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Biology of Northern Krill 1st ed Edition Tarling Digital
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Author(s): Tarling, Geraint A
ISBN(s): 9780123813084, 0123813085
Edition: 1st ed
File Details: PDF, 10.83 MB
Year: 2010
Language: english
Advances in MARINE BIOLOGY
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University of Texas at Austin
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Oregon Institute of Marine Biology
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University of Connecticut
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10 11 12 10 9 8 7 6 5 4 3 2 1
CONTRIBUTORS TO VOLUME 57
Cornelia Buchholz
Alfred Wegener Institute for Polar and Marine Research, Bremerhaven, Germany
Friedrich Buchholz
Janine Cuzin-Roudy
Observatoire Océanologique, Université Pierre et Marie Curie (Paris)-CNRS,
Villefranche-sur-Mer, France
Natalie S. Ensor
British Antarctic Survey, Natural Environment Research Council, High Cross,
Cambridge, United Kingdom
Torsten Fregin
Universität Hamburg, Zoologisches Institut, Hamburg, Germany
Peter Fretwell
Edward Gaten
Department of Biology, University of Leicester, Leicester, United Kingdom
William P. Goodall-Copestake
Michel Harvey
Maurice Lamontagne Institute, Fisheries and Oceans Canada, Mont-Joli, Québec,
Canada
Magnus L. Johnson
Centre for Environmental and Marine Sciences, University of Hull, Scarborough,
United Kingdom
Stein Kaartvedt
King Abdullah University of Science and Technology, Thuwal, Saudi Arabia
Chiara Papetti
Department of Biology, University of Padova, Italy
v
vi Contributors
Tomaso Patarnello
Department of Public Health, Comparative Pathology, and Veterinary Hygiene,
University of Padova, Italy
Reinhard Saborowski
Katrin Schmidt
Yvan Simard
Maurice Lamontagne Institute, Fisheries and Oceans Canada, Mont-Joli, Québec,
Canada and Marine Science Institute, University of Québec at Rimouski,
Rimouski, Québec, Canada
John I. Spicer
Marine Biology and Ecology Research Centre, School of Marine Sciences and
Engineering, University of Plymouth, Plymouth, United Kingdom
Geraint A. Tarling
Konrad Wiesew
Lorenzo Zane
Department of Biology, University of Padova, Italy
w
Deceased
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Brown, A. C. and Odendaal, F. J. The biology of oniscid isopoda of
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Braithwaite, R. A. and McEvoy, L. A. Marine biofouling on fish
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Collins, M. A. and Rodhouse, P. G. K. Southern ocean cephalopods.
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Garcia E. G. The Northern Shrimp (Pandalus borealis) Offshore
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Kerry A. Naish, Joseph E. Taylor III, Phillip S. Levin, Thomas P.
Quinn, James R. Winton, Daniel Huppert, and Ray Hilborn. An
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Hatcheries on Wild Populations of Salmon. pp. 61–194.
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Bridget S. Green. Maternal Effects in Fish Populations. pp. 1–105.
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Mechanisms and Conservation Implications. pp. 107–170.
David W. Sims. Sieving a Living: A Review of the Biology, Ecology
and Conservation Status of the Plankton-Feeding Basking Shark
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Charles H. Peterson, Kenneth W. Able, Christin Frieswyk DeJong,
Michael F. Piehler, Charles A. Simenstad, and Joy B. Zedler.
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Volume 55, 2008.
Annie Mercier and Jean-François Hamel. Introduction. pp. 1–6.
Annie Mercier and Jean-François Hamel. Gametogenesis. pp. 7–72.
Annie Mercier and Jean-François Hamel. Spawning. pp. 73–168.
Annie Mercier and Jean-François Hamel. Discussion. pp. 169–194.
Volume 56, 2009.
Philip C. Reid, Astrid C. Fischer, Emily Lewis-Brown, Michael
P. Meredith, Mike Sparrow, Andreas J. Andersson, Avan Antia,
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Brix, Stephen Dye, Martin Edwards, Tore Furevik, Reidun Gangst,
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Ralph Keeling, Corinne Le Quéré, Fred T. Mackenzie, Gill Malin,
Cecilie Mauritzen, Jón Ólafsson, Charlie Paull, Eric Rignot, Koji
Shimada, Meike Vogt, Craig Wallace, Zhaomin Wang and Richard
Washington. Impacts of the Oceans on Climate Change. pp. 1–150.
Elvira S. Poloczanska, Colin J. Limpus and Graeme C. Hays.
Vulnerability of Marine Turtles to Climate Change. pp. 151–212.
Nova Mieszkowska, Martin J. Genner, Stephen J. Hawkins and David
W. Sims. Effects of Climate Change and Commercial Fishing on
Atlantic Cod Gadus morhua. pp. 213–274.
Iain C. Field, Mark G. Meekan, Rik C. Buckworth and Corey J. A.
Bradshaw. Susceptibility of Sharks, Rays and Chimaeras to Global
Extinction. pp. 275–364.
Milagros Penela-Arenaz, Juan Bellas and Elsa Vázquez. Effects of the
Prestige Oil Spill on the Biota of NW Spain: 5 Years of Learning.
pp. 365–396.
C H A P T E R O N E
An Introduction to the Biology of

Northern Krill (Meganyctiphanes
norvegica Sars)
Geraint A. Tarling,* Natalie S. Ensor,* Torsten Fregin,†
William P. Goodall-Copestake,* and Peter Fretwell*
Contents
1. Introduction 2
2. Morphology and Taxonomy 3
2.1. General morphology 3
2.2. Taxonomy 6
2.3. Phylogeny 7
2.4. Photophores and their function 10
3. Geographic Distribution 13
3.1. General remarks 13
3.2. North Atlantic west coast 15
3.3. Arctic limits 15
3.4. North-east Atlantic and fringes 16
3.5. Mediterranean and south-eastern limits 17
3.6. Distributional trends and changes 17
4. The Sampling of Northern Krill 18
4.1. Net sampling 19
4.2. Active acoustics 23
4.3. Alternative technologies for sampling and observation 25
5. Ecological Role 26
5.1. Benthic interactions 26
5.2. Krill production and predator consumption 27
6. Commercial Exploitation of Northern Krill 28
Acknowledgements 30
References 30
* British Antarctic Survey, Natural Environment Research Council, High Cross, Cambridge, United
Kingdom
{
Universität Hamburg, Zoologisches Institut, Hamburg, Germany
Advances in Marine Biology, Volume 57 # 2010 Elsevier Ltd.

ISSN 0065-2881, DOI: 10.1016/S0065-2881(10)57001-9 All rights reserved.
1
2 Geraint A. Tarling et al.
Abstract
This chapter provides a background to research on Northern krill biology,
starting with a description of its morphology and identifying features, and the
historical path to its eventual position as a single-species genus. There is a lack
of any euphausiid fossil material, so phylogenetic analysis has relied on com-
parative morphology and ontogeny and, more recently, genetic methods.
Although details differ, the consensus of these approaches is that Meganycti-
phanes is most closely related to the genus Thysanoessa. The light organs (or
photophores) are well developed in Northern krill and the control of lumine-
scence in these organs is described. A consideration of the distribution of
the species shows that it principally occupies shelf and slope waters of both
the western and eastern coasts of the North Atlantic, with a southern limit at the
boundary with sub-tropical waters (plus parts of the Mediterranean) and a
northern limit at the boundary with Arctic water masses. Recent evidence of a
northward expansion of these distributional limits is considered further. There
have been a variety of techniques used to sample and survey Northern krill
populations for a variety of purposes, which this chapter collates and assesses
in terms of their effectiveness. Northern krill play an important ecological role,
both as a contributor to the carbon pump through the transport of faecal
material to the deeper layers, and as a key prey item for groundfish, squid,
baleen whales, and seabirds. The commercial exploitation of Northern krill has
been slow to emerge since its potential was considered by Mauchline [Mauchline,
J (1980). The biology of mysids and euphausiids. Adv. Mar. Biol. 18, 1–681].
However, new uses for products derived from krill are currently being found,
which may lead to a new wave of exploitation.
1. Introduction
In 1960, John Mauchline published a seminal work entitled ‘The
biology of the Euphausiid Crustacean, Meganyctiphanes norvegica (M. Sars)’,
which provided detailed descriptions of a number of characteristic traits of
Northern krill, including diel vertical migration (DVM), feeding, growth
mortality rates, maturation and the function of the luminescent organs. The
study was carried out in the semi-enclosed Scottish loch, Loch Fyne (Clyde
Sea), which offered ready sampling-access and the capacity to follow a
population reliably over time (2 years in total). The work was a major
step forward in the understanding of ecology and behaviour of this species,
to add to what was already known in terms of its distribution (Einarsson,
1945; Ruud, 1936) and feeding habits (Fisher and Goldie, 1959;
Macdonald, 1927). Mauchline went on to publish the first major review
of euphausiids, with Leonard Fisher, in 1969, which included sections on
vertical distribution and migration, food and feeding, and growth, maturity,
An Introduction to Northern Krill 3
and mortality that were mainly based on Mauchline (1960). Mauchline

updated this review in 1980.
Fifty years on from Mauchline (1960) and 30 years from the updated
review of euphausiid biology (Mauchline, 1980), we find that the field has
advanced considerably. Since 1980, more than 1000 articles have been
published with a euphausiid species name in the title and more than 3500
with a euphausiid species name in the Abstract or as a Keyword (Table 1.1).
Of these, around 90% have considered some physiological process, more
than 40%, abundance and biomass, and 10%, the role of krill as a prey
item. Furthermore, entire new fields have developed over this time, such as
population genetics and genomics. New acoustic- and video-enabled
instrumentation has allowed unprecedented insights into euphausiid beha-
vioural ecology. State-of-the-art acoustics can remotely identify aggrega-
tions of euphausiids and rapidly assess their distribution and biomass, from
local- to basin-scales. Physiological cycles have now been well described, as
has the capacity of Northern krill to cope with change in temperature and
levels of oxygen saturation.
Everson (2000a) reviewed some of these advances, from a krill (mainly
Euphausia superba) fisheries perspective. Nevertheless, there is still much
recent progress in the understanding of euphausiid biology to be synthe-
sised. A review of the biology of Northern krill, in particular, is timely given
the considerable change in its northern distributional limit. Arctic summer-ice
is now at a historical low and Atlantic species may potentially incur into higher
latitudes. Northern krill is being increasingly reported at latitudes as high as the
Spitsbergen fjords (78 560 N, 12 040 E, Buchholz et al., 2010). The potential
for the establishment of this species further north and the implications this may
have on the Arctic ecosystem can be fully considered only through the study of
all aspects of its biology which the present volume aims to cover.
As a background, we describe in the present chapter some fundamental
details of Northern krill biology. We begin with a description of the general
body plan, highlighting the species morphological characteristics. We then
review its phylogeny and geographic distribution. Sampling methodology is
considered in terms of the diversity of equipment used and its purpose.
Finally, we assess the wider ecological role and the commercial relevance of
Northern krill in the modern world.
2. Morphology and Taxonomy

