F.-W. Tesch Dr. Agr. (Auth.), P. H. Greenwood D.sc. (Eds.) - The Eel - Biology and Management of Anguillid Eels-Springer Netherlands (1977)

THE EEL
THE EEL
Biology and Management of
Anguillid Eels
F.-W. TESCH, Dr. agr.

Bjologjsche Amral! Helgoland, Hamburg
Translated by Jennifer Greenwood
ENGLISH EDITION EDITED BY
P.H. GREENWOOD, D.Se.

Department of Zoowgy,
British Mweum (Natural History )
This first English-language edition of Der Aal by F.-W. T esch, has

been updated throughout by the author, and physiological sections
have been revised and extended by
I.W. HENDERSON, Ph.D.
Department oj Zoology,
University of Sheffield
LONDON
CHAPMAN AND HALL
A Halsted Press Book
John Wiley & Sons, New York
First published, 1973, by Paul Parey, Hamburg and Berlin
under the title Der Aal, written by F.- W. Tesch
© 1973 Verlag Paul Parey, Hamburg and Berlin
This revised and updated, English-language edition,
a translation approved by F.- W. Tesch,
first published, 1977, by Chapman and Hall Ltd.
ISBN-13: 978-94-009-5763-3 e-ISBN-13: 978-94-009-5761-9

001: 10.1007/978-94-009-5761-9
© 1977 Chapman and Hall Ltd.

Soflcover reprint of the hardcover 1st edition 1977
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,
induding photocopying and recording, or in any
information storage or retrieval system, without
permission in writing from the publisher.
Distribution in the U.S.A. by Halsted Press,

a Division of John Wiley & Sons, Inc., New York
Library of Congress Cataloging in Publication Data

Tesch, Friedrich-Wilhelm.
The eel.
Translation of Der Aal.

"A Halsted Press book."
Bibliography: p.
Includes index.
1. Anguilla (Fish) 2. Eel fisheries. I. Title.
Contents
Foreword page xi
Preface Xlll
I. Body Structure and Functions [
l.l Skeleton [
I.l.l Skull [
I. 1.2 Pectoral girdle 6

I. 1.3 Vertebral colwnn 6
J.I.4 Fins 8
1.2 Skin and musculature 9
1.2.1 Structure and function of the skin 9
1.2.2 Scales [2
1.2.3 Pigmentation [4
1.2.4 Teeth [9
1.2.5 Muscuiarure 2[
1.3 Respiratory organs and swimbladder 23
l.4 Feeding and organs of digestion 28
1.4.1 The gastrointestinal tract 28
1.4.2 Pancreas 32
1-4-3 Liver and gall bladder 34
1.5 Circulation of the blood 35
1.5. I Cardiovascular system 35
1.5-2 Blood 38
1.6 Urinogenital system 4[
1.6. I The kidneys 4[
1.6.2 The gonads 43
1.7 The endocrine system 58
1.7.1 The pituitary gland 58
v
VI
I.7.2 The thyroid gland 62

1.7.3 The urohypophysis 63
1.7.4 The interrenal tissue (adrenocortical homologue) 64
1.7.5 Ultimobranchial bodies 64
1.7.6 Corpuscles of Stannius 65
I.7.7 Thymus gland 65
I.7.8 The kidney 66
1.8 Nervous system and sense organs 66
1.8.1 The brain 66
1.8.2 Eye 68
1.8.3 Olfaction 73
1.8.4 Hearing and the lateral line sense organs 79
2. Eel Species, their Developmental Stages and

their Distribution 81
2.1 The European and American eels A. anguilla and
A. roslrata 83
2.1.1 Spawning grounds and larval forms 83
2. 1.2 Continental distribution 102
2.2 Indo-Pacific species IIO
2.2. I The south-east African region: A. marmorara,
A. nebulosa iabiala, A. mossambica and
A. hiealor III
2.2.2 The temperate north Pacific zone: The
Japanese eels A.japonica and A. marmorata 114
2.2.3 The south-east Pacific species from the
temperate zone: A. australis and A. dieffenbachi 118
2.2.4 The tropical eels
A. celebesensis, A. megastoma, A. interioris,
A. ancestralis, A. nebulosa, A .marmorata,
A. reinhardti, A .borneensis, A. biwlor, A. obscura 122
2·3 Zoogeographical relationships 129
3· Post-Larval Ecology and Behaviour 133

3. 1 The glass eel and elvers: their migration on the
Continental shelf and into freshwaters 133
3.1.1 Migration in the sea 133
3.1.2 The movement into freshwater 139
3.1.3 Human intervention 150
3. 2 Migration of pigmented young eels 154
3.2.1 Dependence on environmental factors 154
3.2.2 Human involvement in the ascent of young eels 162
vii
B The 'Yellow eel' stage to the 'Silver eel' stage 169

3.3.1 Feeding 169
The period of activity and food intake-Seasonal
variations in diet-Choice of food and body
size-Choice of food and head width-The
dependence of diet on food availability-Choice
offood
3.3.2 Age and growth 184
Methodological problems-Differences in growth
between males and females and the age and
length at which migration begins- Head width
and growth-Differences in growth due to
environmental factors-Interspecific differences
in growth-Theoretical pattern of growth-
Length and weight
3.3.3 Location, habitat and behaviour patterns 208
Refuge in tubes and hollows, and contact with
members of the same species-Water depth-
Survival in air- Localization and homing
3·4 Silver eel migration 220
3-4. I Seasonal variation 220
3.4.2 Variations in the time of day, and the influence
of light 224
3.4.3 Correlation with lunar phases 224
3.4.4 Hydrological and meteorological factors 226
Low pressure areas-Water level and currents in
rivers-Tidal currents- Wind and currents-
Temperature and depth-Salinity
3.4.5 In subterranean waters 234
3.4.6 In the sea, with special reference to the Baltic 235
3·4· 7 Behaviour and speed 239
4· Fishing Yields 241

4·1 The development of eel fisheries 241
4·2 Annual and regional fluctuations in yield 244
4.2.1 Variations over the whole North Sea region 244
4.2.2 The effect of differing proportions of males and
females on variations in catch size along the
southern coast of the Baltic and in inland waters 244
4.2.3 The relationship between the west wind and
glass eel invasions 246
4.2.4 The influence of seasonal temperature levels 248
viii
4.2.5 Catch variations in Canada 248

4·3 Variation in catch size throughout the year 249
4·4 Population density and catch sizes from an economic
and ecological viewpoint 25 2
4.4. I Regional differences 25 2
4.4.2 The eel in competition with other fishes and
with crustaceans 25 6
4-4.3 Improving yields by stocking 26 5
4.4.4 Summary of yield and stocking success 270
5· Fishing Methods 272

5. 1 Eel traps 273
5. I. I Eel baskets 273
5.1.2 Fyke nets 277
5.1.3 Pound nets 281
5-2 Stow nets 285
5.2.1 Staked stow nets 286
5.2.2 Anchored stow nets 286
5.2.3 Onerboard stow net 288
5-3 Permanent eel traps 292
5.3.1 Eel weirs in rivers 292
5.3.2 Eel traps in streams and rivers 297
5·4 Seines and trawls 300
5·5 Angling and line fishing 304
5·6 Lift nets, spears and other equipment 315
5·7 The use of electricity 318
5. 8 Combination with light 322
6. Eel Culture and Live Storage 328

6.1 Eel farming in Japan 329
6.1.1 The importance of eel farming 329
6. 1.2 Pond systems and yields 330
6. 1.3 Rearing techniques 332
6.1.4 Feeding 333
6. 1.5 Profit and expenditure 335
6.2 Experiments on eel farming in Europe 335
6·3 Live storage 340
7· Diseases, Parasites and Injuries 342

7-I Virus and bacterial diseases 342
7. I. I Cauliflower disease 343
ix
7.1.2 Fresh- and saltwater eel diseases
Freshwater eel disease-Saltwater eel disease 346
7·2 Infestation by fungi, protozoa and metazoa 35 1
7·3 Teratological and other defects 356
8. Trade and Processing 363

8.1 International trade 363
8.2 Prices 364
8·3 Quality 367
References 37 1
Index 423
Foreword
by Dr P.H. Greenwood
British Museum (Natural History), London
Dr Tesch's wide ranging account of anguillid eels impinges on the

interests of many biologists; it is not simply a specialized tome narrowly
aimed at ichthyologists and fishery scientists, rather it provides a
source of primary reference and a comprehensive sununary of informa-
tion that is not likely to be superseded for a long time. It is significant
that the bibliography includes references to learned journals concerned
with physiology, pharmacology, taxonomy, genetics, zoology, endo-
crinology, botany, ecology and environmental interactions. Such is the
breadth of interest in the Anguillidae.
Few fish species have been subjected to as detailed review as Dr Tesch
gives for the (wo Atlantic species of Anguilla. An equally comprehensive
resume of research into the fourteen, rather less well-studied Indo-
Pacific species gives balance and reciprocal illumination to several
biological problems posed by these similar but quite distinctive species.
The German language edition of Dr Tesch's book was well received
and much consulted. The present English edition should greatly increase
its readership, and more especially since the author has revised and
updated the entire text. Those sections concerned with facets of eel
physiology have been expanded and revised in collaboration with
Dr Ian W. Henderson himself actively engaged in investigations into
certain aspects of anguillid physiology. The sections on hormonal
physiology of the eel have received considerable attention, as it is in this
sphere that great progress has been made in recent years. Indeed, studies
of eel systems have contributed greatly to furthering knowledge of fish
physiology generally. The final product is in my opinion an even more
useful and comprehensive volume that was its predecessor, Der Aal.
xi
Preface
A little more than sixty years ago, the life cycle and transoceanic
migrations of the Atlantic eels were first revealed through the work of
the Danish scientist, J. Schmidt. These studies and the way in which
they were conducted are still held in high regard, and Schmidt's work
is recounted in almost every scientific and popular book on eels.
Less widely known, however, are the results of research into the
biology of related eel species from the Pacific and Indian oceans. In this
book data relating to all species of the genus Anguilla have been brought
together for the first time, so that interspecific comparisons can now be
made more readily.
Although research into eels' spawning grounds may once have
appeared the most spectacular contribution to OUf knowledge of these
animals, the eel's anatomical and morphological characteristics are of
equal interest. A creature with such a variety of unusual ecological
characteristics is bound to show a corresponding variety of physical
features, and the number of scientists studying the morphology and
physiology of the eel thus continues to increase. This is the case not
only in Europe and Japan, where the eel is one of the more widely
distributed species of fish, but also in North America, where it has a
much more restricted distribution. A detailed description of the eel's
physical characteristics, its behaviour patterns, and its relationship
with the environment seems urgently required. The eel should not be
used as an experimental animal without an awareness of the special
features in its ontogeny and growth. For example, after a lecture on
respiratory physiology at an international symposium, a member of the
audience asked if the experimental animals were 'yellow' or 'silver'
eels. The lecturer replied that neither of these varieties had been avail-
able, only species of Anguilla anguilla had been used!
Xl1l
xiv
In recent years there has been a marked increase in research on the
physiology of eels. Dr I. W. Henderson is to be thanked for the part he
has played in incorporating this new information into the English
edition.
The sections on sexual maturity and reproduction, pond culture
and pathology, migration and homing ability-the latter two being
my own particular field of investigation-have been greatly expanded
and include references to work published in the last 4 or 5 years as well as
reviews of the older literarure, some of which is not easily accessible.
Finally, I must express my grateful thanks to all those who have
helped in the production of this book. I feel particularly indebted to the
translator, the editor and the publisher, who have worked so
conscientiously to ensure the prompt appearance of the English edition.
Hamburg F.-W.T.
1976
I
Body Structure and Functions
From a physiological viewpoint the eel is a parcicuiarly popular

experimental animal. This is due not only to its extremely marked
resistance to many sorts of experimental conditions, but also to a large
number of unusual characteristics which distinguish the eel from other
species of fish. Included among these characteristics are the osmo-
regulatory capacities of this euryhaline organism; its phases of
differential activity and behaviour patterns; its multistage meta-
morphosis during ontogenetic development; its great endurance, and
its ability to navigate during migration; and lastly, even its unusual
body shape.
The range of publications in these fields of study increased greatly
in the years following the Second World War. The limits of this book
would be exceeded if one tried to deal with all these studies in great
detail, or even if one anempted merely to refer to each separately.
Therefore, in the morphological and physiological parts of this chapter,
particular reference will only be made to those papers which are in
some way connected with the ecology of this interesting fish.
I. I Skeleton
I.I.I Skull
The skull of Anguilla is illustrated in Figs. I and 2. The shape and
structure of the skull are peculiar to the order Anguillifonnes and
I
2 THE EEL
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Fig. I Skull (neurocranium) of A .japonica. A, from the side; BJ from above; C, from
below ; OJ from behind. as, Pterosphenoid ; ba, Basioccipital ; eo, Exoccipitai j
ep, Epiotic; fm, Foramen magnum ; fr, Frontal ; if, Orbit ; os, Orbitosphenoid ;
pa, Parietal ; pr, Prootic; ps, Parasphenoid j pt, Pteroticj pv, Premaxillo-
ethmovomerine bloc; so, Supraoccipital ; sp, Sphenotic (after Matsui &
Takai, (959).
differentiate all eels from members of other orders. For example, the
upper jaw is partly fused with the cranium; the maxilla and the pre-
maxilla-etlunovomerine bloc (Figs. I and 2, pv and rna), have taken
over the function of the upper jaw. Due to the unusual arrangement of
the teeth on this bone, it also provides distinctive features important
BODY STRUCTURE AND FUNCTIONS 3
'1
O' Q'
Fig. , Cranial skeleton (A) with suspensorium, jaws, opercular bones and hyoid
arch (8 ) of A. japonica. See below and Fig. I for abbreviations : aa, Angulo-
articular-retroarticular ; as, Pterosphenoid; bo, Basioccipital ; bh, AnterohyaJ ;
ch, Posterohyal ; de, Demary; gh, Basihyal ; hm, Hyomandibula ; rna, Maxilla;
op, Operculum; po, Preoperculum j pp, Palatopterygoid ; qu, Quadrate ;
su, Suboperculum ; uh, Urohyal (after Matsui & Takai, 1959).
in classification within the family Anguillidae (see p. 100) . The basis-

phenoid bone, which is normally to be found at the base of the skull
above the parasphenoid (Figs. 1 and 2, ps) (Harder, 1964), is absent in
all Anguilliformes. Unlike other members of the Anguilliformes an
orbitosphenoid is present in the Anguillidae. A sympletic, which links
the quadrate of the lower jaw to the hyomandibula and thus to the
cranium, is not found in the eel.
In addition to their long bodies, members of the Anguilliformes have
a very narrow head which enables them to burrow in sand and mud, and
to creep into small holes. The narrowness of the head also necessitates
an adjustment in the position of the gill apparatus. The latter has
shifted from below the skull, where it is normally found in bony fishes,
to just behind the skull. As a result! the gill region is very long and the
pectoral girdle is not attached to the skull. Although in Anguilla the
opercular and subopercular bones are very small, the branchiostegal
membrane and its supporting skeleton are well developed (Fig. 2). The
branchial skeleton and its musculature, together with the branchiostegal
apparatus, play an important role in the production of positive and
negative pressures during food intake (Alexander, 1970). Particularly
marked differences in the morphology of the gill arches and other parts
of the branchial region can be seen within the Anguilliformes (see
Nelson, 1966, 1967 ; Smith & Castle, 1972; Kusaka, 1973).
4 THE EEL
A comparative study of the skull and axial skeleton in Anguilla

rostrata, Neoccmger vermijormis, Moringua edwardsi and Pythonichthys
sp. has recently been made by Smith & Castle (1 972). Illustrations and
further, more detailed, information on skull structure in A. anguilla in
connection with 'broad-' and 'narrow-headedness' in eels can be found
in Torlitz (1922). This author also described the skull structure of the
leptocephalus larva and of the glass eel.
The differences between the skull of the glass eel and the adult eel
are only slight. 'In the glass eel, the bones in the anterior cranial region,
the basisphenoid and the pterosphenoid are the least well developed;
large spaces, containing only connective tissue, lie between them.
A mucus canal bone, a self-contained unit, rests on the pterotic; the
anterior section of the mucus canal bone forms the narrow, tubular
process of the pterotic seen in the skull of the adult eel. This process
juts out between the frontal and the sphenotic ... ' (Tertitz, 1922).
The sphenotic is very poorly developed as compared with its condition
in the adult eel.
There are, however, marked differences between the skull of the
glass eel and the skull of the leptocephalus: 'If we compare the total
morphology of the glass eel's head with that of the leptocephalus, we
see considerable differences, particularly with respect to the ratio
between the length of the neural skull and that of the facial part. While
this relationship in the leptocephalus is in the ratio 1 :1, in the glass eel
it is I' 5: 1 . .. Greatest skull width to greatest skull height is in the ratio
1:1, 1'25:1 respectively in the two developmental stages. This differ-
ential growth continues until the adult stage is reached, when the ratio
is 2: I. The increasing length of the neural skull appears to be connected
with the relative increase in brain size; the increasing width of the upper
jaw must be associated with the growth of the musculature.'
'The skull gets wider and wider and, at the same time, becomes
relatively flatter ... The very prominent sphenotics are particularly
influential in bringing about this increase in width - i.e. as structures
which are causally connected with the masticatory muscles' (Torlitz,
1922).
The spatial relationship between the individual parts of the cranium
and the different areas of the brain also change. If one studies these
developmental relationships more carefully, it is hardly surprising to
find that during ontogeny considerable variations are produced.
Previously, the occurrence of the two extreme varieties, 'broad' heads
and 'narrow' heads, led to the belief that there were two genetically
distinct forms of European eel. Publications by Bellini (1907; see Walter,
1910), who had already discovered differences in the relative head width
of glass eels, were probably largely responsible for this belief.
In Japan more recent studies on A. japonica revealed differences in
the head width of eels described as being at the glass eel stage (Matsui,
1952), although in fact the eels were already strongly pigmemed as can
be seen from a photograph of a specimen used in this study. A frequency
distribution curve of the various head widths in these young eels plotted
against body length has, however, only one peale The same was seen
to be the case in larger Japanese eels. Similar findings have been
published by Thurow (1958). In other words, according to these results
these eel populations are not made up of two distinct forms. Had this
been the case, it would have been expressed as two peaks in the frequency
distribution. The fact is that 'broad' heads and 'narrow' heads simply
represent the less usual variations of a modal type.
On the basis of biometric studies on leptocephali, glass eels and
larger eels, Torlitz (1922) is of the opinion t1:.lt 'narrow-' and 'broad-
headedness' first appears during the course of ontogeny. Thurow (1958)
also believes that he has evidence to show that the two types of head
shape can be determined by providing the eels wiJt different types of
prey - i.e. head shape is the result of environmental factors. Since
Thurow only had access to relatively large eels, it is questionable whether
his measurements can be extrapolated to glass eels, and therefore
whether his data can be interpreted in this way. The fact that one or
other of the forms predominates in certain waters, however, supports a
theory of environmental influence. In Heligoland, for example, 'broad'
heads were found almost exclusively (see p. 193). All the authors who
have studied the European eel and the Japanese eel agree that the
'broad-headed' form shows better growth than does the 'narrow-
headed' one (e.g. Bellini, 19<J7, according to Walter, 1910; Bauch, 1954;
Rahn, 1955a; Thurow, 1958; see p. 193).
Marked differences between male and female Japanese eels are to be
seen in the shape of the head. For example, the distance between the
eyes, i.e. the interorbital width, is greater in males than in females.
The diameter of the eye is also larger in males. The distance from the
tip of the snout to the insertion of the pectoral fin is. however, greater
in females than in males. The same applies to the distance from the tip
of the snout to the dorsal fin origin, and the snout to anal fin distance.
Similar differences have been known fonome time in the European eel
(Nordquist, 1917).
The detection of these secondary sexual characteristics is, however,
the result of a statistical analysis of a great deal of data, and none can
6 THE EEL
therefore be considered suitable as a ready means for distinguishing

individual males and females on external features alone. There are,
nevertheless, experienced workers who, using these parameters, are
almost always successful in separating males from females, even in the
European eel.
I. 1.2 Pectoral girdle

The pectoral girdle in the Anguilliformes has a simple form. This is
probably due to the often marked reduction of the pectoral fins which
may be rudimentary or absent altogether. The anguillids have a supra-
cleithrum, a cleithrum, scapula, coracoid and, as a special characteristic
of this family, at least seven fin radials. In comparison with other
Anguilliformes such as Neoconger vermlformis, Mon'ngua edwards': and
Pythonjchthys sp., Anguilla rostrata has a relatively small
supracleirhrwn.
1.1.3 Vertebral column

The vertebral column in eels is particularly interesting from morpho-
logical, functional and systematic points of view. Hardly any vertebrate
order is as polymorphic in this respect as are the Anguillifonnes. This
polymorphism is illustrated by the number of vertebrae. There are
some species with as few as 75 vertebrae, while others may have up to 260
(Berg, 1958). Even within the anguillids the number of vertebrae is one
of the most important diagnostic features at the species level.
Unfortunately there is still no comparative study on the morphology
of the vertebral column in the various species of Anguilla. As has been
shown by Japanese studies on the osteology of other members of the
order such as the Muraenesocidae and the Congridae (Takai, 1959;
Asano, 1962), there are some especially diagnostic characteristics in the
vertebral column.
Figure 3A-C shows various sections of the vertebral column of
A. japonica. The vertebral column of this species, which has a similar
number of vertebrae to the European eel (Table 5), is subdivided as
follows: total number of vertebrae 116; the 44th is the last abdominal
vertebra; the 38th is situated near the anus; pleural ribs occur on the
7th to 38th vertebrae; the haemal arches begin near the region of the
45th vertebra which then is the first caudal vertebra (Fig. 3) ; dorsal
intennuscular bones (epineurals) occur on the 1st to the 86th vertebrae,
ventral intennuscular bones (epipleurals) occur on the 38th to the 86th
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Fig . 3 Anterior (above), centtal (centre) and posterior (be!ow) vertebra of A.japonica. A, Lateral view, 1St to 5th vertebrae; B, Ventral
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view, 1St to 5th vertebrae; C, First vertebra, ante rior view; D, 5th vertebra, anterior v iew; E, Last abdominal and first two
caudal vertebrae; F, 44th vertebra, anterior view (last abdominal vertebra, without intermuscular bones); G, 45th ven ebra,
anterior view (firSt caudal vertebra); H, 46th vertebra, anterior view (wit hout intermuscular bones). at, Anal fin ray;
ce, Centrum ; cr, Caudal fin rays; dr, Dorsal fin ray; ee, Endoskclctal portion o f the fin; e l, Epipleural intennuscular bone;
en, Epineural intennuscular bone; ha, Haernal arch j hp, Haemal canal ; h s, Haemal spinej ih, Anal pterygiophorej in, Dorsal
pterygiophorej na, Neural arch j np, Ne ural canal; n s, Neural spine j pa, Parapophysisj ra, Distal radialj tv, Hypural
(compound) (after Matsui & Takai, 1959).
....
8 THE EEL
TabJe I Comparison of the vertebral columns of A . anguilla and C. conger (after

Ford, 1938)
A. anguilla C. Cl»IglT
Autogenous neural arches : Vertebrae nos. 1St-5th lSI-16th or 17th
R elatively high neural procencs 2nd, and less 1st-16th or 17th

or spines frequently
Vertebrae nos. 3rd -8th
Double structure of the parapophysis

Vertebrae nos. None 2nd-:;th or 6th
Undulating surfa<;e Structure on

the venebral centra Rare Frequent
Parapophyses and ventral posuygapophyses in the

anterior body section Separate Merged
The last abdominal vertebrae and a large number of

caudal vertebrae have lateral apophyses in addition to
ventral parapophyses Not present Present
vertebrae. According to Smith & Castle (1972), the distribution of

epipleurals in A. rostrata is slightly different, i.e. they occur on the
34th to the 87th vertebrae. In the Anguillifonnes these bones are
dearly attached by ligaments to the vertebral colwnn and are, therefore,
not to be confused with true intermuscular bones which occur,
additionally, in some anguilliforms, e.g. in the congrid Anago (Asano,
1962). A comparison of A. anguilla and Conger conger (Table I ) shows
how much the vertebral column differs between two families in the
same order, despite the outward similarity in members of the two
families. Smith and Castle (1972) have compared the vertebrae of
A. rostrata with those of Neoconger vermljormis, Moringua edwardsi
and Py,honich,hys 'p.
Lopez, Lee and Baud (1970) have determined the amount of crystal-
line apatite and amorphous calcium phosphate in the bone of the
vertebral colwnn of the European eel. With increasing maturity of the
gonads, decalcification takes place resulting in a great decrease in the
amount of amorphous calcium phosphate. Lopez (1970) has studied
the bone structure. Deformities of the vertebral column are described
in the section on diseases (see p. 357)·
1.1.4 Fins
The unpaired fins in the eel, the dorsal and anal fins, are confluent with
the tail fin. However, as the caudal section of the vertebral column shows
Fig. 4 Oudine of the pectoral fins in yellow eels (right) and in eels at advanced stages
of maturity (left) (after Wundsch, 1953).
(Fig. 3), elements of the caudal fin skeleton are still present in the fonn
of hypural plates. According to Bertin (1956), in Anguilla these hypural
plates have a fairly complex developmental history. The first formed
hypural plate divides longitudinally, the second to appear remains
single, but of the two plates formed later, one (the last to appear) divides
longitudinally, and the other (the antepenultimate in appearance)
remains single. Thus, at one stage there are five hypural elements. In
some individuals this condition persists in the adult, but in others the
two uppennost elements fuse together.
The only paired fins present in the eel are the pectoral fins, which are
not greatly different from those of many other species of bony fish.
They are interesting, however, because of their change in shape during
the later phases of development in adults. While the so-called yellow
eel has relatively wide, spoon-shaped pectoral fins, these fins become
long and pointed (Fig. 4) shortly before the gonads mature. Further-
more, differences are known to exist between the paired fins of male and
female Japanese eels, A. japonica: the pectoral fin in the female is
shorter and more rounded than that in the male (Matsui, 1952).
1.2 Skin and musculature
1.2. I Structure and function of the skin

