-
dant, mainly paratracheal. Rays 1-3 or occasionally up to
4 cells wide.

LS. Vessels often storied; intervascular and vessel-ray 292. O/earia odorata, bark TS x 60.
pitting alternate, and sometimes slightly elongated. Per-
foration plates Simple. Narrow vessels and tracheids
sometimes with fine spiral thickenings in O. odorata.
Fibres short, storied. Axial parenchyma storied. Rays some-
times storied, especially in O. x haastii; 1-3 or rarely 4
cells wide; 1-30 or more cells high; heterocellular; cells in
RLS procumbent, square and upright.

SPECIES EXAMINED

O. x haastii Hook. f. (0. avicenniifolia x O. moschata);

O. odorata Petrie; Daisy Bush.

126
293. Olearia odorata, xylem TS x 150.
294. Olearia x haastii, xylem TLS x 150, showing storied
vessels and axial parenchyma cells .

296. Olearia x haastii, xylem RLS x 150, showing narrow

and wide storied vessel elements and heterocellular rays.

127
OSMANTHUS and PHILL YREA Oleaceae

Indistinguishable from each other in their root anatomy.

Bark LS. Vessels: intervascular pitting and vessel-ray pitting

Periderm often ragged in appearance or splitting into large alternate, fine; perforation plates simple; spiral thicken-
wedge-shaped blocks; cells thin-walled or sometimes ings often present. Rays 1-2 or rarely 3 cells wide; multi-
with thickened inner periclinal walls. Cortex parenchyma- seriate rays with uniseriate tails and portions, and multi-
tous, with large groups of stone cells near border with seriate and uniseriate portions of similar width in TLS;
phloem, sometimes forming a complete cylinder. Phloem both multiseriate and uniseriate rays heterocellular; cells
rays dilating slightly towards outer ends, fibres absent; in RLS procumbent, square and upright.
stone cells present at border with cortex. Crystals absent.

SPECIES EXAMINED
Xylem
O. x burkwoodii (Burkwood and Skipwith) P. S. Green
TS . Diffuse or semi-ring porous. Growth rings usually (0. decorus x O. delavayi); O. heterophyllus P. S. Green;
fairly distinct. Vessels narrow, solitary, in radial chains or P. latifolia L.
clusters, and often in loose, oblique clusters of 2-several,
in dendritic arrangement. Fibres with narrow or wide
lumina. Axial parenchyma paratracheal. Rays 1-3 cells
wide; cells often as wide as vessels.

128
298. Osmanthus heterophyllus, xylem TLS x 150, show-
ing heterocellular, uniseriate rays with narrow, biseriate
portions.
299. Osmanthus heterophyllus, xylem TLS x 380, show-
ing narrow tracheids and rays.

300. Osmanthus x burkwoodii, xylem RLS x 150, show-

ing fine, alternate vessel and vessel-ray pitting and hetero-
cellular rays .

296. Osmanthus x burkwoodii, bark and xylem TS x 60,

showing ragged outer periderm, a cylinder of small stone
cells on phloem-cortex boundary and dendritic vessel
arrangement .
'1!f1. Osmanthus x burkwoodii, xylem TS x 150.

129
PARROTIA Hamamelidaceae

Bark
Periderm: cells in regular files, flattened, thin-walled,
inner cells often with dark contents. Cortex: parenchyma
cells with dark contents; more or less continuous cylinder
of stone cells and occasional fibres present. Phloem rays
dilating widely in outer half; cells with dark contents;
conducting tissue interspersed with groups of thick-
walled fibres; occasional cells with dark contents. Crystals
solitary, mainly associated with sclerenchyma in cortex.

Xylem
T5. Diffuse porous. Growth rings sometimes indistinct.
Vessels narrow, often angular, mainly solitary and rarely
in clusters and radial chains of 2-6. Axial parenchyma para-
tracheal and in short uniseriate, tangential rows . Fibres
thick-walled. Rays 1-3 cells wide.

L5. Vessels narrow, with oblique end walls; intervascular 301. Parrotia persica , bark and xylem TS x 60.
and vessel-ray pitting oval, often elongated and scalari-
form, with narrow borders; perforation plates scalari-
form, with 8-20 fairly widely spaced and often bifurcating
bars. Rays 1-3 cells wide, sometimes uniseriate or with
uniseriate tails and portions; 3-40 or more cells high;
markedly heterocellular; cells in RLS procumbent, square
and upright.

SPECIES EXAMINED

P. persica C. A. Mey.

130
302. Parrotia persica, xylem TS x 150.
303. Parrotia persica , xylem TLS x 150, showing fibre-
tracheids, and heterocellular rays with abundant starch
granules .

304. Parrotia persica , xylem RLS x 150, showing vessels

with scalariform pitting and perforation plates and hetero-
cellular rays with abundant starch granules and scalariform
vessel-ray pitting.

131
PASS/FLORA Passifloraceae

Bark
Peridenn very irregular; outer layers often forming
wedge-shaped blocks. Cortex parenchymatous, with
occasional groups of thin-walled lignified cells and thick-
walled stone cells, and also groups of 1-30 fibres, some-
times gelatinous. Phloem rays dilating towards cortex;
conducting tissue in wedge-shaped or triangular areas in
IS, interspersed with groups of fibres, particularly in
outer region. Crystals: cluster crystals abundant in cortex
and phloem.

Xylem
TS . Diffuse porous. Growth rings indistinct. Vessels both
narrow and very wide, mainly solitary and occasionally in
groups of up to 5. Fibres with walls of variable thickness.
Axial parenchyma diffuse. Rays 1-8 cells wide; markedly
heterocellular. 306. Passiflora sp., bark TS x 150, showing phloem with
scattered fibre blocks and cluster crystals.
LS. Vessels: intervascular and vessel-ray pitting alternate,
polygonal or sometimes slightly elongated; perforation
plates simple, often horizontal. Rays 1-5 or rarely up to 8
cells wide; 1-50 or more cells high; markedly heterocellu-
lar; cells mainly tall in ILS, mainly square and upright or
occasionally procumbent in RLS .

SPECIES EXAMINED

Passiflora sp., Passion flower.

132
 306. Passif/ora sp., xylem TS x 150.

307. Passif/ora sp., xylem TLS x 150.

308. Passiflora sp., xylem LS x 380, showing alternate

vessel pitting .

133
PHILADELPHUS Philadelphaceae

Bark
Periderm very thick, with alternating rings of thin-walled
cells and thick-walled cells. Cortex absent or rarely pres-
ent. Phloem usually extending to periderm; cells thin-
walled; fibres and stone cells absent; rays not dilating
towards outer ends. Crystals absent.

Xylem
TS. Diffuse or semi-ring porous. Growth rings fairly dis-
tinct. Vessels solitary, in tangential and radial pairs. Fibres
moderately thick-walled. Axial parenchyma scattered,
sparse. Rays 1-4 cells wide.

LS. Vessels: intervascular and vessel-ray pitting fine, alter-

nate or opposite or occasionally slightly elongated; per-
foration plates scalariform with 20-35 or more very fine,
occasionally forked, bars. Rays 1-4 cells wide, 5-50 cells
high; markedly heterocellular; cells in RLS square and
procumbent and occasionally upright.

SPECIES EXAMINED
310. Phi/ade/phus coronarius, xylem TS x 150.
P. coronarius L., Mock Orange.

309. Phi/ade/phus coronarius, bark TS x 150, showing

alternate layering of thick- and thin -walled cells of
periderm.

134
311. Philadelphus coronarius , xylem TLS x 150.
312. Philadelphus coronarius, xylem RLS x 150, showing
fine alternate vessel and vessel -ray pitting , ascalariform
perforation plate and heterocellular rays .

313. Philadelphus coronarius, xylem RLS x 380, showing

a vessel with a scalariform perforation plate .

135
PIERIS Ericaceae

Bark
Periderm: cells often with dark contents; sometimes thick-
walled. Cortex narrow in P. formosa; discontinuous
sclerenchyma cylinder present at junction with phloem;
in P. japonica sclerenchyma consisting mainly of large
groups of thick-walled stone cells; in P. formosa scleren-
chyma consisting mainly of fibres. Phloem narrow; cells
mainly thin-walled; rays dilating very gradually at outer
ends. Crystals: solitary and double crystals present in
cortex and sometimes in phloem; often associated with
sclerenchyma.

Xylem
TS. Diffuse porous. Growth rings often indistinct. Vessels
narrow, crowded, often angular in P. formosa; solitary
and occasionally in pairs and clusters of up to 4. Fibres
often with wide lumina. Axial parenchyma fairly abun- 314. Pieris formosa var . forrestii, bark and xylem
dant; mainly paratracheal and occasionally apotracheal. TS x 150.
Rays 1-3 cells wide.

LS. Vessels: intervascular and vessel-ray pitting opposite

and sometimes elongated. Perforation plates oblique,
usually scalariform with 20-40 or more thin bars, or rarely
simple. Rays 1-3 cells wide, 1-100 or more cells high;
often with uniseriate tails and portions; markedly hetero-
cellular; cells in RLS procumbent, square and upright.

SPECIES EXAMINED

P. formosa D. Don, var. forrestii Airy Shaw; P. japonica

D. Don.

136
315. Pierisjaponica, bark and xylem TS x 60, showing
abundant stone cells in cortex and phloem .
316. Pieris formosa var . forrestii, xylem TS x 150, show-
ing narrow, angular vessels .

317. Pieris formosa var. forrestii, xylem TLS x 150, show-

ing short, heterocellular rays .
318. Pierisjaponica, xylem LS x 150, showing vessels
with a scalariform perforation plate.

137
PITTOSPORUM Pittosporaceae

Bark
Periderm often narrow, consisting of regular layers of
rectangular cells in radial files, with outer layers peeling
in thin, uniseriate strips; cells often with slightly thick-
ened walls. Cortex parenchymatous; secretory canals
present, lined with thin-walled epithelial cells. Phloem
rays narrow for most of length, dilating towards cortex;
sclerenchyma absent. Crystals: large, solitary, in cortex.

Xylem
IS . Diffuse porous. Growth rings absent. Vessels solitary
and in pairs and radial chains of up to at least 8 cells, and
occasional clusters of up to 5 cells. Fibres with wide
lumina. Axial parenchyma sparse or absent. Rays 1-4 cells
wide.
320. Pittosporum tenuifolium, bark TS x 150, showing
LS. Vessels: intervascular pitting and vessel-ray pitting parenchymatous cortex with wide secretory canals .
fine, alternate; pits oval; vessel elements long, often with
faint spiral thickenings; perforation plates simple. Fibres
septate. Rays 1-2 or sometimes up to 4 cells wide; uni-
seriate rays 1-20 cells high, multiseriate rays 6 to 150 or
more cells high; heterocellular; cells in RLS procumbent,
square and upright.

SPECIES EXAMINED

P. tenuifolium Gaertn., P. tobira Ait.

319. Pittosporum tenuifolium, bark TS x 150.

138
 321. Pittosporum tenuifo/ium, xylem TS x 150.
322. Pittosporum tenuifo/ium, xylem TLS x 150, showing
narrow vessels, septate fibres, and mainly narrow rays .

323. Pittosporum robira, xylem LS x 380, showing vessels

with fine alternate pitting and simple perforation plates.

139
PLATANUS Platanaceae

Bark very few axial elements; heterocellular; cells in RLS pro-

cumbent, square and upright.
Periderm: outer layers usually peeling, some cells with
dark contents. Cortex narrow, with occasional scattered
groups of thin-walled fibres with large lumina. Phloem SPECIES EXAMINED
rays dilating widely towards cortex; phloem fibres in
large, triangular blocks on outer side of conducting P. x acerifolia (Ait.) Willd. (P. orientalis x P . occidenta/is);
tissue; fibres with wide lumina. Crystals solitary, frequent P. occidentalis L.; Platanus sp.; Plane.
in phloem and cortex, often associated with fibres.

Xylem
T5 . Diffuse porous. Growth rings often indistinct. Vessels
wide, crowded, solitary and in pairs, infrequently in
radial multiples of 3; tyloses sometimes present. Fibres
thin-walled with wide lumina. Axial parenchyma para-
tracheal and diffuse. Rays rarely uniseriate, but mainly
4-14 cells wide; cells large and rectangular; rays forming
a considerable portion of xylem. Crystals solitary, some- 324. Platanus x acerifolia, bark TS x 80, showing widely
dilating ray and triangular blocks of phloem fibres.
times present in ray cells.
325. Platanussp., bark TS x 150, showing solitary
L5. Vessels: intervascular and vessel-ray pitting opposite, crystals in ray.
many pits horizontally elongated and scalariform; perfor-
ation plates simple and scalariform, the latter with 5-25
bars, variably spaced, occasionally with trabeculae. Rays:
uniseriate rays rare; mainly 4-14 cells wide; 15-150 cells
high, occasionally with uniseriate tails of up to 15 cells;
rays in ILS occasionally with end of ray closely adjacent
to end of ray above or below, the two rays separated by

140
326. Platanus x acerifolia, xylem TS x 80.
3X1. Platanus x acerifolia, xylem TLS x 80, showing tall,
wide, heterocellular rays.

328. Platanus sp ., xylem RLS x 150, showing alternate

and elongated vessel pitting, a scalariform perforation
plate, and a heterocellular ray.
329. Platanus sp., xylem TLS x 380, showing scalariform
pitting in tracheary elements .

141
POL YGONUM Polygonaceae

Bark
Periderm consisting of square or rectangular thin-walled
cells, occasionally with dark contents, sometimes in more
than one layer, deep-seated. Cortex often absent, difficult
to distinguish from phloem. Phloem very wide; rays not
dilating towards cortex, consisting of thin-walled paren-
chymatous cells; conducting tissue consisting of thin-
walled cells alternating with large blocks of phloem
fibres. Crystals: cluster crystals of various sizes frequent
throughout phloem and cortex; long solitary crystals
(styloids) also occasionally present.

Xylem
TS. Diffuse or semi-ring porous. Growth rings indistinct.
Vessels solitary and in clusters of up to 10; ranging in
width from narrow to very wide, circular or oval. Fibres
thin- or thick-wallt':. Axial parenchyma sparse, para-
tracheal. Rays up to 20 cells wide. Crystals: cluster crystals
frequently present in ray cells.

LS. Vessels: intervascular and vessel-ray pitting alternate,

polygonal, vestured; perforation plates simple; narrow
vessels and tracheids often storied. Fibres short and some-
times storied. Rays 5-20 cells wide, up to several hundred
cells high; heterocellular; cells irregularly shaped.