2.1. General morphology
The morphology of M. norvegica (Northern krill) was first described by
M. Sars (1857) as follows:
Table 1.1 ISSN-referenced articles published on major euphausiid species/genera since 1980
Meganyctiphanes
norvegica Euphausia superba Euphausia pacifica All Thysanoessa spp.
1980– 1990– 2000– 1980– 1990– 2000– 1980– 1990– 2000– 1980– 1990– 2000–
1989 1999 2010 1989 1999 2010 1989 1999 2010 1989 1999 2010
Number of ISSN articles published
1. Number with respective 26 28 54 403 221 257 25 31 41 39 24 22
species name in title
2. Number with name in title, 86 164 323 599 696 1071 60 79 136 130 145 208
abstract or keywords
Percentage of ISSN articles dealing with the following specific disciplines
3. Genetics 12% 14% 13% 5% 6% 9% 8% 3% 2% 10% 8% 5%
4. Population dynamics 27% 68% 46% 15% 32% 34% 4% 32% 56% 31% 54% 64%
5. Biomass and distribution 38% 54% 50% 20% 45% 50% 16% 52% 59% 49% 54% 41%
6. Physiology and process 92% 86% 96% 80% 85% 74% 92% 100% 90% 95% 92% 91%
studies
7. Behaviour 15% 54% 52% 8% 16% 18% 24% 32% 29% 10% 17% 23%
8. Krill predators 4% 7% 22% 2% 10% 13% 4% 10% 22% 0% 8% 0%
Audit carried out through applying search terms to an on-line scientific database (Web of Knowledge, wok.mimas.ac.uk). 1, involved a search for all articles in which the
species or common name was contained in the title. 2, species or common name in the title, abstract or keywords. 3–8, species or common name in the title and discipline
related key words in the title, abstract or keywords. Percentages relate to the number of articles relative to category 1. An article may fit more than one of the categories 3–8.
Whitish translucent, spotted above with red. Cephalothorax equalling a

third to three-eighths of the whole body in length; with the forehead very
short and truncated, not reaching the slender stalks of the very large eyes,
with the median anterior edge angular (rostrum rudimentary), with a sharp
spine on both sides of the triangle. . ..only seven pairs of gills with none on
the first pair of feet, very large on the hindmost. The gills consist of short
double curved branches, with a single row of simple tufts in the six anterior
pairs, covered with secondary tufts of pinnate embellishments in the last pair
(Translation from Latin; Mauchline and Fisher, 1969)
M. norvegica is one of the largest of the 86 described species of euphausiid,
and reaches a total body length of between 40 and 50 mm (Baker et al., 1990;
Falk-Petersen and Hopkins, 1981), which is surpassed only by a small number
of Thysanopoda species and E. superba (Antarctic krill). The basic body plan of
all euphausiids is relatively similar, being divided into two main regions—the
cephalothorax and the abdomen (Fig. 1.1). The cephalothorax contains the
head and thoracic segments and is completely covered by a carapace. It extends
for about one-third of the animal. The abdomen consists of six segments
terminated by a telson (Mauchline and Fisher, 1969). The head contains one
pair of stalked eyes and two pairs of antennae, which have tactile and olfactory
functions. Excretory organs open to the exterior at the bases of the second
antennae. The mouthparts consist of a single labrum, paired mandibles, labia,
maxillules, and maxillae which filter, macerate, and manipulate food to the
mouth. There are only seven pairs of biramous thoracic limbs, the eighth being
absent in Northern krill. The exopodites of the thoracic limbs produce
currents for the oxygenation of the external gills associated with the thoracic
limb bases. The endopodites of these limbs form a food basket that filters
particles out of the water. The first five abdominal segments each have one pair
of biramous pleopods, with which the animal swims. The sixth abdominal
segment bears one pair of biramous uropods. The uropods along with the
Cephalothorax Abdomen
Antennular flagellum
Eye
Antennal
scale Telson
Gills
Uropods
Exopodites
Thoracic limbs
Pleopods
Antennal flagellum
Figure 1.1 Adult Meganyctiphanes norvegica (total body length around 40 mm) indicat-
ing the main morphological features (Illustration by J. Corley).
telson form a ‘tail fin’. The first pair of abdominal limbs of the adult male
develops at maturity to form male organs called petasmae (Fig. 1.2). The
female organ for collecting sperm, the thelycum (Fig. 1.3), is located ventrally
on the sixth thoracic segment.
Euphausiids have two distinct spawning strategies (Gómez-Gutiérrez
et al., 2010); about 26 of the species are sac-spawners, where the just-spawned
eggs are held into an ovigerous sac supported by the eighth periopod until
they hatch. The others are broadcast-spawning species, where eggs are
released freely into the water column at the point of spawning. M. norvegica
belongs to the second of these categories. Euphausiids can also be divided
between those species with spherical eyes and those with bilobed eyes and
1 or 2 pairs of elongated thoracic limbs, of which M. norvegica belongs to the
former. Features that uniquely distinguish M. norvegica from other euphausiid
species have been provided by Baker et al. (1990) as follows: ‘Seventh
thoracic legs consisting of two elongated joints, with the seventh thoracic
exopod being present. Strong post-ocular spines and long recurved anten-
nular (first antennae) lappets are present’.
2.2. Taxonomy
M. norvegica was originally described by Michael Sars in 1857 (Sars 1857) as
Thysanopoda norvegicus, from four specimens from Florö and Söndfjord in
Norwegian waters and a further nine specimens from the guts of two Norway
0.5 mm
Figure 1.2 Petasma of an adult male Meganyctiphanes norvegica, which is a modification

of the internal ramus of the first pleopod (Illustration by J. Corley).
Figure 1.3 Thelycum of an adult female Meganyctiphanes norvegica with 2 attached

spermatophores. The view is from the ventral side of the cephalothorax (6th thoracic
sternite). The thelycum is approximately 1 mm across (Illustration by J. Corley).
haddocks (Sebastes norvegicus). The genus Nyctiphanes was erected by G. O Sars

report in 1883 (Sars, 1883) which reported on the Schizopoda collected during
the HMS Challenger expeditions of 1873–1876. In their report of the schizo-
poda of the north-east Atlantic slope, Holt and Tattersall (1905) decided that the
species differed sufficiently from the others of the genus Nyctiphanes to warrant
the erection of a separate genus Meganyctiphanes to contain it. Hansen (1908)
accepted Holt and Tattersall’s new genus Meganyctiphanes and put forward a
generic difference detectable in the males, namely the form of the clasping
organs (petasma) on the first pair of pleopods. Those of M. norvegica are
essentially similar to those of Thysanopoda and differ from those of Nyctiphanes.
Meganyctiphanes is presently a single-species genus. However, Colosi
(1918) categorised specimens from the Mediterranean as Meganyctiphanes
calmani based on differences in the structure of the petasma. This was dismissed
as natural variation resulting from the state of maturation by Ruud (1936) in
his report on the Euphausiacea collected in the Mediterranean and adjacent
seas. No further specimens of M. calmani have since been reported.
2.3. Phylogeny
The Euphausiacea have no unequivocal fossil representatives ( Jarman,
2001; Maas and Waloszek, 2001). Indeed, the Malacostraca, in which
Euphausiacea is placed, is also poorly represented in terms of fossils
compared to many other major arthropod groups. Consequently, phyloge-

netic relationships between the Euphausiacea and other orders of Malacos-
traca have been investigated using morphological (Medina et al., 1998) and
molecular ( Jarman et al., 2000) characters found among extant taxa. Medina
et al. (1998) examined the sperm ultrastructure of M. norvegica and found
there to be a thick two layered capsule in close contact with the plasma
membrane, similar to that found in Euphausia ( Jamieson, 1991). This
resembles the sperm of the decapods Sergestes arcticus, Aristaeomorpha foliacea
and Stenopus hispidus. Medina et al. (1998) concluded that Euphausiacea are
close to primitive decapods, in line with the assertions of previous phyloge-
netic studies (Gordon, 1955; McLaughlin, 1980; Schram, 1986). Jarman
et al. (2000) used nuclear ribosomal (28S) DNA sequences to examine
Malacostracan phylogeny and concluded that the mysidacean suborder
Mysida was the sister taxon of the Euphausiacea with the Anaspidacea
being another closely related but more basal group. Decapoda were found
to be more distantly related to krill. A recent total evidence (morphology
and molecules) Malacostracan phylogeny by Jenner et al. (2009), which
included M. norvegica, revealed conflict between morphological and molec-
ular evidence. The best supported higher-level clade to emerge from their
analysis grouped Euphausiacea with the Anaspidacea, Decapoda, and Sto-
matopoda. Jenner et al. (2009) stressed the importance of obtaining new
molecular data sources to better resolve the phylogeny of the Malacostraca.
Relationships among genera within the Euphausiacea have been
examined using morphological and ontogenetic characters (Casanova,
1984; Maas and Waloszek, 2001) and through molecular investigations
(Bargelloni et al., 2000; D’Amato et al., 2008; Jarman, 2001; Patarnello
et al., 1996; Zane and Patarnello, 2000). Casanova (1984) examined 13
morphological characters including the shape of the eye, endopodite struc-
ture in the second and third thoracic limbs, the degree of differentiation
between the male and female antennules, the level of instar variation during
development, and the reduction in the sixth and seventh thoracic limbs.
From this, Casanova (1984) deduced a pattern of morphological change and
constructed a phylogenetic tree (Fig. 1.4a). At the base of the tree was the
most archaic taxa, the genus Bentheuphausia. Euphausiidae were defined at
the next stage up, and comprised five main lineages that ranged from the
most primitive (1) Thysanopoda, through (2) Euphausia, (3) Meganyctiphanes,
Nyctiphanes, Pseudeuphausia, and Thysanoessa, (4) Tessarabrachion to (5)
Nematoscelis, Nematobrachion, and Stylocheiron, the latter of which was
considered the most advanced.
Maas and Waloszek (2001) used literature data and scanning electron
microscope (SEM) examinations of selected ontogenetic stages of E. superba
to determine phylogenetic relationships within the Euphausiacea. The SEM
analysis revealed morphological characters that are either missing in, or
significantly changed towards, the adult. Together with adult features,
A B
Bentheuphausia Bentheuphausia
Thysanopoda Thysanopoda
Euphausia Nematobrachion
Meganyctiphanes Meganyctiphanes
Thysanoessa Euphausia
Nyctiphanes Pseudeuphausia
Pseudeuphausia Nyctiphanes
Tessarabrachion Thysanoessa
Nematoscelis Nematoscelis
Nematobrachion Tessarabrachion
Stylocheiron Stylocheiron
C D
Thysanopoda
Nematobrachion
Pseudeuphausia
Tessarabrachion
Euphausia Euphausia
Nyctiphanes Nyctiphanes
Meganyctiphanes Meganyctiphanes
Thysanoessa Thysanoessa
Nematoscelis Nematoscelis
Stylocheiron Stylocheiron
Figure 1.4 Phylogenetic trees of Euphausiacea genera derived from phylogenies and
descriptions in (A) Casanova (1984), (B) Maas and Waloszek (2001), (C) Jarman
(2001) and (D) Patarnello et al. (1996), Zane and Patarnello 2000, Bargelloni et al.
2000 and D’Amato et al. 2008. With the exception of a few molecular trees, support
values were not presented in the original phylogenies. Apart from tree (C), all of the
trees are rooted.
these ontogenetic characters were used in cladistic analysis to generate a