Eels survive in many diverse, often harsh environments. Such ability
SIems at least in part from the possession of a tough durable integument.
Both the epidennis and, especially, the dennis (corium) are thick.
10 THE EEL
Fig. 5 Structure and proportions of the epidermis (ep) and the corium (cor) of
A. anguilla (above) and Platichthysjlesus (below). sc, Sensory cell ; ce, Club cell;
sqc, Squamous cells; 5g, Stratum germinativum ; me, Mucous or goblet cells;
sub, Subepithelial blood vessels; vi, Vascular loop; s, Scale (after Jakubowski,
196oa, b),
Jakubowski ( Ig60b) compared the skin of seven teleost species, among

them the eel, the flounder (Plalichthys flesus ) and the weather fish
(Misgurnus fossilis ) . Only the latter species displayed an epidermis
thicker than the eel. The epidermis of the eel is 0-263 nun thick, while
that oftbe flounder is 0-036 mm Oakubowski, 1960b; Fig. 5). In the eel,
unlike other teleosts, the corium, particularly that on the head, is
thicker than the epidermis. Numerous, extremely well developed dub
cells occur over the whole epidermis and apparently secrete many
substances which have a protective function (Harder, 1964). Such cells
may be as large as o· I 50 rrun in diameter by 0'025 rrun high in the eel,
BODY STRUCTURE AND FUNCTIONS II
contrasting with species such as the armoured catfish Corydoras palaea-

lus in which they are about 0'025-0'035 by 0'013 mm (Pfeiffer, 1960).
Henrikson and Matoltsy (1968a, b, c) have described the ultrastructure
of the eel integument, and the ontogeny from larval to silver eel stages
has been studied by Aust (1936). Major changes also take place in the
skin during the fourth larval stage of metamorphosis (Table 2).
The thick epidermis, a robust protection against mechanical damage,
is also relatively impermeable to water and electrolytes. Indeed,
Bentley (1962) estimated that 1 ml of water would take 5 years to pass
through 1 cm2 of eel skin at a I'ressure difference of 1 atmosphere!
Transfer of eels from freshwater to brackish water results in an increased
epidermal thickness (Thurow, 1957), and fish chronically adapted to
freshwater have thinner skins than those adapted to brackish water;
moreover, the skin made up 8'5 % of the body weight of freshwater
adapted specimens, compared with 9'4% in brackishwater eels.
Repeated transfer experiments produced equivocal results, however,
and Thurow (1957) suggested that the induced excessive secretion of
mucus evenrually exhausted the cells producing this substance. It was
suggested that the thinner skin of freshwater eels requires regeneration
of mucous cells before the fish can successfully enter seawater. Other
data (Portier, 1938, from Remane & Schlieper, 1958; Portier & Duval,
1922) are also relevant. Eels transferred from low to high envirorunental
osmolarities adapted less well if the skin was rubbed with a cloth to
remove the mucus. In particular, hyperosmolarity of blood and deranged
ion balance occurred. It was concluded that the skin - especially its
mucus secretion - acted as a barrier against fluxes of water and electro-
lytes along osmotic and diffusion gradients. Recently, the secretion
of mucus and its relationship to N-acetylneuramine have been examined
(Lemoine & Olivereau, 1971 ) and prolactin, from the adenohypophysis,
has been implicated as a factor controlling the structure and function
of the skin (Olivereau & Lemoine, 1971).
Another physiological aspect of the eel's integument is its possible
role in gaseous exchange (see p, 23). Jakubowski ( 196oa), citing Krogh
(1924), and Jeuken (1957), reported that fishes such as the eel, and the
equally thick-skinned Misgurnus jossilis, me.!t virtually all their oxygen
needs cutaneously. Furthermore, Byczkowska-Smyk (1958) in a study
of branchial respiration also concluded that a large part of the eel's
oxygen requirement must be met from cutaneous respiration (see also
p. 23). Indeed in air, species such as the eel survive far better than do
purely branchial breathers (Murygin & Anokhina, 1967),
The thick epidermis, therefore, does not seem to prevent gaseous
12 THE EEL
exchange or small exchanges of electrolytes. Jakubowski (I960b)

suggested that the secretory cells - both mucus producing and club
cells - lying between the blood vessels of the dermis and epidermis,
contain sufficient amounts of water to permit ready diffusion of oxygen.
Bolognani-Fantin and Boiognani (1964) and Seuner et al. (1970) have
discussed at length the cytological and chemical bases for the production
of mucus by eel skin. Mucus is of great adaptive significance, not only
when the animals are in water, hut it may also serve to prevent dehydra-
tion when the animals undertake their brief excursions on land, and
may aid survival at low temperatures (Gadeau de Kerville, 1918).
1.2.2 Scales
Unlike other families in the Anguilliformes, the Anguillidae possess
scales. These scales are, however, rudimentary - at least in comparison
with those of other species offish. The scales are relatively well embed-
ded in the upper layers of the corium below the epidermis (Fig. 5), and
are not arranged regularly in overlapping rows as they are in other
fish, but are irregular and, in some places, distributed like parquet
flooring. In general, one row of scales lies at right angles to the next,
although the row of scales immediately above and below the lateral line
lie at an angle of approximately 45 0 •
In Anguilla species the first scales do not develop immediately after
the larval stage is over - as is normal in other bony fish - but appear
much later. Opuszynski (1965) and Matsui (1952) have shown that,
in A. anguilla and A. japonica, individuals measuring 15 em or less do
not have scales, whereas scales are present in most individuals more
than 17 em long. It seems likely that other species of eels also develop
scales very late in onIogeny, though Panrulu (1956) reports their earlier
appearance in A. nebulosa, where specimens of 1 I or 12 cm have
already developed scales. It seems likely that in A. anguilla the formation
of scales is not an age-dependent process, and this has been demonstra-
ted in A. japonica (Matsui, 1952). As regards the region of the body
where the scales first develop, it seems that there may be further
differences between A. nebulosa and the so-called 'northern' eels of
the lemperate regions. In A. anguilla this primary region has been
located only indirectly, by establishing which part of the body had
scales with the greatest number of annual rings (Rahn, 1957). Accord-
ing to Rahn the primary region is in the anal area, directly above and
below the lateral line. In A. japonica too, the first scales appear in this
region, though slightly further posteriorly (Matsui, 1952); this is
<
Fig. 6 Position and distribution of the first scales in A. nebulosa (after Pantulu, 1956).
probably also the case in A. anguilla. From the anal region the
zones of scales develop and spread forwards and backwards along the
lateral line as well as dorsally and ventrally j in nonnally developing
eels two or three years may elapse between the appearance of the first
and last scales (Gernzq,e 1906; Matsui, 1952). The lips of the upper and
lower jaws, the throat, and, it appears, the pectoral fin bases (00, all
remain scaleless. Figure 6 shows the area where the first scales appear
Fig. 7 Photographs of eel scales (after Rahn, 1955a).
in A. nebulosa . It is quite clear that this area lies more caudally than
that indicated by research on the Japanese and European eels.
It seems quite natural that the first scales should develop in the
caudal regions and not in the middle of the body. Other developmental
14 THE EEL
changes that progress in a caudo-rostral direction can be seen in the

formation of the fin supports and fin rays (Bertin, 1956), the appearance
of internal pigmentation during the development of the leptocephalus
and later, the appearance of meianophores in the glass eel (Gilson, 1908).
In other teieosts too, scale formation first begins caudally in the region
of the lateral line (Tesch, 1971).
The morphology of the scales has been described in many papers on
the growth of the eel (see p. 186). The superficial structure of the scale
is rather unusual (Fig. 7). Its contours suggest it is a cycloid scale. It
has, however, a very elongate-oval shape, though there are many
variations. So-called circuli (concentric lines) are also seen in the scale
of the eel. These are not made up of smooth or regular 'furrows' and
'grooves' but from rows of plates which resemble strings of beads.
1.2.3 Pigmentation
The development of pigment provides the most useful means of
recognizing the different ontogenetic stages in the eel. This not only
applies to subepidermal, external pigmentation but also to the internal
pigment of the larval phases. During early development, as is the case
in many other species of fish, the internal pigment also acts as a means
of separating different species. In the leptocephalus, the first internal
pigment develops along the notochord, while at the beginning of
Stage II (see Table 2) it spreads in a caudo-rostral direction (Schmidt,
1906; Gilson, 1908). In comparison to the first, external chromatophores,
which are brownish in colour, the internal chromatophores are black
and relatively large. Species differences appear to exist in the ontogenetic
development of internal pigment. According to Table 2, pigmentation
of the notochord during Stage II begins at the posterior end in A. anguil-
la, whereas in A.japonica pigmentation begins anteriorly (Egusa, 1972).
External pigment also develops during the leptocephalus stage and
is visible as a dark patch on the fin rays of the tail fin during Stage III.
This patch remains recognizable during further development and is the
only form of external pigmentation present until Stage V A (Fig. 8,
Table 2). A characteristic of this stage is that developmental changes are
largely internal; if, however, the temperature is raised, large numbers
of melanophores appear along the whole length of the body.
Developmental differences in external pigmentation are evident
in A. japonica and the European eel. In the Japanese eel the caudal
patch does not develop until after the beginning of Stage IV, and not
in Stage III. In contrast, in A. japonica, the 'skull spot' (see below) and
BODY STRU CTURE AND FUN C TIO NS 15
Fig. 8 T ip of the tail showing the caudal spot in a glass eel at Stage V A; this spot is
even more noticeable at Stage V B (after Gilson, 1908).
a certain degree of rostral pigmentation are already in evidence in

Stage III (Egusa, 1972). These recently recognized differences may
also occur in the other Indo-Pacific species, thus making it possible
to distinguish between the various metamorphic stages of the different
specIes.
The beginning and, to a certain extent, the end of Stage V (also
called V B) is marked by the formation of the so-called 'skull spot', the
Table 2 Development of pigmentation in A . anguilla (abridged from Strubberg,

191 3, and adapted from Bertin, I956)
Stagt Characteristics
I Larva, fully grown lep tocephalus (Fig. 36.1)

II Semilarva, pigmentation on the posterior end of the spinal chord
III semilarva, pigmentation on the nerve chord becomes more extensive, skin pigment
also seen on the tip of the caudal fin
IV Semilarva, pigmentation on the nerve chord reaches the head
VA Metamorphosis complete, ed~ likc in form, no e)(ternal pigment (glass eel) e)(cept
the caudal spot (Fig. 8)
VB No pigment on the back, body or tail region, except for the skull, caudal spots and
some rostral pigment
Development of pigmentation along the whole dorsum, postanal dorsolateral
pigment develops, no clear mediolateral pigment (Fig. 93)
V I All No preanal ventrolateral pigment. Postanal development of mediolateral pigment
(Fig·9b)
No preanal ventrolateral pigment. Clea r preanal development of mediolateral
pigment, postanaliy over almost entire dorsum, pigment rows along the myosep ta
and, in places, doubling of the mediolate ral mc1anophores (Fig. 9C)
Clea r development of preanal ventrolateral pigmentation. Initially, in places, a
doubling of the mediolateral melanophores in the preanal region (Fig. 9'1), postana l
pigment betw~n the myosepta in the ventral region (Fig. 9C), and finaliy, similar
changes in the preanal region (Fig. 9f)
VI B Pigment rows along the myosepta becoming indistinct. Lateral line still recognizable,
as are the individual melanophores on the head, behind and below the eyes, and on
the lower jaw (Fig. 9g)
16 THE EEL
appearance of which is certainly of great physiological significance; this

spot can still be seen in older eels though it does become less distinct.
As Gilson (1908) has shown, this pigment is not produced in the
corium - as is the caudal spot and most of the pigment that appears
after it - but in a sort of fontanelle. Later OD, this hole is occluded as a
result of dermal bone fonnation. In older eels the skull spot is found
in the meninges under the frontal and parietal bones.
According to studies on other species of fish, the melanophores in
this region of the head act as regulators cootrolling the amount of
incoming light (Nicol, 1963). The pineal organ (the epiphysis) is
photosensitive and must be protected from excess illumination. On the
other hand, young fishes, including the leptocephalus, are relatively
insensitive to light and receive very little protection from their melano-
phores (Breder & Rasquin, 1950). As research on the salmon has indica-
ted (Hoar, 1955), the almost [Otal absence of morphological
differentiation in the pineal organ in young fish is probably partly
responsible for this insensitivity.
The first appearance of the skull spot indicates an important step
in the life of the eel. I t forms at a time when the eels arrive in the coastal
waters and abandon their purely pelagic existence (see p. 133). The eel's
behaviour during this first phase of pigmentation indicates a marked
sensitivity to light (Tesch, 1965). Melanophores then begin to develop
along the entire body length and a transient phase of reduced sensitivity
to light begins j from time to time glass eels even swim along the banks
near the surface.
A list of characteristics can be drawn up which describes the extent
of pigmentation during various advanced stages of development
(Table 2j Fig. 9) (Strubberg, 1913j Grassi & Calandruccio, 1913).
According to this system Stage VI (Schmidt, 1906) has been divided
into substages A and B. Stage VI B essentially indicates the end of
pigmentation, and Stage VII represents the fully pigmented, benthic,
young eel (Gilson, 19(8). Thus, in Stage VI B the eel loses its glass-like
transparency.
In addition to the black colouring provided by the melanophores,
other pigments also begin to appear and are already in evidence towards
the end of Stage VI A. In particular a green coloration becomes re-
cognizable as a result of the formation of yellow pigment. Heldt and
Heldt (1929a) attribute this to the beginning of food intake (see p. 148),
whereby lipoids are fonned, thus providing the basis for the yellow
colour. Water-soluble flavones are particularly involved in the formation
of yellow pigment in the eel- a characteristic which distinguishes
BODY STRUCTURE AND FUNCTIONS '7
•
Fig. 9 Development of the subepidermal pigment in the glass eel. a, Stage VI AI;
b, Stage VI All; c, Stage VI Alii; d-f, Stage VI A lv ; g, Stage VI B (after
Strubberg, 1913).
the eel from other, less euryhaline fishes (Fontaine & Busnel, 1939).
Stage VI A is further divided into a number of subsections of which
only the main divisions, VI AI-VI A IV, are given in Table 2. According
to Strubberg (1 913) each of these four subsections is made up of a
18 THE EEL
further one to four subdivisions; this system is based on the fact that
pigmentation starts caudally and dorsally and proceeds rostrally and
ventrally.
In German, regardless of the degree of pigmentation, young eels are
referred to as 'Aalbrut', 'Montee' or, if they are not too darkly coloured,
as 'Glasaal' (and in French they are called 'civelles'). In English,
however, one distinguishes between 'glass eels' and 'elvers' j unpigmen-
ted young are referred to as 'glass eels', while pigmented young are
called 'elvers'. When pigmentation is complete the yellow eel stage
is reached; there are no major external changes after this until the eel
returns to the sea. In German, the small, fully pigmented eels, which
are not yet suitable for marketing, are called 'Satzaale', literally 'eel fry'.
Strictly speaking, the name 'yellow eel' is not correct, because, although
many eels do vary from yellow to white on the underside, a large number
have almost completely white bellies that change to a light grey on the
flanks. However, the term 'yellow eel' has now been universally adopted
and is used to distinguish this stage from the silver eel stage. The
yellow eel's dorsal surface varies from dark green or brownish-green
to black, the former colours giving rise to the term 'Griinaal' or 'green
eel'. A similar duality of terms occurs in France; both 'Anguille jaune'
and 'Anguille verte' are frequently used; the use of 'green' or 'vert'
might also imply inunaturity. In English the term 'yellow eel' is com-
monly used, but from time to time one comes across the expression
'golden eel'. However, this use of metallic colour terms can be misleading
because a metallic shinuner is the distinguishing characteristic of the
'Blankaal' ('silver eel' or 'bronze eel') or of eels without black pigment
(Xanthochromatism) (see p. 359).
Naturally, coloration in eels from various biotopes can be very
different and must depend, to a great extent, on the background.
Fishermen involved in marking and transplanting eels in the North
Sea have reported (Tesch, 1967a), that transplanted eels recaptured
outside their original habitats were a quite different colour from those
that normally lived in the area. Thus these eels had retained their
coloration for days, weeks even, without showing adaptation to their
new environment. Experimental studies by Neill (1940) and Odiorne
(1957) have also shown that eels take a very long time to change colour;
as many as 20 days may elapse before one extreme condition of the
melanophores will change [0 another. In the wrasse ( Crenilabrus ) ,
on the other hand, the change begins within a few hours. Hormonal
or humoral processes are probably responsible for colour changes in the
eel. whereas in Crenilabrus and many other species offish, these changes
appear to be under neural control.
If one believes that the eel's coloration tones with the substrate,
this can only be true for a particular tonal range. Most eels which
originated from underground river areas were an unusually light colour.
Thus lack of light obviously produces contraction of melanin in the
melanophores. (See Anomalies in colour, p. 359.)
With the growth of the body and the gonads (see p. 169), a change in
colour takes place. During migration to the spawning grounds in the
Atlantic the eel also, as it were, readjusts itself in other, morphological
respects to its new environment. Thus an open water fish develops
from a benthic fish. Starting on the side of the body the eel develops
a silver tint which gradually spreads ventrally. The eel has then reached
the 'silver' or 'bronze eel' stage as it is so aptly called. On the back and
dorsolaterally the eel becomes darker and darker in colour until it looks
almost black. The pectoral fin also becomes black. Until the silver
tint completely covers the surface area of the belly, the eel is referred
to as being 'half-silver'. There are, of course, many transitional phases
starting with the onset of the silver eel stage and ending with the fully
coloured silver eel, and it is often difficult to decide from external
features whether a particular individual has already reached this stage
physiologically. The best indication of whether the animal is physio-
logically a fully developed silver eel is the method by which it was caught.
Animals which appear 'silver' and which are caught in a downstream
current or in a stow net at low tide, are sure to be fully developed.
However, those eels often caught shortly before their seaward migration
should certainly not be considered mature or to have completed their
change in coloration. An example of such an eel is shown in Fig. 30.
Experiments on male eels, where maturity of the gonads was produced
artificially, have shown that a narrow, whitish-silver band remains on
the ventral side, which is itself flecked with red.
The metallic coloration varies in silver eels from different places.
Some appear a true silver colour while others range from bronze to
coppery tones. It seems certain that this is due not only to differences in
the degree of maturity of the individual but also to its habitat and to
the original colour adaptation to its former biotope, the effects of which
are still visible through the silver sheen.
1.2.4 Teeth
The eel has setifonn teeth on the maxilla, the dentary and the premaxilla-
ethmovomerine bloc. The arrangement of teeth on the maxilla and the
vomerine plate is shown in Fig. 10. In many species of eel the arrange-
ment of the vomerine teeth serves as an important diagnostic feature
(see p. 84 and p. 100).
20 THE EEL
Fig. 10 Teeth of the premaxi1lo.-ethmovomerine bloc in A. anguilla (Photograph :

Freier).
During ontogenetic development the teeth undergo considerable

changes in form. The teeth succeeding the large canines of the larva
are not as large and become progressively smaller towards the back of
the jaw. The larva loses these relatively large teeth during the very first
stages of metamorphosis (Fig. I I ), so that when the glass eel stage is
reached no larval teeth remain. Schmidt (1916) found no differences
between tooth development in A. rostrata and A. anguilla. Setiform teeth
first begin to form at Stage V but do not break through the epidermis
until later (Gilson, 1908; Gandolfi-Hornyo\d, I98Ib); the first teeth
appear on the jaws and on the vomerine plate in Stage VI A. Initially the
very numerous teeth vary considerably in size, and there are no re-
cognizable rows. In Stage VI B too, most of the teeth do not protrude
very far out of the epidennis. There is, however, considerable individual
variability, and the teeth are thus less useful than pigmentation in
classifying the various developmental stages (Gandolfi-Homyold,
1918b). In general, completely fonned rows ofteeth are not seen until
the eels are between 12 and 15 em in length. This is, then, the first time
one can use the teeth to distinguish between species (Fig. 44); in
tropical eel species it may be possible to use this feature at somewhat
earlier growth stages.
Fig. II
, --------------
Head of larval A. anguilla. A, Stage I, with fully developed tooth rows;
B, Stage II, with almost no teeth; C, Stage III, with no teeth (after Schmidt,
19(>6).
1.2.5 Musculature
As far as the author is aware, no comprehensive description of the eel's
musculature has ever been published. Alexander (1969) has mentioned
an unusual feature of the white muscle fibres in A. anguilla as compared
with those of most other species of fish. Their position relative to the
horizontal septum is similar to that in cartilaginous fishes, primitive
22 THE EEL
Fig. I2 Sections through various regions of the body in Conger unlge,.. Ep, Epaxial
muscles; Hyp, Hypaxial muscles; lnf, Infracarinal muscles; Be, Body cavity;
M, Horizontal myoseptum; Sup, Supracarinai muscles; Ve, Vertebral
centrum (after Rlin & Balea, 1955).
bony fishes and members of the Salmoniformes. Considering the

extreme shape of the eel's body, it is highly probable that further
peculiarities will come to light when more research has been carried
out in this field. Blin and Balea (1955; see also Harder, 1964) have begun
to investigate this problem. These authors described the course of the
rnyomeres and the myosepta in many species of fish, from both trans-
verse and longitudinal sections, and have thereby provided a basis for
systematic studies. Unfortunately, however, they have only dealt with
the conger eel (Conger conger) and not with Anguilla anguilla. Since
Conger is not very different from Anguilla, Blin and Balea's (1955)
diagrams have been reproduced here (Fig. 12); for comparison with
other fishes references should be made to the publications cited above.
Nelson (1967) has compared the gill arch musculature of A. rostrata
with that of eels from five other families and genera and found reduction
from the condition in A . rostrata, through that of Conger marginatus,
Moringua javanica, Kaupichthys diodontus, Uropterygius knighti to
that of Gymnothorax petelli.
Research on the chemical composition of eel muscle has been carried
out with regard to concentrations of sodium, potassium, water-
containing ninhydrin-positive particles, free amino acids, ethanol-
amines, cytathionines and carnosines. Results indicated that, after a
month in seawater, silver eels show an increase in amino acid content.
In particular, there were increases in alanine, glutamine, glutamic acid,
glycine, proline, tyrosine and cystathionine. The amount of carnosine,
ethanolamine and other amino acids hardly changed at all (Huggins
& Colley, 1971). The amount of fat, protein, carbohydrates, ash, water
and dry materials is given in Table 42.
1.3 Respiratory organs and swimbladder
In the eel the gill arches, filaments and lamellae conform to the general
teleostean pattern, although minor variations occur. Eels between
32 and 37 cm long, have, on gill arches 1 to 4 of each side, 147, 136, 129
and 119 filaments respectively, giving a total of 1062 filaments for the
four arches; the total length of all the filaments is 3"422 m. Each milli-
metre of gill filament has 30 lamellae, and the total respiratory surface
area per kilogranune of body weight has been estimated at 0·987 m 2
(BYC2kowska-Smyk, 1958).
Such a surface area for respiratory exchange is of intermediate size
as compared with that in other fishes. For example, the pike perch
( Scizosledion lucioperca ) , and the rainbow trout ( Salmo gairdneri) ,
have some 1·800 m2/kg body weight, and the viviparous blenny ( Zoarces
viviparus) only 0·475 m2jkg body weight. This intermediate position
of the eel in terms of branchial surface area is reflected in its slow
swimming, benthic habits and its concomitant low oxygen consumption.
On the other hand, the frequent and wide ranging movements in search
of prey, as well as prolonged migrations at certain phases in the life-cycle,
must often demand sudden and unpredictable oxygen requirements;
furthermore Byzhkowska-Smyk (1958) has calculated that, in water,
the surface area of the gills is insufficient for the respiratory requirements
of eels. A second source of gas exchange is thus relevant, namely the
skin (see p. I I). In a detailed study Berg and Steen (1965) showed that
in water 90% of oxygen was absorbed across the gills while in air some
two-thirds of the oxygen absorption was cutaneous. When the eel is
out of water, its branchial chamber is filled with air, from which there
is a slow absorption of oxygen. At 20 °C, opercular movements replenish
the supply of air by pumping at a rate of 60 movements per hour; in
water, this respiratory rate is 20 times more rapid (Berg & Steen, 1965).
Berg and Steen (1966), in an examination of the ventilatory responses
to separate exposures of the skin or gills to various gas mixtures, found
that reduced environmental oxygen tension increased the breathing
frequency while an environment of pure oxygen markedly diminished
or indeed abolished ventilatory movements. Exposure to pure nitrogen
produced a fourfold increment in breathing movements. Intermediate
rates of breathing were observed when the skin or the gills were separate-
ly exposed to varying gas mixtures. Less than 5% carbon dioxide had
little effect upon either the amplitude of respiratory movements or their
frequency, but had ·a n inhibitory effect when concentrations above 5%
contacted the gills. The authors suggested that hypoxia is ·a major
stimulus for respiratory movements, and, in air at least, carbon dioxide
has little effect.
24 THE EEL
A number of workers have examined the physiological mechanisms

that control the microcirculation of the gills. Gilloteaux (1969) conclu-
ded that both neural and endocrine factors moderate the flow of blood
through the filaments, and hence oxygen uptake, in eels in which the
spinal cord was destroyed (Leicht, 1969). It had been suggested earlier
(Schultze, 1965) that receptors in the head region were responsible for
the regulation of oxygen uptake over a wide temperature range. In
particular these receptors, sited either within the ventral aorta, or on the
afferent branchial arterioles, monitored blood oxygen in the venous
(afferent branchial) blood. and brought about changes in blood volume
and distribution within the branchial vascular bed. The delicate vascular
arrangements within the gills, especially the shunting processes along
the lamellae, are probably of great significance and certainly respond
to various neurohumoral factors (Steen & Kruysse, 1964; Maetz &
Rankin, 1969; Rankin & Maetz, 1971).
The capacity of the adult eel to survive in both air and water is
associated with its ability to use both branchial and cutaneous modes of
respiratory gas exchange. The eel survives better in air than in poorly
oxygenated or polluted water; indeed, its resistance to hypoxic water
is apparently no greater than that of other fishes. Given the choice,
eels invariably select water of high oxygen tension (Hill, 1969), and it is
of interest that when lakes freeze, eels show the highest mortality rate
among the fishes present (Rahn, 1963). The benthic mode of life of the
eel frequently exposes it to hypoxic zones of water which are often at low
temperarures. Low temperatures inhibit the eel's mobility and thus its
well known panic-like escape reflexes. Oxygen supply is therefore not
the only ecological factor affecting the survival of eels, and it seems that
temperature is especially significant in subtropical and temperate
climates (Johansen, 1929; Bernhardt, 1963; Scheer, 1964; Tesch, 1964).
The delicate interplay between oxygen and temperature vis-a-vis
the ecological distribution of eels clearly requires further study.
The gills, in addition to their respiratory function, participate in
osmoregulation. Euryhalinity requires that in seawater the branchial
epithelium extrudes excess ions such as sodium and chloride, while in
freshwater a net uptake of salts takes place. The eel, in particular during
its silver phase, displays a remarkable capacity to effectively change the
polarity of the active transport of ions, and has been the subject of exten-
sive investigations in many laboratories. Aspects of the biochemistry
(Maerz, Nibelle, Bornancin & Motais, 1969), physico-chemistry
(Motais, Isaia, Rankin & Maetz, 1969; House & Maetz, (1974) and
physiology (Henderson & Chester Jones, 1r;67; Maetz, 1971) have been
studied in detail in eels adapted to fresh- and seawater. Many of the
fundamental processes responsible for the salt transfers are under
intimate endocrine control and these too have been elucidated in
part (Henderson, Chan, Sandor & Chester Jones, 1970; Henderson &
Chester Jones, 1972). In seawater, the branchial epithelium displays
a rich population of chloride cells (Keys-Willmer cells; Keys &
Willmer, 1932). These cells, situated at the base of the gill lamellae,
were originally described as the site of chloride transport (Keys,
1931; Schlieper, 1933), and more recently they have been shown to
possess many of the ultrastructural and enzymatic characteristics
of electrolyte transporting epithelia (Masoni & Isaia, 1973; Mizuhira,
Amakawa, Yasmashina, Shirai & Utida, 1969; Shirai & Utida, 1970;
Utida, Kamiya & Shirai, 1971; see also Motais & Garcia-Romeu,
1972). Present evidence thus suggests that the branchial chloride
cells extrude sodium and chloride when eels are in seawater. There
may be more than one type of cell and it is possible that the absorptive
function of the gills in freshwater adapted eels reflects the existence
of a second type. Among the inconsistencies which may argue against
the chloride cell being the unique route of transport is the fact that
immature eels do not have readily recognizable chloride cells when they
are in the sea (Ogawa, 1962).
Whatever the precise sites of water and electrolyte movement across
the branchial epithelium, the gills play a crucial role in the adaptive
processes. As the environmental sodium concentration increases, the
gills progressively extrude the sodium gained by drinking (Maetz
& Skadhauge. 1968; Mayer & Nibelle. 1970). and their water penne-
ability changes (Keys, 1931). Such adaptive responses are under
endocrine control but may be influenced by other factors such as
branchial haemodynamics (Maerz & Rankin, 1969; Kirschner, 19693)
and may be liable to seasonal influences (Utida et ai., 1967a; Utida
et al., 1969).
The molecular and enzymatic mechanisms have been studied in
preparations of eel gills. For example the extrusion process in seawater
is inhibited by materials that block protein synthesis (Maetz et ai. , 1969)
or inhibit the enzyme, sodium-potassium-activated adenosine tri-
phosphatase. The latter is present in greater amounts in the gills of
seawater adapted eels than in those of freshwater adapted fish (Milne,
Ball & Chester Jones, 1971j Kamiya & Utida. 1968, 1969; Jampal &
Epstein, 1970; Forrest et al. , 1973; Motais, 1970) although Kirschner
:1969b) was unable to demonstrate such a difference.
There are two types of euryhalinity according to Motais (1967). One
26 THE EEL
typified by species such as Fundulus, a killifish, is termed 'osmoconfor-

mer' and differs from the second, or 'osmoregulator' type in that a
large portion of the sodium turnover across the gill membranes results
from an exchange diffusion mechanism (Evans, 1969; Maetz, 1971;
Motais, Garcia-Romeu & Maetz, 1969).
A final ftmction of the gills - that of nitrogenous waste excretion - is
related to acid-base balance as well as to ionic regulation. Ammonium
ions pass freely along the gradient from the blood to the surrounding
water. Subsequent work has confirmed Keys' (1931) original suggestion
of an NH;/Na + exchange across freshwater eel gills (Garcia-Romeu &
Morais, 1966) and further, that although small amounts of ammonia
may be produced in the gills, it largely emanates from the liver (McBean,
Neppel & Goldstein, (966).
Much morphological and physiological information is available on
the swimbladder of the eel. To paraphrase Dorn (1961) ' ... (the
swimbladder of the eel) is a physostomous type with a large duct from
the oesophagus, although the opening itself is tiny. The duct is expansive
and highly flexible and can give the appearance of a primitive lung,
indeed it subserves a resorptive function. In other respects the (eel)
swimbladder has many physoclistous characteristics. There are two
highly developed capillary bundles ( retia mirabilia) dorsal and ventral
to the opening into the swimbladder (secretory bladder), which are
separated from the gas gland, itself a special feature and part of the total
epithelial formation.'
Figure 13 shows the general structure of the eel swimbladder in situ.
A most striking feature is the relatively large anterior section - the
pneumatic duct - which has a resorptive function (Steen, 1963b, c),
and, when full, clearly represents a large gaseous reservoir. Histo-
logically the pneumatic duct may, in many respects, be taken to resemble
a primitive vertebrate lung, a fact that adds strength to the theory that
this zone could aid aerial respiration in the eel (Dorn, 1961). It is,
however, only partly true that 'pulmonary' respiration has replaced
branchial respiration when the eel is out of water. When breathing aerial-
ly the eel absorbs only half as much oxygen as it does when breathing in
water, and further, the respiratory rate is much reduced (Berg & Steen,
1966). During the first hour in air only one-third of the oxygen conswned
is derived from branchial respiration, and after 20 hours at moderate
temperatures an oxygen deficiency becomes apparent. The swim-
bladder thus supplies a certain amount of oxygen, but it is in no wayan
accessory respiratory organ.
These facts are very relevant to the husbandry and transport of eels.
,
•
Fig. 13 Swimbladder of A. anguilla in the body cavity. A, Pneumatic duct;
B, Secremry section of the swimbladder; C, Rete mirabile; D, Vein of the
pneumatic: duct; E. Swimbladder artery; F. Swimbladder vein; G, Dorsal
vein; H, Sphincter between the pneumatic: duct and the oesophagus;
I, Oesophagus; K, Dorsal aorta (modified after Steen, 1963b).
Certainly eels can be transported for days without water, providing the
humidity is high, and perhaps more important, that the temperature is
kept low: 7 °C is suitable but 15 °C may be fatal.
Light and electron microscope studies on the histology of the
eel swimbladder have been published by Dorn (1960, 1961). Krogh
(1924) examined the capillary networks, especially those within the
retia mirabiiia, and Steen (1963a. b, c) related the structure to gaseous
exchanges, especially the oxygen, carbon dioxide, lactic acid and blood
pH interrelationships. A brief summary of the findings to date must
include the facts that the swimbladder receives arterial blood which
fonns a capillary net at a localized area on the inner surface of the
swimbladder, and that these capillaries are drained retrogressively by
veins running in parallel with them. A key feature is that the capillary
system of arterial and venous origin intertwines in such a way that
arterial capillaries are surrounded by venous ones and vice versa to fonn
a rete. This arrangement ensures an optimal diffusional state between
arterial and venous blood.; furthermore, the flow in the capillaries is of
the counter-current type. In the eel the rete contains about 100000
capillaries of each type, and the distance between arterial and venous
blood is about 1 Ilffi. Although the organ itself is only 3 or 5 mm in
diameter the aggregate length of capillaries is about 900 m with an area
of 210 mi l Such volume:surface area relationships. together with the
28 THE EEL
counter-current exchange situation may readily explain the respiratory)

absorptive, secretory and acid-base relationships observed in the eel
(Stray-Pedersen & Nicolaysen, 1975).
1.4 Feeding and organs of digestion
1.4. I The gastrointestinal tract