SPECIES EXAMINED 330. Polygonum ba/dschuanicum, bark T8 x 60, showing

P. baldschuanicum Regel. blocks of fibres in phloem.

142
331. Polygonum baldschuanicum, xylem TS x 150.
332. Polygonum baldschuanicum, xylem TLS x 40, show-
ing tall, wide rays .

333. Polygonum baldschuanicum, xylem TLS x 150,

showing short vessel elements with alternate pitting and
tall, heterocellular rays with cluster crystals .
334. Polygonum baldschuanicum, xylem RLS x 150,
showing a heterocellular ray with abundant cluster
crystals.

143
POMOIDEAE Rosaceae
Chaenomeles, Cotoneaster, Crataegus, Cydonia, Malus, Pyracantha, Pyrus, Sorbus
Members of this group are not readily distinguishable
from one another using root anatomy, although they can
differ in a few characters.

Bark
Periderm with outer layers peeling off in short strips, cells
rectangular, sometimes containing tannin, occasionally
with thickened outer periclinal walls. Cortex with groups
of 2-50 or more thick-walled fibres, fibres sometimes very
sparse. Phloem rays dilating slightly towards outer ends;
conducting tissue interspersed with rectangular or
irregularly shaped groups of up to 50 thick-walled fibres,
often in concentric rings; occasional stone cells present.
Crystals: solitary and double crystals often present in
cortex and phloem and in longitudinal chains, mainly
associated with fibres.

Xylem
TS. Diffuse porous. Growth rings often indistinct. Vessels
solitary or rarely in pairs; circular or tending to be angu-
lar. Fibres thin- or thick-walled. Axial parenchyma varying
from sparse to relatively abundant, paratracheal and
scattered, sometimes in short, uniseriate tangential
bands. Rays usually 1-3 cells wide or more rarely up to 7
cells wide.

LS. Vessels: intervascular pitting and vessel-ray pitting

alternate, polygonal and circular; perforation plates
simple; spiral thickenings rarely present. Rays 1-3 cells
wide or rarely up to 7 cells wide; varying in height from
2-60 cells; often fairly homocellular or slightly hetero-
cellular, with cells in RLS square, procumbent or upright.

SPECIES EXAMINED

Chaenomeles japonica Spach, Japanese or Chinese Quince;

Cotoneaster horizontalis Dcne.; C. lacteus W.W.Sm.;
C. salicifolius Franch.; Crataegus x lavallei Herinq. (c. crus-
galli x stipulacea); C. monogyna Jacq.; C. oxycanthoides L.,
Hawthorn (all Crataegus); Cydonia sp., Quince; Malus
pumila Mill., Crab Apple; M. x purpurea Rehd. (M. x
atrosanguinea x niedzwetzkyana); M. toringoides Hughes;
Pyracantha coccinea Roemer; P. sp., Firethorn; Pyrus
amygdaliformis Vill.; P. communis L., Pear; Sorbus aria
Crantz., Whitebeam; S. aucuparia L., Rowan or Mountain
Ash; S. hybrida L.; S. intermedia Pers.

144
335. Cydonia sp., bark TS x 150, showing groups of
fibres in cortex and phloem.
336. Malus pumila, bark and xylem TS x 45.
:rD. pyracantha sp., bark and xylem TS x 55.

145
 338. Sorbus hvbrida, bark and xylem TS x 150, showing
groups of fibres in cortex and phloem, and scattered,
solitary crystals .
339. Chaenome/esjaponica, xylem TS x 150.
340. Cotoneaster horizonta/is, xylem TS x 150.

146
341. Crataegus x lavallei, xylem TS x 150, showing
abundant starch granules in axial parenchyma and rays.
342. Cydonia sp ., xylem TS x 150.
343. Pyrus communis, xylem TS x 150.

147
 344. Sorbusaria, xylem TS x 150.
345. Pyrus communis, xylem TLS x 150.
346. Chaenome/esjaponica, xylem TLS x 150.

148
347. Cotoneaster lacteus, xylem TLS x 150, showing
narrow rays .

348. Sorbus hvbrida, xylem RLS x 150, showing alternate

intervascular and vessel -ray pitting , and a heterocellular
ray.

149
POPULUS and SALIX Salicaceae
Indistinguishable from each other using root anatomy.

Bark para tracheal. Rays 1 or very rarely 2 cells wide; often

Periderm: cells thin-walled, occasionally with dark con- sinuous round vessels or vessel clusters.
tents; sometimes peeling in thin strips. Cortex often nar-
row, with small groups of up to about 20 fibres; stone LS. Vessels: intervascular pitting alternate, polygonal;
cells usually absent or rarely present in small groups. vessel-ray pitting oval or polygonal, slightly larger than
Phloem rays dilating slightly, sometimes widely at outer intervascular pitting, without obvious borders; perfor-
ends; conducting tissue with tangentially elongated ation plates simple. Rays uniseriate, or rarely biseriate;
groups of 2-50 thick-walled fibres dispersed as in cortex; usually homocellular in Populus (with cells in RLS square
fibre groups often stratified; stone cells rarely present; or slightly procumbent) often heterocellular in Salix (with
cells with dark contents (tannins) sometimes present in cells in RLS procumbent, square and upright) though this
both phloem and cortex. Crystals: numerous solitary distinction not always clear.
crystals associated with fibre groups in phloem and
cortex; occasional cluster crystals usually also present.
SPECIES EXAMINED

Xylem P. alba L.; P. balsamifera L.; P. x canescens Sm. (P. alba x

tremula); P. nigra L.; P. tremula L.; P. trichocarpa Hook.; P.
TS. Diffuse porous. Growth rings usually distinct. Vessels sp.; Salix alba L.; S. babylonica L.; Salix sp.; Poplar (Populus
usually abundant and crowded, in clusters or radial spp.); Willow (Salix spp.).
chains of 2-6 and occasionally solitary; thin-walled,
usually flattened on walls adjacent to other vessels, so
that vessel clusters often oval in outline. Fibres thin-
walled with wide lumina. Axial parenchyma very sparse,
349. Populus nigra, bark and xylem TS x 40.
360. Salix sp., xylem TS x 60.

150
 362. Salix sp ., xylem TLS x 150, showing solely uniseriate
rays, and short, wide vessel elements with alternate
pitting.
353. Populus t(emu/a, xylem TLS x 380, showing a vessel
with coarse, polygonal pitting.
351. Populus t(emu/a, TLS x 150, showing solely 354. Populus t(emu/a, xylem RLS x 380, showing coarse
uniseriate rays and coarse, alternate vessel pitting. vessel-ray pitting .

151
PO TEN TIL LA Rosaceae

Bark
Periderm: cells in very regular radial rows, with character-
istic rounded or oval shape in TS, 2-4 layers of thick-
walled cells alternating with 1 or more layers of thinner-
walled cells; several layers usually peeling off together in
long strips; layers of periderm often forming at several
levels in bark. Cortex absent, or present between layers of
periderm as parenchyma and large groups of fibres;
fibres sometimes forming complete cylinder in younger
stems, shed in older stems. Phloem consisting entirely of
thin-walled cells in regular, radial files, or sometimes
with blocks of fibres at outer edge; rays not dilating
towards outer ends. Crystals: solitary crystals present in
phloem, cortex and periderm.

Xylem
TS . Semi-ring porous. Growth rings distinct. Vessels in
tangential clusters of 2-4, but mainly solitary in late
wood . Fibres moderately thick-walled. Axial parenchyma
mainly paratracheal, also apotracheal, diffuse . Rays 1-3
cells wide.

LS. Vessels: intervascular and vessel-ray pitting alternate;

perforation plates simple. Rays 1-3 cells wide, 1-40 or
sometimes more cells high; heterocellular; cells in RLS
mainly square and upright. 356. Patentilla fruticasa, bark and xylem TS x 150,
showing semi-ring porous xylem .

SPECIES EXAMINED

P. fruticosa L.; P. fruticosa var. tenuifolia; Shrubby Cinque-

foil.

152
356. Potentilla fruticosa, bark and xylem TS x 50, show-
ing semi-ring porous xylem .
351. Potentilla fruticosa, xylem TLS x 150.

358. Potentilla fruticosa, xylem RLS x 380, showing

vessels with simple perforation plates, a heterocellular ray
and alternate vessel-ray pitting.
359. Potentilla fruticosa, xylem LS x 380, showing fine,
alternate vessel pitting.

153
PRUNUS Rosaceae

Bark ings sometimes present. Rays 1-8 cells wide; uniseriate

Peridenn consisting of radial files of rectangular cells, rays 1-20 cells high, multiseriate rays 8-60 cells high;
sometimes with dark contents; outer layers often irregu- multiseriate rays sometimes with uniseriate tails; hetero-
lar, peeling away in strips or large blocks. Cortex usually cellular; cells in RLS predominantly procumbent and
square.
narrow, containing scattered, thick-walled fibres, solitary
or in small groups, and occasional large, thick-walled
stone cells. Phloem rays dilating slightly towards cortex;
SPECIES EXAMINED
conducting tissue in rectangular or triangular blocks con-
taining scattered groups of fibres . Crystals: cluster crys- P. avium L., Wild Cherry; P. cerasifera Ehrh., Cherry
tals usually abundant in phloem and cortex; solitary Plum; P. cerasus L., Sour Cherry; P. communis Arcang.,
crystals usually absent, but present in some species. Almond; P. domestica L., Plum; P. insititia L., Bullace;
P. laurocerasus L., Cherry-Laurel; P. lusitanica L., Portugal
Laurel; P. persica Batsch., Peach; P. serrulata Lindl., Jap-
Xylem
anese Cherry; P. spinosa L., Blackthorn; Prunus sp.
TS. Diffuse porous. Growth rings indistinct. Vessels soli-
tary and in pairs, also sometimes in clusters and radial
chains of up to 5 cells. Fibres with wide lumina in some
species. Axial parenchyma paratracheal and apotracheal,
scattered. Rays 1-8 cells wide.
360. Prunus cerasus, bark and xylem TS x 150, showing a
LS. Vessels: intervascular pitting and vessel-ray pitting few cluster crystals in phloem.
fine, alternate; perforation plates simple. Spiral thicken-
361. Prunus laurocerasus, bark TS x 150, showing groups
of fibres and cluster crystals in cortex .

154
362. Prunus insititia, xylem TS x 150.
363. Prunus cerasus, xylem TLS x 150.

364. Prunus cerasus, xylem RLS x 150, showing a hetero-

cellular ray.
366. Prunus avium, xylem LS x 380, showing vessels with
alternate pitting.

364

155
PTEROCARYA Juglandaceae

Bark
Peridenn consisting of regular, radial files of rectangular
cells, some with dark contents; outer layers peeling in
wide strips. Cortex narrow, parenchymatous. Phloem
wide; rays dilating gradually at first, then widely towards
cortex; conducting tissue in triangular areas including
alternating blocks of gelatinous fibres. Crystals: cluster
crytals abundant in phloem and cortex.

Xylem
TS. Diffuse porous. Growth rings distinct. Vessels mainly
very wide, thin-walled; solitary and in pairs and radial
chains and clusters of up to 6. Fibres thin-walled, with
wide lumina. Axial parenchyma scattered and in incom-
plete tangential uniseriate bands; occasional cells adjac-
ent to vessels. Rays 1-3 cells wide.

LS. Vessels: intervascular and vessel-ray pitting coarse,

alternate; pits oval or polygonal; perforation plates
simple; tyloses sometimes present. Rays 1-3 cells wide;
uniseriate rays 1-25 cells high, multiseriate rays 5-45 cells
high, often with uniseriate portions and tails; heterocellu-
lar; cells in uniseriate portions and tails taller than broad
in ILS; cells in RLS mainly square and upright, with few
procumbent.

SPECIES EXAMINED

P. x rehderiana Schneid. (P. fraxinifolia x P. stenoptera),

Wing-nut.

366. Pterocarya x rehderiana, bark TS x 40, showing

rectangular blocks of phloem fibres and large phloem sieve
elements near cambium.

156
3Ii1. Pterocarya x rehderiana, bark RLS x 150, showing
phloem sieve plates, cluster crystals in chambered cells
and a heterocellular ray .
368. Pterocarya x rehderiana, xylem TS x 30 .

369. Pterocarya x rehderiana, xylem TLS x 150, showing

vessel and vessel -axial parenchyma pitting; narrow hetero-
cellular rays and short thin-walled fibres .
370. Pterocarya x rehderiana, xylem RLS x 380, showing
vessel pitting and vessel-ray pitting .

157
QUERCUS Fagaceae

Bark apotracheal, sometimes in short, uniseriate, tangential

bands especially in late wood, although bands sometimes
Periderm peeling in long or short strips. Cortex narrow,
with fibres and stone cells present in blocks, occasionally indistinct. Rays uniseriate and multiseriate, up to 40 cells
forming a more or less complete cylinder. Phloem rays wide; multiseriate rays often aggregates of numerous
uniseriate rays, sometimes with fibres and parenchyma
dilating slightly at outer ends, sometimes with large
between them.
masses of fibres and stone cells, or fibres in small groups;
conducting tissue interspersed with blocks of 4-20 fibres
L~ . . Vessels: intervascular pitting alternate; vessel-ray
and occasionally with stone cells. Crystals: solitary
crystals associated with blocks of fibres in phloem and pIttmg elongated, often orientated in several directions'
cortex; cluster crystals present in cortex and sometimes in pe~foration plates simple; tyloses often present. Rays uni~
serIa~e a~d multiseriate; uniseriate rays 1-20 cells high;
phloem.
m~I~ISerIate r~ys up to 40 cells wide and up to several
~Iihmetres hIgh, often aggregates of uniseriate rays;
Xylem shghtly heterocellular; cells in RLS often square, more
rarely procumbent or upright.
TS. Diffuse or semi-ring porous. Growth rings often in-
distinct. Vessels solitary, sometimes arranged in loose
oblique lines; circular or oval; usually wide (up to 400 J.lm SPECIES EXAMINED
diameter) or relatively narrow in Q. ilex. Fibre wall thick-
ness variable. Axial parenchyma abundant, para tracheal or Q. dentata Thunb.; Q. ilex L.; Q. robur L.; Quercus sp.;
Oak.

371. Quercus sp., bark and xylem T S x30.

372. Quercus sp., bark TLS x 380, showing solitary
crystals in chambered cells adjacent to fibres and numer-
ous cluster crystals in phloem parenchyma .

158
 373. Quercus ilex, xylem TS x 50, showing uniseriate rays
and one multiseriate ray .
374. Quercus sp., xylem TS x 40 .