phylogenetic hypothesis for the Euphausiacea (Fig. 1.4b). The results obtained
confirmed many of the traditional taxonomic groups and, additionally, were
used to propose three novel groups named ‘Euphausiini’ (Euphausia, Pseudeu-
phausia), ‘Nematoscelini’ (Nyctiphanes, Nematoscelina) and ‘Nematoscelina’
(Nematoscelis, Thysanoessa, Tessarabrachion, Stylocheiron). M. norvegica formed an
unresolved trichotomy with the Euphausiini and Nematoscelini. Maas and
Waloszek (2001) also questioned the monophyly of Euphausia, Nematoscelis,
Pseudeuphausia, and Tessarabrachion and went as far as to suggest paraphyly for
Thysanopoda and inferred paraphyly for Thysanoessa.
In terms of recent molecular studies, M. norvegica has been included in
molecular phylogenies in (at least) five publications. These have concerned
the evolutionary history of E. superba and closely related species (Bargelloni

et al., 2000; Patarnello et al., 1996; Zane and Patarnello, 2000), the bioge-
ography of species in the genus Nyctiphanes (D’Amato et al., 2008) and the
evolutionary history of major Euphausiacea lineages ( Jarman, 2001). Prob-
ably, the best resolved study with respect to M. norvegica was that carried out
by Jarman (2001) which considered nuclear ribosomal (28S) DNA analysed
using maximum parsimony and maximum likelihood methods of phyloge-
netic analysis. Jarman (2001) found M. norvegica to be a sister to Thysanoessa
macrura and more distantly related to the ‘archaic’ Thysanopoda (Fig. 1.4c).
Other relationships among major Euphausiacea lineages were less clear due
to differences between the results obtained from the parsimony and likeli-
hood analyses. In the other molecular phylogenies, which were all based on
mitochondrial DNA (16S and ND1 gene sequences) analysed using parsi-
mony, likelihood and Bayesian methods, M. norvegica did not form a
terminal pairing with Thysanoessa but was resolved as sister to a group
containing the genera Thysanoessa, Nematoscelis and Stylocheiron (Fig. 1.4d).
Taxon sampling was narrower in these mitochondrial DNA studies than in
the nuclear DNA study by Jarman (2001), thus comparisons at deeper
phylogenetic levels were not possible.
On the whole, there is support from both morphological and molecular
studies that M. norvegica represents a distinct Euphausiacea lineage. This
appears to be more closely related to Thysanoessa than to the more ‘archaic’
genus of Thysanopoda. However, the position of M. norvegica within
Euphausiacea, like that of Euphausiacea within Malacostraca, is far from
fully resolved. In order to better resolve the phylogenetic position of
M. norvegica, it will be necessary to follow the approach advocated by
Jenner et al. (2009) for higher-level, Malacostraca phylogenetics. This
involves looking beyond the most commonly used sources of data, namely
nuclear ribosomal and mitochondrial DNA sequences, to new sources of
data such as nuclear coding genes.
2.4. Photophores and their function

M. norvegica translates as ‘Norwegian bright shiner’ signifying its notable
capacity to bioluminesce spontaneously. Its bioluminescence is produced by
10 separate light organs (photophores): one on each eyestalk, two pairs on
the ventral thorax and four separate organs under the abdomen (Herring
and Locket, 1978; Fig. 1.5). At a microscopic level, the photophores appear
as highly organised, bell-shaped structures (Grinnell et al. 1988; Herring &
Locket, 1978; Petersson, 1968). Light is produced via a luciferin–luciferase-
type of biochemical reaction that is under hormonal and neural control in
the light-generating cells (Herring & Locket, 1978).
Serotonin (5-hydroxytryptamine, 5-HT) (Kay, 1962) and several analo-
gues (Herring and Locket, 1978) are stimulants for bioluminescence in krill
Figure 1.5 The pattern of photophore illumination in Northern krill (Meganyctiphanes

norvegica) as viewed ventrally. The image was photographed in darkness, exposure time
40 s/400 ASA. The natural colour, a narrow band blue of 480 nm wavelength, is visible
to the human eye with background irradiation dimmer than 10 2 mW/cm2. Note that
parts of the appendages are illuminated.
when added to water in aquaria. Light production can continue for several
hours after single injection of Serotonin into the haemolymph and can
continue for many days on repeated injection of the substance (Fregin and
Wiese, 2002).
Krönström et al. (2007) showed that the interaction between nitric oxide
and Serotonin plays a modulatory role at several levels in the control of light
production in M. norvegica. Inside the photophores, numerous capillaries
drain haemolymph into the light-producing structure (lantern). Filamentous
materials around these capillaries act as sphincters. The sphincters are con-
trolled by nerves containing 5-HT. When exposed to muscle-relaxing
substances (papaverine and verapamil), krill respond with luminescence,
suggesting that the sphincter structures are functionally involved in the
control of light production (Krönström et al., 2009), probably through
restricting the flow of oxygenated haemolymph to the light-generating cells.
Light emission is directional, with a dark pigment masking emission in

most directions, and a highly reflective lining focusing it. By means of small
muscles, the organs can be pointed in different directions and the intensity
of the bioluminescence can be regulated to fit the current ambient situation
(Fregin and Wiese, 2002; Hardy and Kay, 1964; Herring and Locket, 1978;
Kay, 1965; Mauchline, 1960).
In terms of patterns of bioluminescence, there is a lack of meaningful in
situ observations and most studies have been based on the reaction of the
organisms to capture or post-capture manipulation. Macdonald (1927)
stated that freshly caught M. norvegica spontaneously flash during the first
12–24 h but cease to do so quickly after catch. Hardy and Kay (1964) wrote
that the number of flashes was highest in the first 10 h after capture, after
which it decreased quickly but could be observed up to 50 h after capture.
Mauchline (1960) noticed that exposing krill to high intensities of white
light for 20 min (room lights) and then transferring the animals to total
darkness (‘light–dark-experiments’) will trigger the bioluminescence in
60–80% of animals. Hardy and Kay (1964) used this effect to collect the
most responsive animals for their experiments. The first onset of the light
production becomes visible after 20 s and lasts for up to 120 s (Fregin and
Wiese, 2002).
Little has been concluded about the true function of bioluminescence in
krill. During vertical migration experiments within plankton wheels, Körte
(1964) reported that the krill glowed for around 40–80% of the time,
suggesting that bioluminescence may be related to specific behaviours.
Clarke (1963) considered that, under well-lit surface waters, the visual
contrast of krill to an upward looking predator may be countershaded by
the photophores. Light production may also be a form of communication
between individuals. Mauchline (1960) found that individuals can stimulate
each other to flash in a kind of ‘chain reaction’ if he placed individual krill in
adjacent jars. Fregin and Wiese (2002) report that artificial light flashes
presented to a group of krill evoked a signalling behaviour, in which the
animals pointed the beams of light from their photophores for a fraction of a
second at conspecifics at the same depth level. There was a fixed delay
between the signal and the response. Such light signals can probably be
observed at a distance of 10 m or more in clear water (Fregin and Wiese,
2002). Through observations made in situ by a submersible, Widder et al.
(1992) found that the scattering layers, in which M. norvegica was one of
the main emitters of bioluminescence, rarely emitted light spontaneously
but did so once disturbed. They proposed that the light displays were a
defensive function that may distract or blind a predator, warn conspecifics,
or expose primary predators to secondary predators. Increased numbers of
in situ observations are likely to provide better information with
which to ascertain the full functional attributes of bioluminescence in
Northern krill.
3. Geographic Distribution
3.1. General remarks
Zoogeographically, M. norvegica is classified as a boreal species (Dalpadado
and Skjoldal, 1991; Einarsson, 1945; Mauchline and Fisher, 1969).
It characteristically inhabits shelf–slope regions and waters between coastal
banks and deep basins (Melle et al., 2004). Within these habitats, it usually
occurs in areas with bathymetries greater than 100 m (Hjort and Ruud,
1929; Melle et al., 1993) probably because its preferred depth during the day
is between 100 and 500 m from which it performs a DVM to the surface
layers at night (Mauchline and Fisher, 1967).
M. norvegica has been reported to alter its distribution seasonally. Glover
(1952) found that it moved towards coastal areas during the period January
to May, whereas it spreads towards more oceanic areas between June and
December. Mauchline (1960) reports a similar winter dispersion in the Loch
Fyne (Clyde Sea) population. The population movements may be
associated with breeding, with the spring aggregations being a precursor
to mating and spawning. Once these have been effected, dispersion takes
place (Mauchline and Fisher, 1967).
Einarsson (1945) proposed that temperature is the factor that mainly
limits the distributional extent of breeding, with the 5 C isotherm at 100 m
in May limiting the northern breeding extent and the 15 C surface
isotherm in February, the southern breeding extent. Most subsequent
studies have supported these proposed limits, although a functional link
between breeding and temperature remains to be elucidated. The spawning
area is generally more restricted than the overall distribution of the species.
Fowler et al. (1971) found that 18 C approximated a lethal upper tempera-
ture limit for large adults and that the species altered its vertical distribution
in waters off Monaco as a consequence. In the north, the species has been
reported in waters as cold as 2 C (Einarsson, 1945; Hollingshead and
Corey, 1974; Mauchline and Fisher, 1969). Einarsson (1945) considered
the younger stages (the adolescents) may be relatively stenothermic, since
they are generally not found in waters less than 6 C. However, he also
considered the larvae to be more eurythermic, extending well beyond the
spawning limits. As well as temperature, Einarsson (1945) noted that the
breeding area is also a function of the winter distribution, which is generally
more limited than that in the summer. Adult Northern krill are considered
to be euryhaline, with a lower salinity tolerance limit of around 24 PSU,
while the larvae are even more tolerant of low salinities (Buchholz and
Boysen, 1988).
The following section discusses some of the more regional limits within
the wider distributional range of Northern krill (Fig. 1.6).
Figure 1.6 The distribution of Northern krill (Meganyctiphanes norvegica). Data was obtained from three sources: shaded areas were
transposed from Mauchline and Fisher (1967), including any records of larvae or point samples. Blue dots represent records of M. norvegica
present on the Ocean Biogeographic Information System (OBIS—www.iobis.org) as of January 2010. Red dots represent further novel
records obtained in a recent literature search by the present authors. Surface isotherms were extracted from data provided by the NASA
Moderate Resolution Imaging Spectroradiometer (MODIS) representing a mean value for May (2003–2010): the 2 and 18 C isotherms
approximate the distributional limits, with a few exceptions; the 5 and 15 C isotherms represent the notional breeding limits, following
Einarsson (1945).
3.2. North Atlantic west coast