As in all physostomes, the gastrointestinal tract (Fig. 14) receives the
swimbladder duct, which opens into the oesophagus (Kukla, 1954).
The stomach is narrow and cone-shaped (Fig. IS) with an obvious
diverticulum, giving it a Y-shaped form (Harder, 1964). There are no
pyloric appendages such as frequently occur in other teleosts.
The mucous membranes of the oesophagus and stomach are thrown
into long straight folds, which are fully developed only after the glass
eel stage (Berndt, 1938). The folds of the mid-gut are more irregular
and zig-zagged. The vascularization, and the ontogeny of the gastro-
intestinal tract have been described by Kukla (1954) and Berndt (1938)
respectively.
The relative length of the alimentary tract varies with the stage of the
life-cycle. It is, in relation to body size, greatest up to the end of the
leptocephalus stage, gradually reducing during the glass eel phase and
Fig. 14 Cross-section through A. anguilla. ab, Swimbladder; c, Visceral cavity;

g, Gut; gf, Genital fold; ms, Mesentery; msn, Mesonephros (modified after
Harder, 1964).
BODY STRUCTURE AND FUNCTIONS
2. Pyl.-KI.
1. Pyl.-Kl.
P.pylorico
Fig. IS Digestive tract of the eel. P. pylorica, Pyloric part of the stomach; Pyl. KI.,
Pyloric valve (after Pernkopf from Harder. 1964).
then progressively increasing until the silver eel phase, after which there
is a slight decrease in length. During the glass eel stages VI A1to VIlli
(after Strubberg, 1913 ; Table 2) there is a slower intestinal growth rate
in seawater than in freshwater (Vilter, 1945b). A marked increase occurs
at the beginning of voluntary food intake (Vilter, 1945a).
In addition to these gross changes in size there are considerable
changes in the relative proportions of the individual gut segments.
Although the stomach is readily recognizable in the larval forms, there
is tendency for the oesophagus, stomach, mid- and hind-guts to be
relatively undifferentiated. With the emergence of the glass eel the
stomach and mid-gut enlarge concomitantly with changed post-
metamorphic feeding patterns. The leptocephalus feeds on plankton,
while after metamorphosis a more omnivorous habit is characteristic
(see p. (48).
The slightly diminished gut size of silver eels is perhaps to be expected
as it is then that the prolonged fast begins. This, however, occurs
alongside many other, more fundamental changes in gastrointestinal
structure and function. Thus convolutions in the gut disappear and
30 THE EEL
"00
"00
" 00
"00
"00
"00
"lOO
.00
800
roo
600
..,
500
lOO
200
'00 "''''....
..- --- .... ...... -
~
ltptoce~QkJs GlasHei Pigm~ted P'~mtnted y.lkrw ~ Silwr ttl

7·5cm 6·gem ttl II)Scrn ttl 62, .. nan n<,.
Fig. 16 Thickness of the circular ( - - ) and longitudinal (- - - - -) muscle layer

of the midgut at particular developmental stages of the eel (in ,:.w mm)
(after Berndt, 1938, VEB Gustav Fischer Verlag, lena).
radical histological cbanges, especially in the oesophagus and hind-gut,

are apparent. The intestinal musculature begins to degenerate, a process
initially obvious at the ends of muscle fibres, but eventually involving the
disintegration of the nuclei. Figure 16 illustrates that the gut mus-
culature reaches a maximal thickness shortly before the onset of the
silver eel stage, after which there is a sharp and rapid decrease in thick-
ness. One of the more dramatic changes that was thought to take place
was the complete closure of the anus (Fig. 17) and the occlusion of the
gut lumen in advanced silver eels (Schnakenbeck, 1940). However, no
closure of the anus or invasion of the gut lumen is reported by recent
workers (Boetius & Boetius, 1967a; D'Ancona 1959a) nor was gut
occlusion mentioned by Villani & Lumare (1975).
Functionally, the gut of the eel is of interest in that the stomach
and its diverticulum lie parallel and very close to one another. After
feeding, the stomach is distended and greatly compresses the intestine,
but when it empties it becomes almost occluded by the full intestine.
Food intake, therefore, is only possible at infrequent intervals (Wunder,
1970).
In addition to its function in feeding, the gut plays a very important
role in osmoregulation especially in seawater environments. This
function was first demonstrated by Smith (1930). Eels in which the anus
had been occluded were kept in seawater containing phenol red.
Fig. 17 Section through the anus of an advanced silver eel caught in the sea. a, Original
site of anus; P, Peritoneal canal, genital opening (modified after Schnakenbeck,
1934).
Mter 20 hours the gut contents were examined and the phenol red
concentration measured. The dye was more concentrated in the gut
than in the surrounding water, and calculation revealed that some 10 rol
of fluid had been absorbed by the gut. Over the same period, 2'3 rol of
urine were produced and the animals lost 1'3 g in body weight. Thus
fluid loss occurred extrarenally. Moreover, Keys (1933) showed that
occlusion of the gut is a fatal procedure in eels adapted to seawater
(see also p. 41).
Although the gut plays an imponant role in water and electrolyte
balance in seawater, it should be emphasized that in freshwater signi-
ficant quantities are imbibed (Hirano, 1967; Sharrat, Bellamy &
Chester Jones, 19643-; Keys, 1933; Maetz & Skadhauge, 1968;
Skadhauge, 1969). It is also noteworthy that during ontogeny the rates
of movement of water and solute from lumen to blood increase and may
also show seasonal change (Utida el al., 1969). It has been suggested
that these changes in rates of transport and the obligatory absorption
in freshwater are adaptive features preparatory to the onset of the pro-
longed marine migration of the silver eel.
The Japanese eel has been studied extensively with a view to elucidat-
ing the mechanisms of water and ion transport across the teleostean
alimentary tract (aide & Utida, 1967b, 1968; Utida, Isono & Hirano,
1967 j Utida & Isono, 1967; Utida et ai., 1967a) as well as to identifying
factors that govern rates of, and stimuli for, drinking (Hirano, 1974;
32 THE EEL
Hirano, Satou & Utida, 1972). Alkaline phosphatase and especially

adenosine triphosphatase activities are greater in seawater adapted
eels than in those adapted to freshwater, and these enzymes have been
implicated as basic features in the osmoregulatory role of the gut (Utida
et al., 1968; ]ampel & Epstein, 1970). The functional integrity
of the gut appears to be related to adrenocortical steroids and
possibly prolactin (Gaitskell & Chester Jones, 1970, 1971; Utida,
Hirano, Oide, Ando, Johnson & Bern, 1972). Attempts to identify
specific cells responsible for ion and water movement (analogous to the
branchial chloride cells) on the basis of ultrastructUre (Yamagishi et al.)
1969) indicate that ultrastructural laminae are correlated with marine
environments, and most probably with osmoregulation. Further studies
are still required, however, especially with regard to the ontogeny of
marine adaptation.
1.4.2 Pancreas
Both endocrine and exocrine functions of the pancreas change with the
developmental stage of the eel. The differing amounts and quality of
food consumed at the different stages are, of course, primary reasons.
Clearly the exocrine function of producing digestive enzymes will
be governed by both quality and quantity of food (herbivorous, omni-
vorous, etc.) available for digestion. During the various developmental
stages also, metabolic demands change, e.g. the voraciously feeding
omnivorous yellow eel must be contrasted with the migratory, fasting
silver phase. Such metabolic changes are related to or dependent upon,
endocrine pancreatic Mction.
The pancreas of the eel, unlike that in many other fishes, is a compact
structure surrounding the portal vein on the mid-gut. It is readily
distinguishable from the gut by its darker colour. In 35 to 40 em long
eels, the pancreas is 6 to 7 mm wide and 55 to 60 mm long (Kukla, 1958).
Excluding a slightly tapered portion which extends as far as the bile
duct, the pancreas begins at about the point where the stomach joins
the intestine and it extends as far as the gastric caecum. Figure 18, taken
from Kukla (1958), illustrates the pancreatic arrangement, including
the islets of Langerhans. The same author has described the general
morphology of the pancreatic vascularization, and notes that islets of
Langerhans are less numerous in the anterior portions of the pancreas
and that no extrapancreatic islets are found. Theret and Palayer (1967)
have compared the pancreatic ·cytology of the eel with that of other
teleosts, and Epple (1969) compared the pancreatic morphology of the
eel with that in other fishes.
.. .. J i
00,
~ 0 • ,<> •
,
o
"
,.
Opma
-
·'CJ·o
,
,
0
'0
o,
,
,.. , ~" '-'
•
, ,
"
'0
0
,
'"
.,
." '" 0.,0 g, " .' Q O. , .
,
o , , 0. . 0
0
o t . '0 ~ <> , • Oil'
,. 0
. ,", Oo~ o< ., " " ,

,•
0
,
"
•
"
."
," 0
•,
O~$
• ,
•~ 0
," .
.
'
<>
o .0 "
• "
Fig, 18 Distribution of the Islets of Langerhans in the pancreas of the eel. Dpma,
Ductus pancreaticus major; Dpmi, Ductus pancreaticus minor; In, Islets
of Langerhans; P, Pancreas; Vp, Vena porta (after Kukla, 1958).
Hyperphagia and starvation markedly alter the size of both the

endocrine and exocrine pancreas, Starvation or indeed injection of
thyroxine, for example, reduce the volumes of both pancreatic compo-
nents in Anguilla japonica, although Honma (1966) described islet
hyperplasia in Anguilla japonica during its catadromic migration,
The endocrine pancreas also changes correlatively with the stage
of development (Fig. 19), The volume generally increases up to the
time of maximal feeding activity - i.e. in the yellow eel. In larger yellow
eels, reaching the end of their intensive feeding phase the volume is
reduced, an effect especially marked in wimer, The volume reaches a
34 THE EEL
Voju,", of thu llOOcrIf'lt poncrecs
in reiatlOl'l \0 OOdy ~~t
• ---------
, ,, ------ .... _-----
, -- ,,
Small
yellow
."
L".
yelfow
otl.
l •
AUlumnjSilVtf
Spriflq n l
Fig. 19 Development of the endocrine pancreas during the various developmental

stages of the eel (modified after Paiayer, 1967).
minimum in silver eels, although activity is still required for gonadial

function as well as, possibly, for osmoregulation in the sea (Epple &
Lewis, 1975).
1.4.3 Liver and gall bladder

Unlike other fish the eel possesses a unilobular liver with but a slight
indentation at the posterior end. Aberrant types, such as double liver,
have been described (Reichenbach-Klinke, 1952), although in such
cases only one gall bladder is present.
The participation of the liver in metabolism has been the subject
of some controversial studies regarding nitrogenous metabolites. Thus
the eel contains much glutamate dehydrogenase in the liver but only
small amounts in muscle tissue (McBean et aI. , 19(6). Significant
transdeamination is unlikely to take place in muscle therefore, and
Janicki & Lingus (1970) have observed in vitro transdeaminase activity
in the liver of eels. Kenyon (1g67) found that hepatectomy was without
effect upon ammonia production, although some amino acid levels
were elevated in the blood. The liver does not seem to be essential
for amino acid deamination therefore, although excess of these
materials cannot be deaminated in the absence of the liver (see Watts &
Watts, 1974). Hepatectomized eels suffer from hypoglycaemia although
they survive for periods of up to 63 days (Inui, 1969). Like other fishes,
starved glass eels display reduced hepatic cell size and an eventual
exhaustion of the intracellular glycogen granules (Inui & Egusa, 1967).
The precise physiological status of the liver vis-a.-vis amino acid
deamination, gluconeogenesis, and protein synthesis in the eel requires
funher srudy; the availability of this animal and its marked survival
after much surgery suggests that it could prove a suitable subject for
research into the general understanding of hepatic function.
Other aspects of hepatic function including the presence of fatty
acids and their esters (Hata & Hata, 1967) and characteristics of bile
salts (Hatakeyama & Ukamara, 1928; Hirofuji, 1966) have been
srudied in eels. The latter subject has been extensively reviewed (Hasle-
wood, 1969; Tammar, 1974).
1.5 Circulation of blood
1.5.1 Cardiovascular system

Figure 20 shows the heart in the pericardial cavity, and Fig. 2 I schemati-
cally represents the pacemaker system. Skramlik (cited by Wunder,
1936) suggests that the eel is unusual with respect to the number and
distribution of pacemakers. Three (Bielig, 1931) or four (Grodzinski,
1954) have been suggested, the major one being associated with the
atrium (Fig. 21 AB,a). The latter, while controlling ventricular function,
may itself be regulated by the sinus venosus. The ventricle may also
have a semi-autonomous function (Grodzinski, 1954). Electron
•
,
Fig. 20 Position of the eel's hean in the pericardial cavity. Left: with the heart in its
natural position. Right : with the ventricle (V) raised so that the transition
(Si-o) of the sinus venosus (Si) into the auricular canal can be seen : A, Atrium;
B, Connective tissue threads by which the atrium and [he ventricle are
attached to the pericardial sac; P, Pericardial sac; T , Truncus arteriosus
(modified from Wunder, afler Troemer, 1932).
THE EEL
• •
Fig. u The pacemakers (hatched) in the eel's hean. A, according to Bielig, 1931 ;
B, according to Grodzinski, 1954. a, Auricle; ba, Bulbus arteriosus;
ca, Conus arteriosus (also called the Bulbus cordis); dc, Ductus cuvieri;
sv, Sinus venosus; v, Ventricle ; vh, Vena hepatica.
micrographic techniques have been applied to the atrial musculature,

but have failed to produce unequivocal information on its pacemaker
activities (Conteaux & Laurent, 1957). Randall (1968, 1970) reviews
these aspects of cardiac function in eels and other fishes generally.
Cardiac weight relative to body weight is less in eels than in other fishes,
and Torlitz (1922) has related heart weights to head shape in eels.
'Broad-headed' eels have hearts (including the bulbus arteriosus)
weighing some 1'2 % of the body weight, while in 'narrow-headed' eels
(see also p. 5) this weight is 0'8 % . When the bulbus arteriosus is
excluded the weights are respectively 0'9% and 0'6 % . Wunder (1936)
imaginatively suggests that the 'broad-headed' variety with its maraud-
ing behaviour requires a large heart, in contrast to the more peaceful
'narrow-headed eels'.
Cardiac activity and ECG, first described by Gitter (1933) and
Wunder (1 936), and different influences upon them, have been ex-
tensively studied. Cardiac rate both in v£vo and in vitro increases with
increasing temperature; thus intact yellow eels have a cardiac rate of
2o/min at 10 °C, and 95/min at 30 °C (Grodzinski, 1954). Above the
latter lemperature heart rate decreases sharply, indicating an optimal
temperature at this level.
Increases in hean rate also occur in glass eels transferred from
freshwater into seawater, although the opposite transfer has little effect
(Lhotte, 1945). Chloride ions in seawater were shown to activate the
parasympathetic nervous system, which dilated the peripheral blood
vessels and hence cardiac output. Many drugs have been tested on
cardiac function in the eel. Among them, adrenalin may increase cardiac
rate by 50%, while noradrenalin and dopamine have slightly less
activity. The over-all conclusions are, however, that the eel heart is
singularly insensitive to drugs of this sort (Hinge & Ostlund, 1954;
Mott, 1957).
The electrocardiogram of the eel, originally considered unusual
among fishes (Oets, 1950), resembles that of other anamniote hearts.
The differences are explicable on the basis of times of depolarization,
and intervals between the QRS and T components (Noseda, Chiesa
& Marchetti, 1963 j Randall, 1970). (See p. 362 for environmental effects
upon the ECG.)
The amounts of N-acetylhistidine, a substance possibly involved
with acetylcholine synthesis, found in the heart of eels adapted to
freshwater are similar to those in the hearts of eels in seawater or
brackish water (Hanson, 1966). The levels observed were similar to
those of stenohaline marine teleosts, which characteristically are lower
than stenohaline freshwater fishes. As with other physiological compa-
risons the eel then belongs to a marine type rather than a freshwater
one, and is unlike members of the Salmonidae and Osmerus eperlanus.
The latter displays the high cardiac N-acetylhistidine of a freshwater
fish.
Various regional vascular beds have been described and include the
digestive tract (Kukla, 1954), skin (Jakubowski, 1960a), olfactory bulb
(KaluZa, 1959) and swimbladder (Steen, 1963a, b, c). Mott (1950, 195 I)
has given a detailed angiographic description of the general vascular
system of the anaesthetized eel, and describes some actions of vasoactive
materials on blood pressure and flow.
Eels possess a well developed lymphatic system, which requires
accessory pumping structures or lymph hearts. The lymphatic system
(Duna;ewski, 1930; Florkowski, 1930) and the caudal heart situated
at the tip of the tail (Tright, 1913j Mislin, 1960; Harder, 1964) are
morphologically well documented, but their physiology requires
further study. The caudal heart, originally described as a 'ventricle
THE EEL
and valve' system by von Leeuwenhoek, 1695 (see Mislin, 1960)

situated in the tail fin posterior to the last vertebra, has proved suitable
for studying the effects of various environmental and phannacological
agents. Temperature (Jankowsky, 1968) has been shown to affect
caudal heart rate, and Mislin (1960) showed that cardiac rate affected
caudal heart rate. The caudal heart reached rates of 380 beats/min
at 34 °C while the cardiac rate was at a maximum (see above). Caudal
heart rate always exceeds the cardiac rate at all temperatures. Other
agents shown to act upon the caudal heart are those emanating from the
caudal neurosecretory system (Chan, 1971). Caudal heart rate seems to
be governed by both peripheral and central nervous mechanisms.
1.5.2 Blood
The major characteristics of adult European eel blood include:
1'4 x 10& red cells/mm3; haematocrit 38% ; haemoglobin 63-70 g/Ioo
ml; blood volume 3'2% of body weight; red cell dimensions 13 x 10 j..lIll
with surface area of 199 j..lIll2 (or 2'9 dm2/ml blood) (Zaitzewa, 1966,
1967; Schlicher, 1927; Wunder, 1936; Steen & Berg, 1966). The
values for erythrocyte dimensions and concentrations are considerably
higher than those observed in other teleosts (Swarts, 1969), although
it should be pointed out that considerable ranges are reported in the
publications cited above.
Blood makes up 1"1 to 1"15% of the body weight of glass eels, and
haemoglobin concentrations increase in proportion to body weight:
0"22 g glass eels had a haemoglobin content of 4"7 g/Ioo ml, up to 1 g
they had 10'2 g/IOO ml and young eels more than 1 g in weight had a
haemoglobin concentration of 11"7 g/Ioo ml (Zaitzewa, 1(67).
An interesting case of severe anaemia in the eel has been described
by Steen and Berg (1966), and a comparison was made with the haemo-
globin-free ice fishes Chaenocephalus aceratus and Champsocephalus
esox. The anaemic eel had a blood volume somewhat greater than normal
(1"3% body weight cf 0"8 % ), although swimhladder gas content and
blood pH were relatively normal. The oxygen tensions in both arterial
and venous blood were considerably below normal (see also Itazawa,
1970), but there were no obvious defects in general behaviour. It is
possible that the rate of blood clearance through the gills is adequate
to satisfy the oxygen demands of the fish, although experiments upon
the ability of such animals to undergo periods of oxygen debt require
study. I t should be pointed out that the leptocephalus larva of the eel
is without blood pigment (Schnakenbeck, 1955). The ontogeny of
haemoglobin synthesis in eels thus requires further study, and Kawa-
moto (1929) suggested that oxygen partial pressure/saturation/dis-
sociation relationships in the eel are unique. The latter has, however,
been disputed by Riggs (1952), while the presence of the Root effect
in eels (Forster & Steen, 1969) could explain earlier findings.
The general haematology, including numbers of red and white cells,
is affected by temperature (Durairaj, 1970), and by the general state
of the fish's health, including the presence of certain cutaneous and
systemic infections (Enomoto, 1969a, b). The effects of sewage upon
blood cell counts are discussed on p. 362.
Environmental salinity also affects the number of red cells, although
considerable variations occur (Fontaine, Delattre & Callamand, 1945);
after 8 days in seawater the red cell count fell from the original value
on transfer, 1'7 x 106 ceUsjmm3, to about 1 x 106 cells/mm 3 . It should
be remembered that body fluid volumes are altered in the contrasting
environments, and the red cell numbers are expressed as a concentra-
tion; haemodilution or haemoconcentration as a result of changed
water fluxes could thus playa major role. Nevertheless more red cellsj
unit volume occur in freshwater teleosts and this could reflect differing
oxygen demands and usage in the extreme conditions. A primary change
in haematopoetic function has not been excluded; indeed the general
phenomenon of haematopoeisis deserves attention.
Transfer of eels to seawater, in addition to reducing the numbers
of red cells, also decreases plasma protein concentration ; the latter,
unlike the former, is a less reversible process (Firly & Fontaine, 1932),
and the acid-base balance and pH of the blood must clearly be affected
by the changed environment (Fontaine & Boucher-Firly, 1933).
Moreover, the longer eels are kept in seawater, the more the blood
carbon dioxide decreases. Thus the buffering capacity of blood falls
in a medium which itself displays a greater buffering capacity.
The electrOlyte concentrations in blood plasma of eels adapted to
seawater are generally lower than those of truly marine fishes, although
when adapted to freshwater, eel plasma has concentrations of electro-
lytes similar to those of stenohaline freshwater species (Holmes &
Donaldson, 1969). The total osmolarities of plasma in both freshwater
adapted and seawater adapted eels are similar to stenohaline species in
these respective environments.
The regulation of electrolyte concentrations by neuroendocrine
mechanisms, as well as the regulation of blood pressure in the eel, are
considered on pp. 58-66.
Examination of blood albumins in the eel has yielded information
THE EEL
on interrelationships among the various species of the genus Anguilla,

especially the controversy surrounding the specific distinctness of
A. rostrata and A. anguilla. Sindermann and Krantz (1968) observed
that while most fishes have not more than two erythrocytic antigens the
eel has many. Other characteristics of the isoaglutinin system of eels
are discussed by Sinderman and Krantz (1968).
Related studies on the electrophoretic patterns of Atlantic eel
haemogiobins confirm the existence of two species (Sick et al., 1967).
The haemoglobin of the European eel displayed a single electrophoretic
peak, and whilst this peak was also shown by 93·5% of North American
eels, the latter also exhibited three additional peaks. Three variations
are thus present, two being characteristic of co-dominant alleles and
the third an heterozygous genotype. The haemoglobins of Japanese
(Yoshioka et al., 1968) and European eels (ltida et a/., 1970) have also
been studied using ion-exchange chromatography. Their general
biochemical and physical properties are reviewed by Christomanos
(1968).
Other electrophoretic applications employing radioactively labelled
serum transferins A, B and C revealed differing frequencies of the
three types in the European and North American species (Fine et al.,
1967). There is considerable heterogeneity in the electrophoretic
pattern of serum proteins in Japanese eels (Sick, Westergaard & Fryden-
berg, 1962), and other electrophoretic studies, including inununological
analyses, have provided equivocal data as to the discreteness of the
serum proteins of European and North American eels (Drilhon el al.,
1967; Pantelouris el al. , 1970, 1971). It is, however, relevant to note
that the chromosomal patterns of A. rostrata appear to differ from those
of A. anguilla (Ohno el ai., 1973).
The protein content of eel blood shows seasonal variation (K vasova,
1966) and probably also varies with sexual maturity; furthermore,
as many as 12 different antigens have been identified (Fine, Boffa, &
Drilhon, 1963; Drilhon. Boffa & Fine, 1962; Fine & Drilhon. 1961).
Of especial interest is the occurrence of the green chromoprotein.
thought to be largely biliverdin (aide & Utida, 1967a; Yamagushi,
Hashimoto & Matsuura, 1968). This material shows an annual cycle
and is probably associated with nutritional status. Larsson and Fiinge
(1969) have shown that there are changes in certain chemical consti-
tuents of the blood (cholesterol, fatty acids, haemoglobin, urea and
proteins) during the metamorphosis from the yellow to the silver eel
stages.
Finally. some mention should be made of the relatively high toxicity
of eel's blood as compared with that of other fishes. The so-called
'ichthyotoxin' causes muscular cramp ; small doses injected into mam-
mals accelerate the rate of breathing and the heart beat. o. I ml of eel
blood/kg is sufficient to kill a rabbit; 0.2 ml/kg will kill a dog (Mosso
after Guibe, 19.58). Wunder (1936) stales mat: 'If placed on mucous
membranes the poison produces inflammation which can last for days;
one must be particularly careful that eel blood does not contact the
human eye while the fish is being killed. The toxin is destroyed by
heating to 58-70 °C, and mere is no need to fear any effects of me
poison after preparation of the fish ... '. The effects have been classified
into two components: neurotoxic and cytolytic by Bertin (1956), who
also gives further characteristics and a brief history of this substance.
Rocca and Chiretti (1964) have more recently examined the toxic
nature of the serum of eels.
1.6 Urinogenital system
1.6.1 The kidneys

Original descriptions of the kidney in eels include those of Haller
(1908) and Jourdain (1859) (see also Harder, 1964; and Bridge &
Boulenger, 1958). The kidney is primarily a paired organ which shows
degrees of caudal fusion, the pronephric portions remaining separate
as lymphoid, haematopoetic structures containing few nephroi. The
posterior kidney, a mesonephros, perfonns the usual excretory and
endocrine functions. A large part of the blood from the caudal circula-
tion enters the kidney directly while some can effectively bypass the
kidney and enter the hepatic portal system.
The histology and histochemistry of the eel nephron have been
examined (Gerard, 1958; Guglielmone, 1966, 1967a, b), and it has been
argued mat the general fonn of the nephron in eels resembles that of a
stenohaline freshwater teleost. More particularly a distal (diluting?)
segment is present (Grafilin, 1937), and Pequignot and Senary (1965)
suggest that the respiratory responses bom of the gills and the kidneys
are similar to those offreshwater teleosts. These physiologically 'steno-
haline freshwater' characteristics of the eel are, however, unusual (see
pp. 24 and 25). Olivereau (1971 ) has related renal histology to water and
electrolyte composition as well as to the endocrine status of eels. The
kidney plays a major role, both in seawater and freshwater, in assisting
in the maintenance of body fluid homeostasis. Thus in freshwater,
eels have much higher glomerular filtration rates than they do in seawater
42 THE EEL
(Sharratt, Chester Jones & Bellamy, 1964b). In the former environment

the urine is very dilute with respect to plasma, as a result of intense
tubular reabsorptive activity. especially for sodium and chloride
(Chester Jones, Chan & Rankin, Ig6ga). On adaptation to seawater
there is an acute reduction in the glomerular filtration rate; at the same
time, tubular function, although still carrying out reabsorptive activity,
shows divalent cation secretion (Chester Jones et al., 19693; Oide &
Utida, Ig67b; Butler, 1969). Renal mechanisms are largely responsible
for the volwne adjustments as reflected in body weight changes that
occur in eels when transferred from freshwater to seawater or vice
versa (see p. 208 ).
The kidney of eels contains within its vasculature granular epithe-
lioid cells, homologous with the renin-producing juxtaglomerular
cells of other vertebrates (Sokabe & Ogawa, 1974; Nishimura & Ogawa,
1973 ; Krishnamurthy & Bern, 1969). Renal extracts contain a renin-
like enzyme which interacts with a substrate in plasma (Chester Jones
et az', 1966 ; Sokabe et al., 1966). The function of renin in eels is un-
known but blood pressure regulation and the regulation of glomerular
filtration rate have been suggested (Chester Jones et aJ., 1966; Sokabe
et aI., 1973). The changes that occur in plasma renin activity and renal
content are at present difficult to interpret, although the observations
noted in the publications cited argue against an exactly homologous
function between the mammalian renin-angiotensin system and that
of the eel.
Many histological, histochemical, physiological and pharmacological
data suggest close interrelationships between the endocrine system and
kidney function in eels adapted both to marine and to freshwater
environments. Among the hormones which affect renal form
and function are those of the neurohypophysis (Chester Jones,
Chan & Rankin 1969b; Henderson & Wales, 1974), the adrenal cortical
homologue (Chan, Chester Jones & Rankin, 1969), the adenohypo-
physis (Olivereau & Ball, 1970), the urohypophysis and the corpus-
cles of Stannius (Chester Jones, Chan & Rankin, 1969b). These endo-
crine glands have close associations one with another ; the interactions
between the pituitary gland and the adrenocortical homologue are
perhaps the most finnly established (Ball et aI., 1971; Hirano 1969 ;
Henderson, Sa'di & Hargreaves, 1974 ; Hanke & Chester Jones, 1966).
Other tropic and synergistic reactions of hormones in regulating renal
structure and function in the eel are considered by Chester Jones,
Chan, Henderson & Ball (1969), and by Chester Jones, Ball, Henderson,
Baker & Sandor (1974).
1.6.2 The gonads
Figures 14 and 22 show the general position and form of the gonads
in male and female eels. In adults, the gonads lie freely suspended
almost throughout the entire length of the body cavity. The testes,
first described by Syrski (1876) are even less conspicuous than the
ovaries (and were sometimes referred to as the organs of Syrski before
their true identity was realized).
The location, and determination of the gonads as male or female,
is often difficult in immature specimens) fatty tissue, shown in Fig. 22, is
sometimes misidentified as gonad! A useful, although not totally
reliable method for macroscopic identification of the gonad was sugges-
ted by Walter (1910); alcohol is placed on the surface of fresh tissue
and this coagulates the albumen to give a more conspicuous, less
transparent, appearance to the gonads.
The gonads are of unequal size, with the right extending further
forward (about 1 em in 30 em eels), and the left reaching further
posteriorly (about 2 em behind the cloaca in 30 em eels). The left
gonad is about 2-3 % longer (Tesch, unpublished observations), is
heavier and concains more eggs than the right gonad (Matsui, 1952).
Kochnenko (1959) related head width to ovarian width; broad-headed
specimens had gonads about 5-6 nun across, while eels with more
pointed heads had ovaries about 9 nun wide. The earliest stage at which
it is possible to distinguish male from female gonads macroscopically is
when the fish reaches a length of about 20 em (Walter, 1910), although,
according to some authors, definite identification cannot be made until
a length of30 em has been reached (Sinha & Jones, 1966; Kochnenko &
Gorowaja, 1965; Satoh, Nakamura & Hibiya, 1962).
The developme-Tlt of male and female gonads is similar in European
and Japanese eels. Protrusions appear on the genital ridge or undifferen-
tiated cord of the potential gonad; these finally come to overlap one
another and are referred to as 'lobes' (Fig. 23). In the female the
developing folds come to resemble a tight frill, as ruffles develop along
the gonadial cord. Indeed, the female gonads are sometimes referred
to as the 'frilled organ' (Fig. 23). In the yellow and earlier stages of the
silver eel phases, testes and ovaries are insignificant-looking structures,
but in the fully mature animal they arc impressive structures (Fig. 24),
all too rarely observed by zoologists.
Fatty degeneration of gonadial tissue (Schnakenbeck, 1953) may give
an outwardly ovarian appearance to the gonads; a microscopic check is
necessary in such cases. The gonadial cells of degenerated ovaries are
very immature, and an unusually large number of fatty globules
44 THE EEL
Flg. n Above left: longitudinal section through the body of the female eel. a, Right
ovary; b, Left ovary; c/d, Section leading from the right/left ovary; e, Dividing
wall between the two gonads; C, Anal opening; g, Urinary bladder; h/i, Fat
on the right/left side which is often mistakenly thought to be the testis;
6, Fat on the stomach; k, Stomach; I, Pylorus; m, Liver; n, Gallbladder;
0, Pectoral fins. Below It/t: Anal region ofa female eel. a, Intestine; b, Fissura
recto-vesicalis; c, Urinary bladder; d, Anal-genital opening ; e, Dividing wall;
f, Urinogenital opening; g, Orifice of the genital opening into the urethra.
Above right: Longitudinal section through the body of the male eel. alb, Right/
left testis; c/d, Section leading from the right/left testis; e, Dividing walJ
between the two testes; f, Vas deferens; g, Bursa seminalis; h, Anal pit;
i, Urinary bladder (largely covered by the bursa seminalis); k, I, k', Fat on the
right/left side and on the stomach ; m, Stomach; n, Pylorus; 0, Liver (dis-
placed slightly to reveal the inner side adjacent to the oesophagus and the
stomach); p, Gallbladder; q, Pectoral fins. Below right : Anal region of a male
eel. a, Intestine ; b, Fissura recto-vesicalis (covered by the external wall of the
bursa seminalis); c, Opening of the anterior and posterior part of the vas
deferens into the bursa seminalis; d, Urinary bladder; e, Anal opening;
f, Dividing wall; g, Urinogenital opening; h, Orifice of the genital opening
into the urinary duct (after Syrski, 1876).
surround the oocytes. Such anomalies are possibly more common than
was first thought and some instances of eels in supposedly advanced
stages of maturity may indeed be examples of fatty degeneration
(Rasmussen, 1951 j Wundsch, 1953). In the studies cited, the fish were
Fig. 23 Male gonad from a 42 em long eel (above). Female gonad from a 57·5 em long
eel (below)(enlarged x 2)(frorn Kuhlmann, 1975).
THE EEL
Fig. %4 Male eel, 34 em long, in an advanced stage of maturity, caught in Danish