.• 1

\\7\
375. Quercus robur, xylem LS x 60, showing vessel and
I vessel-ray pitting, and a tall aggregate ray.
I , ~~ 376. Quercus sp ., xylem RLS x 150, showing vessel-ray
I!' pitting .
I

1.

159
RHAMNUS Rhamnaceae

Bark
Periderm: cells often thick-walled and with dark contents.
Cortex parenchymatous, cells sometimes with slightly
thickened walls; interspersed with large groups of up to
40 or more thick-walled fibres. Phloem rays not dilating
widely towards cortex; conducting tissue with occasional
groups of fibres towards outer edge, as in cortex. Crystals:
both solitary and cluster crystals abundant in cortex and
phloem, solitary crystals mainly associated with groups
of fibres.

Xylem
TS. Diffuse porous. Growth rings distinct. Vessels solitary
and in loose, oblique clusters of up to 8, often in dendritic
arrangement. Fibres frequently gelatinous. Axial paren-
chyma mainly paratracheal. Rays 1-2 cells wide.
m. Rhamnus cathartica, bark TS x 50, showing cavities
LS. Vessels: intervascular and vessel-ray pitting alternate; in cortex and phloem which once contained fibre blocks.
spiral thickenings present; perforation plates simple.
Fibres occasionally septate. Rays 1-2 cells wide; 1-35 cells
high, often with uniseriate tails, cells in RLS procumbent,
square and upright.

SPECIES EXAMINED

R. cathartica L., Buckthorn.

160
378. Rhamnus cathartica, xylem TS x 50, showing
dendritic arrangement of vessels .
379. Rhamnuscathartica, xylem TLS x 150, showing
uniseriate and biseriate rays.

380. Rhamnus cathartica, xylem RLS x 380, showing

alternate vessel and vessel-ray pitting , anarrow vessel
with spiral thickenings and ray cells containing starch
granules .

161
RHODODENDRON Ericaceae

Bark
Periderm consisting of regular, radial files of rectangular
cells, often in alternating bands of thick- and thin-walled
cells, some with dark contents; outer layers peeling, usu-
ally in thin strips. Cortex very narrow, parenchymatous.
Phloem rays dilating towards cortex; conducting tissue
narrow, sclerenchyma absent. Crystals absent.

Xylem
TS. Diffuse porous. Growth rings sometimes indistinct.
Vessels crowded, solitary and in pairs, angular in outline.
Fibres thick-walled. Axial parenchyma paratracheal, abun-
dant. Rays 1-6 cells wide, some cells in multiseriate rays
as wide as narrower vessels; sometimes noded at growth
ring boundaries.

LS. Vessels: vessel elements long; intervascular and

vessel-ray pitting opposite and in places horizontally
elongated; perforation plates scalariform with up to 30
381. Rhododendron sp., bark TS x 150.
closely spaced bars. Rays 1-6 cells wide; uniseriate rays
1-26 cells high, multiseriate rays 7-50 cells high, some
with uniseriate tails; heterocellular, cells in RLS square,
procumbent and upright.

SPECIES EXAMINED

R. calophytum Franch. 'Cynthia' (R. catawbiense Michx. x

R. griffithianum Wight); R. decorum Franch. ; R. falconeri
Hook. f.; R. ponticum L.; R. vernicosum Franch.; Rhododen-
dron sp.

162
384. Rhododendron vernicosum, xylem RLS x 150, show-
ing vessel pitting, scalariform perforation plates and a
heterocellular ray.
386. Rhododendron ponticum, xylem RLS x 380, show-
ing alternate vessel pitting and scalariform perforation
plates.

163
RHUS Anacardiaceae

Bark
Periderm: outer cells often with dark contents; inner cells
in regular radial files, sometimes with slightly thickened
walls. Cortex parenchymatous, fibres and stone cells
absent; interspersed with numerous, often wide, sec-
retory canals. Phloem rays not dilating; sclerenchyma
absent. Crystals: cluster crystals present in cortex.

Xylem
TS. Ring porous. Growth rings distinct. Vessels solitary
and in radial chains and clusters of up to 15 or more. Fibres
angular with wide lumina. Axial parenchyma very sparse.
Rays 1-3 or rarely 4 or more cells wide.

LS. Vessels: intervascular pitting alternate and sometimes

elongated; vessel-ray pitting often elongated; perforation
plates simple; spiral thickenings present. Rays 1-3 or
rarely 4 or more cells wide; 1-30 cells high, multiseriate
rays often with uniseriate tails; markedly heterocellular;
cells in RLS procumbent, square and upright.

SPECIES EXAMINED

Rhus sp.

387. Rhus sp ., xylem TS x 80.

386. Rhus sp., bark and xylem TS x 50, showing

secretory canals in cortex and phloem.

164
 1 ~ r~
lil ~~r' ~
II

\ ) II\:id~1
1\'\';-t
II !I' :J,
r
ll'
~:j"
I

I I

I~
IJ I' I'i
~I~~ 'I
I
:
'~\fl
t. ~
i ~
~\;'
r( Hi

I ~~
, It U~ Ii~\I .... '

388. Rhus sp., xylem TLS x 150, showing vessel elements

with alternate pitting, and short, narrow rays.
389. Rhus sp ., xylem TLS x 380, showing vessels with
alternate pitting and spiral thickenings.

390. Rhus sp., xylem RLS x 380, showing narrow vessels

with spiral thickenings, one simple perforation plate and
vessel-ray pitting .

165
RIBES Grossulariaceae

Bark
Periderm consisting of radial files of rectangular cells, cells
usually narrow radially; some cells with dark contents;
outer layers often distorted and peeling in thin strips or
large blocks, sometimes giving outer surface an uneven
outline. Cortex parenchymatous. Phloem rays usually
remaining the same width as in the xylem; conducting
tissue lacking fibres. Crystals sometimes absent; when
present, cluster crystals abundant in tangential lines in
phloem cells alternating with cells not containing crystals.

Xylem
TS. Diffuse porous. Growth rings indistinct. Vessels
narrow, sometimes very crowded, usually angular in out-
line; solitary and in pairs. Fibres with wide lumina. Axial
parenchyma infrequent, scattered, virtually indistinguish-
able from fibres. Rays 1-10 cells wide.

LS. Vessels: intervascular and vessel-ray pitting fine, alter-

nate or opposite; pits oval or elongated; perforation
plates scalariform, with up to 16 bars. Fibre-tracheids pres-
ent. Rays uniseriate and 2-10 cells wide; uniseriate rays
1-12 cells high, multiseriate rays 8-135 or more cells high,
sometimes with uniseriate tails of tall cells; marginal ray
cells often much taller than broad, forming a sheath;
heterocellular, consisting mainly of square and upright
cells.

SPECIES EXAMINED

R. nigrum L., Blackcurrant; R. sanguineum Pursh, Flower-

ing currant; R. uva-crispi L., Gooseberry. 391. Ribes uV8-crispi, bark and xylem TS x 60.

166
392. Ribes uva-crispi, xylem TLS x 150.
393. Ribes uva-crispi, xylem LS x 380, showing elongated
vessel pits .

394. Ribes uva-crispi, xylem RLS x 600, showing a

scalariform perforation plate.

167
ROBINIA Leguminosae

Bark
Periderm consisting of radial files of thin-walled, rectangu-
lar cells; outer layers peeling in narrow strips. Cortex
narrow or absent, with blocks of 20 or more fibres, often
gelatinous. Phloem rays remaining same width for most of
length; conducting tissue alternating with more or less
rectangular blocks of fibres and occasional stone cells.
Crystals: solitary crystals present in phloem and cortex,
often associated with fibres .

Xylem
IS. Diffuse porous. Growth rings indistinct. Vessels very
wide, or narrow. Wide vessels mainly solitary and in
pairs, narrow vessels also in clusters of up to 7, some-
times adjacent to wide vessels; thin-walled tyloses and
gum deposits present in some vessels. Fibres in blocks,
usually with gelatinous inner walls, often narrower than
axial parenchyma cells. Axial parenchyma paratracheal,
sometimes abundant. Rays 1-6 cells wide.

LS. Vessels: narrow vessels and tracheids storied; inter- 396. Robinia pseudoacacia, bark and xylem TS x 40.
vascular and vessel-ray pitting alternate or sometimes
elongated, vestured; perforation plates simple; spiral
thickenings sometimes present on narrow vessels and
tracheids. Axial parenchyma storied, cells usually fusiform
with single division. Rays 1- 6 cells wide; uniseriate rays
1-30 cells high, multiseriate rays 8-100 or more cells high,
sometimes with uniseriate tails; heterocellular; cells in
RLS mainly square and procumbent.

SPECIES EXAMINED

R. pseudoacacia L., False Acacia or Locust Tree.

168
396. Robinia pseudoacacia, xylem TS x 80 .
397. Robinia pseudoacacia, xylem TLS x 150.

398. Robinia pseudoacacia , xylem TLS x 150, showing

storied vessels, axial parenchyma and rays.

169
ROSA Rosaceae

Bark
Peridenn consisting of regular, radial files of thin-walled
rectangular cells, some with dark contents; outer layers
peeling in narrow strips. Cortex narrow, sometimes with
scattered groups of 1-10 thin-walled fibres. Phloem rays
dilating widely towards cortex; conducting tissue in rec-
tangular or triangular blocks with blocks of 1-50 or more
thick-walled fibres with wide lumina. Crystals: solitary,
often abundant in phloem and rays .

Xylem
TS. Diffuse porous. Growth rings indistinct. Vessels wide
and narrow, mainly solitary, and in pairs. Fibres moder-
ately thick-walled with narrow lumina. Axial parenchyma
paratracheal and scattered. Rays uniseriate to more than
25 cells wide, widening outwards.

LS. Vessels: intervascular and vessel-ray pitting alternate,

fine; perforation plates simple; spiral thickenings some-
times present. Rays uniseriate to 25 or more cells wide;
uniseriate rays 1-15 cells high, multiseriate rays 30-130
cells high, usually tapering abruptly; heterocellular, cells
in RLS mainly square and upright.

SPECIES EXAMINED

R. canina L., Rosa sp., Rose.

399. Rosa sp., bark and xylem TS x 60.

170
400. Rosa sp ., xylem TLS x 60, showing unusually
narrow rays.
401. Rosa sp., TLS x 150, showing fine alternate vessel 402. Rosa sp., xylem TLS x 380, showing vessel with
pitting and two wide rays. fine, polygonal, alternate pitting.
403. Rosa sp., xylem RLS x 380, showing alternate
vessel-ray pitting.

171
ROSMARINUS Labiatae

Bark
Periderm: outer layers irregular, often forming wedge-
shaped blocks; cells of outer layers often thick-walled,
inner layers thin-walled. Cortex parenchymatous, with
groups of 2-20 thick-walled fibres. Phloem rays narrow,
not dilating towards cortex. Crystals: cluster crystals pres-
ent in cortex and phloem.

Xylem
TS. Diffuse porous. Growth rings indistinct. Vessels
mainly solitary and in tangential pairs; occasionally in
clusters of 3. Fibres thick-walled. Axial parenchyma para-
tracheal and sometimes occurring in large areas between
vessels. Rays mainly uniseriate and occasionally 1-3 cells
wide.

LS. Vessels: intervascular and vessel-ray pitting alternate;

perforation plates simple. Rays mainly uniseriate and
occasionally 1-3 cells wide; 2-20 cells high; heterocellu-
lar; cells in RLS procumbent, square and upright.

SPECIES EXAMINED

R. officinalis 1., Rosemary.

172
404. Rosmarinus officina/is, bark TS x 150.
406. Rosmarinus officina/is, xylem TS x 150.
406. Rosmarinus officina/is, xylem TLS x 380, showing
vessels with fine, alternate pitting and simple perforation
plates.

173
RUBUS Rosaceae

Bark
Periderm consisting of layers of rectangular cells, outer
cells often with dark contents. Cortex parenchymatous.
Phloem rays dilating widely towards cortex; conducting
tissue in triangular blocks, with a few scattered thin-
walled fibres with wide lumina. Crystals: solitary and
double crystals, and sometimes abundant cluster crystals
present in phloem and cortex.

Xylem
TS. Diffuse porous. Growth rings indistinct. Vessels wide,
solitary and in pairs, and clusters and radial chains of up
to 5 cells. Fibres moderately thick-walled. Axial paren-
chyma paratracheal and diffuse, sparse. Rays 1-12 cells
wide.

LS. Vessels: intervascular and vessel-ray pitting alternate,

fine; perforation plates simple. Rays 1-4, but more com-
monly 5-12 cells wide. Narrow rays 1-18 cells high, wide
multiseriate rays usually more than 40 cells high, usually
without uniseriate tails; heterocellular; cells in RLS
mainly square and upright.
407. Rubus sp ., bark and xylem TS x 50, showing wide
vessels in xylem, wide rays dilating in phloem and
scattered solitary and cluster crystals.
SPECIES EXAMINED

Rubus sp., Blackberry.

174
408. Rubus sp., bark and xylem TS x 60, showing a
young root.
409. Rubus sp., xylem TS x 150.

410. Rubus sp., xylem TLS x 50, showing rays of two

distinct sizes: narrow and short, and wide and tall.
411. Rubus sp ., xylem RLS x 150, showing vessels with
fine pitting, simple perforation plates and a heterocellular
ray .

175
SAMBUCUS Sambucaceae

Bark LS. Vessels: intervascular pitting alternate; pits coarse,

Periderm outer surface irregular; outer layers peeling off in tightly packed; vessel-ray pits circular or horizontally
large blocks, consisting mainly of broken and crushed elongated; perforation plates simple. Rays 1-4, occasion-
cells; inner layers consisting of regular files of thin- ally 6, cells wide; uniseriate rays 1-11 cells high,
walled, rectangular cells without contents. Cortex nar- multiseriate rays 6-120 or more cells high; ends of rays
row, parenchymatous, with scattered fibres in groups of tapering, sometimes with uniseriate tails; heterocellular;
1-4. Phloem wide; rays dilating gradually towards cortex, cells in RLS mainly square and upright.
conducting tissue alternating with blocks of 2-30 angular
fibres with wide lumina. Crystals: crystal sand present in
phloem rays and cortex. SPECIES EXAMINED

S. nigra L., Elder.

Xylem
TS. Diffuse porous. Growth rings indistinct. Vessels
crowded, solitary and in pairs and small multiples of up
to 4, thin-walled. Fibres with wide lumina. Axial paren-
chyma indistinct, mainly paratracheal. Rays 1-4, occasion-
ally 6, cells wide.