Along the North American coast, the southern limit of distribution is about
Cape Hatteras (35 N; Hansen, 1915; Leavitt, 1938; Bigelow and Sears,
1939). It occurs in great abundance in the Gulf of Maine and Bay of Fundy
(Bigelow, 1926; Fish and Johnson, 1937; Hansen, 1915; Soulier, 1965;
Whiteley, 1948). In the Pasamaquoddy area (Bay of Fundy), M. norvegica
is known to seek deeper warmer water during the winter to avoid the
chilling down to <1 C that occurs in the upper water layers (Bigelow,
1926; Hollingshead and Corey, 1974). It does not occur in great numbers in
the inner Bay of Fundy, nor onto the southwest Nova Scotia shelf (Kulka
et al., 1982). In the Gulf of Maine, the species is usually found in waters of
between 3 and 4 C. It may congregate around the 100 m contour further
offshore, around Georges Bank, but with seasonal shifts in its distribution
(Bigelow, 1926; Whiteley, 1948). The species is common in the estuary
of the St Lawrence river (Préfontaine and Brunel, 1962), the Gulf of
St Lawrence and Chaleur Bay (Lacroix, 1961), where there has been
detailed descriptions of the interactions between its vertical migration
behaviour and the prevalent 3D advective regimes (Chapter 10). The
species is absent in Hudson Bay and the Hudson Strait marks the northern
limit of distribution along this coast.
3.3. Arctic limits

Disko Bay is the northernmost point that M. norvegica has been recorded on
the west Greenland coast (Dunbar, 1942), those being larvae taken at the
end of May. Nevertheless, there are no records of breeding adults and it is
unlikely that breeding takes place in this region (Einarsson, 1945;
Mauchline and Fisher, 1967). Saunders et al. (2007) found juvenile and
adult size classes in the central and northern Irminger Sea and also in the east
Greenland coastal region, although the relative number of larger individuals
was comparatively less in the Irminger Sea. Larval stages were relatively
scarce although they may have been distributed below the maximum net-
depth of 400 m. Einarsson (1945) considered that no considerable spawning
takes place in the cold less saline water of the East Greenland Current and
that spawning is restricted almost entirely to Atlantic water, which can reach
to the slope of the Greenland continental shelf. This current may be the
origin of small numbers of larvae and adults caught further to the north and
east, in the Denmark Strait (Mauchline and Fisher, 1967). Records are rare
in the Greenland Sea, mainly restricted to larval stages (Ussing, 1938).
Damas and Koefoed (1909) record M. norvegica at 80 030 N, 2 470 E.
Hirche et al. (1994) found M. norvegica to be the most numerous euphausiid
species in the Fram Strait, although absolute numbers were very low.
Around Iceland, adults can occur in numbers comparable to other well-

populated regions (Astthorsson and Gislason, 1997). However, breeding is
almost entirely restricted to the outer parts of the southern continental shelf,
especially on the western side (Einarsson, 1945). For instance, Astthorsson
(1990) found no records of larvae in Ísafjord, northwest Iceland, and
concluded that recruitment came from outside. Astthorsson (1990) also
found that the abundance of adults in the fjord increased during the autumn
and winter, probably as a result of autumn water exchange.
M. norvegica has been found mainly in the southwestern part of the
Barents Sea where its occurrence depends on transport with inflowing
water from the west (Dalpadado and Skjoldal, 1991; Drobysheva, 1979).
Dalpadado and Skjoldal (1991) did not find the species in the central or
northern Barents Sea during their investigations in May–June nor at any
stations during an August cruise to the northern Barents Sea. Drobysheva
(1957) found M. norvegica specimens with spermatophores on both sexes in
the Barents Sea, but larvae were registered only in the western part in the
warm summer of 1954; she therefore assumed that M. norvegica does not
reproduce in the Barents Sea and that recruitment occurs partly from the
warmer Norwegian Atlantic Current. Bjorck (1916) did not record any
M. norvegica in the fjords of Spitsbergen but recently, Buchholz et al. (2010)
found younger specimens of M. norvegica in Kongsfjord, west Spitsbergen.
They found no sign of reproducing adults.
Further to the east, M. norvegica has been recorded in the White Sea and
also off the south-western part of Nova Zemlya and along the Murmansk
coast, although breeding is not believed to occur in this region (Boldovsky,
1937; Drobysheva, 1957, 1961; Zelickman, 1958, 1961; Mauchline and
Fisher, 1967). No one has found M. norvegica east of 52 E, which appears to
be its north-easterly limit apart from one isolated record by Sars (1900) from
81 240 N 125 E.
3.4. North-east Atlantic and fringes

M. norvegica becomes increasingly common heading from the northern to
the southern reaches of the Norwegian coast and the adjacent Norwegian
Sea. At the northern extremities, Falk-Petersen and Hopkins (1981) found a
M. norvegica population in Balsfjorden (69 320 N 19 060 E) containing
adults with spermatophores but no evidence of any larvae. They concluded
that the population in this region depended entirely on recruitment from
external coastal waters, possibly associated with the northward flow of the
Norwegian Coastal Current. On the mid Norwegian shelf (Mre plateau,
Halten, Sklinna, Træna banks), the abundance of M. norvegica reaches >20
individuals per square meter Dalpadado (2006), with the main distribution
being within Atlantic waters of the shelf region. The species is common
off the southwestern Norwegian coast, and in the Skagerrak and Kattegat
and adjacent fjords such as Gullmarsfjord (Heegard, 1948). Heegard (1948)

considered that the species bred in all of these environments apart from the
Kattegat, but breeding and recruitment in deep basins within the Kattegat
has since been reported by Boysen and Buchholz (1984). It is absent from
the North Sea, probably because of its shallow bathymetry (Kramp, 1913).
There have been studies on various life-stages on the Iceland-Fare ridge,
around the Faeres and in the Faere-Shetland Channel (Glover, 1952;
Lindley, 1982a; Lucas et al., 1942; Ruud, 1926). Further detailed studies on
the behaviour and life-history of the species have been carried out on popula-
tions resident along the Scottish west coast (Mauchline, 1960; Tarling, 2003);
Williamson (1956) has studied the local distribution in the Irish Sea and it is
also common off the west and south coasts of Ireland (Frost, 1932). Although
a contributor to the euphausiid community in the south-west reaches of
the British Isles, the species becomes increasingly less common moving into
the open North Atlantic, south of 55 S (Lindley, 1982a).
3.5. Mediterranean and south-eastern limits

M. norvegica is located throughout the Bay of Biscay and in the area westward
of it (Ruud, 1936). Ruud (1936) also records the species off the Portuguese
coast and southwards to Gibraltar as well as in the Bay of Cadiz. Bouvier
(1907) presents records of the species between the Cape Verde Islands and
Tenerife (the Canaries), while Mauchline (1980) reported a record from
A de C Baker that the species was found westwards of the Azores at 37 34.30
N 25 22.00 W and also off the West African coast at 28 04.80 N 14 040 W.
Thiriot (1977) reported that M. norvegica, along with Euphausia krohni,
dominated the euphausiids in the oceanic zone to the north of Casablanca
(Morocco). The species was also found in a region of active upwelling from
Casablanca down to Cape Juby (27 570 N 12 560 W).
M. norvegica occurs throughout the western Mediterranean (Alvarino,
1957; Dion and Nouvel, 1960; Furnestin, 1960; Jespersen, 1923; Ruud,
1936), particularly between 5 and 10 E ( Jespersen, 1923; Labat and
Cuzin-Roudy, 1996). In the eastern Mediterranean, it has neither been
found south of 35 N nor east of 29 E (Mauchline and Fisher, 1967). It has
been recorded in the Sea of Marmara and the Aegean Sea, in small numbers
( Jespersen, 1923; Ruud, 1936). It is more common in the Adriatic and
Ionian seas, particularly in deep waters (Guglielmo, 1979), and also in the
Tyrrhenian Sea (Brancato et al., 2001).
3.6. Distributional trends and changes

There is little information regarding any recent changes in the southerly
limits of this species, which may partially reflect the paucity of appropriate
sampling in these regions. However, there has been particular interest in the
recent extent of the northerly distributional limits, along with increased

incursion of water with Atlantic origin into Arctic waters (Cottier et al.,
2005; Cottier et al., 2007). In particular, inflowing coastal and Atlantic
waters from the southwest are cooled and gradually transformed into Arctic
waters in the Barents Sea (Loeng, 1989; Midttun, 1989). This makes the
Barents Sea a zoogeographical transition zone between the Atlantic-boreal
and the Arctic and its pelagic communities are strongly influenced by
immigration from inflowing currents (Dalpadado and Skjoldal, 1991;
Skjoldal and Rey, 1989).
The first remarks regarding the changing distribution of M. norvegica
in this region were made by Mauchline and Fisher (1967) who stated that
“in the early part of this century, [M. norvegica] was rare but became
common in the southern Barents Sea and extended eastwards up to Nova
Zemlya by the late 1930s. It now appears to be always present, although no
breeding has yet been recorded.” Siegel (2000a) did not consider that this
trend was directly related to increasing temperature but rather to an increas-
ing number of years in which a ‘stronger drift of the North Atlantic Current
carry krill to northern waters’.
Zhukova et al. (2009) examined M. norvegica abundance in the Barents
Sea relative to sea-water temperature using more than 50 years of net-
sample data. They found that the portion of M. norvegica varied considerably
over the time period, from 10 to 20% of the total euphausiids population in
the warm 1950s and 1960s to almost completely disappearing in the 1970–
1990s. The peak of species occurrence (18–26%) took place in the begin-
ning of the warm period (1999–2000) after a succession of cold years.
Nevertheless, there was a subsequent decline in the proportion of
M. norvegica to 7% in 2004–2005 despite the fact that sea-water temperatures
remained abnormally warm. This was believed to be the result of fish
predation, particularly by cod fingerlings. The study of Zhukova et al.
demonstrates that the establishment of a species such as M. norvegica outside
of its previous boundaries depends not only on suitable physical conditions
but also on the food-web dynamics of the new communities into which
they are introduced.
4. The Sampling of Northern Krill