waten (Pristo Fiord, western Baltic, near the Ore Sound) (Schmidt, 11)06).
outwardly mature but the oocytes had diameters of o· 1 to 0·2 em, a size
range no larger than that found in normal immature silver eels. Bezd-
zenyezhnykh (1973, 1974) found resorption of the oocytes in silver
eels which had been prevented from migrating for a period of 10 years;
he considers this resorption to be phagocytic. The oocyres will
occasionally show simultaneous re-growth.
Microscopic examination of a smear or tissue section is of great
- $lEi, _ ~ :::;:;;:;:; ::.:
100m
Fig. 25 Schematic representation of the testes (gross macroscopic appearance) at

seven developmental stages. Stage I represents a normal, untreated silver eel
which was caught at the exit of the Baltic during its spawning migration. The
remaining sections show hormonally treated spttimens of stages 2 to 7
(after Boetius & Boetius, I¢qa).
THE EEL
diagnostic value, particularly since the results of microscopic and

macroscopic examinations do not always coincide (Sinha & Jones, 1966).
The fonner technique demonstrates oocytes in macroscopic lobes
although often only single oocytes occur in the lobes (Penaz & Tesch,
1970). It is possible that testicular tissue could still develop from this
apparently intersexual condition.
Figure 25 schematically shows the macroscopic development of the
testes. The primordial stage ( I ), perhaps the most commonly seen,
Table 3 Histological development of the gonads in the eel
A. anguiila A .japonica
After Rodolico (1934) and Bertin (1958) After Saloh, Nakamura & Hibiya ( 1962)
Glass eels: Cells of the germ~tell Glass eels : Germ tells of the 1st order
epithelium with large nuclei
(protogonia), foremnnen
of the sex tells
From 10 to 12 em Multiplication of the

long: protogonia, which become
increasingly less differen-
tiated deuterogonia
From 14 to 18 em First sexual differentia- Up to 15cm Germ cells of the 1st order
long: tion. Some deuterogonia long: in increasing numbers
become oogonia which later
develop into oocyles, i.e. 18 cm long : Germ cells of the I St order
undifferentiated and female enlarge and become oocytes.
cells co-exist Detectable sexual
differentiation
From 18 to 30 em Other deuterogonia divide

long: and form bands of 99 :ro cm long : Oogonia become recogniz-
spermatogonia, later becom- able, some develop into
ing spermatocytes; i.e. oocytes. It is possible to
male and female sex tells distinguish male gonads
now co-exist giving rise to a externally
condition of juvenile
hermaphroditism. In 10%
of individuals only female 99 25 t 040 cm Oogonia and primary
gonads develop. long: oocytes intermixed
Evenrually, one or other 00 40 em long: Large numben of sperm a-

sex predominates. togonia enlarge, but no
spermatocytes developed
01$ emigrating Spennatocytes present, but

silver eels : no spermatids
99 over 40 em
long: Oocytn grow fast
99 emigrating
silver eels: Yolk cells develop fast
leads to the more advanced stages of maturity. The sketches shown
were made from an eel treated with hormones; the more advanced
stages are rarely encountered in nature.
Matsui ([952) and Kuhlmann ([975 ) (see also Kochnenko & Bezd-
zenyezlmykh, 1973b) have described the histology of the developing
gonad of eels. Immature ova are readily distinguishable from immature
sperm by virtue of their greater size.
The histological stages of gonadial development have been related
to body length, and such information for A. anguilla and A. japonica is
summarized in Table 3. Sexual differentiation in these two species of
eel occurs when the fish is about 30 cm long. Size and gonad development
are probably similarly related in other species of eel. Species in which a
large adult size is reached, for example A. dieffenbachi of the South
Pacific, probably show a similar developmental pattern, but with
relatively larger proportions. The findings presented in Table 3 are,
to a certain extent, contradictory as regards the sexual differentiation
of the young eel. Thus Satoh et ai. (1962) found no intersexuality in the
Japanese eel. On the other hand, reference should be made to Kuhlmann
(1975), who was able to identify an intersexual stage. Ecological and
experimental studies of the European eel (van Oordt & Bretschneider,
1942; J.J. Tesch, 1928; Gandolfi-Hornyold, 1932b; D'Ancona, 19513;
Penaz & Tesch, 1970; Kuhlmann, 1975) have shown that varying
degrees of ambisexuality in young eels can result from environmental
factors (see also Sinha & Jones, 1966) and the subject remains contro-
Fig. 26 Eggs laid by A. rostrata in seawater. The egg in the centre has reached the
most advanced stage of maturity and has a diameter of 1.25 mm. It has oil
globules in the yolk. The smaller eggs (the greatest diameter is 0·60 mm) are
less mature and have a dark ball of yolk (from Edel, 1975a).
50 THE EEL
versial. It should also be pointed out that gonadial growth is more

marked in the autumn than in winter (Sareh et al., 1962).
More information is available on the later development of the
gonads in the male eel once it has left Continental waters; furthermore,
a greater degree of maturity has been induced artificially by hormone
treaunent in this sex (e.g. Boetius & Boetius, 19(73). Boucher, Boucher
& Fontaine (1935) were the first to use this method successfully.
Figure 25 shows, schematically, testicular development in such prepara-
tions. At stage 5 the testes have maximally distended vasa deferentia in
which the spermatozoa are stored. During stage 7 (not shown) spawning
hegins, after which the gonad is barely distinguishable from the first
developmental stage. The hormones used to induce maturity in males
came, in some cases, from the urine of pregnant women (Boucher et al.,
1935; Tuzet & Fontaine, 1937) ; other hormones used include prolan
(Fontaine, 1936; EtieIUle, 1959), carp hypophysial extract, gestyl,
ambion pregnyl (van Oordt & Bretschneider, 1942), prolactin, thyro-
tropic hormone (Olivereau, 1961a), human chorionic gonadotrophin
(Boetius & Boetius, 1967a; Billard & Ginsburg, 1973; Lumare & Villani,
1973 ; Bieniarz & Epler, 1975) and calf spleen extract (solcosplen)
(Meske & Cellarius, 1972). It seems that male eels show little species
specificity to exogenous hormones, but females are more sensitive in
this respect.
In seawater the weight of the mature gonads in male A. anguilla
comprises 3'6% of the body weight, according to Meske & Cellarius
(1973), and 9'6% according to Bieniarz & Epler (1975) ; in freshwater
this percentage varied between 7'3% and 7'5 % . Gonads of untreated
animals weighed between 0'2 and 0'6% of the body weight.
On the basis of electron microscope studies, Ginsburg & Billard
(1972) have described the morphology of sperms in A. anguilla, and
Colak & Yamamoto (1974) have done the same for A. japcmica.
The induction of sexual maturity in female eels using exogenous
hormones has proved more difficult than in males. For example,
research carried out in Denmark on the European eel by Boetius ec al.
(1967) used human chorionic gonadotrophin combined either with a
synthetic oestrogen (diparaoxyphenylhexan) or with oestradiol prop-
ionate from pregnant mares. Oocytes up to 0'56 nun in size and maxi-
mum ovarian weight of 12'6% of body weight were induced. The
corresponding figures in untreated females were 0'2 mm and 1'7 %
respectively. An egg size of at least 1 mm can be expected at spawning.
Fontaine et al. (1964) were the first to successfully induce sexual
maturity in female A . anguilla. They used an extract from carp hypo-
physis and then, three months later, shortly before maturity, desoxy-
corticosterone acetate was also administered. In an earlier experiment,
the use of prolan and oestradiol monobenzoate had not been
totally successful (Fontaine, 1961). After these initial experiments
various other workers managed to produce spawning female eels. At
intervals of 5-10 days Nose (I971 ) administered mammalian gonado-
trophin (synahorin; each dose containing 40 IV) and synthetic oestro-
gen (diethylstilbestrol; each dose containing 0'2 mg), combined on each
occasion with 30 IV of alpha-tacopherol (vitamin E). In just under three
months sexual maturity was achieved. Ochiai el al. (1972, 1974) obtained
sexually mature female A. japonica after four months of treatment
combining follicle hormone (estradiol dipropionate) and gonadotrophin
(synahorin). Various compounds have been used in experiments on
A. anguilla by Italian workers (Villani & Lumare, 1975). The most
successful preparation appeared to be carp hypophysis, backed up by
injections of choriogonadotrophin; deoxycorticosterone was less
effective. Sexual maturity was attained after four months. It seems
that extract of hypophysis from fishes contains the most effective
hormone for producing sexual maturity ; Edel (1975a) reached a
similar conclusion in experiments on A. rostrala (see also Yamamoto
el al., 1974a). Edel used ground carp hypophyses which had been dried
in acetone; this extract produced spawning after It months, a much
shorter period than in the other species mentioned.
Injections can be administered both intramuscularly or intra-
peritoneally (Boetius et al. , 1962). Maturity is accompanied by decalci-
fication of the bones (Lopez et al., 1970), mainly through a reduction
in calcium phosphate.
The diameter of the eggs is 0'9-1 '4 mm in A. anguilla, 1'0-1'3 mm in
A . japonica and 1'1 mm in A. rostrala. In addition to the yolk granules,
the mature eggs (Fig. 26) of all three species also contain droplets of
oil (Fontaine et ai., 1964; Yamamoto el ai. , 1974bj Edel, 1975a).
Gonad weight comprises some 35-45 % of the total body weight.
According to Vallani & Lumare (1975), the gonads of untreated silver
eels (Anguilla anguilla ) comprise 1'0-1 '85 % of the body weight,
those of A. rostrata, however, comprise 2'4-6.3 % (Wenner & Musick,
1974), or 5'9-6'0% (Edel, 1975a). Boetius el al. (1962) also found that
the gonads of untreated animals are heavier in A. rostrala (4'5-6'8 %)
than in A. anguilla (1'7%). The diameter of the oocytes at the silver
eel stage is also correspondingly larger in the American eel (0'45 mm)
than it is in the European eel (0'20 mm) (Boetius et al., 1962), indicating
a more advanced developmental stage of the west Atlantic species at the
52 THE EEL
beginning of the spawning migration . It should be noted here that silver

eels from southern North America have larger, and therefore more
mature eggs than do animals from Canada. In addition, it is important
to point out that, in a comparison of eels of the same age but different
sizes, larger individuals have more fully developed ovaries (i.e. their
oocytes are larger in diameter) than do smaller eels (Kochnenko &
Bezdzenyezhnykh, 1973a). Japanese experiments (Yamamoto et ai.,
19743-) have shown that hormone injections cause the body weight of
female eels to increase by varying amounts. In one case body weight
also showed a marked increase in A. rostrata (Edel, 1975a).
Oogenesis in eels does not vary greatly from that in other species of
fish (Yamamoto er aI., I974b). The number of eggs in a mature A. rostra-
ta weighing 560 g totalled 1'3 to 1'5 million (Edel, 1975a). Similar
figures have been published for silver eels ( A. rostrata) from Chesapeake
Bay (Wenner & Musick, 1974). The number of oocytes ( Y ) can be
calculated from the body length ( X ) as follows:
log Y ~ - 4.2951 + 3.74418 log X
By producing male and female A. japonica that reached sexual
maturity simultaneously, Yamamoto and Yamauchi (1974) were the
first to successfully carry out artificial fertilization of eel eggs. This
experiment involved 20 mature males treated with synahorin, and five
females, which had been injected with extract of salmon hypophysis
(8 hypophyses, extracted in acetone, per kilogram of eel body weight;
see Yamamoto et al., 1974a). Fertilization was effected in salt water at a
temperature of 23 °C. The larvae began to hatch 38-45 hours later and
survived for five days, during which time they grew from 4'8 nun to
6'2 nun in length. On the third day the mouth and anus were open and
the pectoral fins appeared. After five days the larvae had developed
53 pre-anal and 48 post-anal myomeres, i.e. a total of 15 myomeres
fewer than the number of vertebrae recorded in A.japonica (Table 5).
An important and controversial issue from both scientific and practical
view points is the question of sex determination. It is clear that micro-
scopically recognizable 'intermediate' (intersexual) stages occur in
eels. Thus the gonadial anlage passes through phases during which
either male or female types could develop, and the direction of develop-
mentmay be influenced by envirorunental factors. This does not exclude
genotypic sex determination and the presence of sex chromosomes
(Lieder, 1963); Anguilla japom'ca, A. rostrata and A. anguilla all
contain 19 pairs of chromosomes (2n = 38) (Chiarelli el al., 1969 ; Sick
et al., 1962) but the proportions of different types (acrocentrics, meta-
centrics, etc. ) vary, and diploid components clearly differ (Ohno et ai.,
1973; Passakas & Klekowski, 1973). Indeed, intersexuality may only be
displayed by 10% of eels within a population (Rodolico, 1934; Bertin,
1956, 1958). Such eels are syngamic females in which genetically
determined sex is uninfluenced by, or overrides, envirorunental
influences. Gene actions vis-a-vis sex determinations can be modified
by environmental factors, although precisely how these effects are
elicited requires more detailed study.
It has neither been possible to induce intersexuality experimentally
nor (0 itemize exactly the environmental effects upon sex determination.
Several attempts have been made to influence the sex of differentiated
and undifferentiated eels by different environmental conditions. These
studies have been widely discussed (Bertin, 1956, 1958; Sinha & Jones,
1966; Penaz & Tesch, 1970) and merit a brief consideration. Eels, less
than 25 em long and caught in the Ijsselmeer, where normally 96 % of
the catch are males, were put in a 16 m 3 tank with running water and
were fed for several years (].J. Tesch, 1928). A sample of the animals
was tested after a year and all were clearly males. At the end of the
third year the eels were 30 to 45 em long and all possessed female sex
organs. Gandolfi-Hornyold ( I932b) carried out similar studies using
90 Grande Briere yellow eels of which less than 10 % were found to be
female. They were held in a I m 3 tank and fed raw meat. After 3-4 years,
420fthe surviving eels (all 23 to 36 em long) were sexually indeterminate,
10 were females and 15 to 17 were males. Thus, this experiment pro-
duced similar but less convincing results than the other studies
mentioned.
D' Ancona (1950), in Italy, has investigated the effects of population
density on sex determination by using different numbers of glass eels
held in ponds 2500 m 2 in area. After three years it was found that the
less dense the population the more females there were. Schnakenbeck
(1944, 1953) also suggests that in general females develop from un-
differentiated eels and from male eels when the latter originate from a
crowded population and then enter waters in which there is less competi-
tion from other eels. In the study cited, eels from the thickly populated
areas of the Lower Elbe were used to stock the ponds. After transfer,
the animals grew well and most had female gonads. (For further
examples, see Kuhlmann, 1975.)
Ecological studies also indicate a link between sex determination
and environmental factors, In the Japanese eel pond industries, a total
of8558 individuals held in crowded conditions contained only 50 female
specimens (Matsui, 1952). Similarly, Egusa and Hirose ( 1973) found
54 THE EEL
that out of 611 Anguilla anguilla, 30-40 em long, held in ponds, 92-4%
were males. In contrast, in the less densely populated Japanese lakes
and rivers 729 female eels were present in a sample of 2534 individuals.
Muller (1967), after introducing some glass eels (presumably of the
same origin) into a .pond, found a huge preponderance of males. If no
selection took place before the glass eels were caught, then the eels had
not sought out their biotope (i.e. the pond) according to genotypically
specific characteristics of males and females.
D'Ancona (1958, 1959b, 1961) suggested that the differential
distribution of the sexes in river estuaries, in inland waters, and in the
east and west Baltic, can be attributed, primarily, to the differential
levels of migratory drive correlated with the sex genotypes. As a result,
males, with a low migratory drive, remain for example in the estuary of
the Elbe, while the more active females migrate further up the Elbe
to produce the well established preponderance of females in the Upper
Elbe, its tributaries and nearby lakes. This hypothesis is supported
by many studies on the distribution of the two sexes (see Table 4;
Walter, 1910; Gandolfi-Homyold, 1925; Ehrenbaum, 1930; Callamand,
1943)·
More recently Penaz and Tesch (1970), however, have been unable
to confinn such differential sex distribution in the Elbe area. It is
indeed true that in the Elbe the proportion of males is at its greatest
just before the river enters the North Sea. It is also true that the propor-
Table 4 The proportion of female A.anguilla and A.roSlrala in different water

bodies; arranged according to length
No. of eels Proportion i.mg"
Locality studied af females (em) Aut'\'or
('Yo )
South-eastern North SlQ ana L~r Elk
'"
He1igoJand OS 30--76 H eineke, 1894
Wadden Sea near Seherrebek, Denmark ,60 '3 20--30 Marcus, 1919
Wadden Sea near Seherrebek, Denmark
Lower Elbe ",
37
"7
30-40
20-30
Marcus, 1919
Ehrenbaum &
Marukawa, 1914
Lower Elbe "7 3' 30-", Ehrenbaum &
Marukawa,1914
Slor (tributary of the Lower Elbe)
Sior (tributary of the Lower EJbe)
Niederelbe
'"
'7
443
9
",
20-30
30-40
20-30
Marcus, 1919
Marcus, 1919
Marcus, 1919
NiedereJbe
Lower Elbe
8,
3'
,
'9 30-40
28-30
Marcus, 1919
Wundseh,1954
"
Lower Elbe ,8 31-34 Wundseh, 1954
Alsler 66 9 20-30 Ehrenbaum &
Marukawa,19 14
BOD Y STR UC T U RE AND F UNC TION S 55
T able 4 continued
No. of ells Propor/wn !.mg,h
Locality studied offemaleJ (= ) Author
(%)
Alster '4' ,8 3~'" Ehrenbaum&

Marukawa, 19 14
£1"
Saalc near Calbe 66 93 20-30 Marcus, 1919
Saale near Calbe
Rhin near Fehrbellin
"4
8,
'00
78
30-40
20-30
Marcus,
Marcus,
1919
1919
Rhin near Fehrbellin <0, 97 30-40 Marcus, 1919
WeStr
Weser near Bremerhaven , 20-30 Marcus, 1919
Weser near Bremerhaven '" 80 Marcus, 1919
" 30-40
Balti,
Western Baltic ' 57 8, 30-46 T hurow, 1959
Neuendorfer Wiek and Breeger Bodden (Rugen)
Strelasund "
25-35
30-35
30-35
Schlumpberger,
Lauterbach &
Falk, I964
Greifswalder, Lebbiner, CrolXr Jasmunder Lauterbach &
and Sule Bodden and Wismar Bay 4~60 30-35 Falk, 1964
Kleines Haff, Kleiner Jasmunder Bodden Lauterbach &
and Lower Warnow 70 30-35 Falk, 1964
Trave near Lubeck 460 ,8 20-30 M arcus, 1919
T rave near Lubeck 60 93 30-40 Marcus, 1919
Lake Dassow (an inland lake)
'" 36 20-30 Ehrenbaum &
Marukawa,I914
Lake Dassow (an inland lake) 53 68 30-40 Ehrenbaum &
Marukawa,1914
Lake Schmollen ,60 ,6 20-30 Marcus, 19 19
Lake SchmoJlen 83 99 30-40 Marcus, 19 19
Lake Schmollen 3' 38 20-57 Wundsch, 1916
Lake Aeppel and Paddcnphul 8, 29-46 Wundsch, 1916
.,.,
94
Lakes in Lithuania 65-80 > 35 Malschenis,
Southem North St a and the Rhine t$luary

Ijsselmeer 2584 ., 28-42 J.J. Tesch, 1928
"
Wadden Sea 715 28-42 J. J. T esch, 1928
Haringvliet, Holl. Diep. '00'6 43 28-42 J .J. Tesch, 1928
Wul
Inland waters in Holland
'4
,8, "7' 28-42
28-42
J .J. Tesch,
J .J. Tesch,
1928
1928
Atlanti,
Bermuda' 80 ,. 18-58 Boetius &
Boetius, 1'}67b
EaST roast of North Ameri,a
Newfoundland
New Brunswick '"
,,8
94
80
19-93
21 - 64
Vladykov, 1966
Vladykov, 1966
Nova Scotia
Quebec
37
,6, ""99 20-73
19-128
Vladykov, l966
Vladykov, 1966
'00
Ontario
'<06
" 42- 103 Vladykov,l966
"
Massachusetts <07 26-42 Vladykov,l966
New York 67 23-63 Vladykov,l966
lAo rostrata
THE EEL
tion of males decreases up-river. In the upper waters there is an extreme-

ly high proportion of females, but the proportion of females also
increases from the river estuary towards the sea. Around Heligoland,
for example, there is an extremely high proportion of females. The
proportion of males roughly correlates with population density:
high densities with a large proportion of males (as well as hennaphro-
dites) occur in the estuary. Population density gradually decreases
upriver since the influx of glass eels is gradually reduced in these more
distant regions. The density of eels also decreases near the sea, and here
again there is an increased proportion of females. [The population
density decreases near the sea because few of the invading glass eels
remain in that region, most moving inland (Creutzberg, 1961; Tesch,
1971).] The proportion of males also drops towards the sea in certain
cases: the population of the Ijsselmeer comprised 94% males but there
are only 70% in the Wadden Sea which is nearer the open sea (J.J. Tesch,
1928). In small rivers on the east coast of North America it has been
shown that males predominate, although the proportion of females
increases in the estuaries (Winn et ai., 1975). Other studies in lakes
(Rasmussen, 1952; Frost, 1950; Rep. Lough Neagh Minstr. Agn·cult.
Belfast, 1966) also relate male predominance to increased population
density. In these studies large numbers of glass eels were introduced
into lakes and the proportion of males increased, as the total catch
increased, from 0% to 62%. Research in the Camargue, the estuary
area of the Rhone, revealed vastly different figures for the proportions
of males in three nearby waterways in the Rhone plain (Gandolfi-
Hornyold, 1930, 1935). The male to female ratios in the three places
were 45 :55, 32 :88 and 6 :94. Thus a different migratory drive is an
inadequate explanation for the sex distribution. These few, of many,
examples therefore suggest that the eventual sexual differentiation of
eels is not entirely dependent upon genotype. Other results in this
field need further critical investigations. For example Bellini (1907; see
also Walter, 1910; and Kuhlmann, 1975) observed two different sized
groups of glass eels, and from a breeding experiment concluded that
the larger animals were females. More recently Vladykov (1966) has
shown two peaks of size frequency among glass eels, which seem to be
more related to geographic origin. Temperature, length of migration,
etc. could well explain the observed size differences among a population
(see also p. 134 et seq.).
The relation between a high population density and a high proportion
of males or hermaphrodites suggests that competition for food and
hence relative malnutrition stimulates maleness (see also Burnet, 1969a).
This is supported by the fact that males grow more slowly (Bertin,
1958; Sinha & Jones, 1967c; Penaz & Tesch, 1970) and are smaller at
sexual maturity. The slower growth rate is found largely in those
younger age groups in which sexual differentiation takes place. (Fig. 75;
Sinha & Jones, 1966, 1967a; Penaz & Tesch, 1970). For example,
Heligoland eels, with a high proportion of females, grew more rapidly
during the first three years of their stay in Continental waters. On the
other hand, the older (already differentiated) eels showed a normal or
poor rate of growth, similar to English eels.
Laboratory experiments, however, do not support these ecological
observations (Meske & Cellarius, 1973; Kuhlmann, 1975) ; Egusa and
Hirose (1973) found that pond cultured males and females grow at the
same rate until they have reached a length of 40 cm.
In goldfish, malnutrition, by affecting hypophysial gonadotrophic
function, inhibits gonadial growth (Bogenschutz & Clemens, 1967).
In eels, it may be that ovarian growth is inhibited in an analogous
fashion. Work by Kuhlmann (1975) suggests that sex determination
in Anguilla anguilla may be influenced by the quality of its food;
feeding elvers on cod ovaries (Gadus morhua ) resulted in a higher
proportion of females.
It has been suggested that sodium ion concentrations might affect
the release or action of gonadotrophic hormones, which differentially
influence the ovaries rather than the testes. In addition, che increased
salt concentrations effectively lower the water content of the organism
and again, by an action upon the hypophysis, might favour maleness.
For example male Gambusia holbrooki have a lower water content than
do females. Such interpretations require further study in particular with
regard to prolactin function which has important effects upon water and
electrolyte metabolism, but as yet its gonadial effects are incompletely
understood (Ball, 1969). No difference in sex ratio could be detected
between eels grown in fresh- and saltwater ponds (Egusa & Hirose,
'973)·
Environmental salinity cannot have direct, decisive actions on sex
determination in the eel, however, since males and females occur
together in large numbers at both high and low salinities (Sinha &
Jones, 1966; Penaz & Tesch, 1970).
No clear-cut relationship between temperature and sex determination
can be established from Kuhlmann's (1975) experiments on the
European eel. Exposure to moderate temperatures (20 ° and 23 °C)
resulted in a preponderance of males, low and high temperatures
seemed to favour female production, but extremely high temperatures
produced no definite bias towards either sex.
It is noteworthy that the same experiments suggested that the
THE EEL
geographical origin of the elvers might have some influence on sex

determination; elvers from the Tyrrhenian Sea produced more males
than did those from the North Sea.
1.7 The endocrine system

The eel has proved to be an eminently suitable fish for studies of
endocrine mechanisms. It is able to adapt to large alterations in environ-
mental salinity, both acutely and chronically, and can withstand massive
surgical interventions. The pituitary gland, interrenal (adrenocortical
homologue) tissue, thyroid gland, gonads, corpuscles of Stannius,
ultimobranchial bodies and thymus have all been subjects of study.
Particular attention has been paid to endocrine factors involved in
osmoregulation, although other aspects have begun to be examined,
including intermediary metabolism, growth and reproduction.
1.7. I The pituitary gland