412. Sambucus nigra, bark TS x 55, showing blocks of

thin-walled fibres in phloem and cortex.
413. Sambucus nigra, bark TS x 380, showing crystal
sand in some phloem parenchyma cells.
414. Sambucus nigra, xylem TS x 60.
415. Sambucus nigra, xylem TLS x 150, showing hetero-
cellular rays .
416. Sambucus nigra, xylem TLS x 380, showing coarse
vessel pitting and scalariform vessel-parenchyma pitting .
417. Sambucus nigra, xylem RLS x 150, showing coarse
vessel pitting, horizontally elongated vessel-ray pitting and
a heterocellular ray.

176
177
SARCOCOCCA Buxaceae

Bark
Peridenn narrow, consisting of thin-walled rectangular
cells in regular files, breaking into wedge-shaped blocks.
Cortex parenchymatous with occasional small groups of
1-5 small, thick-walled stone cells. Phloem rays dilating
gradually towards cortex; conducting tissue with stone
cells occasionally present, solitary and in small groups.
Crystals absent.

Xylem
TS . Diffuse porous. Growth rings indistinct. Vessels nar-
row, angular, solitary, and in groups of 2-3. Fibres with
wide lumina. Axial parenchyma diffuse. Rays 1-4 cells
wide; ray cells often similar in width to vessels.

LS. Vessels: intervascular and vessel-ray pitting opposite

or horizontally elongated or scalariform; perforation
plates very oblique, scalariform, often with 40 or more
bars. Rays 1-4 cells wide, 1- 50 or more cells high,
markedly heterocellular; cells in TLS very irregular in
shape, often taller than broad, in RLS mainly square and
upright.

SPECIES EXAMINED

S. ruscifolia Stapf, Sweet Box.

418. Sarcococca ruscifolia, bark TS x 60 .

178
419. Sarcococca ruscifo/ia , xylem TS x 150, showing
narrow, angular vessels and ray cells of similar w idth .
420. Sarcococca ruscifo/ia, xylem TLS x 150, showing
heterocellular rays.

421. Sarcococca ruscifo/ia, xylem RLS x 150, showing

vessels with fine pitting and scalariform perforation plates .

179
SENECIO Compositae

Bark
Peridenn very irregular in appearance, with irregularly
shaped cells. Cortex parenchymatous, with occasional
groups of fibres; secretory canals present. Phloem tissue
thin-walled; rays sparse, dilating slightly towards outer
ends, sometimes with secretory canals. Crystals absent.

Xylem
TS. Diffuse to semi-ring porous. Growth rings sometimes
indistinct. Vessels crowded in some areas (early wood),
very sparse and narrow in other areas (late wood); soli-
tary and in radial chains and clusters of 2-6. Fibres thick-
walled. Axial parenchyma abundant in early wood; rela-
tively sparse, paratracheal elsewhere. Rays infrequent,
1-4 cells wide.

LS. Vessels storied; intervascular and vessel-ray pitting

alternate, fine; perforation plates oblique, simple. Fibres
short, storied. Axial parenchyma often storied. Rays 1-4
422. Senecio greyi, bark and xylem TS x 60, showing
irregular periderm, secretory canals in phloem and very
cells wide, 2-60 or more cells high; markedly heterocellu-
narrow vessels in xylem.
lar; cells tall in ILS and usually upright or occasionally
square in RLS.

SPECIES EXAMINED

S. greyi Hort.

180
 ' -'U"~Nl,
IlIll"~: ~

l~
J Il
U ~~~
~
,~ ;r.;~rr:i!