Historically, much of what we have learnt about Northern krill has
been obtained through the deployment and analysis of trawled-nets. Net
sampling remains the standard method with which to determine the size-
structure of krill populations and to obtain specimens for incubation and
experimentation. Nevertheless, net sampling alone has limitations in ascer-
taining distribution patterns and true levels of biomass since Northern krill
can avoid on-coming nets quite effectively given their fast swimming speed
and high visual acuity. Active acoustics are now frequently used alongside
net-sampling deployments in carrying out such surveys, giving the added
benefits of greater coverage and minimal avoidance. Direct visual observa-
tions via manned-submersibles or remotely operated vehicles (ROVs) are
also providing useful insights into the behavioural ecology of krill species.
The use of each of these techniques and their intercomparison are discussed
in the following sections.
4.1. Net sampling

Nets that have achieved the greatest success in obtaining samples of
Northern krill have incorporated a number of attributes, of which the
main ones are:
Being as large as possible
Being propelled relatively fast at a constant speed
Remaining at least 85% efficient throughout the haul
Being free of obstructions forward of the mouth, such as tow lines
Generally being dark coloured with no shiny metal components
Some of the earliest nets were simple ring nets, with diameters generally
between 0.5 and 3 m (e.g. Einarsson, 1945; Mauchline, 1960; Table 1.2).
They either contained meshes of between 200 mm and 7 mm, or stramin, a
coarse canvas of hemp, with 500 threads to 100 cm. Oblique tows at speeds
of 0.5–1 m s 1 have been the most common deployment method.
Kaartvedt et al. (2002) customised this simple device with a strobe light to
temporarily startle the krill and decrease avoidance. The plummet net,
which is a ring net packed with lead shot around its rim, decreases avoidance
through a rapid vertical approach to krill from above (Hovekamp, 1989).
The rapidity of the deployments and the fact that little pressure is placed on
the cod-end makes plummet nets effective at maintaining captured speci-
mens in a healthy state for further process work. A commonly used alterna-
tive for this purpose is the Isaacs-Kidd Midwater Trawl (Isaacs and Kidd,
1953) which has a mouth opening of 1 m2 and a mesh size of 1.5 mm. The
net is easy to deploy and capable of rapid retrieval, which minimises the
amount of time captured animals remain in the cod-end.
To examine the depth distribution and behaviour of krill populations,
multinets have been mainly deployed, which are systems based on the
principle of opening and closing a series of individual plankton nets in
succession. Two widely used types are the BIONESS (Sameoto et al.,
1980), and the MOCNESS (Wiebe et al., 1985). The systems are towed
horizontally or obliquely with the nets being incremented at desired
intervals. Best results have been achieved through sampling mainly
during the upward trajectory. The BIONESS is towed at higher speeds
Table 1.2 A synthesis of the different types of nets deployed to capture Northern krill (Meganyctiphanes norvegica) categorised according to the
purpose of the net deployment
Purpose of net deployment Net and deployment parameters
Abundance/ Population
biomass dynamics Horizontal Towing
Net sampling geographic and and vertical Behaviour Live Accompanying Number of speed
device surveys biochemistry distribution and feeding specimens acoustic data Mouth Mesh nets (knots) Haul Speed
Simple nets
WP2/Conical 4 1, 2, 4, 5, 4 6 (with 3, 7, 11 Area—0.5 m21, 200 mm2,3, 1 42 1 m s 11
net/Nansen 8, 9 strobe 2 m25, 1.2 m28 300 mm1, 0.75 m s 110
Net/ring trawl light),10 Diameter— 500 mm2,7,
0.6 m2, 7 mm4,
1 m2,11,6,7,10 2 mm5,9,10
2 m4, 3 m4
Bongo nets 12, 15 13, 14 12, 16 12 120 kHz Area—0.2 m213 200 mm13, 2 2.514,15 10 m min 114,15
echosounder: 0.05 m213 243 mm16,
16 Diameter— 335 mm14,15,
0.6 m12,14,15, 500 mm13,
0.75 m16 505 mm12
Stramin Net 4 4, 17 4, 17 17 Diameter— Coarse canvas 4 (surface, 417
(conical) 1 m17, of hemp, 50 m,
1.5 m4, 2 m4 500 threads 100 m
to 100 cm and
150 m)17
Isaacs-Kidd 5, 8, 19, 18, 26, 33 6, 23, 25 23, 27, 300 kHz Area—0.6 m2 500 mm25,30 1 1.7518
Midwater 20, 31, 28, ADCP: 21, 20,21,22,23, 1500 mm 2.022,26
Trawl Isaacs 32 29, 22 24,25,28,29 20,21,22, 2.0–2.519
and Kidd 30, 38 and 2 m218 23,27,28,29
(1953) 34, 120 kHz 9 m226 2 mm8
35 echosounder: Diameter— 4 mm18
24, 25 3-feet6 1 cm26
Ori net Omori 19 1 2.0–2.5
(1965)
Beyer’s Low 13 Area—0.7 m2 1 mm 1
Speed
Midwater
Trawl
Multiple nets
BIONESS 39, 40 41 36, 37, 36, 37, 42 3 12, 50, 122 and Area—1 m2 500 mm36,38 12 2–337 1.5–2 m s 136
electronic 38, 42 200 kHz 250, 333 438
mulitnet echosounder: mm37,39
Sameoto et al. 41
(1980) 38 and 120 kHz
echosounder:
39, 40, 42
153 and
300 kHz
ADCP: 42
MOCNESS 44, 57 5, 9, 44, 2 45, 48, 49, 50, 55 153 kHz Area—1 m2 333 mm 9 1.5–544, 0.5 m s 146
Wiebe et al. 45, 51, 52, 53, ADCP: 2,9,44,45,46, 2.545,55
(1985) 54 56, 58 46, 47, 52, 58 ,47,48,49, 346
300 kHz 52,54, ,56
ADCP: 49 2 mm
9,45,47,48,49,51,
52,53,54,55,56
Methot net 59 59 59 60 Area—1.5 m259 500 mm/ 259 1.5–2.059
Diameter—2 m60 1.2 mm/60 160
1.5 mm59
Combination 61, 62 1 m2, 8 m2 0.33 mm, 2
Rectangular 3 mm
Midwater main mesh,
Trawl Baker 1.5 mm cod
et al. (1973), end
Roe et al
(1980)
Benthic nets
Macroplankton 63 63 0.2 m2 564 mm 1
net attached to
bottom trawl
Trawls
Pelagic trawls: 4, 57 4 4 25, 64, 65, 38 and 120 kHz Area— 20 cm at 1
‘Young-fish’25 66 echosounder: 100 m225,57 opening
‘Åkra’57,64 25,66 and 1 cm at
(continued)
Table 1.2 (continued)
Purpose of net deployment Net and deployment parameters
Abundance/ Population
biomass dynamics Horizontal Towing
Net sampling geographic and and vertical Behaviour Live Accompanying Number of speed
device surveys biochemistry distribution and feeding specimens acoustic data Mouth Mesh nets (knots) Haul Speed
‘Fipec’65 18, 38, Vertical rear

‘Petersen’s 70, 120, and opening— end25,57
young-fish’4 200 kHz 30 m57,64 3200 mm at
echosounders: front to
64 20 mm in
38, 120 and cod-end64
200 kHz 5 mm65
echosounders: Stramin (500
65 threads to
100 cm)4
38 mm mesh
cod end net
with a 20 m
long
inner-net of
16 mm
mesh57
High speed
samplers
Continuous 67 280 mm Continuous Variable 3 m3 of water
plankton spool ship filtered per
recorder speed- 16 km of tow
Gulf III sampler 68 68 44
Information on net deployment parameters (where detailed in the study) are also provided. Italicised numbers relate to the following articles: 1, Timofeev (2002); 2, Boysen and Buchholz (1984); 3, Mayzaud et al. (2005); 4, Einarsson (1945); 5, Albessard and
Mayzaud (2003); 6, Kaartvedt et al. (2002); 7, Torgersen (2001); 8, Albessard et al. (2001); 9, Cuzin-Roudy and Buchholz (1999); 10, Onsrud and Kaartvedt (1998); 11, Conti et al. (2005); 12, Kulka et al. (1982); 13, Falk-Petersen and Hopkins (1981);
14, Astthorsson (1990); 15, Astthorsson and Gislason (1997); 16, Sameoto (1976); 17, Mauchline (1960); 18, Hassan (1999); 19, Labat and Cuzin-Roudy (1996); 20, Thomasson (2003); 21, Liljebladh & Thomasson (2001); 22, Strömberg et al. (2002); 23,
Spicer et al. (1999); 24, Everson et al. (2007); 25, Klevjer & Kaartvedt (2006); 26, Sardou et al. (1996); 27, Schmidt et al. (2004); 28, Cuzin-Roudy et al. (2004); 29, Thomasson et al. (2003); 30, Båmstedt and Karlson (1998); 31, Hollingshead and Corey (1974);
32, Matthews (1973); 33, Guglielmo (1979); 34, Spicer and Strömberg (2002); 35, Salomon et al. (2000); 36, Brancato et al. (2001); 37, Harvey et al. (2009); 38, Sardou and Andersen (1993); 39, Simard and Lavoie (1999); 40, Simard and Sourisseau (2009);
41, Cochrane et al. (2000); 42, Sourisseau et al. (2008); 43, Sourisseau et al. (2008); 44, Dalpadado and Skjoldal (1991); 45, Tarling (2003); 46, Buchholz et al. (1995); 47, Tarling et al (1999a); 48, Tarling et al. (1999b); 49, Tarling et al. (2002); 50, Saborowski
et al. (2002); 51, Tarling and Cuzin-Roudy (2003); 52, Tarling et al. (2001); 53, Lass et al. (2001); 54, Virtue et al. (2000); 55, Saborowski et al. (2000); 56, Tarling et al. (1998); 57, Dalpadado et al. (1998); 58, Zhou et al. (2005); 59, Saunders et al. (2007);
60, Dalpadado et al. (2008); 61, Mauchline (1985); 62, Lindley et al. (1999); 63, Zhukova et al. (2009); 64, Kaartvedt et al. (2005); 65, Cotté and Simard (2005); 66, Klevjer and Kaartvedt (2003); 67, Lindley (1982a, 1982b); 68, Dalpadado (2006)
(3–4 knots) than the MOCNESS (1.5–2 knots) and may be more efficient at
capturing faster moving organisms such as krill. Nevertheless, the
MOCNESS has been used effectively in a number of research efforts on
euphausiids (Table 1.2). Most such systems collect a suite of environmental
parameters during deployment, particularly temperature, salinity, down-
welling irradiance and fluorescence.
Some studies have employed trawl nets, designed for larval fish surveys,
to capture Northern krill (Table 1.2). In many instances, the surveys were
oriented towards the capture of fish and have obtained krill as a by-catch.
Nevertheless, such catches have provided a great deal of useful information
on distribution and population structure. Similarly, samples of krill obtained
from the epibenthic layer have mainly been obtained from surveys with
other sampling priorities. For instance, Zhukova et al. (2009) performed a
valuable study on Northern krill using a macroplankton net attached to a
bottom trawl.
4.2. Active acoustics