Figure 27 shows the position and topographic relations of the pituitary
gland, situated at the base of the midbrain behind the optic chiasma
Sir.
•
•
lub. Ie ....
lin.y
, Sir. tin,.onl .lp.-slr.
JHllI.
v--c.._fin.ltr·OP.-llr.
\~,L/ 1111.y
Iin,. lat . lp._lIr. -"'-J
Fig. "7 The eel's brain. A, from below; B, Corpora striata from the side; C, Corpora
slriala from above. ep.-str., EpistrialUm; fiss. str., ep.-str., Fissura striata-
epistriatica; fiss. y., Fissura ypsiJiforrnisj h, Hypophysis; ling. am. ep.-str.,
Lingua anterior epistriatica; ling. lat. ep.-str., Lingua lateralis epistriaticaj
pall., Pallium; str., Striatum; s.v. Saccus vasculosusj tub. taen., Tuberculum
laeniae j VIII nerves (after Lissner, 1923).
and in front of the saccus vasculosus. I ts connections and generalized
topography with regard to the hypothalamus and ventricular system
conform with the overall pattern of the vertebrate hypothalamo-
hypophysial system (Holmes & Ball, 1973). The weight of the hypo-
physis varies considerably; for example Callamand (1943) found an
average hypophysial weight of 3'4 mg (range 1-7 mg) in a group of
10 eels whose mean body weight was 520 g.
The component parts of the pituitary body are distinguishable on
superficial examination into anterior and posterior sections. On closer
histological examination the two primary divisions are apparent
(Fig. 28). The central neurohypophysis interdigitates with the shell-
like adenohypophysis. The adenohypophysis is subdivisible into the
anterior pars distalis, with rostral and proximal parts, and the posterior
pars intermedia. The neurohypophysial portions come into intimate
contact with the pars intermedia; their communications with the pars
distalis are less obvious. Olivereau (1967) summarizes the extensive
studies of the cell types contained within the eel pituitary, but reference
should also be made to Florentin and Weiss (1931), Honma (1957),
Jazinski (1961 ) and Vollrath (1966). Ball and Baker (1969) review the
literature on the cytology and function of the hypophysis of the eel and
other teleosts.
The cytology of the cell types - gonadotroph, thyrotroph, somatot-
roph, eta cells, alpha cells and neurohypophysial fibres - changes
within the individual's life history, as well as with the salinity of the
environment, adrenocortical status, sexual state, etc. Such changes have
contributed much to contemporary allotment of function to the parti-
cular cell types (Bernardi, 1942, 1948; Olivereau, 1954, 1966a; D'An-
cona, 1951b; Olivereau & Fontaine, 1966; Evans, 1940; Kathuria,
1972). Neurohypophysial peptide content as well as the degree to which
this system innervates the pars distalis also varies with stage of develop-
ment (Holder, 1968; Knowles & Vollrath, 1965b, 19(6).
Pituitary function may be related to the haiotropism of eels. In
particular the migrating glass eel, which moves from seawater into less
saline environments (Deelder, 1958, 1960b; Tesch, 1971 ), has been
shown to be especially sensitive to hypophysectomy. If the pituitary is
removed in Stage VI A IV (see p. 15; Suubberg, 1913) 70% of the
animals fail to move from tanks of seawater into freshwater when offered
the choice; control animals all made the excursion within two hours
(Vilter, 1946b).
Fontaine, Callamand and Olivereau (1949) were among the first to
recognize pituitary involvement in osmoregulation of the eel. The
60 THE EEL
Fig. 28 Structure of the pituitary gland of A. anguilla. A: Agr, Anterior lobe; Brc,
Connection with the base of the brain; Cav, Cavity in the anterior lobe;
Hyp R. Recessus infundibularis and its diverticulum; Hyp F. Opening into
the posterior lobe (with diverticula) of the neurohypophysis; Mgr, Middle
lobe; 0, Acidophil cells; e, Basophil cells; Nl, Neurohypophysis;
Pgr, Posterior lobe (after Vivien, 1958, modified from Kerr, 1942). B, the
anterior lobe (rostral pars distalis) : c, Corticotropic cells; p. Prolactin cells;
t, Thyreotropic cells (black). The pars distaJis: g, Gonadotrophic cells (finely
stippled); s, Somatotropic cells (coarsely stippled); i, Pars intermedia;
N, Neurohypophysis (modified after Olivereau, 1967).
changes that occur in body fluid homeostasis after removal of the

pituitary depend in part upon environmental salinity. Thus, hypo-
physectomized eels in freshwater display hyperhydration, hypona-
traemia and disrupted calcium and magnesium metabolism. In seawater
adapted eels, hypernatraemia and hypophydration are characteristic
symptoms. In other words the routes for exchange of water and electro-
lytes have been modified such that the animal is unable to maintain the
gradients between its body fluids and the environment. Renal, branchial
and gastrointestinal functions are abnonnal after removal of the
pituitary. Electrolyte transport and membrane penneabilities are
affected (Hirano, 1967; Hirano el al. , 1967; Chester jones, Henderson &
Butler, 1965 ; Butler, 1966; Chan, Chester jones, Henderson & Rankin,
1967; Henderson & Chester jones, 1967; Chan, Chester jones &
Mosley, 1968). Many of the changes may reflect directly the absence of
pituitary factors especially prolactin, but others result from secondarily
impaired adrenocortical, thyroidal and other endocrine functions
(see Olivereau & BaU, 1970).
Prolactin has been one of the most thoroughly studied hypophysial
hormones. The eel differs from cyprinodont fishes in that it does not
die after hypophysectomy in freshwater (BaU, 1969). In this medium,
body fluid homeostasis is, however, compromised as a result of both
renal and extrarenal defects (Chester jones et al' 1969a,b). There is a
J
slow and progressive reduction in plasma sodium, potassium and

calcium concentrations, and ovine prolactin is able to prevent or retard
the diminutions (Olivereau & Chartier-Baraduc, 19(6), a finding
contradicted by Butler (1967). Cytological changes in the prolactin-
producing eta cells (Ball & Olivereau, 1964) of the pituitary correlate to
suggest a role for prolactin in the hyperosmoregulation of eels in
freshwater. I t is now generally accepted that a primary action of prolac-
tin in eels is to sustain reduced branchial water permeability and
sodium fluxes (Maetz, Mayer & Chartier-Baraduc, 1967). It seems also
that prolactin and cortisol (or ACTH) act in concert to maintain plasma
composition of freshwater adapted eels (Chan, Chester jones & Mosley,
1968).
The anatomical relationship between neurosecretory fibres and
pituicytes of the hypophysis suggest a functional connection between
the hypophysis and the hypothalamus (Knowles & VOllrath, 1965a).
The activity of the hypothalamus-neurohypophysial complex is
influenced by environmental changes such as shock from electric
fishes, transfer from freshwater into saltwater, and alterations in light
intensity. All these changes influence the whole or part of the complex
(Leatherland & Dodd, 1969) ; Holder (1970) describes temporal
transport rates of uterotonic (oxytocic) hormones along the preoptico-
hypophysial tract of hypophysectomized eels. The relationships
between hypothalamic nuclei and the pituitary gland, as well as putative
62 THE EEL
hypothalamic releasing and inhibitory hormones have heen discussed

by Hawkins and Ball (1973) and Ball, Baker, Olivereau and Peter (1972).
Two neurohypophysial peptides in eels - 4-serine-8-isoleucine-
oxytocin (isotocin) and 8-arginine-oxyrocin (arginine vasotocin) -
have been described by Follett and Heller (1964), and Holder (1968,
1969, 1970). Neither peptide has been identified in plasma of eels kept
in air, in seawater or in freshwater (Bentley, 197 I). The renal and
extrarenal actions of these peptides (Chan & Chester Jones, 1969;
Chester Jones et al., 1969b; Henderson & Wales, 1974) as well as the
changes that occur in neurohypophysial cytology with season and
salinity CArry, Fontaine & Gabe, 1959; Leatherland, Budtz & Dodd,
1966) require the demonstration of changed circulatory concentrations
of those hormones before finn physiological conclusions can be reached .
1.7.2 The thyroid gland

The thyroid gland of eels (see Hagen, 1936) is not a discrete strucrure,
and does not show marked macroscopic changes during the various
stages of development. The gland is situated rostral to the origin of the
afferent branchial arteries, lying under the urohyal bone (see Fig. 29).
Anguilla anguilla differs somewhat from A. rostrata in the distribution
of thyroidal tissue (Fig. 29). The histology of thyroidal follicles changes
dramatically with the stage of life history. In particular the number of
follicles and their diameters increase tenfold during metamorphosis so
that in eels 15 to 20 em long the gland may appear lobular and be pigmen-
A B
Fig. 29 Position of the thyroid gland relative to (he gill arteries. A, in A. rostrata
(after Gundematsch, 191 I); B, in A. anguilla (after v. Hagen, 1936).
t, Bifurcation of the ventral aorta into the first gill artery; II, Second gill
artery (VEB Gustav Fischer Verlag, Jena).
ted (Sugiyama & Sato, 1960; Sklower, 1930; Murr & Sklower
1928), while thyroid activity may be reduced when feeding begins
(Vilter, 1943).
The migratory behaviour of young eels appears to have limited
connections with thyroidal activity. However, processes such as the
increase in relative length of the intestines during the glass
eel stage and at metamorphosis may be influenced by thyroxin
(Vilter, 1945a). In no way, however, is pigmentation controlled by en-
vironmental factors such as putting the eel into freshwater. During that
period of development in older yellow eels towards the silver stage, the
activity of the thyroid gland increases (Leloup, 1959). The structure
and function of the thyroid gland is under strict if not complete control
of the hypothalamo - hypophysial system (Olivereau, 1955a, b, c, 1962;
Knowles & Vollrath, 1966; Baker, 1969). Other interactions, including
those with the corpuscles of Srannius (Leloup & Leloup-Hatey, 1964),
with environmental temperatures (Leloup & Fontaine, 1959), as well as
those with prolactin, have been discussed (Hopkins & Baker, 1968;
Olivereau, 1966a, b).
1.7.3 The urohypophysis

The caudal neurosecretory system (urohypophysis) is apparently
concerned with osmoregulation in the eel (Sano, 1961). In Japanese
eels up to 7 em long this organ is, in contrast to the neurohypophysis,
still relatively underdeveloped. Later, however, differentiation occurs
very quickly and the total amount of neurosecretory materials in
individuals of only 7'5 em is about the same as in adult animals (Katsu-
toshi, 1965a). The development of the caudal neurosecretory system
is controlled, presumably, by internal (neural) factors and not by
external osmotic changes.
Ishibashi (1962) has studied the nerve supply to the eel's urohypo-
physis and its electrophysiological connections. Katsutoshi (1¢i5b) and
Inano (1959) have described how this organ regenerates. The character-
istics of the urohypophysial vasoactive materials have been studied by
Chan, Chester Jones & Ponniah (1969), and their influences on the
caudal circulation have been discussed by Chan (1971 ) who more
recently (1975) has suggested that two hormones emanate from the
urohypophysis, one of which is active in seawater and the other in
freshwater environments. They have their effects at both vascular and
renal sites (see also p. 42).
THE EEL
1.7.4 The interrenal tissue (adrenocortical homologue)

Interrenal tissue occurs in the head kidney about the walls of the
posterior and anterior cardinal veins near the ducts of Cuvier. The
tissue can be surgically removed (Chester jones, Henderson & Mosley,
1964) so that it is possible to examine the function of the honnones
in the absence of endogenous production. The major corticosteroid of
the eel is cortisol, although other materials have been described (Sandor,
Vinson, Chester jones, Henderson & Whitehouse, 1966j Sandor,
Lanthier, Henderson & Chester jones, 1967 j Sandor, 1969; Hargreaves
& Porthe-Nibelle, 1974). The production rates of cortisol are greater
in eels adapted to seawater than in animals in freshwater (Leloup-
Hatey, 1974; Henderson, Sa'di & Hargreaves, 1974; Henderson,
jackson & Hargreaves, 1975). The functional and structural capacities
of the interrenal tissue is dependent upon pituitary ACTH (Hanke &
Chester jones, 1966j Olivereau & Fromentin, 1954j Henderson et al.,
[974)·
Cortisol is active in maintaining both renal and extrarenal metabolism
of sodium in fresh- and seawater environments (Mayer el al., 1967 j
Chan, Chester jones & Rankin, 1969; see also Henderson et al., 1970j
Chester j ones, Chan, Henderson & Ball, 1969 j Chester jones, Bellamy,
Chan, Follett, Henderson, Phillips & Snart, 1972). Other aspects of
corticosteroid physiology are less well known, although primary or
secondary effects upon the cardiovascular system, reproduction and
intennediary metabolism of eels have been indicated (see Chester
jones, Chan, Henderson & Ball, 1969 ; Chester Jones, Bellamy, Chan,
Follett, Henderson, Philips & Snart 1972 ; Henderson el al., 1970).
1.7.5 Ultimobranchial bodies

The ultimobranchial bodies, situated close to, although histologically
clearly distinct from, the thyroid gland are almost certainly the site of
calcitonin production in the eel. They may be involved with calcium
metabolism although the evidence thus far is indirect. Ultimobranch-
ialectomy, injections of heterologous calcitonin preparations, removal
of the corpuscles of Stannius or injections of oestrogens have produced
data which are not physiologically coherent, although histophysio-
logical studies do relate ultimobranchial bodies to calcium turnover
and bone deposition (Chan, 1969, 1972 ; Lopez et ai., 1968; Chan,
Chester jones & Smith, 1968). An unequivocal role for calcitonin has,
however) yet to be shown.
1.7.6 Corpuscles ofStannius

The corpuscles of Stannius are paired bodies lying on the posterior
kidney of the eel and embryologically are derived from the pronephros.
Removal of one corpuscle leads to hypertrophy of the other (Pettit,
1896; Lopez & Fontaine, 1967), and Callamand (1943) noted a partial
correlation between corpuscular size and the gono-somatic index in
yellow and silver eels. In spite of earlier claims for a steroid-secreting
function for these organs (Fontaine & Leloup-Hatey, 1959; Cedard &
Fontaine, 1963), other biochemical, histological and ultrastructural evi-
dence argues against such a role (Chester jones el ai., 1965; Chieffi &
Botte, 1963; Hanke & Chester jones, 1966). The current views are that
corpuscles of Stannius produce a honnone, most probably a peptide,
which is intimately concerned with calcium metabolism. The identity
of this principle, its site, and mode of action remain elusive.
The presence of pressor materials, resembling renin (Sokabe el ai.,
1970; Chester jones el al., 1966) mayor may not have significance with
regard to blood pressure regulation in the eel. The broad conclusion
that the corpuscles of Stannius are involved with osmoregulation
(Leloup-Hatey, 1964a, b; Chester jones el al., 1965; Fontaine, 1964,
1967) is perfectly acceptable. It is a moot point whether this is a direct
action or an action via other endocrine glands such as the interrenal
(Olivereau, 1963; Hanke & Chester jones, 1966; Hanke, Bergerhoff &
Chan, 1967; Leloup-Hatey, 1970), the pituitary (Olivereau & Fontaine,
1965), the gonads (Fontaine, 1960; Lopez, 1969; Olivereau, 196Ib),
the ultimobranchial bodies (Lopez, DeVille & Bagot, 1968; Chan, 1972)
or indeed the thyroid (Leloup & Leloup-Hatey, 19(4). Such a gamut
of possibilities reveals our real ignorance of the nature and functions
of the corpuscles of Stannius secretion(s).
1.7.7 Thymus gland

Hagen (1936) described the thymus and Hassal corpuscles of eels.
This system is situated dorso-Iaterally in the gill chamber abutting
onto the opercular cover. As in other vertebrates, thymic histology
changes markedly with age. Thus in glass eels, when the thymus
begins to contain myoid ceUs and Hassal bodies, the gland is extensive,
but later shrinks. During metamorphosis the gland enlarges by a
factor of 7 between the leptocephalus Stage I and the glass eel (p. IS).
In eels 25 em long, the gland is diminutive and often only fatty remains
are distinguishable.
66 THE EEL
l.7.8 The kidney

By virtue of at least one secretion, renin, the kidney can be classified
as an endocrine gland. Such appears to be the case in the eel, in which
juxtaglomerular cells, generally considered to be the source of renin
in other vertebrates, are present (Sokabe & Ogawa, 1974; Sokabe,
Ogawa, Oguri & Nishimura, 1969), and renin-like materials are present
in the circulation (Nishimura & Ogawa, 1973). The role of renin in the
physiology of water and electrolyte metabolism, in particular its
actions upon the kidney, have been discussed (Sokabe, 1974). In some
ways rather paradoxically, renin activity is greater in eels adapted to
seawater, than in eels adapted to freshwater (Henderson, Jackson &
Hargreaves, 1975; Henderson, Jotisankasa, Mosley &, Oguri, 1976;
Nishimura, Sawyer & Nigrelli, 1976). It is possible that an endocrine
axis exists between the interrenal secretions and the renal renin-
angiotensin system (Borriraja, Henderson & Chester Jones, 1972).
1.8 Nervous system and sense organs

1.8. I The brain
Infonnation on the brain of eels is limited and is available only for
particular pans. Lissner (1923), in a general morphological description
(see Fig. 27), concisely outlines the structures as follows:
'The brain of the eel is as remarkable as its life. The very thick
olfactory nerves gradually swell to terminate in two extremely large,
egg-shaped olfactory bulbs situated on ridged corpora striata, which,
particularly in the ventrocaudal region, are enonnous. The epiphysis is
a short, cone-shaped structure, wider distally. Caudal to this are flat,
roundish, optic lobes which are smaller than the corpora striata. The
optic lobes have a ridge that runs laterally, from the upper regions
behind, diagonally to the lower regions in front. The part of the optic
lobe that lies caudally under this ridge is slightly flatter than the hulk of
the lobe. This, and the origin of the ridge can be explained by the fact
that the large nerve bundles of the trigeminofacialis group (V and VII ),
which run diagonally forwards, are pressed against the optic
tectum. The cerebellum has a curious shape. It is a smooth
rectangular structure with rounded edges that widen posteriorly,
and lies flat on top of the medulla, revealing a deep, longitudinal
furrow on the side thus creating the impression that the
cerebellum has been folded on itself as a result of insufficient space, and
that areas of contact have merged with one another. According to
Victor Franz the cerebellum of all teleosts has a ridge on the side which
indicates the line of embryological dorso-ventral fusion between the
granular layer and the ventral part. This fusion does not occur in
Anguilla, and the section of the corpus cerebellum under the ridge is
the border of a granular layer that stands out quite clearly. Also there are
no eminentiae granulares in this region because their homologue is to be
found in the 'granular border'. Franz describes the actual details very
clearly 'The brain of Anguilla is, however, so unusual in its entirety that
one cannot resist thinking that extensive secondary changes have taken
place. The valvula is rather small. In the region of the medulla oblongata,
under the cerebellum, there is a short fissure, the rhombencephalon,
which is surrounded by two ledge-like protrusions that rapidly become
narrower towards the rear. Posteriorly the medulla increases slightly in
size, and then continues into the spina dorsalis. Ventrally, long, thin
optic nerves, round in cross-section, and sturdy discrete ventral inferior
lobes occur on either side of the midline. The lateral lobes are separated
by a shallow oculomotor groove. The hypophysis is large and flat
with its front surface forming an obtuse angle, while its caudal surface
is rounded. '
The brain of Anguilla is very similar to that of the Moray eels
(Muraenidae) also described by Lissner (1923). Ito (1974) has compared
the fine structure of the torus semicircularis in eels with the same organ
in carp ( Cypn"nus carpio) , loach (Cobitidae) and rainbow trout ( Sairno
gairdneri) . The neuroanatomy and the relationships of the medulla
oblongata have been described by Ishida (1958). The parasympathetic
nervous system affects osmoregulation, although in the eel, in contrast to
other freshwater fishes, hormonal control dominates neural mechanisms
(Pequignot el ai., 1968). Richard (1955,1956) studied neural influences
on the different locomotory reactions of the eel (see also Bickel, 1897 ;
Gray, 1936, and B1ancheteau, 1972, 1973). Eels continue to show
movements despite such manipulation as skin removal, brain destruc-
tion or indeed sectioning of the posterior nervous system. The latter
contributes to neuromuscular activities. If the spinal cord is severed
1 em behind the pectoral fins, electrical stimulation produces rhythmi-
cal, snake-like movements. It appears that the brain has an inhibitory
influence on movement in intact eels. In fact, excitation appears to be
controlled by the tail seclion. In any event it is generally known that
eels may be inunobilized by wrapping them in a towel, particularly
around the tail region. In addition, optic lobe activity inhibits mobility,
as illustrated by the fact that feeding and migration take place mostly
at night.
68 THE EEL
The eyes are not the only regions of the eel sensitive to light. Blinded
animals will show flight reflexes when the head or tail is exposed to light,
and this reaction still occurs when afferent nerves to the skin have been
severed (Motte, 19<>3, 19(4).
The subcommissural organ and Reissner fibre complex in the eel
are thought to transmit optic signals to the motor nucleus of the spinal
cord, and have been described by Leatherland & Dodd (1968) who
showed that changes in the brightness of the background stimulated the
complex, but that other environmental factors had no such effect.
1.8.2 Eye
Literature on the most important part of the eel's eye, the retina, is
available from as far back as 1882 (Denissenko, 1882; Virchow, 1882).
Infonnation from more recent research on the retina is considered
below.
A peculiar feature of the eel's eye, in comparison with that of other
fishes, is its iris. In contrast to other species, the iris is relatively well
developed and, furthennore, is able to regulate the amount of light
entering the eye by appropriate degrees of expansion and contraction
(v. Studnitz, 1933). In bright light the pupil contracts, in dim light it
expands; in other words it simply reacts according to the amount of
illumination present - in contrast to humans where the diameter of the
pupil has a much more complicated form of adaptation. As has also been
shown in the isolated iris of '. he eel, the adjusnnent of the iris muscle
occurs as the result of the direct influence of light on the iris. The
greatest reaction is seen by using light that corresponds to the absorp-
tion maximum of rhodopsin (Seliger, 1963). As will be described below,
rhodopsin makes up most of the pigment in the rods of the eel's retina .
I t is therefore assumed that, in order to activate the iris muscle, the
same pigment composition is present in that muscle.
The ventral attaclunent of the lens is primitive in the eel as compared
with other teleost fishes (Stramke, 1972); the lens is not attached by
means of a muculus retractor lentis, but by a broad ligament. Due to
the absence of lens muscles, accommodation is not possible. Further-
more, the eel does not have a corpus chorioidae nor a musculus ciliaris.
The blood supply to the vitreous humour and the retina also differs from
those in other teleosts.
Biometric studies on the nucleus of the oculomotor nerve have shown
that the yellow eel probably makes relatively little use of its eyes
(Kirsche, 19(6). All other species of fish studied, including the burbot
BODY STRUC TURE AND FUNCT IONS
Fill. 30 Abow : Head of a female eel with enlarged eyes, caught

in M editerra nean
costal waters near Martigu e/Marseilles in Februar y, t969. 8tlfJW:
Normal
silver eel (Photog raph: Marschall).
(Lota Iota ) - which is well known as a 'non-v isual' anima l-

have a
larger nucleus than the eel. Just before its spawning migration the
eel
has, in comparison with other fish, relatively small eyes (Wund er, 1936).
This suggests a similarly reduce d visual capacity just before the eel's
marine period. With metamorphosis into a silver eel the diame ter of the
eye increases in size (Figs. 30 and 31; Matschenis, 1965) ; this growth
should be accompanied by an increase in the efficiency of the eye.
Simultaneously with this increase in eye size, the degree of maturi ty
of the gonads also increases, as has been experimentally verified with
hormonally induce d sexual maturi ty in male eels (Fig. 31 ). It thus
appears that the degree of maturi ty of migrating eels caugh t in the sea
can be determined. from the size of their eyes. However, one also finds
enlarged eyes in eels which have been prevented from migrat ing sea-
wards (Wund sch, 1953). In this case it is highly questionable wheth
er
an increased degree of sexual maturi ty is also attained. (Schnakenbec
k,
'953; see p. 43)·
At the same time as these external changes are taking place, morph o-
logical and physiological changes are occurring within the eye. Despit
e
the increased size of the eye in the silver eel, the retina still retains
its
70 THE EEL
/,
-
;j
rj " f" , T
0.. " 20 42 !
0
v" " x" " "-
"
f- -
I I I I I
1. 2. ~ 4. S. ~ 7.
!ltvtlopmrntol StQ<jf of U,. ttstis
Fig. 31 Eye size in male silver eels, which were treated with hormones (see p. 50). in
relation to the developmental stages of the testes. Vertical lines: standard
deviation ; figures : number of specimens (after Boetius & Boetius, 1967a).
disproportionately large size - i.e. it does not become smaller in relation

to the surface area of the body as is the case in all other species of fish
that have been studied (Teiciunann, 1954). In advanced stages of
maturity and with large increases in the diameter of the eye, the retina
shows a fourfold increase in its surface area. However, the number of
rods does not remain constant as was suggested by D'Ancona (1927,
1929), but increases considerably, i.e. the original rod density is actually
retained) as was shown in studies by Stramke (1972). On the basis of this
discovery, Stramke concluded that there was an increase in photo-
sensitivity in the silver eel eye at the expense of resolution.
It has also been noted (Wunder) 1936) that) in comparison with other
fishes which usually live in well-lit areas, the yellow eel stays in dimly
lit or dark places and possesses a retina that is very sensitive to half-light.
On a strip of yellow eel retina measuring 80 J.Ull, 143 rods were found.
BODY STRUCTURE AND FUNCTIONS 7'
Fish which are active in daylight - e.g. carp, pike, perch and many
others - have only between 10 and 50 rods on a similar sized strip;
only nocturnal fishes and fishes that live on the boltom, such as the
burbot ( Lota Iota ) and the barbel ( Barbus barbus) have more rods than
the eel. The cones in the eel's retina are extremely small ; this indicates
reduced capabilities where orientation by means of the visual system
in daylight is concerned. Negative phototaxis increases with develop-
ment from the yellow eel to the silver eel stage, as has been demonstrated
in studies with migrating eels and barriers of light (see p. 320).
An experiment to test the eel's colour vision produced negative
results - in contrast to similar experiments with other species of fish
(Betge et al., 1965).
The ability to see well in crepuscular conditions is not equally
developed at all ontogenetic stages. This became clear when infonnation
on the ntunber of rods in eels living in caves became available (Vilter,
1951). Such eels develop proportionately more rods in the lower side of
the retina than on the upper side. This is completely reversed in eels
living in surface waters. In addition, the number of rods in cave-
dwelling eels is about seven times that of surface-dwellers. However,
if one examines the ntunber of rods in the eye of a moderately pigmented
glass eel, their distribution is seen to be similar to that of rods in cave-
dwelling adults. Vilter (1951) is of the opinion that, in cave-dwelling
individuals·, the rods, which in these circtunstances would not be
influenced by light, would remain ntunerically constant during develop-
ment into the yellow eel stage. In surface-dwelling individuals, however,
the rods on the underside of the retina must have decreased in number
because this part of the retina lies in the pathway of light coming from
above, and light inhibits the development of rods. On the upper side of
the retina the rods develop in relatively large ntunbers because this
region is less well illtuninated.
Similar correlations with light can be observed between the centri-
fugal and centripetal migration of melanin in the retina, and the pig-
mentation stages in the young eel. Pigmentation and sensitivity of the
retina increase in parallel. At the same time the migrational activity
of the young eel is seen to decrease eVilter, 1943, 1946c). Vilter (1942,
1946c) has also shown that the hypophysis influences the distribution
of melanin in the retina; in glass eels centrifugal migration of the
pigment also takes place as the result of the influence of the pars inter-
media of the hypophysis. This change can be revoked by adaptation -
i.e. through the sympathetic nervous system. According to studies by
Thornton and Howe (1974) visual factors appear to influence the
72 THE EEL
hypophysis. These authors found structural changes in the secretory

cells of the pars intermedia in the hypophysis when the background
colour changed between black and white.
In addition [0 the morphological changes already mentioned,
during the second metamorphosis of the eel biochemical-physical
peculiarities and transformations have also been observed in the eye.
It is well known that animals develop retinal pigments that are best
suited for the absorption of the predominant wavelengths in their
particular environments. Fishes that live on the sea bed and in deep
water are mainly provided with rhodopsin which principally absorbs
the kind oflight that penetrates [0 greater depths; the wavelength of this
light is in the range 480-490 nm. In freshwater fishes, on the other
hand, porphyropsin predominates; this pigment absorbs light of over
500 nm (Wald, 1958). The presence of vitamin AI in deep sea fishes
and of vitamin A2 in freshwater fishes corresponds to the different
light-absorbing properties of the retinas in these two types of fishes.
Both vitamins are found in anadromous and catadromous species - e.g.
trout, charr, salmon, and of course the eel. However, in the eel, in
contrast to the salmonids, there is a predominance of vitamin AI and
thus of rhodopsin as well, which can be attributed to the eel's marine
origins.
Research into the wavelengths of rhodopsin in the European eel
( A. anguilla ) during different stages of development has shown
(Carlisle & Denton, 1959) that the colour of the pigment in the retina
changes during development from the yellow eel to the silver eel
stage. Yellow eels have a purplish coloured retina which is not unlike
that seen in other freshwater fishes. Silver eels have a golden shimmer to
the retina which is similar to that of deep sea fishes. These authors
call this pigment chrysopsin. Comparison with the retina of the conger
eel ( Conger conger ) reveals a great similarity between the two species.
The results of Carlisle and Denton's work on the European eel have
also been found to apply to the American eel (Beatty, 1975). Data on
A. rOSlrata show that yellow eels have a mixture of rhodopsin P50I
and porphyropsin P5232; silver eels at an early developmental stage
have a mixture of the two rhodopsins P4821 and P501 as well as a
small quantity of porphyropsin P523 2 • More advanced silver eels did
not appear to have the latter pigment, but showed a predominance of
rhodopsin P4821.
In conclusion, one can sum up the capabilities of the silver eel's
eye as follows: The enlarged eye and the correspondingly large pupil
make the organ more sensitive to light. The influence oflight is rendered
more effective because the photosensitive pigment of the retina is
better suited to deep sea conditions.
Pathological changes and anomalies in the eel's eye have been
described. Macrophthalrnus, for example, is known in the eel (Mercier &
Poisson, 1927). In another case itwas found that the left eye had develop-
ed in the lower jaw (Drooglever & van Leyden, 1917).
1.8.3 Olfaction
In contrast to its visual capacities, the eel has a well developed sense
of smell. While in most species of fish the two nostrils of each nasal
cavity lie close to one another, in the eel they are unusually far apart
(Fig. 32). This is probably due to the very elongate shape of the olfactory
rosette which, in other species, is virtually round or forms only a slightly
elongated oval. The rosette stretches almost from the anterior margin of
the upper jaw to the anterior border of the eye, where the posterior
nostril is situated. The anterior, inflow, nostril lies at the anterior
margin of the nasal cavity; it is tubular in shape and points downwards.
The olfactory rosette is formed from a great nwnber of olfactory
folds. The burbot ( Lota Iota ) , which also has a well developed sense of
smell, has about 30 olfactory folds, and the rainbow trout ( Sa/rno
gairdneri) , which has a mediocre sense of smell, has about 15 olfactory
folds, but the yellow eel possesses between 50 and 70 such folds, and,
in some cases, may have as many as 100 (Laibach, 1937; Teichmann,
1954; KaluZa, 1959). In the glass eel, and especially in the leptocephalus,
these laminar structures are considerably fewer in number (Fig. 33) and
are of a different shape. In other words, the olfactory rosette and the
nasal cavity in the early stages of the eel's development are similar to
those of the adults in other species of fish.
A further noticeable characteristic of the nasal cavity of the eel is
that it and particularly the posterior part of the rosette, are well provided
with cilia; these enable a strong flow of water to pass through the cavity,
and, in conjunction with the special shape of the latter, also increase
contact between the olfactory epitheliwn and the water (Teichmann,
1959). The histolOgy of the epithelium at various stages of development
has been described by Laibach (1937) who found increases in the size of
the various cell elements during ontogeny. Kaluza (1959) investigated
the blood-vascular system of the nasal region and of the olfactory
folds (see also Gorovaya, 1973).
Comparison with other species of fish shows just how much the eel
must rely on olfaction in sensory perception. In most species of fresh-
74 THE EEL
, '"
'"
...
o lmm
~
.. , .~;,
'."
"<~''''
Fig. 3z Snout of the eel. A. Head from above; B, Head from the side; the nasal
cavity exposed to show the olfactory rosette. CI, Lateral line canal; Co, Lateral
line canal opening; An; Anterior nostril; Po, Posterior nostril (after Kaluza,
1959).
water fish studied, the olfactory surface area, measured as a percentage

of the retinal area in the eye, is between 14 and 140% ; in the eel,
however, this proportion reaches 623 % (Teidunann, 1954). The eel is
thus the most extreme example of an 'olfactory animal' yet found among
fishes. A further indication of the eel's well developed sense of smell lies
in its relatively large olfactory bulb which, because of its position,
belongs to the 'salmonid type'; the eel also has large corpora striata
which playa part in the processing of olfactory stimuli (Fig. 27) (Harder,
1964 ; Gorovaya, 1969, 1973).
This dependence on a sense of smell accords well with the eel's
almost unbelievably refined ability to differentiate between various
Fig. 33 Olfactory cavity at different developmental stages. A, Leptocephalus; B, Glass

eel; C, Pigmented eel 8-12 em long; D, Yellow eel4S em long. P.N., Posterior
nostril ; A.N., Anterior nostril; R, Raphe; O.L., Olfactory lamellae; U.L.,
Unpaired lamella (after Laibach, 1937, VEB Gustav Fischer Verlag, Jena).
scents. In painstaking conditioning experiments it was shown that the

eel can perceive the scent of roses CP-phenylethyl alcohol) even when
the latter is diluted by I :2.857 x 10 18 pans (Teiciunann, 1957). Such
a degree of dilution corresponds to a solution of I ml of scent in a volume
of water 58 times that of Lake Constance (the Bodensee). In addition,
there are only about 1800 scent molecules in I ml of water. The minnow
(Phoxinus phoxinus) , which is also very sensitive to smell, needed a
THE EEL
concentration of IOJ(l times that strength in order to perceive rose scent.