'lo
IJ
, ~! j

~M
~
~ ~
n~
it ~ "- ~ ~(~~m~
~
tJ~ ~j
. ~~.. t(~~
r~

i,~1-,..~~~~
~ ~ ~

~K '
c~ ~
~~¥U
-:.1i ~~ :'!'.
~ I> ~
n
1M"
rlIL ,~r ~
{:~;
H !:.Q:.
~~
"" ~ ]iii
~~ ~

~~~~
n ,. ~~ ~

~ """"'l
~
~ ' . e , ~([
-, ~
. ~ ~'r
.&.
rr.J,
~

-
--)- ~ r1..
"'~.,.
~,..

~~
F"'I
r~"•
po ~
~ ~J '\,;l

: ~J!
I!! ~ •
,~ ~ t:: I' ~ ,-;
r ~~ ~
~ ~

l~,.
~rt
11 •• ~
,
423. Senecio greyi, xylem TS x 150.
p..~ ~t"
423 j ~ttlI( :L~ I rJl 424. Senecio greyi, xylem TLS x 60, showing narrow, tall
~ )!~ • '-'lI"a,." heterocellular rays .

425. Senecio greyi, xylem RLS x 150, showing storied

vessels with fine pitting and simple perforation plates,
storied axial parenchyma, and fibres .

181
SKIMMIA Rutaceae

Bark
Peridenn: outer layers peeling in long strips; cells often
with thickened periclinal walls and occasionally short,
tangential rows of cells with thick, lignified walls; cells
arranged in regular radial files, rectangular in IS. Cortex
mainly parenchymatous, narrow. Phloem rays dilating
widely towards outer ends, conducting tissue in long,
triangular or wedge-shaped areas interspersed with
groups of gelatinous fibres . Crystals: sparse solitary crys-
tals present in cortex and phloem.

Xylem
TS. Growth rings: none seen. Vessels crowded, ranging in
size from narrow to wide, mainly in large clusters of up to
12 and long radial chains of up to 10, also occasionally
solitary. Fibres relatively sparse, thin-walled. Axial paren-
chyma abundant, diffuse. Rays 1-5 cells wide.

LS. Vessels often storied; intervascular and vessel-ray pit-

ting alternate, polygonal. Perforation plates simple. Axial
parenchyma storied. Rays 1-5 cells wide, 1-40 cells high;
markedly heterocellular; cells in RLS procumbent, square
and upright.

SPECIES EXAMINED
426. Skimmia japonica, bark TS x 60.
S. japonica Ihunb.

182
 4Z7. Skimmiajaponica, xylem TS x 60.
428. Skimmiajaponica, xylem TLS x 150, showing
vessels with alternate pitting , narrow rays and axial
parenchyma .

429. Skimmiajaponica, xylem RLS x 150, showing vessel

and vessel-ray pitting and a heterocellular ray . 429
183
SOPHORA Leguminosae

Bark
Peridenn consisting of radial files of thin-walled rectangu- parenchyma cells usually storied, many cells fusiform,
lar cells, sometimes with dark contents; outer layers some once or twice divided. Rays 1-4 cells wide; uni-
crushed and distorted, peeling in strips. Cortex narrow, seriate rays 2-4 cells high, multiseriate rays 7-150 or more
parenchymatous, with occasional groups of gelatinous cells high, often with uniseriate tails; heterocellular, cells
fibres. Phloem rays narrow near cambium, dilating widely in RLS procumbent, square and upright.
towards cortex; conducting tissue with alternating bands
of fibres. Crystals: solitary crystals present in cortex and
phloem, usually in association with fibres. SPECIES EXAMINED

S. affinis Torr. and Gray; S. flavescens Ait.; S. japonica L.;

Xylem Pagoda Tree.
TS. Diffuse porous. Growth rings frequently indistinct.
Vessels solitary, in pairs, radial multiples and clusters of
up to 10; many vessels very wide. Fibres gelatinous, nar-
rower than axial parenchyma cells, and in conspicuous
blocks. Axial parenchyma forming bulk of ground tissue;
mainly paratracheal and in tangential bands 1-3 cells
wide traversing axial tissue; individual cells much thin-
ner-walled than narrow vessels and fibres. Rays 1-4 cells
wide.

LS. Vessels: intervascular pitting and vessel-ray pitting

alternate, pits oval or polygonal where crowded, often
horizontally elongated; pit vesturing not always apparent
with light microscope; perforation plates simple; narrow
vessels storied, sometimes with spiral thickenings. Axial

430. Sophora flavescens, bark and xylem TS x 60.

431. Sophora flavescens, xylem TS x 60, showing wide
vessels and thin-walled fibres.

184
432. Sophorajaponica, xylem TS x 150, showing wide
vessels, blocks of narrow fibres and wider paratracheal
parenchyma.
433. Sophora affinis, xylem TLS x 150, showing storied
axial parenchyma and narrow rays.

434. Sophora affinis, xylem LS x 380, showing a vessel

element with coarse, alternate pits, indistinct vesturing.

185
SPIRAEA Rosaceae

Bark SPECIES EXAMINED

Peridenn consisting of regular, radial files of thin-walled, S. japonica L.f. 'Bumalda Elegans'; S. x sanssouciana
rectangular cells, some with dark contents; outer layers K. Koch (S. japonica x S. douglasii) 'Superbetulifolia';
sometimes peeling in strips. Cortex narrow, up to about 5 S. veitchii Hemsl.
cells wide, many cells with dark contents; inner cortex
and outer phloem consisting of an almost continuous
cylinder of small stone cells and fibres up to 20 or more
cells wide. Phloem rays dilating towards cortex, some
outer ray cells consisting of thin-walled stone cells with
wide lumina; conducting tissue in more or less rectangu-
lar blocks without sclerenchyma. Crystals: solitary and
cluster crystals present in phloem, rays and cortex, many
associated with sclerenchyma cylinder.

Xylem
TS . Diffuse porous. Growth rings distinct. Vessels mainly
solitary and in pairs, sometimes in groups of up to 4;
vessels wide in S. veitch ii, narrow and angular in outline
in other species. Fibres with wide lumina. Axial paren-
chyma paratracheal and diffuse . Rays 1-7 cells wide.

LS. Vessels: intervascular and vessel-ray pitting very fine,

alternate; perforation plates simple. Rays 1-7 cells wide;
uniseriate rays 1-20 cells high, multiseriate rays 8-95 cells
high, sometimes with uniseriate tails; heterocellular, cells
in RLS procumbent, square and upright.

435. Spiraea veitchii, bark and xylem TS x 80 .

436. Spiraea veitch/i, bark RLS x 150, showing a hetero-
cellular ray, scattered solitary crystals in phloem and many
small stone cells with wide lumina in cortex .

186
437. Spiraea veitchii, xylem TS x 150.
438. Spiraea x sanssouciana 'Superbetulifolia', xylem
TS x 150.
439. Spiraea japonica 'Bumalda Elegans', xylem
TLS x 150, showing tall, narrow vessel elements, short,
uniseriate rays and taller, multiseriate rays .
440. Spiraea japonica 'Bumalda Elegans', xylem
TLS x 380, showing vessels with fine, alternate pitting,
and heterocellular rays.

439
187
STYRAX Styracaceae

Bark
Periderm irregular; cells with dark contents. Cortex inter-
spersed with mainly solitary fibres and rarely groups of
stone cells. Phloem: cells mainly thin-walled, with
occasional solitary fibres; rays dilating slightly towards
outer ends. Crystals: solitary and cluster crystals very
abundant in cortex and phloem.

Xylem
TS . Diffuse porous. Growth rings indistinct. Vessels
mainly in clusters and radial chains of up to 20 or more
cells, rarely solitary. Fibres mainly with moderately wide
lumina and thin-walled. Axial parenchyma apotracheal
and in uniseriate, tangential bands. Rays 1-5 cells wide.

LS. Vessels: intervascular and vessel-ray pitting fine,

opposite to alternate, and sometimes slightly elongated;
perforation plates scalariform, with usually 4-8 widely
spaced bars. Rays 1-5 cells wide; 2-60 or more cells high; 441. Styrax hemsleyana, bark and xylem TS x 60.
multiseriate rays often with uniseriate tails and portions;
markedly heterocellular; cells in RLS procumbent, square
and upright; mainly upright in uniseriate portions.

SPECIES EXAMINED

S. hemsleyana Diels, Storax.

188
442. Styrax hems!eyana, xylem TS x 60.
443. Styrax hems!eyana, xylem TLS x 60 , showing rays .

444. Styrax hems!eyana, xylem RLS x 150, showing a

heterocellular ray.
446. Styrax hems!eyana, xylem RLS x 380, showing a
vessel with fine pitting and a scalariform perforation plate ,
and ray cells containing starch granules.

189
SYMPHORICARPOS Caprifoliaceae

Bark
Peridenn consisting of radial files of square or rectangular
cells, some tangential rows of thick-walled cells. Cortex
narrow or absent. Phloem rays not dilating or slightly
dilating towards cortex; conducting tissue occasionally
with small groups of 1-10 thin-walled fibres, with wide
lumina. Crystals: cluster crystals present in phloem and
rays.

Xylem
TS. Ring porous or semi-ring porous. Growth rings dis-
tinct. Vessels angular, solitary, in pairs and clusters of up
to 5; vessels often crowded in early wood, sparse in late-
wood. Fibres with slightly thickened to thick walls. Axial
parenchyma sparse, paratracheal and diffuse. Rays 1-5, or
rarely up to 10 cells wide.

LS. Vessels: intervascular pitting and vessel-ray pitting

fine, alternate and occasionally opposite and scalariform;
perforation plates mainly simple, scalariform perforation
plates very infrequent, with up to 25 bars. Fibres occasion-
ally with spiral thickenings. Rays 1-5 or occasionally up to
10 cells wide, 10-50 or more cells high, occasionally with
uniseriate tails of tall cells; markedly heterocellular; indi-
vidual cells of variable shape, marginal cells sometimes
forming incomplete sheaths; cells in RLS mainly square
and upright.
446. Symphoricarpos occidentalis, bark and xylem
TS x 80, showing ring porous xylem.
NOTE

See Appendix, Table 4.

SPECIES EXAMINED

S. albus Blake, Snowberry; S. x chenaultii Rehd. (5. micro-

phyllus x S. orbiculatus); S. occidentalis Hook., Wolfberry;
S. rotundifolius C. Koch.

190
447. Symphoricarpos albus, bark and xylem TS x 50,
showing ring porous arrangement of vessels .
448. Symphoricarpos occidentalis, xylem TLS x 150,
showing heterocellular rays.

449. Symphoricarpos albus, bark and xylem LS x 380,

showing cluster crystals in phloem, and vessels with fine
pitting and a simple perforation plate .

460. Symphoricarpos occidentalis, xylem RLS x 380,

showing a vessel with fine pitting and a scalariform
perforation plate, and a heterocellular ray.

191
SYRINGA Oleaceae

Bark
Periderm consisting of thin-walled cells; groups of cells
sometimes split into blocks. Cortex with groups of stone
cells and occasional fibres; sclerenchyma often in concen-
tric, discontinuous, or rarely continuous, rings. Phloem
with rings of sclerenchyma as in cortex; rays dilating
slightly towards outer ends. Crystals absent.

Xylem
T5. Diffuse or seml-rmg porous. Growth rings often
indistinct. Vessels mainly solitary, and in clusters of up to
6. Fibres with wide lumina. Axial parenchyma absent or
very sparse, paratracheal. Rays 1-3 cells wide.

L5. Vessels: intervascular pitting and vessel-ray pitting

alternate, sometimes slightly elongated; perforation
plates simple; spiral thickenings sometimes present. Rays
1-3 cells wide, 2-30 cells high; multiseriate rays some-
times with uniseriate portions and tails; heterocellular;
cells in RLS procumbent, square and sometimes upright.
Crystals: small, solitary crystals rarely present in ray cells.

SPECIES EXAMINED

5. villosa Va hi; 5. vulgaris L.; 5. yunnanensis Franch.;

Syringa sp.; Lilac.

451. Syringa vil/osa, bark and xylem TS x 60.

192
452. Syringa vil/osa, xylem TLS x 150, showing narrow
vessels with alternate pitting and narrow, heterocellular
rays.
453. Syringa vil/osa, xylem RLS x 150, showing vessel
elements with simple perforation plates, heterocellular rays
with predominantly procumbent cells, alternate vessel-ray
pitting, and small solitary crystals in some ray cells .

454. Syringa vil/osa, xylem RLS x 380, showing vessels

with simple perforation plates and vessel-ray pitting.

193
TAMARIX Tamaricaceae

Bark
Peridenn consisting of square and rectangular cells; outer
cells crushed and distorted, outer boundary irregular.
Cortex very narrow, parenchymatous. Phloem rays dilat-
ing widely towards cortex; conducting tissue in rectangu-
lar blocks capped with triangular blocks of fibres, some-
times broken up by parenchyma cells; fibres with wide
lumina. Crystals absent.

Xylem
T5. Diffuse porous. Growth rings indistinct. Vessels soli-
tary and in pairs and clusters of up to 5. Tracheids often
present adjacent to vessels. Fibres with wide lumina. Axial
parenchyma paratracheal. Rays 1-10 cells wide.

L5. Vessels: intervascular pitting fine, alternate; pits poly-

466. Tamarix sp ., bark and xylem TS x 60.
gonal; perforation plates simple; vessels storied. Tra-
cheids storied, with similar pitting to vessels. Rays 1-10
cells wide; uniseriate rays very infrequent, 4-8 cells high,
multiseriate rays 12-150 or more cells high; heterocellu-
lar; cells in RLS square and upright.

SPECIES EXAMINED

Tamarix sp., Tamarisk.

468. Tamarix sp., xylem TLS x 380, showing storied

fusiform axial parenchyma in front of vessel elements,
storied fibres and part of a multiseriate ray .
469. Tamarix sp., xylem RLS x 380, showing vessel
elements with fine, alternate pitting .

194
466. Tamarix sp ., xylem TS x 150.
467. Tamarix sp ., xylem TLS x 50, showing storied
vessels and fibres and tall rays.

195
TILIA Tiliaceae

Bark
Periderm very narrow, consisting of rectangular cells.
Cortex very narrow, consisting of thin-walled paren-
chyma cells; sclerenchyma absent . Phloem rays dilating
widely towards cortex, consisting of square or tangen-
tially rectangular parenchyma cells; conducting tissue in
triangular blocks, including alternating bands of thick-
walled fibres. Crystals: cluster crystals present in phloem
rays and cortex; solitary crystals infrequent or absent, but
occasionally present in conducting tissue .

Xylem
TS . Diffuse porous or semi-ring porous. Growth rings
distinct. Vessels in radial multiples of 2-13, and in pairs
and clusters, occasionally solitary; often angular in out-
line. Fibres thin-walled with wide lumina. Axial paren-
chyma often forming large part of axial tissue, diffuse and
often in radial chains alongside rays . Rays 1-6 cells wide.

LS. Vessels: intervascular and vessel-ray pitting fine, alter-

nate; perforation plates simple; spiral thickenings
present. Axial parenchyma conspicuous, slightly storied.
Rays uniseriate and up to 6 cells wide; uniseriate rays 1-30
cells high, multiseriate rays 9-75 cells high, often tall and
460. Tilia sp ., bark and xylem TS x 80, showing phloem
narrow; cells thin-walled; heterocellular; cells in RLS
with widely dilating rays, triangular blocks of alternating
mainly procumbent and square. fibres and conducting tissue and scattered cluster crystals;
xylem with vessels in radial multiples and clusters.

SPECIES EXAMINED

T. x europaea L. (T. cordata x T. platyphyllos); Tilia sp.;

Lime.

463. Tilia sp ., xylem RLS x 150, showing a heterocellular

ray and vessel-ray pitting .
464. Tilia sp ., xylem LS x 380, showing vessels with fine,
alternate pitting, simple perforation plates and spiral
thickenings.

196
461 . Tilia sp ., xylem TS x 80 .
462. Tilia sp., xylem TLS x 150, showing vessels with
fine, alternate pitting and spiral thickenings, thin -walled
fibres and axial parenchyma, and narrow rays .

197
ULMUS Ulmaceae

Bark
Peridenn: outer layers peeling in short strips; cells often
with dark contents. Cortex parenchymatous, interspersed
with large mucilage-containing cells, and occasional
small groups of fibres. Phloem rays not dilating widely
towards cortex; conducting tissue interspersed with
small groups of up to 20 thick-walled, often gelatinous,
fibres. Crystals: solitary crystals present in cortex and 466. Ulmus sp ., bark TS x 150, showing phloem with
phloem. several large mucilage-containing cells, and scattered
gelatinous fibres .
Xylem 466. Ulmus procera, bark and xylem TS x 60, showing
phloem with large mucilage-containing cells and scattered
IS . Diffuse porous. Growth rings indistinct. Vessels both
fibres .
narrow and wide, crowded, solitary and in radial chains