Acoustics has perhaps now overtaken net sampling as the preferred means of
investigating krill distribution and abundance (Hewitt and Demer, 2000;
Macaulay, 2000). Some view nets as secondary tools, whose main purpose is
to verify the results of acoustic studies (Pauly et al., 2000).
Acoustic surveying has the advantages of being non-invasive and capable
of covering large areas rapidly. Nets nevertheless remain the primary tool
for obtaining krill specimens for population and experimental work (Nicol
and Brierley, 2010).
Modern day acoustic systems consist of a number of different frequency
echosounders, ranging between 18 and 500 kHz. The difference in scattering
strength of targets across these frequencies gives information on their body size
and tissue density which, through modelling and in situ validation, has become
a means of inferring target identity remotely. Technologies and analytical
techniques continue to develop (e.g. Korneliussen and Ona, 2002) but one
well-established approach towards locating aggregations of Antarctic and
Northern krill is to compare the scattering strength of aggregations at
38 kHz and 120 kHz (Madureira et al., 1993a,b). All acoustic approaches to
estimating numerical density and biomass of krill rely on an estimation of the
target strength (TS) of individual specimens. There has been relatively little
consideration of Northern krill TS compared to Antarctic krill. Nevertheless,
there is much overlap in morphology, size, and tissue density to allow TS
estimates to be interchanged. For instance, in an acoustic survey of a Northern
krill population in Gullmarsfjord, Sweden, Everson et al. (2007) used a TS of
76.1 dB (range 6.2) at 120 kHz and 85.1 (range 6.5) at 38 kHz which were
values reported for E. superba by Foote et al. (1990). Foote et al. (1990)
estimated E. superba TS from a cage experiment where the size range of
specimens (30–39 mm) was almost identical to that of the fjordic Northern
krill. TS has been estimated during in situ observations of Northern krill using a
moored upward looking split-beam echosounder by Klevjer and Kaartvedt
(2006). Although their mean values differed little from those used by Everson
et al. (2007), of particular interest was the distinct variability in TS they
observed as a result of the size and behaviour of the krill, which could account
for a change in TS of the order of 10 dB.
It is well recognised that there is often a discrepancy between net-catch
and acoustic survey estimates of krill biomass, with the general belief that net
sampling may underestimate true biomass by orders of magnitude because of
escapement (Hamner and Hamner, 2000; Watkins, 2000). Echosounders
also sample much larger volumes than the nets, for example, 50-fold more at
150 m range. Nevertheless, over broad temporal and spatial scales, Atkinson
et al. (2009) found there to be a general agreement in biomass estimates of
Antarctic krill made by nets and acoustics. Sameoto et al. (1993) used a
combination of echosounders, an optical particle counter, cameras attached
to ROVs and a BIONESS net equipped with a strobe light to examine the
level of net avoidance by Northern krill. They concluded that net-avoidance
was significant but reduced by the strobe lighting. Simard and Sourisseau
(2009) carried out a comparison on fine temporal and spatial scales during a
3-day experiment at a fixed location, using strobe-equipped BIONESS and
echosounders, and found that diel effects altered both net catch and acoustic
estimates of Northern krill biomass. Night-time catches in the krill scattering
layer (SL) were 15 times the acoustic estimates, but the situation was reversed
during daytime, when the acoustic estimates in the SL were 5 times larger
than the catches. Net avoidance explained the daytime anomaly while, at
night time, it was believed that a change in krill orientation, affecting TS,
led to an underestimation of biomass by the echosounder. The effect of
orientation is a major consideration in krill TS model development
(Conti and Demer, 2006).
There has been comparatively little use of echosounders to perform
large-scale biomass surveys of Northern krill. By contrast, efforts to survey
biomass distributions of Antarctic krill have involved simultaneous multi-
ship international efforts (Hewitt et al., 2004). This may reflect the greater
interest in stock sizes of the latter species. Echosounders have nevertheless
revealed interesting details with regards local-scale distribution patterns of
Northern krill. In the Gulf of St Lawrence, for instance, combined net and
echosounder surveys found a number of vertical and horizontal distributional
features of Northern krill scattering layers in relation to upwelling coastal
currents, 3D circulation patterns as well as evidence of predator–prey interac-
tions, particularly with regards the feeding behaviour of whales (Simard et al.,
1986; Sourisseau et al., 2006, 2008). Moored echosounders have provided
revealing insights into vertical migration behavioural patterns (Onsrud et al.,
2005) as well as estimates of swimming speed (Klevjer and Kaartvedt, 2003)
and TS (Klevjer and Kaartvedt, 2006). Moored acoustic Doppler current

profilers (ADCPs) have obtained time-series information at coarser resolution
levels. Tarling et al. (2002) for instance, was able to detect a normal DVM in
Northern krill plus an additional secondary midnight sinking behaviour in its
prey, Calanus finmarchicus. ADCPs are principally designed to measure current
speeds and are particularly suited to documenting the interaction between krill
distributions and the horizontal advective regimes (Liljebladh and Thomasson,
2001; Zhou et al., 2005). The devices also provide an estimate of the vertical
velocity of particles and Tarling et al. (2001) used this parameter to estimate the
relative vertical migration speed of individuals within Northern krill swarms.
4.3. Alternative technologies for sampling and observation

Nets and acoustics remain the most tried and tested sampling technologies
for the study of Northern krill. In their review, Nicol and Brierley (2010)
consider that this will remain so in the study of all krill species into the future.
Nevertheless, they also advise that new technologies be embraced and used
in parallel, with their effectiveness assessed through cross-comparisons. This
is particularly so when krill occupy areas where sampling by traditional
methods is limited.
Ship-based acoustics systems have trouble in sampling the very surface of
the water column, the 3–15 m of water which resides above the downward
looking, ship-borne transducers. Surface reflectance means that this zone is
also unresolved by upward looking moored systems. Northern krill is one of
several euphausiid species that notably aggregate within surface swarms,
either through the action of tides and upwelling or as a result of their own
motivation to mate and spawn (Nicol, 1986). Surface swarms offer unique
sampling opportunities because the population is concentrated in the surface
layer and is often highly visible (Nicol and Brierley, 2010). The swarms can
be adequately viewed from ships, as appreciated by the first International
krill biomass experiment (FIBEX) in the Southern Ocean, which developed
standard protocols to measure surface-swarm parameters such as area, shape,
spacing, and even colour (Hampton, 1982). More synoptic data can be
obtained from cameras mounted to aircraft, as reported by Nicol (1986) for
Northern krill in the Bay of Fundy. Satellite imagery is now approaching
similar levels of resolution which may greatly facilitate data coverage and
acquisition (Nicol and Brierley, 2010). At smaller scales, it is feasible for
divers to enter surface swarms and examine their structure first hand, as has
been achieved for Euphausia pacifica by Hanamura et al. (1984) and Endo
et al. (1985), and for E. superba by Hamner (1984). These studies have
reported swarm densities that are orders of magnitude higher than estimated
through either nets or acoustics (Hamner and Hamner, 2000).
The epibenthic layer is another region that is poorly sampled by pelagic
nets and badly resolved by acoustics. In shelf and upper slope areas,
euphausiids can have access to the seafloor and M. norvegica has been caught
in these regions by hyperbenthic sledges (Beyer, 1992) and bottom trawls
containing additional macroplankton nets (Zhukova et al., 2009). This layer
may be a daytime residence to escape predation as well as a feeding
environment (see Chapter 10 and Chapter 5). Our knowledge of the
biology of krill in this layer is further supplemented by observations from
manned submersibles and ROVs. For instance, Youngbluth et al. (1989)
observed Northern krill feeding on epibenthic particles while Jonsson et al.
(2001), Hudson and Wigham (2003) and Rosenberg et al. (2005) described
the swimming behaviour of Northern krill in the epibenthic layer and their
interaction with benthic fauna. The potential for these kinds of observations
is continuously increasing, as ROVs become more common and increas-
ingly capable of carrying out deeper and longer dives. For instance, from
ROV observations, Clarke and Tyler (2008) were recently able to report on
E. superba feeding benthically at 3500 m. More such studies in Northern
krill habitats would be valuable.
5. Ecological Role
M. norvegica can make up a significant part of the biomass of the
euphausiid community over large parts of its distributional range (see
above). This makes it both an important consumer of productivity at
lower trophic levels (Chapter 5) and an important prey item for many higher
predators (Chapter 10). Beyond these well-considered roles, however,
Northern krill has a number of other ecological functions that have impacts
on other communities and contribute to wider biogeochemical processes.
5.1. Benthic interactions