The eel is thus almost as sensitive to smell as the dog, which is not
surpassed by any other animal. Teichmann (1959) has calculaled that
'at the threshold concentrations indicaled there is only a single scent
molecule in one of the two olfactory organs at anyone time. The specia-
lized structure of the olfactory organs appears suited to act as a quanti-
tative filter for the scent molecules in the waler that flows through the
nasal cavity. At threshold levels of concentration, central excitation of
the olfactory areas is only obtained when the arrival of two or more
stimuli in the nose are registered. The temporal interval between
individual stimuli can probably be as much as three seconds or even
slightly longer. However, if this interval is exceeded, then central
excitation does not occur. The stimulus is subliminal.'
Other scents are just as efi'eclive if they are detected in slightly
higher concentrations. Among these is the scent of prey animals, which
the eel locates by smell rather than by sight. One of the eel's preferred
prey is the tubiculous blood worm ( Tubifex) . Its smell was still perceiv-
ed by the eel even when five Tulnfex with a total weight of 25 mg were
finely ground and diluted in 6.67 x 10 12 ml of water. In view of the eel's
olfactory capabilities it is not surprising that hungry eels would, after
a short time, swim up to and bite at a hand that had previously been in
contact with meat (Schiemenz, from Wunder, 1936). If the other hand,
which had not come into contact with the meat, was held in the water,
it excited no interest.
The eel's ability to detect such low concentrations of scent is most
marked in late winter and high summer. In late autumn and in early
winter the concentration of scent must be raised by 10' before the eel
can perceive it (Teichmann, 1959).
The eel's sense of smell enables the animal to follow a scent trait.
'Orientation is achieved by trial and error (through the successive
perception of differences) and not by a topotactic adjustment (the
simultaneous perception of differences). In addition to orientation in the
horizontal plane, orientation in the vertical plane plays a very significant
role in the life of the eel' (Teichmann, 1959).
The eel's use of smell in orientation has frequently been exploited
by baiting fish traps. The effectiveness of wicker eel baskets is increased
several fold when bait is placed inside them. In this type of trap, the
eel is not led in by the channelling effect of a barrier, nor as a result of
protection from light, nor by the need for contact (i.e. negative photo-
taxis and thigmotaxis), but as a result of osmotaxis (attraction by smell)
(Tesch, 1970; see p. 276).
Attempts have also been made to develop artificial baits which would
appear attractive to the eel without its being specifically conditioned
to them (Schoeniger, 1951). However, none of the many substances
tried, most of which were similar to camphor and aniseed in smell, had
any decisive effect. Only Cibeton can be considered as being at aU
attractive. In practice, however, Cibeton is used only occasionally
because of its high price. There are also substances that serve to drive
the eel away - such substances are of practical importance, for example,
in the conservation of nets (Mohr, 1966, 1969). Carbolineum has a
repellent effect as long as it is not leached out of the netting; diesel oil
used as an emulsion, tobacco smoke, ammonia (over 0'0002 %), copper
acetate (over 0'001 %) and phenacyl chloride (over 0'0002 %) also serve
to repel eels. Net preservatives such as Racorit, Jolasteen and black
varnish have no effect once they have hardened.
Since many of the eel's prey animals, for example the smelt ( Osmerus
sp.) and various Crustacea, have been frequently observed to act as a
strong lure not only to yellow eels but also to elvers, it is obvious that
there are certain scents to which the eel responds, not as a result of
experience or conditioning, but through innate reaction. Proof of this
is seen in the fact that even glass eels show reactions that suggest a
specific olfactory sensitivity (Creutzberg, 1961; Miles 1968b). Glass
eels swimming around in seawater show increased activity when
freshwater is added to their container. If one adds tapwater, however,
no such response is seen. Natural surface water from a lake, for example,
proved, on the other hand, to have great effect. If, however, one passed
this water through a carbon filter, it lost its 'attractiveness'. Glass eels,
arriving stra'jght from the sea, cannot yet have had any experience of the
various scents contained in freshwater. The capacity to detect these
scents, despite their marked dilution by seawater, must be innate.
In a study of four rivers, each was found to have water that was
differentially attractive to glass eels. One river in particular had an
~xtremely good luring effect. Oddly enough the glass eels from this
river were particularly easy to activate with water from other rivers. A
:omparison of the chemical characteristics and 'attractiveness' of the
rivers revealed that the strength of attraction increased with the pH
v-alue of the water. However, merely altering the pH alone had no effect;
me pH value seems to be more of a secondary characteristic in this
:ontext since there are indications that a higher pH value coupled with a
~reater degree of fertility in the area from whence the water came, resul-
:ed in increased 'attractiveness'. This theory is supported by the fact
:hat river water from an area consisting of predominantly granitic
:ocks is the least attractive.
Water in which glass eels have been kept appears to be less attractive
THE EEL
than water which has not been in contact with glass eels; older eels, on
the other hand, did not seem to have this effect on the water, and indeed
appeared to increase its attraction for glass eels (Miles, 1969b). Nordeng
(197 I) has suggested that these results indicate that glass eels may even
be attracted by substances (pheromones) given off by older members of
their species.
Substances attractive to eels have been shown to contain dissolved
and undissolved (particulate) organic material, to be destroyed by
bacteria and thereby made ineffective, to be unaffected by heat, and
finally, that they are not volatile (Miles, 1969b). Scents attractive to
eels are in no way similar to those attractive to salmon. Homing salmon
orient by means of substances which are mostly volatile and which are
destroyed by heat (Hasler & Wisby, 1951; Fageriund ec ai. , 19(3).
Japanese studies on the nature of these attractants have produced
more conclusive results (Hashimoto ec al., 1968). Among the bio-
chemical components of a mussel that have been tested - e.g. amino
acids, quaternary ammonium bases and nucleotides - only amino
acids have proved to be attractive. However, they were only effective
when used in combination with one another. Not one of the 18 amino
acids studied could act as a lure by itself. Among the effective amino
acids were taurine, aspargine acid, threonine, serine, glutamic acid,
glycine and alamine.
The eel's extremely refined sense of smell has naturally raised
widespread interest and has led to the assumption that the eel uses its
olfactory organs not only to locate food but also to locate different places.
In land animals this is a well known phenomenon - for example in the
dog, which is considered equal to the eel in its olfactory capabilities
(Teichmann, 1959). Other species of fish, such as the salmon, provide
good. evidence that aquatic animals, too, can follow scents. Indeed, the
orientation of glass eels towards freshwater is thought to be a form of
orientation by smell. This is, however, only a general orientation to
scents from any natural source of inland water flowing into the sea. It is
highly unlikely that individual inland waters have any specific effect
(Miles, 1968b); in other words glass eels are attracted by the smell of
any river. I t could, however, still be argued that eels migrating to their
Atlantic spawning grounds use olfactory cues for orientation, and that
yellow eels transported some distance away find the way back to their
old home by using a sense of smell (Mann, 1965; Tesch, IC)67a j Deelder
& Tesch, 1970; Tesch, 1970). When discussing the use of smell for
orientation by salmon in rivers and also by the eel in running water - if,
that is, the processes involved are the same for the two species - it
should be noted that a 'scent trail' is followed and that the possible
direction of this trail is narrowed down by the sides of the river, thus
making it easier to follow. A difference in scent only exists for a very
short while at the junction between two rivers where also the scent of the
contluent rivers is sharply contrasted. Such dearly contrasted trails
are not available in the sea. A displaced eel that is homing many kilo-
metres to some point in Heligoland, for example (Tesch, 1967a, 1968b),
would be confused if it came across widespread 'scent waves' from
Heligoland, presuming such scent waves were present at this point.
Keeping this image in mind, and also if one remembers the difficulty
eels have in following a scent trail a few centimetres long in an aquarium,
then the idea of their differentiating between huge, diffuse waves of
scent becomes improbable. Besides, hydrographic factors make it
unlikely that scent from Heligoland would reach Cuxhaven, for example,
which is 65 kIn away. Locating the Gulf Stream would be equally
difficult using a trial and error approach.
Experiments in which eels with blocked nasal cavities were transpor-
ted to the North Sea coastal areas, showed that such eels display the
same homing trends as control eels (Tesch, 1970). Thus location by
olfaction is unlikely, at least as far as the detection of distant goals in the
sea is concerned. Evidence that olfaction may playa role in orientation
in the sea and in large inland lakes is, as yet, not available for other
5pecies of fish either (Royce et at., 1968 j McCleave, 1967 j lahn, 1969).
1.8.4 Hearing and the lateral line sense organs

The Anguillidae do not belong to the Ostariophysi - i.e. eels do not
have a Weberian apparatus connecting the swimbladder with the inner
~arj therefore direct transmission of vibrations from the swimbladder
:0 the auditory organs cannot occur. As is generally agreed, such non-
)stariophysan fishes have less acute auditory perception. This is
:ertainly the case in the eel (see below).
Attempts have been made to determine the centres of sound percep-
ion in Anguilla anguilla (Diesselhorsr, 1938). Destruction of the two
luditory labyrinths resulted in a reduction of acuity from about 600 Hz
:0 400 Hz. Above 600 Hz, according to these studies, the eel's perceptual
:apacity shows a sharp drop anyway. The same apparent loss of sensiti-
lity was produced not only by the removal of the pars inferior of the
nner ear but also, in contrast to other species, by removal of the pars
uperior. Thus in the eel, the site of auditory perception in the frequency
'ange already mentioned cannot be located more precisely and it
80 THE EEL
appears that the eel is able to detect sounds below 400 Hz through
some other sense organ.
Diesselhorst (1938) suggests that the tactile sense of the skin is the
receptor mechanism for sounds below 400 Hz. According to studies by
Schrieven (1935), the lateral line sense organs, which are also able to
detect vibrations, are not activated by tones above 150 Hz.
More refined methods of study have shown (Stepanek, 1968) that
the eel's capacity for detecting higher sound frequencies is not quite as
limited as was suggested above. If one makes a distinction between a
weak response and an abnormal reaction, then the 600 Hz frequency
mentioned above as the upper limit of hearing should be taken to be an
abnormal reaction for young eels 8 em in length. At 800 Hz weaker
responses can still be seen in such animals. Older eels (30 em in length)
show a strong reaction at 2500 Hz and will even show a weak response to
9000 Hz. In addition, eels display an escape reaction to sounds of only
50 Hz, while at sounds of 400 Hz they merely become restive. Whether
this is evidence for the existence of different receptor organs for different
frequency ranges is a theory that needs closer investigation.
A comparison with other species of fish shows that as far as auditory
acuity is concerned, the eel is relatively insensitive. Out of a total of
twelve species studied, four - the trout, the milU1ow, the grayling
and the pike - show much greater sensitivity, particularly in the higher
frequency ranges (Stepanek, 1968). In the lower ranges, too, the eel is
less sensitive; its lower limit of perception is between 50 and 70 phon,
while the minnow ( Phoxinus phoxinus ) for example, and the mormyrids
can perceive sounds as low as 20 to 30 phon. Similar findings are
recorded for interval discrimination; this stands at an octave in the eel
and is therefore rather crude.
Alnaes (I973a, b,c) has studied the lateral line organ and itsconnec-
tions with the central nervous system. These studies were an attempt
to identify the complicated system of sensory cells and their afferent
and efferent nerves by using an electrophysiological and histological
approach.
2
Eel Species, their Developmental
Stages and their Distribution
In some respects it is very difficult to distinguish between different

species of eels. This is particularly true in the case of larval forms;
these, as is well known, show continuous cbanges in body form and
pigmentation, so that the characteristics of individuals at different
ontogenetic stages are not comparable. Only a very few characters can
be seen in all stages before and after metamorphosis. Even such
basic and important features as coloration and markings can only be
used after the metamorphosis of glass eels into yellow eels, and then
only until the final metamorphosis into the silver eel. The so-called
marbled patterning of certain species only becomes evident when the
eels are over 20 em long; in the silver eel stage it disappears again
(Ege, 1939; Jubb, 1961). Three features do remain the same through
:III stages from larva to silver eel, namely the number of vertebrae, the
(lumber of myomeres (the latter count is always one higher than the
former), and the relationship between the lengths of the dorsal and
mal fins (Figs. 34 and 35). All other characteristics can only be used
:omparatively if the individuals are of almost the same age and are at
the same stage of development.
In addition to the specific characteristics listed in Tables 5 and 7,
md in Figs. 34 and 35 (which, it should be noted, are used mainly for
iistinguishing between individuals at more advanced developmental
>tages), at the larval stage one can also use the position of the
nelanophores and the vertical blood vessels of the gut (Fig. 35), the
81
82 THE EEL
Fig. 34 Body measurements in the eel. a~d, Total length; a-b, Predorsal length;
a-c, Preanal length; a-e, Head length (after Ege, 1939; reproduced from
Dana Report, No. 16).
,
I rllyom"es~
Fig. 3S Body measurements in the eel larva (leptocephalus) showing standard

length, the position of the pre- and postanal myomeres, and the position of the
three vertical blood vessels aCthe gut in relation to the myomeres ( 15,38 and
43). The figured specimen is of a short-finned eel, which has only a few
preanal myomeres (3) from the beginning of the larval dorsal fin fold to the
vertical through the origin of the anal fin fold (Jespersen, 1942; reproduced
from Dana Report, No. 22 ) .
dentition (Fig. I I) and the shape of the anal fin. However, these features
can only be used if a complete record is available of metamorphosis in
the various species. This is rarely the case with the numerous and often
sympatric species of the Indo-Pacific region (Castle, 1963). The identi-
fication of these species thus causes quite considerable difficulties. In
such instances the number of myomeres, fin length and the geographical
distribution are of no help in deciding to which of the two or three
possible species a specimen may belong.
The genus Anguilla Shaw (Gen. Zool., 4, IS; 1803) is classified as
follows: Class, Teleostomi j Subclass, Actinopterygii; Order,
Anguilliformes; Suborder, Anguilloidei; Family, Anguillidae. The
family Anguillidae has the following features in conunon with most
other members of the Anguilliformes: the swimbladder is connected
to the gut by a duct. Pelvic fins absent. Scales cycloid. No mesocoracoid
or post-temporal bones. The premaxillae are fused with the ethmoid.
Paired orbitosphenoid present. Teeth on the maxillae. No basisphenoid
EEL SPECIES
or sympletic. Parapophyses and most neural arches fused with the

vertebrae. No myodome. Gill openings narrow. The dorsal and anal
fins very long and confluent with the greatly reduced caudal fin. Bone
cells present in the bones.
A total of 28 fossil and living families make up the Anguilliformes
(Berg, 1958; for funher information on the skeleton, see p. I and
Table I). More recent sources indicate a total of 26 families (Greenwood
et al., 1<)66).
The following are diagnostic features of the family Anguillidae and
thus of the genus Anguilla as well: the pectoral fin is supported by 7-9
radialia (up to I I in young); the mouth is terminal, the lower jaw slightly
longer than the upper; the teeth are small and arranged in several
rows on the jaws and palate. The lateral line is well developed. The gill
openings are not confluent. A 'tongue' is present and the lips are thick.
The frontal bones are paired. The palatopterygoid arch is well developed
(Berg, 1958).
The characteristics of the individual species are described in Tables
5 and 7, in Fig. 44 and in the following sections; for specific synonymies
see Ege (1939).
2. I The European and American eels

A. anguilla and A. rOSlrata
2.1.1 Spawning grounds and larval forms

A fish named Leptocephalus brevirostn's, on account of its small, narrow
head (Fig. 36), has been known since 1856 (Kaup), when it was first
caught in the Straits of Messina in the Mediterranean Sea. The body
was narrow with a high back, and resembled a willow leaf in shape.
The correct identity of this creature was not discovered until 40 years
later (Grassi & Calandruccio, 1897a & b; Grassi, 1896). These two
Italians realized that the fish called Leptocephalus was really a larval
stage of the river eel. Further investigations led them to believe that the
eel's spawning grounds would be found in the depths of
the Mediterranean. These authors also suggested that the leptocephali
lived on the sea bed and were drawn to the surface by the eddy effect
of the Straits of Messina; they were not, however, able to prove this
hypothesis, particularly since they did not catch any smaller eels at
earlier larval stages. The smallest larvae caught were already 6-8 em
long. In addition, they were unable to find adult eels which were ready
to spawn - none has been found to this day, even in other areas.
Table 5 Characteristics of the different species of Anguilla Shaw (after Ege, 1939) Principal diagnostic features are italicized Z'
Distance
No, of verttbrat betwten the
Length of vomerine
verticals
Species lOoth band as a % of
jrombegin-
tht lenglhof liu maxil-
ni'Wof
lary loolh band
Total Precautial dorsal fin
to emus,
expressed .9 a.
in % oj
aa
IOtal
length
A, ce/tbesen.sis 103'4 39'6 9·0 Vomerine tooth band 86-87

A , mqastlmUl 112'J 41'7 11'1 parrower than the 80-81
A. interioNS 105'4 4°'7 13'0 } band of teeth on
A . ancestralis 10J'J 38'8 10-11 Skin the maxilla
A . nebulosa nebulosa 109'1 4°'7 11'7 marbled Tooth rows on the 70
A . nebulosa labiala 111'3 40'9 11'9 73- 74
} maxilla even
.......'"
A, marmorala 105 '6 41 ' 1 16'J } Dorsal 70 ...
A, reinhardti 107'8 42 'S 10,8 fin Uneven 74- 75
A. borneen.sis lOS'S 40'6 II 'S long
A . japonica 115'8 43'6 9·' Vomerine looth band 8, 8,
A . dieffenbach; 1I2'7 443 11 '1 wider than the band 8,
A . mossaml,ica 102'9 40'5 t4'6 of leeth on the 73-74
A . rostrala 107'2 4 2 '8 9·' maxilla Vomerine tooth band 69-7 0
A. anguilla 114'7 4S'2 10'2 Skin uni- narrows before 80 79
A. bicolor pacifica 107 ' 1 43 ' 1 formly } the middle of 82-83
A . bicolor bicolor 109'5 43'3 0,8 Dorsal coloured the plate 85-86
A. obscura 104'0 41 'S? 3'6
A. australis australis 112'6 46 '2
O'}
1'2
fin
short
7'
70
A . australis schmidti 111 '7 46 '1 , ·6 Vomerine [Ooth band 70
} narrows afler the
middle of the tooth
plate
EEL SPECIES 85
Fig. 36 Development of the eel from the larva (leptocephalus) to the glass eel. (J) Fully
grown larva 75 mm long (Stage I) ; (2-6) Metamorphosis of the larvae (re-
sembling a willow leaf in shape) into one with a typical eel shape; (7) Glass eel
at the earliest developmental stage; (8) Eel at the beginning of pigmentation,
still very similar to the glass eel (after Schmidt, 19Q9a).
86 THE EEL
The situation changed suddenly when, in 1904, the Danish research

ship 'Thor' caught a leptocephalus larvae in the Atlantic, west of the
Faroe Islands, the first larva to he caught outside the Mediterranean
(Sclunidt, 1912). A further report in the same year from an Irish ship
that had caught leptocephali outside the Mediterranean led the Danes
to begin systematic srudies, in 1905, into the question of the distribution
of the larvae. Using a relatively new fishing technique, the Petersen
young-fish trawl, Schmidt (1906) succeeded in June 1905 in catching
in the upper water layers hundreds of fully developed leptocephalus
larvae west of the 1000 m depth line. In the following year Schmidt
tried to catch still younger fishes but had no success at all. In fact,
later in the year, in August and September, more advanced larval
stages were caught even closer to the coastline; these larvae were
approaching the glass eel stage.
Later, the youngest larvae were to be found furthest from the coast
and over greater depths, i.e. beyond the 4000 m line. These and subse-
quent investigations (Schmidt, 1909a) indicated that the spawning
grounds must lie far out in the Atlantic, north of the equator, and that
the earliest larvae are not to be found on the sea bed but near the surface
of the water.
In 1912, Schmidt put forward certain views and evidence in a
convincing argwnent against the idea that the spawning grounds of
A. anguilla were in the Mediterranean, viz. :
( I) Vertebral counts of 1700 Atlantic eels (from Iceland and Madeira)
gave an average count of 114'736 vertebrae. One thousand eels
from the Mediterranean had an average of 114"731 vertebrae - i.e.
this traditional taxonomic character corresponded exactly in the two
groups of eels.
(2) Between 1908 and 1910, only 5% of the eels caught in the
Mediterranean east of longitude 3 0 W were less than 70 nun long,
i.e. were leptocephalus larvae. West of this point, however, 60% of
all larvae caught were at this early stage of development. In other
words, the further east one went from Gibraltar, the more mature
were the larvae. Furthennore, the number of larvae in the region
around Gibraltar was significantly greater than the nwnber found
further east. In the eastern Mediterranean basin, i.e. east of Italy,
no larvae were caught at all.
(3) Near Gibraltar the larvae were most evident in winter; in the
mid-Mediterranean near Messina, on the other hand, they were
predominant in spring and summer. This temporal difference
corresponds to the drifting time required for the larvae to be carried
EEL SPECIES
from Gibraltar to Messina. The larvae drift along with a surface

current which flows eastwards from Gibraltar.
(4) In Italy, as in western Europe, glass eel fisheries are only found
along the west coast; further east there is none (which does not
mean, of course, that glass eels do not occur, for example, on the
coasts of the Adriatic) (see Heldt & Heldt, 1929b).
(5) In the Straits of Messina there is a fishery that uses cordons of
lights; the migrating eels are caught in nets operated by hand.
These eels, whose direction of passage is easy to detect, travel north-
wards, i.e. towards the Straits of Gibraltar. Near Messina silver
eels are also caught later in winter and spring than in the eastern part
of the Mediterranean from whence they would have to start migrating
earlier in order to pass Messina by the tum of the year.
(6) Eels only seldom occur in the Black Sea. If eels had spawning
grounds in the eastern Mediterranean, one would expect them to be
more abundant in the Black Sea.
Arguments in favour of spawning grounds within the Mediterranean
are, in the main, attributable to Grassi and Calandruccio (e.g. 1897)
(see Schmidt, 1912). These authors maintained that particularly large
numbers of eels with enlarged eyes occur in the Straits of Messina.
Such individuals, whose large eyes were thought to indicate an advanced
stage of sexual maturity, are found only rarely in northern Europe
(see pp. 43, 69). Schmidt (1912) agreed that eels with large eyes are
more frequent near Messina but more recent, unpublished observa-
tions by the author on eels sent from the Mediterranean to markets in
Hamburg show that the frequent occurrence of eels with enlarged eyes
is not restricted to the Straits of Messina. In a shipment of eels from
the Rhone Delta (near the Mediterranean coast) in February/March
of I1}68, the majority of individuals had, to a greater or lesser extent,
enlarged eyes (Fig. 30). However, the state of ovarian development in
these fishes hardly differed from that of silver eels from north-western
Europe; the same appears to be the case for eels from Messina (Schmidt,
1912). Although enlarged eyes undoubtedly are an indication of fairly
advanced maturity, as experimental investigations have shown (see
p. 70), they are only a secondary characteristic which, preswnably, can
develop faster at higher temperatures and a corresponding increase in
hormone (thyreotropic) production (Fontaine, II}6I ), than can the
gonads (which are dependent on gonadotrophic and oestrogenic
hormones).
Further, and quite recent evidence apparently favouring an isolated
spawning ground in the Mediterranean might be derived indirectly
88 THE EEL
from differences in the composition of serum albumin of Atlantic and

Mediterranean eels (Drilhon et al., 1967; Drilhon & Fine, 1968;
Drilhon & Fine, 19(9). These authors found differences in the serum
transferin composition of eels from the Atlantic coast of France and
eels from the Mediterranean. It must be noted here that differences
were also found between areas within the Mediterranean, e.g. between
eels from the French, Greek and Turkish coasts. Although this evidence
suggests that there may even be different spawning populations re-
presented within the Mediterranean, it does not prove that there are
different genetic groups (see also Pantelouris et al., 1970; 1971), nor
that the spawning grounds are in the Mediterranean.
The fact that leptocephalus larvae measuring 30 mm have been
caught in the Mediterranean could strengthen the hypothesis that
there are one or more spawning grounds in the Mediterranean. Also
it has been noted that hardly any silver eels have been caught at the
point where the Mediterranean opens into the Straits of Gibraltar
(Tucker, 1959). In fact this is hardly surprising as only a very small
percentage of migrating silver eels have been caught, for example, on
the German coast of the Baltic, because, until recently, the traps
scarcely extended more than 10 m down into the water (Martinkowitz,
1961 ). The Straits of Gibraltar are, moreover, over 200 m deep, and the
traps used there are unsuitable for catching eels at this depth 0 ones,
1959). If there is no migration away from the area aroWld Gibraltar,
then, as Ekman (1932) suggests, the Mediterranean should be consi-
dered a kind of huge trap.
Further extensive studies in the Atlantic by Schmidt (1923; 1925a),
located an area where larvae measuring as little as 5 nun were to be found.
Since the larvae are probably about this length on hatching, the spawn-
ing groWld must have been very close by . The centre of the area where
larvae measured 10 nun or less was calculated to be at 26 0 N and 56 0 W
(Fig. 37).
During this same period the larvae of the American eel ( A. rostrata )
were also recorded in most of the samples taken by a number of ships
operating in the western Atlantic . In contrast to the practically non-
existent differences in vertebral numbers between Atlantic and
Mediterranean eel populations, the American eel differs considerably
from A. anguilla. Anguilla rostrala has a mean myomere count of
108"170espersen, 1942 ; according to Vladykov and March, 1975, this
figure is 105 "13) - i.e. 7"5 less than A. anguilla, which has 115"58
myomeres (Jespersen, 1942 ; Vladykov and March, 1975, give a mean of
EEL SPECIES
'0'
Fig. 37 Limits of occurrence of Anguilla larvae of various sizes ( 10, IS, 25 and 45 mm).
Continuous lines indicate the areas of occurrence of [he European eel (A .
anguilla), discontinuous lines of the American eel (A. roscrata). For the
American eel the outer boundary line (01) indicates the absolute limits of
distribution of the larvae. Five to seven millimetre long larvae, i.e. relatively
recently hatched European eel larvae, were caught within the hatched area,
and thus the spawning grounds are thought 10 lie within this region (after
T!ning, 1938).
III '76 myomeres). Only 1 '5% of individuals fall within the intersection
of the two distribution curves (Fig. 38).
A total of only about 2300 American eel larvae (called Leptocephalus
grassi by Eigerunann and Kennedy, 1902) were taken in the various
samples. In contrast, 12000 leptocephali of A. anguilla were recorded.
This difference may have been partly (or even totally) due to the fact
that Schmidt's investigations were concentrated mainly in the spawning
area used by A. anguilla, and partly because the larvae of A. rostraca
prefer a greater depth. It is also possible that the collections were made
at a time of the year unfavourable for A. roslrala (Vladykov, 1964).
On the basis of results from these samples) Schmidt (1923) came to
THE EEL
Af>lJ<lilla Dllgvilla
",,2715
"
Fig. 38 Frequency distribution of vertebrae in A . anguilla and A. rostrata (data from

Schmidt, 1913; and E@e, 1939).
the conclusion that the spawning grounds of the American eel are
probably to be found further west and slightly more to the south,
about 15 0, according to his figure (see Fig. 37). The relative proportions
of A. rOSlrata and A. anguilla in the catches recorded by Sclunidt (i.e.
in the ratio of almost I :6) were mostly brought about by the (act that
too few samples had been taken in the most westerly areas. More
recent hauls made in the Straits of Florida and to the north of the
Bahamas produced 270 A. roslrata and 15 A. anguilla, thereby shifting
the proportions completely in favour of the American eel (Smith, 1968).
Vladykov and March (1975) investigated a larger region that also lay
EEL SPECIES 91
mainly in the A. rosrrala area of distribution j their collection yielded
790 A. rostrata and 433 A. anguilla, i.e. it contained a relatively large
proportion of European eel larvae.
As additional support for his observations, Schmidt indicated that
commercial eel catches during the period, recorded as 2000 tons a
year for A. roslrala, were only one-fifth the size of those for A. anguilla
( 10000 tons), and thus corresponded to the relative proportions seen
at the larval stage (Schmidt, 1923; see also Table 25 and Vladykov,
1966).
However, more recent results do not basically alter the picture
created by Schmidt's ideas and research on A. rOSlrara. As Smith
(1968) has pointed out, during July 1966 records from traps in the
southern Caribbean sea between Panama and Trinidad did not include
a single specimen of an Anguilla species among the many larval anguilli-
forms which were caught. However, in the well-known area of distri-
bution just to the north, large numbers of Anguilla larvae were recorded
during this period. Further west, in or around the Gulf of Mexico,
there is no indication that eels occur in large numbers (Eldred, 1971).
According to data from Schmidt (1925a), only 2% of eels sold on the
American market come from the coastal areas, i.e. from around the
Gulf of Mexico; nowadays this percentage is even smaller (Vladykov,
1966). All other eels are caught along the east coast of America. It
seems that the eels are carried along by the strong, dominant current
which orginates in the northern equatorial stream, and flows through
the Straits of Florida into the Gulf Stream (Fig. 39). There is, in other
words, no reason at all to believe that the spawning ground of the
American eel is much further south than Schmidt first suggested. The
question still remains unsolved as to how A. rostrala can manage, in
spite of the northern current, to get to its southern-most point of
distribution near Trinidad and Guayana (Vladykov, 1964).
In contrast, it is easy to explain how, in general, the two species of
eel reach their northern areas of distribution. The chief means of
transport is the Gulf Stream; the Florida Stream and the North
Atlantic Stream also play imponant roles in the distribution of American
and European eels respectively (Fig. 39). Because the North Atlantic
Stream comes into contact with Iceland but flows straight past Green-
land (according to Le Danois, 1938), Iceland is populated exclusively
by the European eel. Greenland, on the other hand, has populations of
American eels, but it remains a mystery how, in reaching Greenland,
larvae or glass eels of A. rostrata pass through the Labrador Stream,
which flows against their direction of travel (Smith, 1968).
1•
i
1
,.
1
1
1
EEL SPECIES 93
80 80
60
• Al)gvtl'a al)gvilla
II Angvlfla rostrata
I 'roo
Scnm,dt
o Al)gvd'a rostrata from 20