and large clusters of up to 25. Fibres in conspicuous 467. Ulmus procera, xylem TS x 40, showing diffuse
blocks, often gelatinous. Axial parenchyma often forming porous arrangement of wide and narrow vessels .
bulk of ground tissue. Rays 1-6 cells wide.
468. Ulmus procera , xylem TLS x 50, showing storied
vessels, tracheids and axial parenchyma, short, uniseriate
LS. Vessels: intervascular and vessel-ray pitting alternate, rays and tall, multiseriate rays.
polygonal or circular; perforation plates simple. Small
vessels and tracheids storied and often with faint spiral 469. Ulmus sp ., xylem TLS x 380, showing two narrow
thickenings. Rays 1-6 cells wide, up to 70 or more cells vessel elements with alternate, polygonal pitting and
high; height and width very variable; not markedly simple perforation plates, vessel to axial parenchyma
heterocellular; cells in RLS mainly procumbent and pitting and a short ray .
square. 470. Ulmus x sarniensis, xylem RLS x 380, showing a
homocellular ray with alternate vessel-ray pitting.
SPECIES EXAMINED

U. glabra Huds.; U. procera Salisb. ; U. x sarniensis Ban-

croft (U. angustifolia x U. ho/landica); U. stricta Lindl.;
Ulmus sp.; Elm.

198
199
VIBURNUM Caprifoliaceae

Bark
Peridenn consisting of radial files of thin-walled rectangu-
lar cells, sometimes alternating with bands of thick-
walled cells, breaking into blocks on the periphery. Cortex
narrow, sometimes with large thick-walled stone cells,
with wide or narrow lumina; fibres absent. Phloem rays
dilating widely towards cortex; conducting tissue often
without sclerenchyma, but large thick-walled stone cells
or small groups of 1-20 fibres sometimes present.
Crystals: cluster and solitary crystals often abundant.

Xylem
T5. Diffuse porous. Growth rings sometimes indistinct.
Vessels angular, solitary, in pairs and occasional radial
multiples and clusters of up to 5. Fibres with wide,
occluded lumina. Axial parenchyma para tracheal and
diffuse. Rays 1-4 cells wide.

L5. Vessels: intervascular pitting and vessel-ray pitting 471. Viburnum tinus, bark and xylem TS x 60.
fine, opposite and in places scalariform or alternate; per-
foration plates strongly oblique, scalariform, long, with
15-30 or more bars. Fibres occasionally with spiral thick-
enings. Rays uniseriate and 2-4 cells wide, uniseriate rays
1-20 cells high, individual cells taller than broad; multi-
seriate rays 10-50 cells high, sometimes with uniseriate
tails, individual cells of varying shape; heterocellular;
cells in RLS square and upright, occasionally procum-
bent.

NOTE

See Appendix, Table 4.

SPECIES EXAMINED

V. x bodnantense Aberconway (V. farreri x V. grandi-

[Iorum); V. buddleifolium C. H. Wright; V. faTTeri Stearn;
v. foetens Decne.; V. lantana L., Wayfaring Tree; V.
rhytidophyllum Hemsl.; V. tinus L.

474. Viburnum farreri, xylem RLS x 150, showing vessels

with scalariform perforation plates and heterocellular rays
with mainly opposite and scalariform vessel-ray pitting.
475. Viburnum farreri, xylem TLS x 380, showing a vessel
with opposite and scalariform pitting.

200
472. Viburnum farreri, bark and xylem TS x 60 .
473. Viburnum farreri, xylem TLS x 60.

201
VITACEAE
Ampe/opsis, Parthenocissus, Vitis
Very similar in their root anatomy. Xylem
TS. Usually diffuse porous or occasionally semi-nng
Bark porous. Growth rings indistinct. Vessels crowded, solitary
Periderm: outer edge often peeling in short strips; often and in clusters of 2-5 and radial chains of up to 8; both
several layers thick; cells rectangular, sometimes with narrow and very wide and occasionally with intermediate
thickened periclinal walls, and occasional cells with dark sizes. Fibres sometimes with large lumina. Axial paren-
contents. Cortex narrow in some cases where phloem chyma sparse, paratracheal. Rays 1-20 cells wide;
extending almost to periderm; parenchymatous, cells heterocellular; cells often square.
occasionally with dark contents; large thin-walled sec-
retory cells often present; stone cells rare. Phloem rays LS. Vessels: intervascular and vessel-ray pitting scalari-
dilating widely towards outer ends; conducting tissue in form or occasionally opposite; perforation plates mainly
wedge-shaped blocks, sometimes extending almost to simple, rarely scalariform; faint spiral thickenings some-
periderm, often divided by short phloem rays, and con- times present on small vessels and tracheids; tyloses
sisting of thin-walled tissue alternating with blocks of sometimes present. Rays very wide, 1-15 or up to 20 cells,
fibres. Crystals usually abundant but occasionally sparse; and often several millimetres high; heterocellular, includ-
solitary crystals often present around phloem fibres and ing occasional larger cells sometimes containing rap hides
in phloem rays and cortex; cluster crystals often present or cluster crystals, and occasional cells with dark con-
in cortex and phloem rays and rarely around phloem tents; cells in RLS mainly square, also procumbent and
fibres; raphides sometimes present in cortex and phloem upright.
ravs.

202
476. Parthenocissus inserta , bark TS x 30.
477. Ampe/opsisarborea, bark TS x 50.
478. Vilis vinifera, bark and xylem TS x 80 .
479. Vilis vinifera, bark TS x 80, showing phloem with
blocks of fibres associated with solitary crystals .
480. Parthenocissus inserta, xylem TS x 80, showing
wide and narrow vessels and wide , multiseriate rays.

SPECIES EXAMINED

A. arborea Koehne; A. brevipedunculata Trautv. , Boston

Ivy; P. inserta K. Fritsch; P. quinquefolia Planch., Virginia
Creeper; V. coignetiae Planch.; V. vinifera L.; Vitis sp.;
Vine.

203
 . 481. Vitis vinifera, xylem TS x 80, showing centre of root.
482. Ampe/opsis arborea, xylem TS x 80 .
483. Vitis coignetiae, xylem TLS x 50, showing tali,
multiseriate rays.

204
484. Vitis coignetiae, xylem TLS x 150, showing narrow
vessels and tracheids with scalariform pits and a tall, wide,
multiseriate ray.

485. Vitis vinifera, xylem TLS x 60, showing wide rays

composed of small and larger cells .
486. Parthenocissus inserta. xylem TLS x 150. showing
the tapering end of a wide ray. some ray cells containing
cluster crystals .

205
WISTERIA Leguminosae

Bark
Peridenn consisting of regular radial files of rectangular
cells, some with dark contents; peeling in strips. Cortex
parenchymatous, sometimes with small blocks of thick-
walled gelatinous fibres. Phloem wide, rays dilating
gradually towards cortex; conducting tissue alternating
with rectangular blocks of gelatinous fibres, fibre blocks
sometimes present only in outer phloem. Crystals soli-
tary, in phloem and cortex, often associated with fibre
blocks.

Xylem
TS. Diffuse porous. Growth rings: none seen. Vessels
wide or narrow; vessels solitary, in pairs and occasional
clusters of up to 8; narrow vessels and tracheids of similar
width to axial parenchyma cells, angular in outline. Fibres
in small blocks, narrow, mainly with gelatinous inner
walls. Axial parenchyma paratracheal, in large blocks; indi-
vidual cells much wider than fibres. Rays 1-14 cells wide.

LS. Vessels: intervascular and vessel-ray pits alternate;

perforation plates simple; narrow vessels and tracheids
storied, some with spiral thickenings. Axial parenchyma
cells storied, fusiform in shape, divided once or twice,
same length and width as narrow vessels. Crystals: soli-
tary crystals in vertical chains in chambered axial paren-
chyma cells. Rays 1-14 cells wide; uniseriate rays 1-8 cells
high, multiseriate rays 6-100, occasionally 150 or more
cells high, tapering abruptly; heterocellular; cells in RLS
mainly square and upright.

SPECIES EXAMINED

W. floribunda DC.; Wisteria sp.

206
487. Wisteria floribunda, bark TS x 60.
488. Wisteria sp., xylem TS x 60, showing wide vessels,
blocks of fibres and paratracheal parenchyma .
489. Wisteria floribunda, xylem TS x 60.
490. Wisteria sp ., xylem TLS x 50, showing storied
vessels, tracheids and axial parenchyma; short. narrow
rays and tall, wide rays.

207
ZELKO VA Ulmaceae

Bark
Peridenn consisting of thin-walled cells, some containing
tannin; outer layers peeling in thin short strips. Cortex
narrow, with more or less continuous ring of stone cells,
or in some cases stone cells sparse. Phloem rays dilating in
outer part of phloem; conducting tissue alternating with
stone cells. Crystals present in cortex and phloem, soli-
tary.

Xylem
TS. Diffuse porous. Growth rings usually indistinct.
Vessels mainly solitary and in pairs, and narrow vessels
and tracheids in clusters of up to 12 or more. Fibres thin-
walled with narrow lumina. Axial parenchyma para-
tracheal, in tangential bands and blocks. Rays 1-6 cells
wide, consisting of radially elongated rectangular cells.

LS. Vessels: intervascular pits coarse, alternate, poly-

gonal; vessel-ray pitting oval; perforation plates simple;
spiral thickenings present on some narrow vessels and
tracheids; tracheids storied. Rays uniseriate and 2-6 cells
wide; uniseriate rays 1-15 cells high, multiseriate rays
5-110 cells high; cells in RLS procumbent in large
portions of rays, square and upright particularly at ray 491. Zelkova carpinifolia, bark and xylem TS x 60.
margins; some ray cells containing solitary crystals.

SPECIES EXAMINED

Z. carpinifolia K. Koch; Caucasian Elm.

208
492
492. Zelkova carpinifolia, xylem TS x 150.
493. Zelkova carpinifolia, xylem TLS x 150.

494. Zelkova carpinifolia, xylem TLS x 380, showing

vessel elements with alternate pitting and a mainly
biseriate ray.
496. Zelkova carpinifolia, xylem RLS x 380, showing
alternate vessel-ray pitting.

209
Family descriptions
of Gymnosperms
ARAUCARIA Araucariaceae

Bark
Peridenn consisting of numerous, regular layers of narrow
elongated thin-walled cells. Cortex parenchymatous,
interspersed with large, irregularly shaped astrosdereids,
each with numerous minute crystals embedded in their
outer walls; axial resin canals present in both cortex and
phloem. Phloem consisting of thin-walled parenchyma-
tous cells, rounded in TS, and sieve cells, thin-walled and
often crushed; occasional solitary thick-walled fibres
interspersed. Crystals: minute crystals present in outer
walls of astrosdereids and fibres, and also sometimes in
parenchyma cells.

.
Xylem
TS. Growth rings present. Axial parenchyma sparse or • •
absent. Rays uniseriate. Resin canals absent.
.. •
. • • ••"... .....
o
LS. Tracheids with 1-3 rows of polygonal bordered pits • tit

.-
mainly on radial walls and in characteristic alternate
arrangement; cross-field pitting cupressoid. Rays uni-
seriate; radial resin canals absent.
••• ~

•
.'

SPECIES EXAMINED

A. araucana K. Koch, Monkey Puzzle Tree; A. bidwillii

Hook. 496. Araucaria bidwillii, bark and xylem TS x 50, showing
cortex with large astrosclereids and phloem with scattered
fibres .

212
4fl7. Araucaria araucana, bark and xylem TS x 60, show-
ing a resin canal in cortex, and scattered fibres in phloem.
499. Araucaria araucana, xylem RLS x 150, showing
498. Araucaria bidwil/ii, xylem TLS x 50, showing short, tracheids with alternate, polygonal bordered pits and rays
uniseriate rays . with cupressoid cross-field pitting .
500. Araucaria bidwi//ii, xylem RLS x 380, showing
tracheids with alternate, polygonal bordered pits and rays
with cupressoid cross-field pitting .

213
CUPRESSACEAE
Chamaecyparis, x Cupressocyparis (Cupressus x Chamaecyparis), Cupressus,
Juniperus, Thuja.

Bark Xylem
Peridenn usually narrow, irregular, often deep-seated and TS. Growth rings present. Axial parenchyma present. Rays
present in outer phloem layers, so cortex sloughed off. uniseriate. Resin canals absent.
Cortex absent or sometimes present (in Chamaecyparis
nootkatensis, C. thyoides, x Cupressocyparis leylandii, Juni- LS. Tracheids with 1-3 rows of bordered pits mainly on
perus communis), parenchymatous and with axial resin radial walls and in opposite arrangement; cross-field
canals. In Juniperus communis cells occasionally thick- pitting cupressoid, sometimes taxodioid in Chamaecy-
walled and with dark contents. Phloem consisting of paris. Rays uniseriate; radial resin canals absent.
tangential bands of one layer of thick-walled fibres alter-
nating with several layers of thin-walled cells, the thin-
walled cells consisting of alternating layers of sieve cells SPECIES EXAMINED
and rounded parenchyma cells, often with dark contents;
Chamaecyparis lawsoniana ParI., Lawson Cypress; C.
axial resin canals present (not seen in phloem in x
nootkatensis Spach., Nootka Cypress; C. thyoides Britt.,
Cupressocyparis, Chamaecyparis lawsoniana, C. thyoides).
Sterns and Pogg.; x Cupressocyparis leylandii Dallim.
Crystals absent, or present in Juniperus communis, Thuja
(Cupressus macrocarpa x Chamaecyparis nootkatensis),
plicata, where numerous minute crystals lining inside of
Leyland Cypress; Cupressus macrocarpa Gord., Monterey
walls of occasional ray cells.
Cypress; Juniperus communis L., Common Juniper; Thuja
plicata D. Don, Western Red Cedar; T. standishii Carr.

501. Chamaecyparis lawsoniana, bark and xylem TS x 60,

showing phloem with uniseriate tangential lines of fibres
and xylem with distinct growth rings .
502. Chamaecyparis thyoides, bark and xylem TS x 50,
showing phloem with uniseriate, tangential lines of fibres.

214
 503. Chamaecyparis nootkatensis, bark and xylem
TS x 150, showing a resin canal in cortex, and uniseriate,
tangential lines of fibres in phloem .

504. Cupressus macrocarpa, bark and xylem TS x 60.

506. Juniperus communis, bark TS x 60 .

506. Thuja p/icata, bark and xylem TS x 60, showing
several resin canals in phloem .

215
 507. Chamaecyparis nootkatensis, bark RLS x 150, show-
ing phloem fibres, sieve tubes and parenchyma.
508. Juniperus communis, xylem TS x 60.
509. Chamaecyparis nootkatensis, xylem TLS x 150,
showing short and tall uniseriate rays.

509

216
 t .;.- '!
. 1-, n
~
r.
r?; .
(~ ' f·-

· .. i,::'
,~i
·
·
I'
. . .. .
I

.
,

.·
I~ I ~:
I'" _

[ ', oJ
I;'
"

. I : I""
I'"
r-: I-
I?~
I", ; 1'-
,
I ;~
. ~I, ,t j
~, ~

t io-"-
:-
,-- ..
,,)
" i
510 1 ~
,
510. Thuja standishii, xylem TLS x 150, showing an axial
parenchyma strand and short, uniseriate rays.
511. Juniperus communis, xylem RLS x 380, showing ray
cells with one row of large, cupressoid cross-field pits .

512. Thuja standishii, xylem RLS x 150, showing

tracheids with one or two rows of bordered pits on radial
walls and ray cells with two or three rows of cupressoid
cross-field pits .
513. Thuja plicata, xylem RLS x 380, showing cupressoid
cross-field pitting .

217
GINKGO Ginkgoaceae

Bark
Periderm wide; outer layers often forming wedge-shaped
blocks, cells thin-walled. Cortex parenchymatous, inter-
spersed with occasional small groups of thick-walled
stone cells and unlignified gelatinous fibres. Phloem inter-
spersed with thick-walled unlignified gelatinous fibres.
Crystals: large cluster crystals present in cortex and
phloem.

Xylem
TS. Growth rings present. Axial parenchyma sparse. Rays
uniseriate. Resin canals absent.

LS. Tracheids with 1-2 rows of bordered pits in opposite

arrangement, on radial walls; cross-field pitting cup-
ressoid. Rays uniseriate; radial resin canals absent.

SPECIES EXAMINED

C. biloba L., Maidenhair Tree.

514. Ginkgo biloba, bark TS x 60.

517. Ginkgo biloba, xylem TLS x 60, showing very short,

uniseriate rays.

518. Ginkgo biloba, xylem RLS x 150, showing tracheids

with a single row of bordered pits on radial walls and ray
cells with small, cupressoid cross-field pits; some ray cells
containing starch granules.

218
 .. -

?J~
~ I'Jj~
~'fflll~