The behaviour and biomass of M. norvegica, particularly in near shore waters,
may have a profound impact on benthic and epibenthic communities. The
species can be potentially a major source of biogenic material (faecal strings,
moults, carcasses) for these communities, can act as a bioturbator and trans-
porter of resuspended and flocculent sediments, and can be a major prey
item for demersal predators. This contribution to benthic communities was
examined in detail by Youngbluth et al. (1989), who made observations
with a submersible within the Wilkinson and Jordan basins in the Gulf of
Maine and in canyons south of Georges bank. The submersible observed
high densities of krill faecal strings in a 5–24 m thick layer coincident with
the pycnocline (at depths of around 15–30 m) which were collected and
later examined for sinking rate and organic-matter content. Youngbluth
et al. (1989) calculated sinking rates to be 204 m d 1 24 S.E and
organic matter transport to the bottom to be 7–12 mg C m 2 d 1. This

represented 1–6% of the daily primary production in the surface mixed layer
of that region. The packaging of organic material within fast sinking faecal
strings not only provides biogenic material to the benthic community but
also means that carbon is rapidly exported out of the surface mixed layers, so
increasing its likelihood of ultimate carbon sequestration. This effect was
also examined in E. superba in the Southern Ocean by Tarling and Johnson
(2006), where krill were estimated to transport 2.3 1013 g C to the ocean
interior each year.
Nevertheless, in the case of Northern krill, the transport of carbon may
not always be downwards. Youngbluth et al. (1989) observed particularly
dense aggregations of M. norvegica (up to 104 individuals per cubic meter)
close to benthic boundary layer. Consumption of biogenic detritus in the
epibenthic region by these krill may account for considerable repackaging
and bioturbation. Assuming that these krill feed more or less continuously,
individuals that then migrate vertically each night may introduce recycled
biogenic and inorganic materials back into the surface mixed layer when
they release faecal strings.
The dense epibenthic aggregations may be particularly targeted by
groundfish such as cod, hake, pollack as well as squid. The density of
these aggregations may also be sufficient to supply the daily food rations
of fin whales. Sourisseau et al. (2006) considered how the predictable nature
of krill to form high density aggregations in certain locations influences the
behaviour of whales. The sea bottom boundary may limit the potential for
escapement by krill during feeding forays by whales, so increasing whale
foraging efficiency.
5.2. Krill production and predator consumption

For understanding the quantitative role of Northern krill in the transfer of
organic material from primary production to higher trophic levels, consid-
eration of its productivity must be made. Siegel (2000b) considered the
production to biomass (P/B) ratios of M. norvegica was relatively low
compared to temperate and warm water species. Typical values were
around 3:1–4:1. E. pacifica in Japan and E. superba in the Southern Ocean
have similarly low values, ranging between 1:1 and 4:1 (Siegel, 2000b).
There is notable regional and temporal variability in levels of productivity
in M. norvegica. For instance, Lindley (1982a) estimated that its annual net
production at 10 m depth in the North Atlantic varied between 0.80 and
4.31 mg m 3 y 1 in 1966 and between 1.62 and 18.74 mg m 3 y 1 in 1967.
The range in values, particularly in the second year, was mainly explained by
the comparatively high but variable levels of production in Norwegian
waters. In other North Atlantic regions, productivity was lower and less
variable. This variation does not necessarily translate into accompanying
Exploring the Variety of Random
Documents with Different Content
FRONT ELEVATION
ATTIC PLAN
PLATE XLVI.
PAIR OF COTTAGES.
SEE PAGE 47.
Plates xlv. and xlvi. give plans, and Plate xlvii. the view of a pair of
three-storey cottages of about the same accommodation, the left-hand
having the following:—
Ground Floor.
Dining Room, 11 ft. 6 ins. × 18 ft., with French window. Drawing Room, 12 ft. 6
ins. × 15 ft., with deep bay. Small Sitting Room, 7 ft. × 11 ft. 2 ins. Working
Kitchen, 11 ft. 2 in. × 12 ft. 6 ins. Larder and China Pantry, Porch and Hall. w.c.,
Coals, Tools, and Enclosed Yard.
Bedroom Floor.
First Bedroom, 12 ft. 6 ins. × 15 ft., and deep bay. Second Bedroom, 11 ft. 6 ins.
× 16 ft. Third Bedroom, 10 ft. 6 ins. × 11 ft. 2 ins., with oriel. Bathroom, with
Lavatory, w.c. Two Attics and Large Box Room.
PLATE XLVII.
PAIR OF COTTAGES.
SEE PAGE 47.
The accommodation in the two houses differs owing to the aspect. If

the two plans were identical, a considerable portion of the right-hand
garden would be shut off from the south, and the larder would not face
the north. Stress has already been laid on the necessity of considering
aspect.
The left-hand is a corner house, and the projecting out-houses
answer the double purpose of screening the garden from the road and
protecting the house from the north wind.
In the adjoining house there is no small sitting room, but an extra
attic. The outbuildings are attached to the main building, and do not
project into the garden; the principal room is thus left open to the
south.
Materials.—The houses are built of common bricks whitewashed,
with a tarred plinth. There are half-timber porches, and the spaces
between the bays and under the dormers are covered with rough-cast
and decorated with parquetry. The rainwater head in front is picked
out in vermillion, the introduction of a very little bright colour giving a
pleasant jewel-like effect. Peake’s hand-made tiles, of dark colour, are
used for the roofs, with half-round ridging, and ornamental iron stays
support the gutters, which are of 6 in. half-round iron.
A pair of houses erected to a similar plan to that of the right-hand
house in 1904 cost £610 each. The cost of the examples given work
out more owing to the fall in the land, which necessitates very deep
footings, and also to the plans differing in order to suit aspect and site.
PLATE XLVIII.
PAIR OF THREE-STOREY COTTAGES.
PLATE XLVIII.
PAIR OF COTTAGES.
SEE PAGE 48.
Plate xlviii. gives the view of a pair of houses similar to the last, but
somewhat reduced in size, and the treatment varied. Brindled bricks
are used for the ground floor, and rough-cast for the upper storeys.
PLATES XLIX., L., LI., LII., LIII., LIV., AND LV.
TWO PAIRS OF COTTAGES.
These plates illustrate two pairs of cottages of two storeys each,

almost identical in plan, but differently treated.
FRONT ELEVATION
GROUND PLAN
BEDROOM PLAN
PLATE XLIX.
PAIR OF COTTAGES.
SEE PAGE 49.
PLATE L.
PAIR OF COTTAGES.
SEE PAGE 49.
The accommodation of the pair shown in Plates xlix. and l. is:—

Ground Floor.
Dining Room, 13 ft. 6 in. × 16 ft. and bay. Drawing Room, 13 ft. 6 ins. × 16 ft. 3
ins., including ingle and bay window. Kitchen, 10 ft. 6 ins. × 12 ft. Hall, with
storm doors, 12 ft. 6 ins. × 10 ft. Scullery, Larder, w.c., Coals, and Tool House.
Frontage, 15 yds.
Bedroom Floor.
First Bedroom, 12 ft. × 16 ft. 3 ins., and bay. Second Bedroom, 12 ft. 4 ins. × 13
ft. 6 in. Third Bedroom, 10 ft. × 10 ft. 2 ins. Fourth Bedroom, 9 ft. × 13 ft. 6 ins.
Boxroom, 8 ft. × 10 ft. Bathroom, with Lavatory, and w.c.
The dining room is lighted by a small east window and a west bay
window, the latter being covered by the roof of the verandah, which
terminates in the bay window of the drawing room. Although the
kitchen is a small one, it has the advantage of not being a passage
room, the door from the hall to the kitchen and that from the kitchen to
the scullery being arranged beside one another in the same wall. In
these houses the windows have wooden frames and wrought-iron
casements.
The principal rooms occupy the full width of the back, and the hall is
therefore extended to admit of the doors of the two rooms being
conveniently placed.
Ingle Nook.—The ingle which results from this arrangement has a
beam with a shelf above continuing the line of the architrave, and the
ceiling of the ingle is only 6 ft. 6 in. high. There is a small light on one
side.
PLATE LI.
COTTAGE INGLE.
SEE PAGE 49.
PLATE LII.
DETAIL VIEW.
SEE PAGE 49.
The ingle nook is shown on Plate li., and a view of the oriel on Plate
lii.
PLATE LIII.
PAIR OF COTTAGES.
SEE PAGE 50.
PLATE LIV.
PAIR OF COTTAGES—BACK.
SEE PAGE 50.
The pair of cottages shown in Plates liii. and liv. have outer
porches, whereby the size of the hall is reduced. A separate view of
one of them is given on Plate lv.
PLATE LV.
PORCH.
SEE PAGE 50.
In this example, as in the former also, the outlook at the back of the
house is to be preferred to that in the front, and as should always be
done when the aspect is favourable, the principal rooms are placed at
the back. There is in this instance a west prospect, with a delightful
view of undulating woodland and distant hills. The forecourt affords a
pleasant outlook from within the house. The lowness of the eaves has
the effect of giving the pair a very homely and cottage-like
appearance. The height of the bedrooms in the former example is 8 ft.
3 in.
PLATES LVI. AND LVII.

SINGLE COTTAGE.
PLATE LVI.
SINGLE COTTAGE.
SEE PAGE 50.
Plate lvi. gives a single cottage of a plan similar to the last, with
enlarged accommodation and somewhat different treatment, namely:
—
Rough-cast from ground, with tarred plinth; oriel window to first floor,
with the introduction of a little colour in parquetry, which is also applied
round the small window over the entrance, and a half-timber porch
glazed with leaded lights, having coloured centres of rich glass. The
cloak space is here converted into a china pantry.
A separate view of the porch is shown on Plate lvii.
PLATE LVII.
PORCH OF SINGLE COTTAGE.
SEE PAGE 50.
GENERAL NOTES.
The Bath.—The bath, without
which no house is nowadays
regarded as complete, should be
supplied in all cottages, however
small. At Bournville, wherever
there is no bathroom, the bath is
placed in the kitchen, this room
being considered the most
suitable: hot water is here at
hand, and, as there is usually a
fire in winter, it is both more
convenient and comfortable than
in one of the bedrooms, where
the space can be ill-spared,
especially where there are
children. Even in the kitchens of
these small cottages there is
necessarily none too much
space, and various devices have
been employed to prevent the
bath being an inconvenience
when not in use. One way of THE PATENT ADJUSTABLE CABINET
disposing of it is to sink it into BATH.
the floor near the hearth, the
boarded covering serving as a standing or draining board when the
bath is in use. Another way, where there is a little more room to spare,
is to fix it on the usual floor level, and make its cover serve as a settle
or table. The introduction of the Patent Adjustable Cabinet Bath,
however, is better than either of these methods. In this arrangement
the bath is hinged at the bottom of one end in order that it may be
easily lowered from and raised back into the cabinet, where in its
vertical position it is no inconvenience when not in use. In the hinge a
waste pipe is introduced. With this bath not only is there a gain of
space, but the bath may be used with a saving of time and labour, and
without fear of deluging the floor. Above the cupboard in which the
bath is kept are convenient shelves. The cost of the bath and cabinet
is about £3 5s. The illustration on the last page shows a bath of this
kind fitted in one of the Bournville cottages.
Another patent bath
used at Bournville in
cottages of larger size but
not sufficiently large to
admit of a bathroom is
Cornes’ Combined
Scullery-Bath-Range and
Boiler. The patent utilises
to the fullest extent the
heat of the kitchen, so
that, in addition to the
economy of space, there
is a further economy of
fuel to the householder.
The heating and cooking
range forms a great part
of the division between
the kitchen and scullery-
bathroom, the flue being
coursed over the head of
the bath. In the centre of
the range is the grate,
with an oven on one side
and on the other a
CORNES’ PATENT BATH. twelve-gallon boiler, in
which water is kept hot
for domestic purposes. Boiling water can be obtained by raking down
live fuel into a small secondary grate under the boiler through a small
hole made for the purpose. If desired, clothes can be boiled in the
boiler and access to it from the scullery may be gained by opening a
curved door. Owing to its open construction there is no risk of
explosion. Further developments have been made in the way of
providing a folding door in front of the range, which will shut off the
boiler from the kitchen when necessary. The scullery-bathroom, which
contains about 36 superficial feet, is fitted with a full-sized iron
enamelled bath, supplied with hot water through a pipe from the range
boiler and with cold water from the cistern, or through a shower-bath
sprinkler fixed overhead, so that this latter luxury can be enjoyed by
simply turning the tap. The introduction of White’s Patent Steam
Exhaust effectually prevents the steam from permeating the other
rooms of the house. An illustration is here given showing Cornes’
patent fitted up.
The Ingle Nook.—Like many old-time features which have been
revived during the last few years, the ingle nook has perhaps been a
little overdone. The ingle is intended to serve as a cosy retreat in a
spacious room, and it should not be introduced in a room the size of
which is insufficient to warrant its existence. On this account it is
usually undesirable to provide ingle nooks in cottages, except in those
with the large living-rooms. Comfort should always be the object in
view in the construction of the ingle, but in many modern examples
this is sacrificed to over elaboration and that straining for effect which
shows that it was designed for ornament and not for use. No doubt an
effect is sometimes gained, but the usefulness of the ingle is so far
sacrificed that not infrequently one of most inviting appearance will be
found to possess inadequate seating accommodation even for a
single person.
ELEVATION
PLAN
SMALL COTTAGE INGLE.
The ingle, to be comfortable and useful, should not be less than 10