Srn iln (19681
III V VII IX XI III V VII IX XI I III V VII IX XI I II I
Fig. 40 Length increments in the twO species of Atlantic eel. Continuous curve
A . allguilla; discontinuous curve A. rostrata (data from Schmidt, 1923; and
Smith, 1968).
The means whereby such clear partitioning of the two species

between America and Europe is achieved, is, to some extent, understood.
The American eel develops into a glass eel within one or two years,
but the European eel requires, on average, three years to reach this
stage. Various pieces of evidence confirm this difference in develop-
mental times. In both species hatching takes place in spring. While, in
winter, not a single A. rOSlrata larvae was caught (Smith, 1968) and
only the older larvae of A. anguilla were in evidence (Schmidt, 1923),
from April onwards 10 nun larvae of both species were caught in large
quantities. These larvae. which could already be specifically differentia-
ted by the number of muscle segments, showed a monthly increase in
average length (Fig. 40). Here the American eel has a definite headstart
which is probably due to the fact that this species spawns four weeks
earlier. During the summer this advantage is further consolidated. The
American eel is no longer in evidence in late summer and autumn.
By this time the larva has reached its greatest length (Le. Stage I, see
Table 2). After this, eels are first caught in the coastal areas and in river
estuaries; at this stage they have completed metamorphosis and,
apart from pigmenration, are fully developed. As Figs. 36, 40 and 57
show, they must have gone through a certain decrease in length.
The larvae of the European eel continue to be caught well into
autumn. A few are even found the following spring in the central
regions of the spawning areas. The great majority, however, are pro-
ducts of the new generation. For example, at two stations west of 50 0 W,
484 new larvae (age group 0 ) but only 3 one-year old larvae (age group I
94 THE EEL
"
AgtQroupO(AGOI
25 25
20 ZO
f
•:1 IS
"
10
5 5
" llngth(mml "

Fig. 41 Length frequency distribution of European eel larvae caught in a trap west
or
of 50c W in June, 1920 . Only the upper and lower length groups age groups
o and I are represented here, in order to provide a clearer picture of the size
intervals (data from Schmidt, 1923).
or 'AG 1') were caught. As one goes further north-east, the proportion
of AG I eels increases. Both generations, however, can easily be dis-
tinguished from one another, as Fig. 41 shows. Even further north-east,
eels from AG 0 disappear altogether and, in addition to larvae of AG I
which measured 55 mm in June, other, even larger, young are in
evidence not far from the Continental Slope. These latter are already
almost 80 mm long and must therefore be regarded as fully grown
leptocephali at Stage] of development (see Table 2). This, then, is the
third generation (AG II) of eels which is travelling towards Europe and
which will finally, in the late summer, change into glass eels, reaching
the coasts of Europe in winter or spring.
VI.dykov and March (1975) confirm Schmidt's (1923, 1925')
data on the three-year duration of the larval period in A. anguilla. They
are of the opinion, however, that their data indicate that A. rostrata
spends more than one year in the sea. Of course, the time spent in the
sea, whether one or three years, must not be regarded as invariable.
EEL SPECIES 95
On the basis of larval material collected recently in the eastern Atlantic
(Tesch, unpubl.), the author suggests that sometimes at least two
generations of eels can be found together, but in differing nwnbers, on
the European Continental Slope. The same may be the case for A.
rostraca, with the result that the average age at metamorphosis is
difficult to determine.
The fact that the American eel needs only one or two years to reach
the coast, and the European eel needs three, is an important factor in
the geographic distribution of the two species. Not only as a consequence
of their western spawning grounds, but also as a result of their shorter
developmental time, the larvae of A. rostrata are able to reach the coasts
of America. If they too needed a three-year period for development,
they might also get caught up in the North Atlantic Drift and arrive off
European coasts; alternatively, if the larval period for the European
eel was only one year, the larvae might make for the coasts after a
single year and thus perhaps reach North America. There seems no
nthpT f>xnh'ln~tinn ~!'I. tn whv Rf'TmlH"i~ i!'l. nnmllMf'fi hv A rmTrfltn
THE EEL
Fig. 42 Pre-leptocephalus larva of A . anguilla, 6 mm long; CD, spinal chord;

H, oil droplets in yolk; I, intestine; M, notochord; My, myomeres; N,
embryonic fin fold ; P, pectoral fin (after Schmidt, 1923 ; and Bertin, 1956).
later years (Schmidt, 1929; Tming, 1938; Bigelow & Schroeder, 1953;
Fontaine et al., 1964), principally because a compact oil globule,
which is present in the earliest larval stages (Fig. 42), was absent in
the ova described by Fish. It is equally unlikely that eggs found at
40 0 N and 69 0 W, in water between 0 and 54 m deep (Evseenko, 1974),
were eel eggs, because correspondingly small Anguilla larvae have
never been found in this region.
The remains of potential parental eels have so far only been found
in the stomachs of predatory animals, e.g. an eel which was found in a
spermwhale near the Azores (Vaillant, 1898) had probably just started
on its journey to the Sargasso Sea. Since the whale cannot descend to
very great depths (up to 700 m), it seems unlikely that spawning eels
migrate at great depths. Eels have been found in the stomachs of sword
fishes in the Straits of Messina, i.e. still some distance from the Sargasso
Sea (Grassi & Calandruccio, 1897a & b). This record would seem to
indicate that migrating, pre-spawning eels move in the pelagic zone.
Two other discoveries were made in predatory fish which had been
caught in trawl nets at a depth of 730 m (Reinsch, 1968). The two
fishes involved, Mora moro and Aphanopus carbo are both bathypelagic
species and had each eaten a female eel of 44 ern in length. Thus it is
probable that the eels were not taken on the sea bed, although at a depth
not far above the bottom. The predatory fish were caught about 100
miles north-west of the Hebrides, i.e. they were also quite far from the
Sargasso Sea. The oocytes in the ovaries of the eels were 0'15
and o· 12 mm in diameter, and were thus no further advanced develop-
mentally than those of normal silver eels. The diameter of the eyes
suppons this conclusion.
Greater numbers of silver eels have been found at lesser depths
(Table 23). Most of these eels were caught in trawl nets used in the
herring fisheries on the Dogger Bank. In areas which are fished in-
tensively it is quite common to come across migrating eels, though,
with the wide-meshed trawl nets used, it is unlikely that many
EEL SPECIES 97
individuals are brought to the surface with the rest of the catch. Almost
all these records are from October and December - that is towards the
end of the migratory season.
The depth of the spawning grounds can only be determined with
the help of information on the occurrence of the youngest larval stage.
To quote Schmidt (1923) on this matter: 'The average length in
June was about 25 nun, and the great majority of larvae of this size
were found to occur near the surface - from a depth of about 50 m to
the surface itself. The younger larvae (7-15 nun long) were taken
somewhat deeper, at depths ranging from about 200 to about 75 m.'
Elsewhere, Schmidt (1929) adds the following: 'Of the European eel,
we have comparatively many prelarvae and quite young larvae, in
which the oil globule of the yolk sac has not yet been resorbed. These
are found somewhat deeper down than the rather older larvae, but
still in what may relatively be called surface strata, about 200-500
metres down.'
On the basis of experiments in which the sexual maturity of male
eels was prolonged, Boetius and Boetius (1967a) conclude that spawning
must take place at a depth of about 150 m. It is highly unlikely that eels
spawn on or near the sea bed, i.e. at a depth of about 6000 m. Here, the
temperature, which may fall below 5 °C, is almost certainly too low.
Relatively high temperatures have been necessary to obtain advanced
sexual maturity in eels and also to effect spawning (Fontaine, 1961;
Fontaine ec al., 1964; Boetius & Boetius, 1967b). In experiments with
female eels the temperatures were initially set at 24-25 °C, but during
ovulation were decreased to 20 °C.
The Sargasso Sea represents a sort of basin in which relatively high
temperatures extend to quite considerable depths (Fig. 43). The
isotherms of 17 °C best define the spawning ground.
The temperature increases as one approaches the Sargasso Sea and
this may help orientate the migrating eels on their journey to the
spawning grounds (Ekman, 1932). If this is the case, then the change in
temperature is bound to be only one of the orientation cues used by the
eel, because we know today that the eel, like migratory birds, probably
has a sense of direction. American silver eels swim south (Miles, 1968a)
and European eels swim west to north-west to reach the spawning
grounds (Maar, 1947; Edelstam, 1965; Tesch, 1972). Orientation
simply by means of following a gradient of increasing temperature is
hardly feasible in the case of eels from the Mediterranean, because the
temperature range there is similar to that in the Sargasso Sea.
Salinity was also considered to be an important environmental
THE EEL
Fig. 43 Temperatures in the spawning ground of the two Atlantic eel species at a
depth of 400 m (annual isotherms). Horizontal hatching = spawning ground
of the American eel ; vertical hatching = spawning ground of the European
eel (after Schmidt, from Ekman, 1932).
factor in the detennination of optimal spawning conditions (Bertin,

1956). Associated with the temperature change is a water mass, in mid-
Atlantic, which is of tropical origin and which has an elevated salinity of
35 parts per thousand; this floats on another water mass (S = 34 parts
per thousand) which is of arctic origin. In the area around the Sargasso
Sea the latter extends downwards like a trough or depression into the
deeper waters (Le D~ois, 1938). Orientation experiments with yellow
eels have led to the conclusion that, at this developmental stage at
least, salt concentration is not a significant orientation cue for the eel
over long distances (Deelder & Tesch, 1970 ; Tesch, 1970).
Other factors used by eels for orientation towards the spawning
grounds have been considered. The Gulf Stream could be influential
in spreading the smell of the Sargasso Sea into northern areas. So, by
perceiving differences in odour between the Gulf Stream and neigh-
bouring waters, the eel has the choice of swirruning in the direction of
the Sargasso Sea (Hasler, 1956). However, it seems that odour is not a
decisive factor behind orientation. I t is clear that yellow eels, which
relocated their home territory after being transported to a region many
kilometres away. did not use their sense of smell as a navigational aid
(Deelder & Tesch, 1970; Tesch, 1970). In addition, the boundaries
between the Gulf Stream and adjacent water masses could hardly
be very distinct, thus making it impossible for the eels to keep on a
EEL SPECIES 99
constant course the whole time. Orientation towards the source of a
smell by using variations in its intensity, can be ruled out from aquarium
observations (Teicrunann, 1959; Hasler, 1966). Finally, orientation by
smell in combination with rheotaxis - i.e. by using simultaneous
changes in current and smell- is improbable, since differential
currents within a large water mass are surely difficult to perceive.
Orientation using visual cues such as the stars or the sun must be
ruled out. Certainly the improved visual capacity of the eye in
the migrating eel suggests an increase in the use of this organ (see p. 69).
However, this change can most certainly best be understood. as an
adaptation to cope with the changed light conditions at greater depths.
Besides, how is a fish travelling at a depth of 700 m going to be able to
follow the direction of the stars? So, as mentioned above, some other
sense of direction remains the most probable means of orientation. The
receptor mechanisms and the environmental cues involved are, as yet,
unknown.
The discussion above shows how the spawning grounds of the
European and American eel were discovered as a result of research by
the now famous Dane, johannes Schmidt. As with the question of
possible spawning grounds in the Mediterranean, so the other conclu-
sions Schmidt drew from his results have also been questioned. The
objections are, in part, of a very basic sort, casting doubt on the existence
of two genetically distinct species of eel in the Atlantic (Tucker, 1959),
and in part deal with side issues such as the exact geographical position
of the spawning grounds of A. rostrata (Vladykov, 1964). These
critics have performed the useful service of triggering off further
research. Many of the questions raised have been discussed above ; the
most important points are summarized briefly below where the critics'
views and, in particular, argwnents put forward by scientists with
opposing ideas, have been taken into consideration (D'Ancona, 1959a;
jones, 1959; Deelder, 19600; Fontaine, 19<)1 ; Bruun, 1963; Sinha &
jones, 1967a).
( I) The number of vertebrae in the two species of Atlantic eel is
relatively constant; a difference of 7 vertebrae sharply separates the
two species. In A. anguilla, for example, the number of vertebrae
remains constant both from year to year and in individuals from the
Mediterranean and the East Atlantic. This suggests genetic
independence between East and West Atlantic eels and also implies
that the Mediterranean eel is conspecific with the East Atlantic eel.
An environmentally determined difference in the number of
vertebrae, which could have arisen in the spawning ground of the
100 THE EEL
...,-
,<~~'>~.
""'f-
/
,v:Y,;'(
A111/1#'10'10 " 't?~'
1~'::\
,.j'I.'
~i i A_Q,,~.trali.
,
A,c.leb_s.".i.
!
,
.............. _-_ ................ _-_ ........ .
Fig. 44 Tooth panerns from the upper jaw of different Anguilla species, and their
phylogenetic interrelationships (after Ege, 1939; reproduced from Dana
Report, No. 16).
American eel as a result of higher temperatures, would, according to

studies on salmonids (Tming, 1952), amount to a difference of less
than one vertebra (Bruun, 1963).
(2) Another taxonomic difference, the relative lengths of the
vomerine and maxillary tooth bands (Table 6, Fig. 44), confirms the
genetic independence of the two species.
(3) Electrophoretic studies of the haemoglobin (Sick et ai., 1967),
serum transferin (Fine el ai., 1967), and malate dehydrogenase in
the liver (Ligny & Pantelouris, 1973) revealed variations which point
towards species differences. However, these differences must be
viewed with reservation since eels from the East Atlantic and the
Mediterranean also show differences in these characteristics (see
p. 88; Drilhon & Fine, 1968, 1969).
(4) If the 19 pairs of chromosomes in the Atlantic species are
compared, morphological differences can be seen which seem
to indicate that both species have been living in reproductive
isolation for a long time (Ohoo et aI., (973).
EEL SPECIES 101
(5) The distance of more than 6000 km from Europe to the Sargasso
Sea could overtax the European eel's migratory capacities, with the
result that it might not be able to travel that far. This idea reinforces
the suggestion that each new generation of European eels arises from
the spawn of the American eel (Tucker, 1959). It can be rejected
because the Azores, for example, where A. anguilla occurs, are not
much further from the spawning grounds than are the coasts of
Canada.
(6) There are a number of arguments in favour of the idea that the
European eel is not capable of crossing the Atlantic. However, at
no time have these arguments seemed particularly well founded.
(a) The American eel is larger than the European eel. It seems
paradoxical that the fish which has to undertake the longer journey
is also physically weaker. There are, however, a large number of
arguments against this sort of reasoning (Bruun, 1963). For
example, it is well known that the smaller eels begin their migration
earlier in autumn than do the larger animals. Thus, to compensate
for their reduced physical capacity, the smaller eels have more
time in which to complete their migration. Also, because of
decreasing population density and the increasing number of
females, the size of the eels increases with the distance from the
spawning ground (Penaz & Tesch, 1970); some very large European
eels have been caught in distant regions such as north-eastern
areas of the Black Sea (Drapkin, 1964) and in the Danube drainage
of Hungary (Telegut, 1955). Similar phenomena seem to occur in
the American eel also (Vladykov, 1966).
(b) At the beginning of their migration, American eels start to
show a certain bronze colouring (Vladykov, 1966). In the European
eel this coloration is only found at advanced stages of development
(Svardson, 1949; Rasmussen, 1952; Wundsch, 1953; Bruun,
1963) ; the same phenomenon was seen in Mediterranean eels
with enlarged eyes (see p. 69). Tucker (1959) mistakenly considers
the bronze coloration to be an earlier developmental stage than
the silver coloration. Contrary to the views of this critic, the slower
development of A. anguilla can readily be seen as a correlate of the
greater distance travelled by this species.
(c) Physiologically the European eel is better equipped than
almost any other organism for a long sea journey. There is no
reason to suggest that it would become weak at an early stage in its
travels. The European eel has a large reservoir of fat, the fat content
increasing from about 4% in males measuring 25 cm, to 29% in
102 THE EEL
males measuring 39 em just before migration starts (Meyer-

Waarden, 1965; Table 43). The migrating eel can compensate for
changes in salt concentration of the seawater more easily than can
younger eels (Fontaine, Ig6I; see p. 24). Hormonally treated male
and female eels with maturing gonads were kept alive for several
months without food and without showing a great loss of strength
(Fontaine, 1961 ; Fontaine et ai., 1964).
(7) It has been suggested that it is easy for the American eel to find
its spawning grounds, which lie south of the continent. The spawning
animals are simply carried southwards in the counter-current of the
Gulf Stream, which runs along the coast of South Carolina, and in
addition, can use rising temperatures and increasing salt concentra-
tion as orientation cues. However, as was pointed out above, it is, on
the whole, not these factors, but some other sense of direction
which appears to guide both species towards the Sargasso Sea, at
least in the early stages of migration.
(8) Doubts have been raised as to whether the European eel can
reach the Sargasso Sea early enough in spring considering the speed
at which it has been observed to migrate. According to calculations
by Jones (1959) and Deelder (1960a), however, this certainly is
possible. Even yellow eels will show migratory speeds of as much as
3 km/h over long distances (Tesch, 1967a). Silver eels have been
followed in the North Sea using telemetric devices; despite the fact
that they were hindered by a transmitter, these animals still showed
swimming speeds of up to 6'5 km/h with the tide and, during un-
interrupted migration, travelled at speeds of 44 km per day (Tesch,
1972, 19743-c). Such speeds, together with the use of advantageous
currents (Deelder, 1960a), should enable the eels to reach the spawn-
ing grounds situated 6000 km away. by spring time after a journey
lasting about five months (see p. 93).
2.1.2 Continental distribution

The geographical distribution of the European eel after metamorphosis
is relatively well documented (Table 6; Fig. 45 ). It is panicularly
important to know the northern- and southern-most limits of distri-
bution because little doubt can then exist about the more central
areas of distribution. For example, eels were caught on the coast of
Munnan and on the northern Cape, so one can be certain that eels are
to be found everywhere to the south and west of the coast. According
to Schmidt ( l909b), there is an extensive area [0 the south that is
EEL SPECIES 103
- .,
.
..
, ",
"
"'
,~
..
,.,. ,,,.'"
Fig. 45 Distribution of the different species of the genus Anguilla.
populated by eels. At present, the Canary Islands represent the most

southern point where eels have been caught. Recent confirmation of this
was obtained by the author from fishermen on Las Palmas and Grand
Canary. Although the author did not personally see any of the eels
caught this far south, the fishermen's statements seem quite feasible
because, in addition to conger eels and morays, European eels appear
to be landed there frequ ently. The question thus arises whether
European eels also occur even further south along the west African
coast.
The western distribution of Anguilla anguilla is represented by the
eastern boundary lines for the distribution of American eels. Since
only a few islands must be investigated, there 3re no difficulties in
establishing the western distribution of A. anguilla. The eastern
distribution poses far more problems. While there are few doubts
with respect to the Baltic and its passageways, information on the
eastern rivers entering the Mediterranean must be viewed critically.
This applies particularly to regions lying south-east of the Suez Canal.
Although there are opportunities for eels to migrate in that direction,
reports of eels caught in the area could be the consequence of human
intervention, i.e. as a result of building the Suez Canal, just as eels
104 THE EEL
found in various places in the Volga have preswnably migrated there

through the network of canals connecting the Baltic with the Volga
(Schmidt, 1909h), Schmidt's (I92Sh) report of eels occurring near
Massaua on the west coast of the Red Sea seems credible, but Ege's
(1939) report of European eels in East Africa, three examples of which
are said to originate from Nairobi, is somewhat more doubtful. Con-
fusion with A. mossambica is possible (Jubb, 1964). Studies by Frost
( I957b) in the same area yielded no evidence to suggest the presence of
A. anguilla in Kenya.
The natural ascent of A. anguilla into the Black Sea can hardly be
doubted. Sdunidt ( l909b) reports many instances of eels being caught
in the nineteenth and even in the eighteenth and seventeenth centuries,
when stocking programmes over wide areas were not commonplace,
as they are today. For example, Schmidt cites Massili who reports that
eels were being caught as early as the seventeenth century near Linz,
Krems and Vienna. His reports on catches in the Upper and Lower
Dnieper are numerous; for the most part they originate from the
nineteenth century and in some cases, regular, annual catches are
described. There are also reports of catches from three different places
in the Sea of Asov during the nineteenth century.
Publications by Drapkin (1964), who provides reports on eleven eel
traps in the north-eastern part of the Black Sea between 1946 and 1965,
and by Telegut (1955 ) and Gyurko (1961 ), who cover the estuary area
of the Lower Danube, show how rare it was to come across eels in the
drainage of the Black Sea after the Second World War. Eels are only
seen from time to time, e.g. in 1900 in the Theiss and Danube (Sterbetz,
1960), though it is not clear from these two examples whether the report
was connected with eel stocking or with an influx of a particularly
strong year class. According to Michailowa (of the Zoological Gardens,
Sofia; pers. comm. ) individual eels are caught from time to time in
Bulgaria, in rivers which flow into the Mediterranean, e.g. in the
Struma, the Mesta and the Maritza. Even more rarely are they caught
in rivers which flow into the Black Sea. Sometimes eels are also found
in lakes along the shores of the Black Sea; these eels have probably
invaded the lakes from the Black Sea. Work by Caspers (1951) indicates,
however, that such reports are the exception rather than the rule. The
eel is not included in the list of fishes found along the coast of Bulgaria,
nor does it feature in the special section on less common, migratory
fishes which are of economic value.
The Black Sea is today, therefore, very much at the outer limits of
the European eel's area of distribution. The fact that relatively many
EEL SPECIES 105
eels are now to be found in European and Russian rivers that flow into
the Black Sea, is solely due to stocking (Orlov, 1966; Kochnenko, 1958,
1967). It is difficult to understand why, but even A. rostrata has been
introduced into waters in central and southern Russia, according to
Kochnenko (1975). The same applies in the drainage of the middle and
upper reaches of the Danube (Volf & Smisek, 1955; Meyer-Waarden,
1964; Schmid, 1962, 1964, 1966; Lassleben, 1966; Wiesner, 1966).
Eels have been caught in southern Germany since 1881 (Wiesner, 1966;
see also Hofer, 1897; Haack, 1879, 1881). These sources reveal that all
reports of eels in the Danube drainage since the end of the nineteenth
century could be the result of stocking. Mter the Second World War eel
stocking took place on a large scale. Near DiHingen on the Danube,
3'5 kg of eels per hectare were caught in 1959 and 1960,5'5 kgjha in 1961
and 8'0 kg jha in 1962. These figures exceed the yields for most areas in
northern Germany. The eel populations of some smaller lakes in
Northern Bavaria are also very large. In the Harunansberg lakes (in the
Rosenheim discrict) over 3 kgjha of eels are caught annually (Schmid,
1966). In the Upper Pfalz region, the greater part of which lies in the
drainage area of the Danube, the annual yields amount to between
3000 and 5000 kg (Dorfner, 1966). In Lake Balaton, 60000-400000
young eels and in addition 2 million glass eels have been stocked every
year since 1g6 I, resulting in very large landings of eels of a marketable
size (Koops, 1967a; Biro & Laszlo, 1970).
In view of these figures it is surprising that so few eels apparently
occur in the lower regions of the Danube. One would at least expect to
find migrating silver eels there. Information from the regions of the
Upper Danube indicates that eels do in fact, leave this area but do not
migrate in a westerly direction, as has been suggested in various
unverifiable publications (e.g. see Fisch u. Fang, 7, p. 196, 1966; the
Elbe-Jeczel Zeitung, p. 3, 9.8.63). An angling society on the Danube
claims, in one of its reports, that about 10 eels are found annually with
turbine wounds, and that many more probably perish without record.
All were large eels which suggests that they were ready to undertake the
spawning migration (Wiesner, 1966). Schmid (1g66) says: 'Experience
has shown that the silver eel migrates from the various lakes of northern
Bavaria at quite different times of the year.' From this information
it seems that there is a normal migration of silver eels in the region of
the Danube. The reason, then, that so few eels are caught in the Lower
Danube, is more likely to be due to a lack of suitable traps than to a
dearth of eels cravelling through the area.
The Bermuda Islands are seen as the boundary between the areas
106 THE EEL
of distribution for the two species of Atlantic eel. Although the larvae
of the European eel are more abundant than those of A. rostrata in the
waters round these West Atlantic islands, only the American species is
to be found on the islands themselves (Boetius & Boetius, 1967b).
Greenland also lies right on the border of the two distributions. The few
studies. that have been made, were based on only seven samples
(Sdunidt, 1909b; Jensen, 1937); further sampling is imperative. It is
quite possible that members of one species wander 'by mistake' into the
area occupied by the other - in fact this does occur, as Bruun (1937)
was able to show when he made counts of the vertebrae in a few hundred
glass eels from the coasts of North Spain. In this investigation one eel
was found which had 108 vertebrae, i.e. taxonomically it belonged to
A. rostrata. The probability of coming across A. rostrata is greater way
out in the Azores. Schmidt ( l909b), however, identified 34 eels from
these islands as A. angu£lla.
The northern boundary lines of Greenland and Iceland were men-
tioned above during the discussion on larval drift (p. 91). The northern
continental distribution of the American eel can be seen from Fig. 45.
Figure 46 shows a comparison of the annual yields from the various
provinces of Canada. According to these figures, the largest source
of eels is the province of Quebec which is supplied by the St. Lawrence
River. This is probably due to the fact that Quebec has, technically,
the most favourable conditions for utilizing weirs and draw nets, as
Fig. 46 Average eel catches (in pounds sterling) over the to-year period 1956-65.
compared with catches in 1965 in the Canadian provin~s of Newfoundland
(NFLD.), Nova Scotia (N .S.), the Prince Edward Islands (P.E.I.), New
Brunswick (N .B.). Quebec (QUE.) and Ontario (ONT.) (after Eales, 1968).
EEL SPECIES 107
well as having the largest freshwater drainage on the east coast of

North America. The catches show a sudden drop as one proceeds
further northwards, e.g. in Newfoundland (see Fig. 46; also Eales,
1968). Investigations into the distribution of A. rostrata in Newfound-
land (Flechtner & Anderson, 1972) have, however, shown that this
species can be found in almost every river system along the entire
coastline. The eel populations of the south-western coast are so extensive
that professional eel fishing has become well established there. It is
concentrated mainly in St. George's Bay, and in 1972 more than 36 tons
of eels were caught. In Labrador, eels are so scarce that professional
fishing is hardly economical. As mentioned before, this may be due to
the coldness of the southerly-directed Labrador current (Fig. 39),
which hinders the distribution of larval and glass eels.
The eel can only penetrate into the Great Lakes as far as Lake
Ontario; here there is an important fishery with catches of up to 136'2
tons per year (1959) (Gonad. Fisherman, 51, NO·3, 1964). Migration into
Lake Erie and the adjacent Great Lakes is prevented by the Niagara
Falls, though a few eels do seem to have managed the passage (Schmidt,
I909b). Their entry appears to have been via the Weiland Canal, i.e.
not by natural means. However, as far as the author can tell, eels are
rare and largely unknown in the Great Lakes.
Lastly, the eel's distribution is furthered by means of the Mississippi.
Around Illinois (Chicago), for instance, the eel is found more fre-
quently in the Mississippi than in Lake Michigan. Over a period of
five years anglers have caught 2 1 eels and the more professional concerns
have caught 16 eels in a lake near the river Illinois, which is itself a
tributary of the Mississippi (Starrett & Fritz, 1965). In its more northern
tributaries, such as the Wisconsin River, eels are even rarer. When
compared with data from Sdunidt (l909b) current figures show that
there has been a sharp decline in the nwnber of eels in this area. This is
probably largely due to the erection of dams. At the tum of the century,
eels were widespread in the Upper Mississippi and in the Missouri.
In the state of Illinois, for example, 20000 lb of eels were caught. In
Wisconsin, the Mississippi fisheries provided 1745 lb in 1902, and even
in the more northerly state of Minnesota 9<X> lb of eels were caught in
the river there.
The eel is still relatively common in the Lower Mississippi and in
other rivers which empty into the Gulf of Mexico. At the turn of the
century, however, catches were already considerably smaller than those
from the east coast of North America; as suggested before, this is
probably correlated with the marine currents which only bring a few
108 THE EEL
Table 6 Posttnetamorphic distribution of the various species of the genus Anguilla
Geographic region Geographic diJlribunon AurhQr
North Atlantic A. anguilla Europe and North Mrica. North- Schmidt,l909b

(Unnaeus) wards : as far as North Cape and also Sorokin &
to the cast of it as far as the Munnansk Konstantinow,
coast, in Kola Bay and in Northem 1960
Dvina. Southwards: as far as Atlantic
coast of MorolX(l and the Canary
Islands, Tenerife, Palma, Gomera
and Gran Canaria. Eastwards : the
whole of the Mediterranean region, the
Black Sea. Westwards: Iceland,
Madeira, the Azores.
A. rostrala North and South America. North- S<:hmidl, 1909b
(Le Sueur) wards: southern Greenland, New- Jensen, 1937
foundland, Labrador. Southwards:
Gulf coast of Mexico as far as Tampico,
and in Panama, GRater and Lesser
Antilles, c.g. Cuba, Jamai!;a, Puerto
Rico, St. C roix, St. Vincent, DominiQl,
Grenada and on the South Ameri!;3n
mainland as far as Guyana. Eastwards:
Bermuda Islands.
N.E. Pacific A.japonica Japan and China. Northwards: Ege, 1939

T emminck & H okkaido, coast of Manchuria, Matsui, t9S1
S<:hlegei Liao-ho River. Southwards: H ainan,
Gulf of Tonkin. Westwards: Bonin
Islands.
Northern A. rubulOfa Ceylon, Indian hinterland, Burma, Ege, 1939

Indian Ocean nebu/osil Andaman Islands in the upper drainage
McClelland region of the riven.
Northern to A. biro/or Nonhwards: Indian hinterland and Ege, t939

Equatorial birolor Burma. Eastwards: Sumatra, Java Jubb, 1961
Indian Ocean McClelland and Nonh-west Australia. Westwards: Kiener, t965
Malagasy and the smaller islands of the
Seychelles, Reunion, coast of East
Afri!;3, i.e. Kenya, Dar es Salam, and
isolated specimens further south as far
as the Cape coast. Occun mainly in
coastal regions.
A. nebu/osalabiala East African coast of Kenya, where it Ege, 1939
Peten is the most common species, as far as Frost, t954,
southern Mm:ambique, in Lake r957a
Malawi, isolated specimens as far as Someren &
the Cape coast, rare on Malagasy; Whitehead, t959
mostly in the upper stretches of the Jubb, 1961
rivers, near Nairobi at a height of Kiener, t965
looom.
A. moSJambica Coast of East Afri!;3, mosl southern Ege, 1939
Peters species, from Zanzibar to the Cape of Frost, 1954
Good Hope, slightly less frequent Kiener, 1965
in Kenya, the most common species on
Malagasy
EEL S PE C I ES 109
Table 6 continued
Geographic region Species Geographic distribution Author
Indian and A. marmora/a Most widely distributed. Westwards: Ege, 1939