~~~..J"~
~ 0::; ':1f(~F:!J. ::(
9'.: ~W~

516 if

515. Ginkgo biloba, bark TS x 150, showing cortex and

phloem with gelatinous fibres and scattered cluster
crystals .
516. Ginkgo biloba, xylem TS x 60 .

219
PINACEAE
Abies, Cedrus, Larix, Picea, Pinus, Pseudo/anx, Pseudotsuga, Tsuga .

Bark
Peridenn usually narrow, sometimes with outermost 2-3 LS. Tracheids with 1-3 rows of bordered pits mainly on
layers of cells thick-walled, lignified. Cortex consisting of radial walls and in opposite arrangement; cross-field
irregular parenchymatous cells, sometimes very large pitting piceoid in Picea, Larix, Pseudotsuga, cupressoid in
and mucilaginous in Abies, Pseudolarix; stone cells Tsuga, taxodioid in Abies, Pseudolarix, piceoid, cupressoid
occasionally present in Cedrus, Larix; astrosclereids and taxodioid in Cedrus, large in Pinus; spiral thickenings
(much branched) present in Abies, Cedrus, Tsuga, Pseudo- present in Pseudotsuga. Rays uniseriate, 1-10 or up to 30 or
tsuga; tannin often present in cortical parenchyma and more cells high; radial resin canals present in Picea, Larix
periderm. Phloem irregular, consisting of rounded paren- and Pinus.
chyma cells and crushed sieve cells and sometimes
occasional fibres, gelatinous in Picea, Larix, Pseudolarix;
resin canals seen in Picea, Pinus. Crystals: in Picea and SPECIES EXAMINED
Pinus occasional cells in cortex containing numerous
Abies grandis Lindl., Grand Fir; A. sp.; Cedrus atlantica
minute crystals; in Cedrus longitudinal chains of small
Manetti, Atlas Cedar; Larix decidua Mill., European Larch;
solitary crystals sometimes present in cortex and phloem.
Larix sp., Larch; Picea sp., Spruce; Pinus cembra L., Arolla
Pine; P. pumila Reg., Dwarf Siberian Pine; P. sylvestris L.,
Xylem Scots Pine; P. sp.; Pseudolarix amabilis Rehd., Golden
Larch; Pseudotsuga menziesii Franco, Douglas Fir; Tsuga
TS . Growth rings present. Axial parenchyma present, canadensis Carr., Eastern Hemlock.
sometimes sparse. Rays mainly uniseriate except where
radial resin canals present. Resin canals both axial and
radial present in Picea, Pinus, Larix, Pseudotsuga; small
axial canals (possibly traumatic) seen in Abies grandis.

220
519. Cedrus atlantica, bark and xylem TS x 60 .
520. Picea sp., bark and xylem TS x 50, showing xylem
with axial resin canals lined with thick-walled epithelial
cells.

521. Picea sp., bark LS x 50, showing horizontal resin

canals .
522. Pinus cembra, bark and xylem TS x 50, showing
xylem with axial resin canals lined with thin-walled
epithelial cells .
523. Pseudolarix amabilis, bark and xylem TS x 60.