ft. 6 ins. in width by 4 ft. 6 ins. in depth. If it is smaller lengthways the
heat from the fire will be too great, while if less deep there will be
insufficient accommodation at the sides for two persons without
projecting the seats into the room, which can only be satisfactorily
done, perhaps, when the side of the ingle is in line with that of the
room. A reasonable height is 6 ft. 6 ins. A pleasing way of treating a
cottage ingle is to introduce a step up of about 3 ins., with an oak
curb, and to tile or quarry the whole recess, as illustrated by the
accompanying drawing. This ingle, which is provided in the pair of
cottages with the large living-rooms dealt with earlier (Plate xx.), is
constructed as follows:-¾ in. match-boarding is nailed to studding,
which has stout angle-posts to support the beam above; along the
side of the latter a 7-in. by 1-in. shelf is carried by small wooden
brackets; and the wood seats are of 1¼ ins. in thickness by 1 ft. 4 ins.
from back to front, at a height of 1 ft. 3 ins. or less from the floor. The
introduction of the ingle here is advantageous because some privacy
is thus afforded in a room which is entered directly from the road. The
match-boarding in this case is continued, and forms a framework for
the tile-surround of the grate, giving an appearance of unity to the
nook, while the simplicity of the material is pleasing and restful. The
insertion of a mantelpiece different in character should be avoided.
Some interest may be given to the centre of the fireplace by inlaying a
little ebony in simple forms.
For drawing-rooms of larger houses the back of the nook might be
panelled, the seat upholstered, and the panels filled in with tapestry.
White wood looks well, and the fireplace might be built up with glazed
brickettes. The ways of treating the more expensive ingles are so
numerous, however, that it would be of little use making definite
suggestions.
The ingle nook of one of the larger cottages is illustrated on Plate li.
Chimneys.—The economy of grouping chimneys, and the
desirability of carrying them to the highest point of the roof to avoid
down-draughts, has already been mentioned. Generally speaking, for
cottages, the simpler the chimneys are the better, and they should all
be of hard burnt bricks, and the top courses built in cement. Diagonal
chimneys are pleasing, but expensive, and on an estate should only
be used occasionally. The Dutch chimneys, built up with corners of
brick and covered with stone slabs or 12-in. drain pipes, as frequently
seen in Holland and Belgium, are picturesque (see Plate xxii.), but
care has to be exercised in their construction. Though they are often
regarded as being liable to smoke, it may be pointed out that in many
cases their employment is the only remedy for a smoky flue. Outside
chimneys, it will be borne in mind, are always expensive. Chimney
pots do not improve the appearance, but sometimes they are a
necessity. In these cases the simple or plain tall-boys are
recommended, and the colour—whether of soft red or buff—should be
chosen to suit the design and colouring of the cottage. As a variation
of these there are the beehive pots, the main idea of which is to keep
down the height.
Windows.—The casement window is cheaper than the sash
window, and if beauty of effect is also to be considered its adoption is
further desirable. Its simplicity and homeliness of appearance render it
extremely fitting for the cottage. The old difficulty of cleaning may now
be obviated by a very simple device introduced at Bournville, that of
causing the window to open upon a pivot in the centre, inwardly as
well as outwardly, which admits of the outside of the fixed pane being
easily reached by the hand.
The sash window, while objectionable in the form frequently used,
may yet be made suitable for cottages; but it should be divided, and
the proportions very carefully studied, say 9-in. by 11-in. panes, and
the bars not less than 1 in. in thickness. It should be brought forward,
showing the full width of the boxing. The sash window, however,
necessitates an additional height to rooms.
Bricks.—As regards bricks, it is well as far as possible to avoid
those which are mechanically made (the pressed stock-brick) and to
use the hand-made bricks from local yards. The brindled Staffordshire
bricks are largely used at Bournville; they are very suitable for cottage
building where the position is not too exposed. A pleasing variety of
colour is introduced at a low cost, the tint being a bright cherry red
blended with blue and purple, the blue being quite different from the
dead blue-black of the vitreous brick. For inside work the common red
wire-cuts are suitable.
It is a mistake to suppose that a good effect cannot be obtained by
the use of the cheaper makes of bricks, a remark which also applies in
the case of the London stock-bricks, so long as they are not uniformly
selected; a good effect may be gained, for instance, by using a few of
the darker ones indiscriminately with the cream-coloured ones. The
splash of dark colour caused by the black ones coming together is by
no means undesirable. A good example of an effective use of these
bricks is to be seen at Brewer’s Estate, London.
Roof Covering.—The materials to be employed in roofing depend
upon the style of cottage, and also upon the locality. The Bangor
slates are cheap, and may be an excellent covering as regards
durability; but unfortunately, in the class of cottages here dealt with, it
is rarely possible to get so good an effect with them as with other
kinds. They may be used, however, in the whitewashed cottage, so
long as the smaller sizes are selected. Hand-made roofing tiles, and
thick Welsh green and rustic Precelly slates may be recommended, as
also the Peake’s & Hartshill hand-made tiles.
Pantiles are cheap, but should only be employed on unbroken roofs
having few valleys, where it is less difficult to keep out the wet. The
roof should be steep, the angle in no case being less than 45 degrees.
Before covering, care should be taken to ascertain whether they are of
good manufacture, and whether they are porous or not. There are
sometimes pantiles of an indifferent quality on the market; and, if this
precaution is not taken, a roof may have to be stripped and re-tiled.
Where they have been used and have afterwards been found to be
bad they may be tarred, as are wood coverings in Norway and
Sweden. It is always essential that the services of a practised layer of
pantiles should be secured.
Gables should have damp courses under the coping to shield them
from frost and wet.
Roof ridging should have careful attention, and it is wiser to
suppress rather than to sharpen, the better to obtain that rustic
appearance suitable to a cottage. Many fantastic ridges, with vulgar
finials, are employed in the building of small suburban villas, of a more
or less sharp-pointed character, and of a depth out of proportion to the
roof, which gives an unpleasant harshness to the general appearance.
With the principle in view that the sky-line should be softened as much
as possible, the brindled hand-made half-rounds should be used. With
green slates, ridges of blue are the most suitable, as the colours
harmonise. Experience will probably show that the red and buff ridges
will not stand the weather so well as other kinds.
Wall Spaces: Rough-cast—Whitewash—Half-timber.—
However strong may be the temptation to introduce a variety of colour
upon exteriors, it is advisable with cottages of the class dealt with to
refrain from so doing. It is best to get the colour in masses, treated
broadly—say, each house, as far as wall surfaces and roof are
concerned, of one colour; for where the cottages stand close together,
or even where they are semi-detached, sufficient contrast or relief is
afforded by contiguous cottages treated differently, and in the case of

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Volume 30, 1994.

Sokolova, M. N. Trophic structure of abyssal macrobenthos.

Rodhouse, P. G., Elvidge, C. D. and Trathan, P. N. Remote sensing

Volume 45, 2003.

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Volume 54, 2008.

An Introduction to the Biology of

Advances in Marine Biology, Volume 57 # 2010 Elsevier Ltd.

and mortality that were mainly based on Mauchline (1960). Mauchline

2. Morphology and Taxonomy

Whitish translucent, spotted above with red. Cephalothorax equalling a

Figure 1.2 Petasma of an adult male Meganyctiphanes norvegica, which is a modification

Figure 1.3 Thelycum of an adult female Meganyctiphanes norvegica with 2 attached

haddocks (Sebastes norvegicus). The genus Nyctiphanes was erected by G. O Sars

compared to many other major arthropod groups. Consequently, phyloge-

these ontogenetic characters were used in cladistic analysis to generate a

the evolutionary history of E. superba and closely related species (Bargelloni

2.4. Photophores and their function

Figure 1.5 The pattern of photophore illumination in Northern krill (Meganyctiphanes

Light emission is directional, with a dark pigment masking emission in

3.2. North Atlantic west coast

3.3. Arctic limits

Around Iceland, adults can occur in numbers comparable to other well-

3.4. North-east Atlantic and fringes

and adjacent fjords such as Gullmarsfjord (Heegard, 1948). Heegard (1948)

3.5. Mediterranean and south-eastern limits

3.6. Distributional trends and changes

recent extent of the northerly distributional limits, along with increased

4. The Sampling of Northern Krill

4.1. Net sampling

Purpose of net deployment Net and deployment parameters

Purpose of net deployment Net and deployment parameters

‘Fipec’65 18, 38, Vertical rear

4.2. Active acoustics

and TS (Klevjer and Kaartvedt, 2006). Moored acoustic Doppler current

4.3. Alternative technologies for sampling and observation

5.1. Benthic interactions

organic matter transport to the bottom to be 7–12 mg C m 2 d 1. This

5.2. Krill production and predator consumption

The accommodation in the two houses differs owing to the aspect. If

These plates illustrate two pairs of cottages of two storeys each,