221
r:~~ ~~ . •

~~~, ~ ~~ ", "-I~.~ .... ".l >- ,..,uon._j

'UIJ .~
i ~ n ,,'.~~n~u~ 7~
:~, ~t • I '~."';)r6.M" &II

~ ~ ~~Rft'~~~au~~

~, ~l f
524 ~ ;~~~""LAI.~)o ~J\~ ~~ ~

524. Abies grandis, xylem TS x 60, showing traumatic

resin canals .
525. Cedrus atlantica, xylem TS x 60.
526. Larix sp., xylem TS x 60, showing several axial resin
canals.
527. Pseudotsuga menziesii, xylem TS x 60, showing
several growth rings with abrupt transition from earlywood
to latewood, and tangential lines of axial resin canals .

222
528. Larix sp., xylem TLS x 50, showing three rays with
horizontal resin canals.
529. Picea sp., xylem TLS x 150, showing several
uniseriate rays and one ray with a resin canal.

530. Pinus pumi!a, xylem TLS x 150, showing axial paren-

chyma, and one ray with a resin canal.
531. Pseudo!arix amabi!is, xylem TLS x 60.

223
532. Pseudotsuga menziesii, xylem TLS x 150, showing
short. uniseriate rays, tracheids with spiral thickenings,
and axial parenchyma strands containing starch granules.
533. Abies grandis, xylem RLS x 380, showing ray cells
with mainly taxodioid cross-field pitting and abundant
starch granules.

534. Cedrus atlantica, xylem RLS x 380, showing large

bordered pits on radial walls of tracheids and ray cells with
cupressoid cross-field pitting .
535. Larix sp ., xylem RLS x 380, showing rays with
piceoid cross-field pitting .

224
536. Pinus cembra, xylem RLS x 150, showing tracheids
with one or two rows of large bordered pits on radial walls
and ray cells, each with one or two large window cross-
field pits.
537. Pseudolarix amabilis, xylem RLS x 150, showing
tracheids with two rows of large bordered pits, ray cells
with taxodioid cross-field pits and some axial parenchyma 538. Pseudotsuga menziesii, xylem RLS x 380, showing
and ray cells filled with tannin. tracheids with spiral thickenings and rays with piceoid
cross-field pitting.
539. Tsuga canadensis, xylem RLS x 380, showing rays
with cupressoid cross-field pitting .

225
TAXUS Taxaceae

Bark
Peridenn narrow. Cortex parenchymatous. Phloem consist-
ing of alternating layers of thin-walled rounded (in TS)
parenchyma cells and thin-walled crystalliferous sieve
elements, with occasional solitary thick-walled fibres
arranged in tangential rows with several cells between
them. Crystals: sieve elements containing numerous
minute crystals lining inside of walls.

Xylem
TS. Growth rings present. Axial parenchyma sparse or
absent. Rays uniseriate. Resin canals absent.

LS. Tracheids with 1-3 rows of bordered pits mainly on ;.;';.!~..t..,,:~·~tj~''''~~~!!"":;~k~~~~~~rti~~~~~~~

radial walls and in opposite arrangement; cross-field pit- ~
ting cupressoid; characteristic spiral thickenings present
on tracheid walls. Rays uniseriate; radial resin canals
absent.

SPECIES EXAMINED

Taxus baccata L., Yew.

540. Taxus baccata, bark TS x 60, showing two uniseriate

bands of fibres in phloem.

226
 '""

-'
•
. ,.I •

• I;,

.'. .
;'.
. .
.
I ~
~

541 . ~: tI
541. Taxus baccata, xylem TLS x 150, showing tracheids
with spiral thickenings and short, uniseriate rays .
542. Taxus baccata, xylem RLS x 150, showing tracheids
with one or two rows of large bordered pits on radial
walls.

543. Taxus baccata, xylem RLS x 240, showing tracheids

with spiral thickenings and rays with cupressoid cross-field
pitting .

227
TAXODIACEAE
Cryptomeria, Sequoia, Sequoiadendron.

Bark
Periderm narrow, irregular, consisting of thin-walled cells,
sometimes in more than one layer, and often deep-seated
so cortex sloughed off. Cortex absent or narrow, paren-
chymatous. Phloem consisting of tangential layers of one
row of thick-walled fibres alternating with several rows of
thin-walled cells, the thin-walled cells consisting of sieve
cells and rounded parenchyma cells; resin canals absent;
phloem rays dilating slightly towards cortex. Crystals
absent.

Xylem
TS. Growth rings present. Axial parenchyma present,
sparse. Rays uniseriate. Resin canals absent.

LS. Tracheids with 1-2 rows of bordered pits mainly on

radial walls and in opposite arrangement; cross-field pit-
ting mainly taxodioid, some pits cupressoid in Sequoiaden-
dron . Rays uniseriate; radial resin canals absent.

SPECIES EXAMINED

Cryptomeria japonica D. Don, Japanese Red Cedar; Sequoia

sempervirens End!., Coast Redwood; Sequoiadendron gigan-
teum Buchholz, Wellingtonia.

546. Sequoiadendron giganteum, bark and xylem TS x

60, showing tangential lines of thick-walled fibres in
phloem.

544. Crypromeriajaponica, bark and xylem TS x 50,

showing tangential lines of thick-walled fibres in phloem.

228
546
546. Sequoia sempervirens, xylem TLS x 60.
547. Sequoia sempervirens, bark and xylem RLS x 60,
showing tracheids with double rows of large bordered
pits .

548. Cryptomeriajaponica, xylem RLS x 380, showing

rays with taxodioid cross-field pitting.
549. Sequoiadendron giganreum, xylem RLS x 380,
showing rays with taxodioid and cupressoid cross-field
pitting .

229
Appendix
 APPENDIX

Tablet Diagnostic characters in root anatomy of Dicotyledon trees. Brackets denote variable character.
Key: +, present; - absent; c, cluster crystals (druses); s, solitary (prismatic) crystals; sand, crystal sand; A, alternate
pitting; 0, opposite pitting; Sc, scalariform pitting; E, elongated pitting (intermediate between polygonal and
scalariform); Agg., aggregate rays.

Bark Xylem

Fibres Stone Crystals Secretory Vessels Ray width Vessel Perforation Spirals Page
cells cells! mainly <number of pitting plates reference
canals solitary cells) scalariform

Acer (+) + s 1-7 A + 10

Aesculus + + c, s 1 A + 12
Ailanthus + c, s 1-4 A 14
Alnus + + c, s 1(-3), Agg 0, fine + 16
Arbutus + 1-3 A, (E) 18
Betula + + c, s 1-3 A, minute + 24
Buxus c, s (+) 1-3 0, fine + 30
Carpinus + + c, s 1-4 (5), A (+) 34
Agg
Castanea + (+) s + 1 A 36
Catalpa + 1-2 (3) A 38
Corylus + + c, s 1-4, Agg A, (0) + (+) 48
Crataegus + + s + 1-3 A (+) 144
Eucalyptus + s + + 1 (2) A, (E) 64
Fagus + + c, s 1-20 0, Sc (+) 68
Ficus + c, s + 1-5 A, (E) 70
Fraxinus + + sand 1-2 A, fine 76
Ilex (+) c, s, sand (+) 1-7 0, Sc + + 94
Juglans + + c 1-5 A 98
Laburnum + + s 1-12 A + 104
Liquidambar + (+) c + (+) 1-3 A, Sc + 112
Liriodendron + (+) 1-5 0, Sc + 114
Magnolia + (+) 1-3 Sc + 118
Malus + + s + 1-3 A (+) 144
Morus + c, s 1-6 A, (E) 122
Nothofagus + + s 1-2 (3) 0, E, Sc + 124
Parrotia + + s (+) 1-3 O,E, Sc + 130
Platanus + s (+) (1) 4-14 0, E, Sc + 140
Populus + (+) c, s 1 (2) A 150
Prunus + + c, (s) 1-8 A, fine + 154
Pterocarya + c 1-3 A 156
Pyrus + + s + 1-3 A (+) 144
Quercus + + c, s + 1,10-40 A, (E) 158
Robinia + + s 1-6 A, (E) + 168
Salix + (+) c, s 1 (2) A 150
Sophora + s (+) 1-4 A, (E) (+) 184
Sorbus + + s + 1-3 A (+) 144
Tilia + c, (s) 1-6 A, fine + 196
Ulmus + s + 1-6 A + 198
Zelkava + + s 1-6 A + 208

233
 APPENDIX

Table 2 Diagnostic characters in root anatomy of Dicotyledon shrubs and climbers. Brackets denote variable
character.
Key: +, present; - absent; R, raphides; c, cluster crystals (druses); s, solitary (prismatic) crystals; sand, crystal sand;
A, alternate pitting; 0, opposite pitting; Sc, scalariform pitting; E, elongated pitting (intermediate between poly-
gonal and scalariform); Agg., aggregate rays.

Bark Xylem
Fibres Stone Crystals Secretory Vessels Ray width Vessel Perforation Spirals Page
cells cells/ mainly (number of pitting plates reference
canals solitary cells) scalariform
Abelia (c) (+) 1-7 (A), 0, E, +, (-) + 8
(Sc)
Ampelopsis + (+) c, s, R + 1-20 0, Sc (+) + 202
Aucuba sand 1-10 A,O, Sc + + 20
Berberis (+) s 1--6 A, fine + 22
Bignonia + + c + 1-20+ A 26
Buddleja 1--6 A, (E) + 28
Camellia (-) (-) (s) + 1-4 E, (Sc) + 32
Ceanothus + + c, s 1-3 A 40
Celastrus (+) c 1-11 A, (E) + 42
Chaenomeles + + s + 1-3 (+) A (+) 144
Clematis + 1-15+ A, fine 44
Comus (+) s 1-4 O,E + 46
Cotinus c, s + 1-2 (3) A 50
Cotoneaster + + s + 1-3 (+) A (+) 144
Cydonia + + s + 1-3 (+) A (+) 144
Cytisus + 1-11 A + 52
Daphne + + c, s 1-4 A + 54
Deutzia + + 1--6 O,E + + 56
Dieruilla (+) (+) (c), (s) 1-4 (7) (A), 0, E, + (+) 58
(Sc)
Elaeagnus + c 1-5 A, fine 60
Escallonia + (+) 1-3 O,E + 62
Euonymus (+) c 1-2 0, fine + 66
Forestiera + 1-2 (3) A, fine 72
Forsythia + (+) 1-3 A, fine (+) + 74
Fuchsia R 1-4 A, (E) 78
Garrya + + + 1--6 (15) A, fine + (+) 80
Hamamelis + s, c 1-3 0, Sc + 82
Hebe + + absent A, (E) 84
Hedera + (+) c + 1-10 0, A, (E) 86
Hibiscus + c 1-4 A, fine + 88
Hydrangea R 1--6 0, Sc + 90
Hypericum + 1-4 A, (E) 92
Jasminum (+) + (+) 1-3 A, fine + 96
Kerria + c 1--6 (10) A, E, (Sc) 100
Kolkwitzia (+) 1-5 A,O,E + + 102
Laurus (+) + 1-5+ A, (E) 106
Lavandula + s, c + 1-3 A, fine (+) 108
Ligustrum (+) (+) (s) (+) 1-3 A, (E) (+) 110
Lonicera + c 1--6 A, (E), fine (+) (+) 116
Mahonia + (s) 1-5 A, fine + 120
Olearia + 1-3 (4) A, (E) (+) 126
Osmanthus + 1-2 (3) A, fine + 128
Parthenocissus + (+) s, c, R + 1-20 0, Sc (+) + 202
Passiflora + + c (+) 1-8 A, (E) 132
Philadelphus + 1-4 A, 0, fine + 134
Phillyrea + 1-2 (3) A, fine + 128

(continued)
234
 APPENDIX

Table 2-continued
Bark Xylem
Fibres Stone Crystals Secretory Vessels Ray width Vessel Perforation Spirals Page
cells cells/ mainly (number of pitting plates reference
canals solitary cells) sealariform
Pieris + + s 1-3 O,E + 136
Pittosporum s + 1-4 A, fine + 138
Polygonum + c, s 1-20 A 142
Potentilla + s + 1-3 A 152
Pymcantha + + s + 1-3(+) A (+) 144
Rhamnus + (-) c, s 1-2 A + 160
Rhododendron + 1-6 O,E + 162
Rhus c + 1-3 (+) A, (E) + 164
Ribes (c) + 1-10 A, 0, fine + 166
Rosa + s + 1-25+ A, fine + 170
Rosmarinus + c (+) 1 (-3) A 172
Rubus + c, s 1-12 A, fine 174
Sambucus + sand 1-4 (6) A, (E) 176
Sarcococca + (+) 1-4 0, E, Sc + 178
Senecio + + 1-4 A, fine 180
Skimmia + s 1-5 A 182
Spimea + + c, s (+) 1-7 A, fine 186
Stymx + (+) c, s 1-5 A, 0, (E), + 188
fine
Symphoricarpos + (-) c (+) 1-5 (10) A, (0), (+) + 190
(E), (Sc)
Syringa + + (+) 1-3 A,(E) + 192
Tamarix + 1-10 A, fine 194
Viburnum (+) (+) c, s (+) 1-4 A, 0, (E) + (+) 200
Vitis + (+) s, c, R + 1-20 O,Sc (+) + 202
Weigela (+) (c) 1-4 (7) (A), 0, E, + (+) 58
(Sc)
Wisteria + s 1-14 A + 206

235
 APPENDIX

Table 3 Main differences in root wood anatomy in four related genera.

Genus Ray width Pitting Perforation plates

(number of cells>

Alnus 1-(3) + aggregate fine, opposite scalariform, with 10-20 bars

Betula 1-3 minute, alternate scalariform, with 5-15 bars
Carpinus 1-5 + aggregate coarse, alternate simple ,
Corylus 1-4 + aggregate medium, alternate (+ opposite) scalariform, with 3-10 bars

Table 4 Diagnostic characters in root anatomy of the family Caprifoliaceae. Brackets denote variable character.
Key: +, present; - absent; c, cluster crystals; s, solitary (prismatic) crystals; vessel pitting - A, alternate; 0, op-
posite; E, elongated pitting (intermediate between polygonal and scalariform); Sc, scalariform; RP, ring porous;
SRP, semi-ring porous; DP, diffuse porous; Fibres - Tk, thick-walled; Tn, thin-walled; Stone cells - L, large;
S, small.

Bark Xylem

Genus Number Phloem Phloem Crystals RPI Vessel Vessel Simple Scalari- Number Fibres, Fibres Ray Rays
of fibres stone SRPI pitting spirals perfor- form of thickl with width with
periderms cells DP ation perfor- bars thin spirals sheath
plates ation walled cells
plates

Abelia 2+ (e) DP (A),O, (+) (+) (+) 10-30 Tk + 1-7 +

E, (Se)
Dieruillal 1,2+ (+) (5) (e), (s) DP (A),O, + 25-40+ Tn, (Tk) (+) 1-4 (7) (+)
Weigela E, (Se)
Kolkwitzia 2+ DP A,O, + 10-25 Tk + 1-5 (+)
E, (Se)
Lonicera 1 + e SRP, DP A, (E) (+) + (+) 1-30 Tk (+) 1-6 (+)
Symphoricarpos 1 + c RP, SRP A, (0), + (+) <25 Tk (+) 1-5 (10) +
(E), (Sc)
Viburnum 1 (+) (L) c, (s) DP (A),O, + 15-30+ Tk (+) 1-4 (+)
E, (Sc)

236
 APPENDIX

TableS Diagnostic characters in root anatomy of Gymnosperm trees. Brackets denote variable character.
Key: +, present; -, absent; As, astrosclereids; s, solitary crystals; c, cluster crystals (druses); M, mucilage sacs; A,
alternate pitting; 0, opposite pitting; Cu, cupressoid pitting; P, piceoid pitting; T, taxodioid pitting.

Bark Xylem

Fibres Stone Crystals Resin Resin canals Tracheids

cells canals
Axial Radial Bordered Cross- Spirals Page
pits field reference
pits

Abies5 + As M (+) 0 T 220

Araucaria4 + As s + A Cu 212
Cedrus 7 +, As s 0 CU,P,T 220
Chamaecyparis 6 + + 0 Cu, (T) 214
Cryptomeria 6 + 0 T 228
x Cupressocyparis6 + + 0 Cu 214
Cupressus6 + + 0 Cu 214
Ginkgo3 + (+) c 0 Cu 218
Juniperus 6 + s + 0 Cu 214
Larix + + (-) (-) + + 0 P 220
Picea + s + + + 0 P 220
Pinus 2 s + + + 0 large 220
Pseudolarix5 + M 0 T 220
Pseudotsuga 1 + As + + 0 P + 220
Sequoia6 + 0 T 228
Sequoiadendron 6 + 0 T, (Cu) 228
Taxus 1 + s 0 Cu + 226
Thuja 6 + s + 0 Cu 214
Tsuga + As 0 Cu 220
1 Spiral thickenings on tracheids in Taxus and Pseudotsuga; these two readily distinguishable from each other by presence of
resin canals in xylem of Pseudotsuga.
2 Large cross-field pitting in Pinus.
3 Cluster crystals in bark of Ginkgo.
4 In Araucaria, alternate inter-tracheid pitting, and astrosclereids with associated crystals in bark.
S Abies bark very mucilaginous (mucilage also present in Pseudolarix).
6 Genera of Cupressaceae and Taxodiaceae not easily distinguishable, except in some cases by cross-field pitting and certain
features of bark.
7 Bordered pits on tracheid walls occasionally with scalloped tori in Cedrus; however this character not reliable, particularly
in partly decomposed material.

237
 Index of scientific names

Abelia 8, 234, 236 pendula 24

chinensis 8 sp. 24, 25
grabneriana 8 Betulaceae 16, 24
schumannii 8 Bignonia 26, 234
spathulata 8, 9 capreolata 26, 27
triflora 8, 9 Bignoniaceae 26, 38
umbellata 8 Buddleja 28, 234
Abies 220, 237 davidii 28, 29
grandis 220, 222, 224 Buddlejaceae 28
sp.220 Buxaceae 30, 178
Acerl0, 233 Buxus 30, 233
palma tum 10 sempervirens 30,31
platanoides 10, 11
pseudoplatanus 10 Camellia 32, 234
Aceraceae 10 cuspidata 32, 33
Aesculus 12, 233 japonica 32, 33
hippocastanum 12, 13 Caprifoliaceae 8, 58, 102, 116, 190, 200,
Ailanthus 14, 233 236
altissima 14, 15 Carpinaceae 34
Alnus 16, 233, 236 Carpinus 34, 233, 236
glutinosa 16, 17 betulus 34, 35
Anacardiaceae 50, 164 Castanea 36, 233
Altingiaceae 112 sativa 36, 37
Aquifoliaceae 94 Catalpa 38, 233
Ampelopsis 202, 234 speciosa 38, 39
arborea 202, 203, 204 Ceanothus 40, 234
brevipedunculata 203 americanus 40, 41
Araliaceae 86 arboreus 40,41
Araucaria 212, 237 griseus 40
araucana 212, 213 impressus 40,41
bidwillii 212, 213 masonii 40
Arancariaceae 212 Cedrus 220, 237
Arbutus 18, 233 atlantica 220, 222, 224
andrachne 18, 19 Celastraceae 42, 66
Aucuba 20,234 Celastrus 42, 234
japonica 20, 21 hookeri 42, 43
Aucubaceae 20 monospermus 42
Chaenomeles 144, 234
Berberidaceae 22, 120 japonica 144, 146, 148
Berberis 22, 234 Chamaecyparis 214,237
x stenophylla 22 lawsoniana 214
vulgaris 22, 23 nootkatensis 214,215, 216
Betula 4, 24, 233, 236 thyoides 214,215

239
 INDEX OF SCIENTIFIC NAMES

Clematis 44, 234 rivularis 58

x jackmanii 44, 45 sessilifolia 58
montana 44, 45 x splendens 58
patens 44 Elaeagnaceae 60
sp.44 Elaeagnus 60, 234
Compositae 126,180 pungens 60, 61
Comaceae 46 umbellata 60, 61
Comus 46, 234 Ericaceae 18, 136, 162
amomum 46, 47 Escallonia 62, 234
australis 46 viscosa 62, 63
chinensis 46 Escalloniaceae 62
controversa 46,47 Eucalyptus 64, 233
macrophylla 46 coccifera 64, 65
nuttallii 46 gunnii 64
officinalis 46 parvifolia 64
sanguinea 46 Euonymus 66, 234
sp.46 europaeus 66, 67
walteri 46 japonicus 66, 67
Corylaceae 48
Corylus, 48, 233, 236 Fagaceae 36, 68, 124, 158
avellana 48,49 Fagus 68, 233
Cotinus 50,234 sylvatica 68, 69
obovatus x coggyria 50,51 Ficus 70,233
Cotoneaster 144,234 carica 70, 71
horizontalis 144, 146 Forestiera 72, 234
lacteus 144, 149 sp. 72, 73
salicifolius 144 Forsythia 74, 234
Crataegus 144, 233 x intermedia 74,75
x lavallei 144, 147 suspensa 74, 75
monogyna 144 Fraxinus 76, 233
oxycanthoides 144 excelsior 76, 77
Cryptomeria 228, 237 Fuchsia 78, 234
japonica 228, 229 magellanica 78, 79
Cupressaceae 214,237 sp.78
x Cupressocyparis 214,237
leylandii 214
Cupressus 214, 237 Garrya 80, 234
macrocarpa 214,215 elliptica 80, 81
Cydonia 144, 234 Garryaceae 80
sp. 144, 145, 147 Ginkgo 2, 218, 237
Cytisus 52, 234 biloba 218, 219
hirsutus 52, 53 Ginkgoaceae 218
multiflorus 52, 53 Grossulariaceae 166
purgans 52 Guttiferae 92
scoparius 52
sp. 52, 53 Hamamelidaceae 82, 130
Hamamelis 82, 234
Daphne 54, 234 virginiana 82, 83
mezereum 54, 55 Hebe 84,234
Deutzia 56, 234 speciosa 84, 85
x maliflora 56, 57 Hedera 86,234
scabra 56, 57 helix 86,87
sp.56 Hibiscus 88, 234
Diervilla 58, 234, 236 syriacus 88, 89
lonicera 58 Hippocastanaceae 12

240
 INDEX OF SCIENTIFIC NAMES

Hydrangea 90, 234 fragrantissima 116

paniculata 90, 91 nitida 116, 117
sp. 90, 91 orientalis 116
Hypericum 92, 234 prolifera 116
androsaemum 92
x inodorum 92, 93 Magnolia 118, 233
sp. 92, 93 campbellii 118, 119
denudata 118,119
Ilex94,233 sieboldii 118, 119
aquifolium 94, 95 x soulangiana 118
sp.94 Magnoliaceae 114,118
Mahonia 120, 234
Jasminum 96, 234 aquifolium 120,121
humile 96, 97 japonica 120
nudiflorum 96 Malus 144,233
officinale 96, 97 pumila 144, 145
x stephanense 96 x purpurea 144
sp. 96, 97 toringoides 144
Juglandaceae 98, 156 Malvaceae 88
Juglaris 98, 233 Moraceae 70, 122
nigra 98 Morus 122, 233
regia 98,99 alba 122
Juniperus 214, 237 nigra 122, 123
communis 214, 215, 216, 217 sp. 122, 123
Myrtaceae 64
Kerria 100,234
japonica 100, 101 Nothofagus 124, 233
Kolkwitzia 102, 234, 236 obliqua 124, 125
amabilis 102, 103
Oleaceae 72, 74, 76, 96, 110, 128, 192
Labiatae 108, 172 Olea ria 126,234
Laburnum 104, 233 x haastii 126, 127
alpinum 104 odorata 126, 127
anagyroides 104 Onagraceae 78
x watereri 104, 105 Osman thus 128, 234
Larix 220,237 x burkwoodii 128,129
decidua 220 heterophyllus 128, 129
sp.220,222,223,224 Parrotia 130, 233
Lauraceae 106 persica 130, 131
Laurus 106, 234 Parthenocissus 202, 234
nobilis 106, 107 inserta 202, 203, 205
Lavandula 108, 234 quinquefolia 203
sp. 108, 109 Passiflora 132, 234
Leguminosae 52, 104, 168, 184, 206 sp. 132, 133
Ligustnum 110, 234 Passifloraceae 132
vulgare 110, 111 Philadelphaceae 56, 134
Liquidambar 112, 233 Philadelphus 134, 234
styraciflua 112, 113 coronarius 134, 135
Liriodendron 114, 233 Phillyrea 128, 234
tulipifera 114, 115 latifolia 128
Lonicera 116, 234, 236 Picea 220, 237
x americana 116 sp. 220, 221, 223
caprifolium 116, 117 Pieris 136, 235
discolor 116, 117 formosa 136, 137
etrusca 116 japonica 136, 137

241
 INDEX OF SCIENTIFIC NAMES

Pinaceae 220 dentata 158

Pinus 220,237 ilex 158, 159
cembra 220,221,225 robur 158,159
pumila 220,223 sp. 158, 159
sp.220
sylvestris 220 Ranunculaceae 44
Pittosporaceae 138 Rhamnaceae 40, 160
Pittosporum 138, 235 Rhamnus 160, 235
tenuifolium 138, 139 cathartica 160, 161
tobira 138, 139 Rhododendron 162, 235
Platanaceae 140 calophytum 162
Platanus 140, 233 cataubiense x griffithianum 162
x acerifolia 140, 141 decorum 162
occidentalis 140 falconeri 162
sp. 140, 141 ponticum 162, 163
Polygonaceae 142 sp.162
Polygonum 142, 235 vemicosum 162, 163
baldschuanicum 142, 143 Rhus 164, 235
Pomoideae 2, 144 sp. 164, 165
Populus 2, ISO, 233 Ribes 166, 235
alba 150 nigrum 166
balsamifera 150 sanguineum 166
x canescens 150 uva-crispi 166, 167
nigra 150 Robinia 168, 233
tremula 150,151 pseudoacacia 168, 169
trichocarpa 150 Rosa 170, 235
Potentilla 152, 235 canina 170
fruticosa 152, 153 sp. 170, 171
Prunus 154, 233 Rosaceae 2, 100, 144, 152, 154, 170, 174, 186
avium 154, 155 Rosmarinus 172, 235
cerasifera 154 officinalis 172, 173
cerasus 154, 155 Rubus 174, 235
communis 154 sp. 174, 175
domestica 154 Rutaceae 182
insititia 154, 155
laurocerasus 154 Salicaceae 2, 150
lusitanica 154 Salix 2,150,233
persica 154 alba 150
serrulata 154 babylonica 150
sp. 154 sp. 150, 151
spinosa 154 Sambucaceae 176
Pseudolarix 220, 237 Sambucus 176, 235
amabilis 220, 221, 223, 225 nigra 176, 177
Pseudotsuga 220, 237 Sarcococca 178, 235
menziesii 220, 222, 224, 225 ruscifolia 178, 179
Pterocarya 156, 233 Scrophulariaceae 84
x rehderiana 156, 157 Senecio 180, 235
Pyracantha 144, 235 greyi 180,181
coccinea 144 Sequoia 228, 237
sp. 144, 145 sempervirens 228,229
Pyrus 144, 233 Sequoiadendron 228, 237
amygdaliformis 144 giganteum 228, 229
communis 144, 147, 148 Simaroubaceae 14
Skimmia 182, 235
Quercus 158,233 japonica 182, 183

242
 INDEX OF SCIENTIFIC NAMES

Sophora 184, 233 Tilia 196, 233

affinis 184, 185 x eu ropaea 196
flavescens 184 sp. 196, 197
japonica 184, 185 Tiliaceae 196
Sorbus 144,233 Tsuga 220, 237
aria 144, 148 canadensis 220, 225
aucuparia 144
hybrida 144, 146, 149 Ulmaceae 198,208
intermedia 144 Ulmus 198, 233
Spiraea 186, 235 glabra 198
japonica 186, 187 procera 198, 199
x sanssouciana 186, 187 x sarniensis 198, 199
veitchii 186, 187 sp. 198, 199
Styracaceae 188 stricta 198
Styrax 188, 235
Viburnum 200, 235, 236
hemsleyana 188, 189
Symphoricarpos 190, 235, 236
x bodnantense 200
buddleifolium 200
albus 190, 191
farreri 200, 201
x chenaultii 190 foetens 200
occidentalis 190, 191
lantana 200
rotundifolius 190
rhytidophyllum 200
Syringa 192, 235
tinus 200
villosa 192, 193
Vitaceae 202
vulgaris 192
Vitis 202, 235
yunnanensis 192
coignetiae 203, 204, 205
sp.192
sp.203
vinifera 203, 204, 205
Tamaricaceae 194
Tamarix 194, 235 Weigela 58, 235, 236
sp. 194, 195 decora 58, 59
Taxaceae 226 floribunda 58
Taxodiaceae 228, 237 florida 58
Taxus 226,237 praecox 58, 59
baccata 226, 227 Wisteria 206, 235
Theaceae 32 floribunda 206, 207
Thymelaeaceae 54 sp. 206, 207
Thuja 214,237
plicata 214, 215, 217 Zelkova 208, 233
standishii 214, 217 carpinifolia 208, 209

243
 Index of common names

Acacia, False 168 Cypress, Lawson 214

Alder 16 Cypress, Leyland 214
Arolla Pine 220 Cypress, Monterey 214
Ash 76 Cypress, Nootka 214
Ash, Mountain 144
Atlas Cedar 220 Daisy Bush 126
Dogwood 46
Barberry 22 Douglas Fir 220
Bay Laurel 106
Beauty Bush 102 Eastern Hemlock 220
Beech 68 Elder 176
Beech, Southern 124 Elm 198
Birch 24 Elm, Caucasian 208
Blackberry 174 European Larch 220
Blackcurrant 166
Blackthorn 154 False Acacia 168
Boston Ivy 203 Fig 70
Box 30 Fir, Douglas 220
Box, Sweet 178 Fir, Grand 220
Broom 52 Firethorn 144
Buckthorn 160 Flowering Currant 166
Buddleja 28
Golden Larch 220
Californian Lilac 40 Gooseberry 166
Camellia 32 Grand Fir 220
Catalpa, Western 38
Gum, Sweet 112
Caucasian Elm 208
Gum Tree 64
Cedar, Atlas 220
Cedar, Japanese Red 220
Cedar, Western Red 214 Hawthorn 144
Cherry, Japanese 154 Hazel 48
Cherry Plum 154 Hemlock, Eastern 220
Cherry, Sour 154 Holly 94
Cherry, Wild 154 Honeysuckle 116
Cherry-Laurel 154 Hornbeam 34
Chestnut, Horse 12 Horse Chestnut 12
Chestnut, Sweet 36
Chinese Quince 144 Ivy 86
Cinquefoil, Shrubby 152 Ivy, Boston 203
Clematis 44
Coast Redwood 228 Japanese Quince 144
Cross Vine 26 Japanese Red Cedar 228
Currant, Flowering 166 Jasmine %

244
 INDEX OF COMMON NAMES

Juniper, Common 214 Redwood, Coast 228

Red Cedar, Japanese 228
Laburnum 104 Rose 170
Larch, European 220 Rosemary 172
Larch, Golden 220 Rowan 144
Laurel, Bay 106
Laurel, Portugal 154 Scots Pine 220
Lavender 108 Shrubby Cinquefoil 152
Lawson Cypress 214 Smoke Tree 50
Leyland Cypress 214 Snowberry 190
Lilac 192 Sour Cherry 154
Lilac, Californian 40 Southern Beech 124
Lime 196 Spindle Tree 66
Locust Tree 168 Spruce 220
Storax 188
Sumach, Venetian 50
Maidenhair Tree 218
Sweet Box 178
Maple, Norway 10
Sweet Chestnut 36
Mock Orange 134
Sweet Gum 112
Monkey Puzzel Tree 212
Sycamore 10
Monterey Cypress 214
Mountain Ash 144
Tamarisk 194
Mulberry 112
Tree of Heaven 14
Tulip Tree 114
Nootka Cypress 214
Venetian Sumach 50
Oak 36, 156 Vine 203
Oleaster 60 Virginia Creeper 203
Oregon Grape 120
Walnut 98
Passion Flower 132 Wayfaring Tree 200
Peach 154 Wellingtonia 228
Pear 144 Western Catalpa 38
Plane 140 Western Red Cedar 214
Plum 154 Whitebeam 144
Plum, Cherry 154 Wild Cherry 154
Poplar 150 Willow 150
Portugal Laurel 154 Wing-Nut 156
Privet 110 Witch Hazel 82
Wolfberry 190
Quince 144
Quince, Chinese 144 Yew 226

245