Pacific Oceans; Quoy & Gaimard Natal, Malagasy (not very common) Jubb, 1959
Equatorial and islands near Malagasy, Mauritius Kiener, 1965
regions and Johanna; in South Africa from Matsui,1952
Port EliUlbeth and beyond. North- Nishi &
wards: Sumatra, Hong Kong, Taiwan, Imai, l969
Marianas Islands, the southem Japanese
islands of Yaku, Philippines. East-
wards : islands in the western Pacific,
e.g. from Samoa to the Marquesa
islands. Southwards : New Caledonia
Pacific, A. bicoior bicoior Insular, eastern part, particularly Ege, 1939

Equatorial Schmidt Celebes and New Guinea.
Regions A . ~Jebesensis Northwards: Philippine Island of Ege, 1939
Kaup Luzon. Southwards : Sunda Islands
Roti and Timor. Westwards: Nias
Island west of Sumaua. Eastwards:
Moluccan island of Halmahcira and
Geelrink Bay in the eastern region of
New Guinea.
A. ancestralis Manado, on the north-eastern part of Ege,I939
Eg' Celebes.
A. borneensis River 50, tributary to the Mahakam Ege, 1939
Popta on Borneo
A. interioris New Guinea: Humboldt Bay, Owen Ege, t939
Whitley Stanley mountains, Hagen
mountains at a height of 1300-1800 m.
A . obscura North-eastwards: Molluccan island of Ege,I939
Gunther Halmaheira. Central region : islands
north-cast of Australia, e.g. New
Caledonia, Fiji Islands and Samoa.
South-eaSlwards: Society Islands,
T ahiti, Cook islands.
South-eastern A. megl$/o»la North-eastwards: Solomon Islands, Ege, 1939

to southern Kaup from there in a south-easterly direction
Pacific to the New Hebrides, New Caledonia,
Fiji islands, Society islands and Cook
Islands as far as Pitcairns in the south
Pacific.
A. reinhardti East Coast of Australia from Cape Ege, 1939
Steindachner York in the north to Port PhiJIip in the
south.
A. awtralis australis South-eastern Australia from Cape Ege, 1939
Richardson Byron on the cast coast to Warrnambool
on the south coast, Tasmania and Lord
Howe Islands.
A . auS/ralis schmidti New Zealand, north of Norfolk Island, Ege, 1939
Phillips New Caledonia. Eastwards : Chatham
Islands. South : Auckland Islands.
A. dieffenbachi New Zealand, Chatham and Auckland Ege, 1939
Gray Islands.
IIO THE EEL
eels to the Gulf of Mexico, and therefore eels are uncommon on the
more southerly coastlines (see p. 91 and Table 6). On the other hand, in
a tributary of Lake Suwann which flows from Florida into the Gulf of
Mexico, large numbers of eels were caught throughout the year (Mellier,
1967), and of course streams from the Lower Mississippi complex are
well populated by eels (Gwming & Shoop, 1962).
As a result of restocking, A. rostrata has not only found its way into
remote regions of North America such as the Saskatchewan River in
Alberta (Radford, 1972), but is also to be found in natural waters in
other regions of the world. For example, Kochnenko (1975) reports
that A. rostrata has been introduced in central and southern Russia.
Healthy adult eels which have been recaptured show that the species has
adapted itself well to the new surroundings.
2.2 Indo-Pacific species

Geographically, the Indo-Pacific eel species are well separated from
the two Atlantic species. Thus there can be no doubt that the fonner
are genetically isolated from the Atlantic species, despite some dose
morphological similarities between them. However, the systematic
separation of the various Indo-Pacific species is not nearly as certain as
is that between A. rostrata and A. anguiJIa. Thanks to Schmidt and the
completion of his work by Ege (1939), the original jwnble of what was
thought to be about 50 species has been reduced to 16; nevertheless,
this field still requires funher investigation.
If one looks at the distribution of eels over the world (Fig. 45), in
addition to the division of the species mentioned above, one is tempted
to distinguish between the south-east African and the Indo-Pacific
species. A general review has led the author to make such a division.
Various studies suggest, however, that there are several instances
where species occur together in regions separated by great distances.
The most remarkable example is that of A . marmorata. This species
occurs over practically the entire Indo-Pacific region populated by
eels. The only place where it is not to be found is the northern temperate
zone occupied by A.japoniea (Figs. 47 and 53). The species A. nebulosa
and A. hieolor are also present in both the south-east African and Indo-
Pacific regions. One of the vital questions concerning this area of distri-
bution is just how far is one justified in distinguishing between species
and subspecies, a problem that is particularly acute in the case of A.
nebulosa and A. hieolor.
EEL SPECIES III
" ., .~ ~ s·
'/" ~
'" ~ ~ s·
I~
s
,'"
V (
I' _(
/ r'
/
18
I >s'
I".
1\ , ./
)
/,Ih I'nIb4bIt l/ICIwni"'lllrounci
.. AI>9<II//tJ t.icolor bicolor
'" 14ngulllo mormorolo
-
I""
,•
1
U••
s s
1-
"'
"~
1\ .. An9ulllo t.icofor bicQIor _

I""
\, , b An9ufllo mI1rmorola
I == A"9uilla MtJvfoso 10blala
loon Al>t}Ullio mouamI>icQ ,_
I'~
1 1 I ~ ~
15" 2!)0 2'!
" "" " 45" W"
ii,. 47 Distribution of the South African eel species. Abow: larval distribution and
SS<> fif' 65"
probable spawning area. Below: coastal and inland dislribution (modified

after Jubb, 1961 ).
.•2. I The south-east Mrican region: A. marmorala,

A. nebulosa fabiara, A . mossambiea and A. bie%r
iigure 47 shows the distribution of larvae and adults of the four species
f eel which occur on the east and south-eastern coasts of Africa; the
,robable spawning ground, north-east of Malagasy, in the path of the
112 THE EEL
South Equatorial current, is also marked. In this area, as in the North

Atlantic spawning ground, there is a zone of higher temperatures at
greater depths Gespersen, 1942). Between 10° and 20° S and at 60° E
warm water at a temperature of 15 °C extends to a depth of about 400 m.
This information, together with the predominant westward current and
the occurrence, north of Malagasy, of relatively small larvae measuring
37 mm has led to the assumption that the spawning grounds are in this
area. There are, then, far fewer clues in this instance than there are for
the Atlantic species, so that any assumptions are made with great
uncenainty. As can be seen from maps showing currents and the distri-
bution of larvae (Figs. 39 and 47), the latter are carried to Malagasy
and the east coast of Africa. Some drift round the northern part of the
island and the rest are carried round the southern part. Due to its
position relative to the various currents, Malagasy is populated by all
four species of eel. The larvae which drift northwards are distributed
mainly along the coast of East Africa. Some drift southwards between
Africa and Malagasy and are either caught up in the wann Mozambique
current or arrive on the coasts of Mozambique and Natal. The
leptocephali which drift around the south of Malagasy are also caught
up in the Mozambique current and later in the Agulhas current, which
continues even further southwards. From Port St. Johns onwards,
cold counter-currents on the coast, and the west wind drift from the
Atlantic, virtually stop the westward drift of the larvae, so that the
various species are unable to spread much further than Cape Town.
It is asswned that those eels which occur further south than 31° S and
which, for example, reach Cape Town, are not glass eels but are pro-
bably elvers which reached the coast the previous year and have pro-
gressed further by active migration.
Like the Atlantic species, the South African eels leave the coast and
ascend the rivers at a particular time. For A. mossambica and A. nebulosa
labiata, this transitional period is during the summer months, namely
January to March (Frost, 1957a; Jubb, 1961). Thus, in contrast to the
Indo-Pacific species, the South African eels probably have a seasonally
detennined spawning time. This assumption is strengthened by the
fact that silver eels of all four species also begin migrating at a particular
time of the year, i.e. between November and March. If the eels spawn in
the area close to the north of Malagasy, then it can reasonably be
assumed that larval development only takes one year. Support for this
assumption comes from the fact that in both A. mossambica and A.
nebulosa the glass eels are only 50 mm long. We know that the American
glass eel from the Atlantic (see p. 93) is also only 50 mm long after a
EEL SPECIES 113
larval period of one year ; in contrast the European glass eel is almost
70 mm long, but it takes approximately three years to reach that stage.
Jubb (1961) asswnes that the three species A. marmorata, A . nebulosa
labiata and A. meolor bieolor spawn in the northern part of the area
described. This would also accord with their distribution on the
continent.
Probably as a result of its position relative to the surrounding currents,
the island of Malagasy is populated by all four species. This makes it
possible to test various hypotheses which have been proposed with
regard to the inland migration and distribution of these species. A.
In'eolor meolor shows the least penetration inland; it is true that this
species is found in freshwaters but it does not leave the freshwaters
near the coast. One exception has been reported in Southern Rhodesia
(Frost, 1957a). A. marmorata migrates a little further inland, up to
altitudes of about 500 m. A. mossambiea and A. nebulosa, which was
recently discovered on Malagasy, penetrate to altitudes of 1000 m and
more inland (Kiener, 1965).
Similar differences in distribution which serve to reinforce taxonomic
features such as differences in the number of vertebrae or the length of
the fins (Table 5), have also been found in the two species of eel from
New Zealand (see p. 118). Such differences are probably due to
variations in environmental influences. Of course the extent of inland
migration or even of vertical distribution cannot always be determined
so easily. According to Jubb (1961), with increasing distance from the
main area of its distribution A. nebulosa labiara, for example, is not
found so far inland and, in general, is found only along the coasdine.
Normally, A. nebulosa labiara occurs far inland. In Kenya this is
the only species of eel which is to be found at heights over 1000 m
above sea level (Frost, 1954). A. nebulosa labiata has populated the
Zambesi River and its tributaries for 1000 km inland and has, in the
process, negotiated a series of waterfalls which span a total difference
in height of 60 m (Frost, 1957b). Jubb (1964) cites the case of a single
specimen being caught above the Victoria Falls. Anguilla nebulosa
labiara overcomes similar difficulties when it penetrates into Lake
Malawi. With many waterfalls and rapids, the outflow from this lake
drops 300 m over a distance of 50 km. As a result of this isolation there
are over 200 endemic fish species in Lake Malawi which is also protected
from invasion by other species from the Zambesi river system. Anguilla
nebulosa, however, is able to overcome this barrier if dry spells do not
cause the outflow to dry up (Schmidt, 1925b; Jackson, 1959). A.
mossambiea. which is the predominant inland species, is found further
114 THE EEL
south, on the edges of the main area populated by A. nebulosa labiata.

In Rhodesia, for example, A. mossambica has been caught 750 km from
the sea at a height of 1400 m ; A. nebulosa labiata first appears 150 kIn
further downstream, and then only infrequently (Jubb, 1961). Anguilla
marmorata is found along the entire coast of Soum Africa, on the
periphery of the main distribution of the other species. Its inland
range is thus slight, though of course there are exceptions. This very
adaptable species has been found in Rhodesia, 1000 km inland (Frost,
1957b), and at another place 1530 m above sea level (Herre, from
Ege, [939).
2.2.2 The temperate north Pacific zone: The Japanese eels:

A. japanica and A. marmorata
The Pacific counterpart to the north-west Atlantic eel is the Japanese
eei, A. japonica. This eel is probably carried, as larvae, from the North
Equatorial current into the Kuroshio current (Fig. 39), a strongly
flowing stream which is directed northwards by the coast, and which
corre~ponds to the Florida and Gulf Streams of the Atlantic. Thus the
distribution of the Japanese eel ranges from 18° N in the south to 42° N
in Ihe norIh (Fig. 48).
There have been innwnerable discussions about the spawning
ground of A. japonica. Unfortunately, only a few larvae have been
caught. One specimen, reliably identified as A. japonjca, was found
near the southern tip of Taiwan over water at least HX>O m deep (Matsui
el al., 19<)8). This 54 mm long leptocephalus was clearly not quite fully
grown, a larva in Stage I of development (Table 2). It is difficult to
deduce anything about the location of the spawning ground from such a
well developed specimen. Matsui et al. (1968) have cast doubt on the
identity of two specimens caught in earlier years (e.g. Matsui, 1957),
and other specimens identified as A. japonica were already in the glass
eel stage and had reached the Continental Slope (Shojima, 1966, 1967).
Thus, very little can be concluded from these specimens, and the
younger larvae described recently by Matsui and Takai (1971) and
Tabeta and Takai (1973) do not throw any light on the problem either.
More specimens of migrating adult eels, however, are available
(Matsui, 1957). These have been caught in a wide variety of gear in the
YelJow Sea, the China Sea, the Straits of Korea and around Japan,
over depths of 70 to 300 m. It appears that the adults were swimming
mostly at not very great depths and that quite a number were caught in
dip nets. The diameter of oocytes in these fishes (see p. 237) ranges
EEL SPECIES II5
120 '
40'N
II Angvll/a japoNca
~ AJ>gViI'" a~.trali•
• AII!!IIII", ce~/)e.en.i•
• Angvil'" IlIIf,,'IQri.
.•
:. .
,~.
•
"1+--\
20'SI+-- - - - - - - - , ..,)/ 'd------H ~.,
'W' 150' E
Fig, 48 Distribution of Anguilla japonica and some of the tropical Indo-Pacific eel
species (for original references, see Table 6),
from 0'17 to 0'27 nun and suggests a relatively advanced state of gonad
maturity. These catches are very interesting from the view point of
establishing the preferred depth of water for migration (and contribute,
indirectly, to our knowledge on the migratory behaviour of the European
eel as well). However, they do not give any indication of the preferred
direction of migration and thus cannot be used to locate the spawning
grounds.
Water temperatures give the only clue to the location of spawning
places, if conditions in the Sargasso Sea are taken as a model. In the
area south of Japan a surface eddy controls the movements of the water
masses; from it the Japanese current and the Kuroshio current
emerge on the west side (Fig. 39). As has been mentioned already,
these two currents are responsible for the northern distribution of the
larvae. There is a trough of warm water, similar to that in the Sargasso
Sea, on the southern periphery of the Kuroshio current (Schmidt,
1925a). Matsui (1957) has confirmed Schmidt's data and, on the
1I6 THE EEL
basis of the temperature distribution at a depth of 400 m (160 isotherm),

he suggests that the spawning grounds lie to the south of Japan between
20 and 28 N and between the islands of Lutschu and Bonin, between
0 0
125 0 and 140 E.

0
According to Matsui ( 1952, 1957) the Japanese glass eel migration

in the lower stretches of the rivers begins two months earlier than that
of the American eel, starting in October and ending in May. The earlier
arrival of the glass eels confinns his suggestion that the spawning
grounds of the Japanese eel must be nearer the coast than are those of
the American eels. Such ad hoc conclusions should not, however, be
accepted uncritically. We know that in the European eel, the ascent of
the southern coastal rivers begins in September (Gandolfi-Homyold,
1934; Heldt & Heldt, I9293,b) and may even last until July in the
north-east (Schmeidler, 1963; Tesch, 19(5); even the less well docu-
mented observations on A. rostrala indicate that the ascent begins. in
January but may also occur in June (Vladykov, 1966; Smith, 1968).
Thus it i~ very difficult to make interspecific comparisons for the times
of ascent because such large local differences occur within each species.
Furthennore, Matsui (1 957) assumes that the spawning period of the
Japanese eel must last a relatively long time; Matsui bases this assump-
tion on the wide time range for the beginning of ascent and the relatively
homogeneous lengths of the glass eels (5 6-58 mm). Thus, with respect
to the duration of its spawning season, the Japanese eel appears to lie
somewhere between A. rOSlrala and the tropical eels, which,
presumably, spawn throughout the year or at least for a large part of it
(Jespersen, 1942). As already mentioned, the spreading out of the period
of ascent is also seen in the Atlantic eels. The main causes of this are
the varying distances travelled to the coasts and variations in premature
warming of the waters involved in the ascent (Tesch, 1971). The
tropical species A. marmorala, which also occurs in the areas populated
by A. japonica, only ascends in the winter half of the year too (see
below) (Nishi & Im.i, 1<)69).
Figure 49 gives some idea of the frequency and distribution of eels
on the islands of Japan. The greatest number of eels is caught on the
south coast, i.e. on the side facing the ocean. The strongest influx of eels
can be expected on this side because of the Japanese and Kuroshio
currents. Catches are considerably smaller on the continental side.
The island of Hokkaido to the north has practically no eels and re-
presents the northern limits of the eel's qistribution (Fig. 48); this
relatively restricted northern distribution may be due to the cold water
of the south-flowing Oyashio current.
EEL SPECIES 1[7
• 17SCIO k~
o HSO kg
6. >37S0kg
Fig. 49 Annual catch by the Japanese commercial eel fishing industry in various
districts (modified after Matsui, I952).
On the mainland of Asia the japanese eel is not nearly as conunon as it

is on the japanese islands; in Korea, however, it is thought to be very
conunon. In and around China it is caught on a large enough scale to be
of economic importance. The japanese eel's southern-most point of
distribution on the mainland of China is the Canton River; in the
Yangtse-Kiang it is caught as far as 600 km upriver (Sclunidt, 1925b).
Unlike the Atlantic species, which are not sympatric, A. japonica
occurs together with A. marmorata, at least in the southern half of its
distribution. Specimens of both species have, for example, been found
near the island ofQuelpart to the south of Korea and along the southern
coasts of the japanese islands (Nishi & Imai, 1969). There were no
difficulties in identifying the two species (Table 5) j in contrast to the
japanese eel, A. marmorata has a marble-patterned coloration and 10
vertebrae fewer than A. japonica. The distance between the beginning
of the dorsal fin and the anus is also much less in the latter. While Ege
(1939, Table 6) reports that A. marmorata has a mean of 105'6 vertebrae,
individuals of the same species from a south japanese island were found
to have a mean of 104'6 (101-107) vertebrae (Nishi & Imai, 1969).
In view of their geographical separation, it may seem unnecessary to
distinguish between Japanese and European eels. However, as a result
II8 THE EEL
of human intervention, like the introduction of European glass eels in

Japan for stocking purposes, the differences between the two species
are assuming a new importance. Both species are very similar as far as
meristic features are concerned (Table 5), though they differ in the
electrophoretic characteristics of their haemoglobin (Sick et al., 1962).
The electrophoretic patterns of lactate dehydrogenase, malate dehydro-
genase isoenzymes, esterases and muscle proteins are particularly
useful as a sure means of distinguishing between the glass eels of the
two species (Taniguchi el al., 1972).
2.2.3 The south·east Pacific species from the temperate zone:

A. ollsrraiis and A. dieffenbachi
Localizing the spawning ground of the south-east Pacific eel species
presents only slightly less of a problem than it does for A. japonica.
The situation may well be similar to that of the south-east African
species.
Sixteen specimens of leptocephalus larvae have been found in this
area. The smallest of these larvae are around 25 nun long, and the
majority come from the area north-west of the Fiji Islands. To the
south of the Equator there is an almost exact mirror image of the current
formations of the Sargasso Sea; the dominant South Equatorial Stream
here always flows in a westerly direction (Fig. 39). The larvae from the
eastern regions must also come from here. An extensive trough of
deep, warm water stretches between 10° and 20° Sand 140° and
170° W (Jespersen, 1942). According to Schmidt ( I925b), this trough
lies a little further south, easily as far as 20° S. With the exception of
one specimen, all the larvae were found to the west of this area. The
great length of the warm water basin, the distribution of the larvae and
the multiplicity of species (besides the species from the temperate zone
there are also four from the tropical Pacific) suggest that the spawning
grounds stretch from west to east in a long series. Castle (1963) also
assumes that the spawning ground lies between the Fiji Islands and
Tahiti, the centre of this area being 170° Wand 18° S.
When one compares the freshwater distribution of A. australis and
A. die/fenbachi with that of the other species (Fig. 50, Table 6), it seems
highly likely that the spawning ground lies far to the east, and is perhaps
the most easterly of them all. A south-westerly current from the
South Equatorial Stream branches off as close as 60° W (Fig. 39).
This must be the most westerly point at which the two species could
still spawn without the larvae drifting too far to the north instead of to
EEL SPE C IES 119
....
J
~, Antull10 ,..",hordt,
&/, A"9u,/IO ou.trall. ou.ll"Olis
11111 Af>'}ul/la ouurol,. schmidti
Fig. So Distribution of the south·eastern to tropical· Pacific eel species (modified

from Ege, 1939).
New Zealand and South Australia, their normal area of distribution.

Bearing this in mind, it is clear that A. dieffenbachi has the longest
journey between spawning ground and coast, for the further east the
spawning ground, the furth er south are the larvae carried. This assump-
tion is supported by the fact that glass eels of A. dieffellbachi have the
greatest average length ; at Stage VI A n (Table 2) they are 64-70 mm
long . The glass eels of both sub-species of A. auslralis are 49-62 mm
long at the same stage (Ege, 1939).
As in other species of the temperate zones, the glass eels only begin
their ascent of rivers at certain times of the year. This is true for New
Zealand at least. There, glass eels of A. australis schmidti and of A.
dieffenbachi were seen ascending between August and January, i.e.
during the spring and early summer months of the southern hemisphere.
I t thus seems reasonable to conclude that these species have a particular
spawning period determined by seasonal changes. A. australis australis,
which is only found in Australia - i.e. in the warmer regions - does
not appear to have a seasonal rhythm of this sort (Ege, 1939) and is, in
this respect, similar to the tropical species.
Such observations playa large part in determining the biological
validity of a species. The taxonomic differences (Table 5) between
120 THE EEL
A. australis australis and A. australis schmidti are very slight and could
raise a certain amount of doubt as to the validity of recognizing two
subspecies. It seems possible that these taxa, unlike the Atlantic pair,
may have an environmentally detennined difference in the number of
myomeres and vertebrae. A difficulty here is, of course, the fact that
the subspecies with the greater number of vertebrae is carried to wanner
zones than the subspecies with fewer vertebrae. Thus an increase in
the number of myomeres as a result of temperature differences is very
questionable.
Populations of both A. australis and A. djejjenbachi are found very
close to one another in New Zealand and on the islands (Fig. 50, Table
6). Anguilla dieffenbachi could well be called the 'New Zealand eel'; in
New Zealand it is, however, called the 'long-finned eel', in contrast to
A. australis schmidt; (see Table 7 for distinguishing characteristics)
which is known as the 'short-finned eel' and is, with the exception of
its occurrence in a few additional islands, also a 'New Zealand eel'
(asswning, of course, that A. australis australis which also occurs in
south-eastern Australia, is really so clearly distinct from A . australis
sclum'dtt) .
The term 'short-finned' applies to four other forms of Indo-Pacific
eels (Table 5), the other species all have long fins. The dorsal fins of the
laner, unlike those of the short-finned species, begin relatively far
forward, so that the distance between the origin of the dorsal fin and
the anus appears very large in relation to body length.
In addition to New Zealand, the islands of Auckland and Chatam
are also populated by these two species. The short-finned New Zealand
eel, A. australis schmidti, is also found in large numbers to the north of
New Zealand, on the Norfolk Islands and New Caledonia, and there is
some indication that it might also occur on the islands of Fiji and
Tahiti (Ege, 1939), The boundary line between the distribution of this
subspecies and A. a. austrai£s appears to lie between Norfolk and Lord
Howe Islands.
In addition to A. austrai£s, the more tropical species A. reinhardti is
also wid'espread in eastern Australia. Both species can be distinguished
easily from one another by several characteristic features (Table 5).
Anguilla australis seems to predominate in soUth-eastern Australia;
according to a newspaper article, 4' 5 tons of eels of this species were
caught in the western part of the State of Victoria and were exported to
Europe (Trade News, Ottawa, 18, 10-1 I. H. 7).
The long-finned eel A. dieffenbachi appears to be predominant in
New Zealand. The press article mentioned above reports that a firm in
EEL SP EC IES 121
Table 7 Differences between the two species of New Zealand eel (modified after
Cairns, 1942a) (see also pp. 198, 208)
A. dieffenbachi A. aus/ralis schmid/ i

(LolIg -fimud eel) (Shor/-fi mud ul )
( I) Dorsal fin considerably longer than ( t ) D orsal fin only slightly longer than
anal fin anal fin
(l ) Vomerine teeth fonn a narrow band (l ) Vomerine teeth fonn a compact,
club-shaped band
(3) Eye in from of and above Ihe corner (3) Eye directly above the corner of the
of the mouth mourn
(4) Lips thick (4) Li ps thin
(5) Head broad (5) Head narrow
(6) O lfactory organ large (6) Olfactory o rgan small
(7) Mouth opening large, strong jaw (7) M outh opening small, small jaw
(S) T ail broad, tail fin well developed (8) Tail narrow, tail fin poorly developed
(9) Pectoral fins large (9) Pectoral fins small
( 10) Grows to over 180 cm in length and ( to) Rarely more than 90 em long and
IS kg in weish t I·S kg in weight
( II) Male silver eels 55 to 65 cm long ( I r) Ma le silver eels 35 to 4S cm long
and 0·9 to I· t kg in weight and an average weight of 0·l5 kg
( Il ) Scales not very distinct ( Il) Scales dearly \'isibl e
Auckland, on North Island, exports 100 tons of silver eels, all A.

die./fenbachi, to Europe each year. However, it is questionable whether
this information gives a true representation of the relative numbers of
the two types of eel. Research has shown that both species occur with
almost equal frequencies on South Island, though this is not obvious
from Ege's (1939) data ; of 2215 eels caught in a trap on the east coast of
South Island 53% were long-finned and 47% were short-finned eels
(Burnet, 1968). However, since long-finned eels are larger than short-
finned eels, the total weight of the former is much greater than that of
the latter. In contrast with these figures, electrofishing in a stream in
the same region produced a larger number of A. australis (64 %) than of
A. dieffenbachi (36%) (Burnet, 1959). The percentage of A. australis in
three streams in the southern regions of North Island varied as follows :
11-88%, 1-21 % and 32-37%. From the point of view of weight,
however, these proportions were mostly much smaller (Burnet, 1952a).
Thus it is easy to see why, commercially speaking, the short-finned eel is
largely overlooked.
According to current infonnation, the short-finned eel (A. australis)
stays mainly in coastal waters, whereas the long-finned eel (A.
die./fenbachl) migrates further inland. This difference in distribution is
attributable not so much to the migratory characteristics of the two
species, but more to the effects of differing environmental pressure .
To quote Burnet (1968): 'The short-finned eel is selective in its choice
of habitat, and is the exclusive inhabitant of some coastal waters. It
122 THE EEL
generally prefers lakes and spring streams. The long-finned eel is

widely distributed, and it is found that almost every water has either
one or both species.' Thus there is no question that the short-finned
eel also inhabits the upper stretches of rivers. On the whole, however,
these eels are very small and it was only the use of electrofishing techni-
ques that revealed the large numbers that sometimes occur there.
Burnet (I952b) assumes that, when the short-finned eels grow larger,
they seek out slower flowing rivers, lakes or coastal waters.
2.2-4 The tropical eels

A. celebesensJ"s, A. megastoma, A. interioris, A. anceslralis,
A. nebu/osQ, A. marmorata, A. reinhardti, A. horneensis,
A. hie%r, A. obscura
Sclunidt and his co-workers had more success in their search for the
spawning grounds of the tropical eels than they had with the eels of
the temperate zones. Immediately south-west of Sumatra about 1300
larvae were caught in a relatively precisely defined area, and north-west
of New Guinea a further 233 were trapped. Presumably these two areas,
together with the one detennined for the species of the south-east
Pacific (see p. 118), comprise the most westerly spawning grounds of
the tropical eels.
A valuable clue to the location of the spawning areas was given by
the way in which the eels are distributed through the islands (Fig. 51).
The south-western coasts of Sumatra and Java which face the deep
sea, and the rivers which open there, are populated with eels but none
are to be found along the north-eastern coasts which are exposed to
shallow waters. The south-western side of Borneo, which likewise
faces shallow water, is also unpopulated by eels. On the east side of
Borneo, however, there are large numbers of eels, as is also the case
over the whole of the Celebes which lies directly in front of the east
coast of Borneo. Here the sea is, in places, well over 1000 m deep, an
imponant prerequisite for the occurrence of leptocephalus larvae.
The islands of the East Indian archipelago present a similar arrangement
to that of the Japanese islands where most eels are to be found on the
coasts facing deep water beyond the Continental Shelf (Fig. 49).
It thus seems likely that the spawning grounds are situated on the
south-western and eastern side of the East Indian archipelago.
Figure 52 shows the number of eel larvae caught at different stations
near Sumatra. Judging from this diagram, the highest concentration
of larvae occurs directly on the Continental Slope, over depths of
.. ...
EEL SPECIES 12 3
.
i
j
\-
\ ' -'lOOm" ."
ArtClSWlt h nls
lI........ s without MIs
. . ,...., Riven ....Ith rtlotlvtly f, .... "Ii
Fig. SI Distribution of Anguilla species on Sumatra, Java, Borneo. Celebes and

neighbouring islands (modified after Delsman, 1929).
between 2000 and 5000 m, just off the central area of the south-western
coast of Sumatra. As in the Sargasso Sea, high concentrations of larvae
(especially of leptocephali less than 20 mm long) and high temperatures
at some depth occur in the same area. In the region off Sumatra the
temperature was 12 °C at a depth of 300 m j it was only 11 °C in the
surrounding walers (Jespersen, 1942). Of the 1300 larvae caught in this
spawning ground the great majority, at least 950 animals, was A.In·color
biwlor. The larvae of this short-finned species are easily distinguishable
from those of the other species. The remainder, all of which were
long-finned, consisted of A. marmoraca, A. nebulosa nebulosa and A.
celebesensis. For the most part it was impossible to distinguish one species
from the other j the most accurate identification possible was of A .
nebulosa nebulosa, which has relatively more vertebrae and myomeres
(Table 5). For this reason larval identification was based mainly on the
distribution of the various species in the neighbouring areas of land
(Figs. 48, 50, 53 and 54).
The situation for determining the outer limits of the spawning
grounds is not so favourable east of the Indo-Malayan archipelago.
124 THE EEL
,,- ,,-
,i
"
Grtclt Nicooor Is. (;)
., ~,--
..• _--- 200", depthhn,
._--.-. l000m depth line
'3 \> 1 3. ," ", ~ --_.- 500Qm depth line
I· \\ . \ ' . ........ }
'v r '
,.f----'---~,'~'--~'4'-\ MALAU ',.-+----1'-
,. •
:'ingapore
v
; .o!\'il
"
i/ (.'\ Chr's\fII(IS I~ ~
. '
(n '
) '~.
/
,,- ,00"
Fig. 52 Number of eel larvae caught at various stations near Sumatra, calculated
per 10 trawl hours. Crosses without numbers .:. no eel larvae caught (modified
after Jespersen, 1942; reproduced from Dana Report, No. 22).
As was mentioned earlier, only 23s1arvae were caught here. Jespersen's

(1942) distribution map shows the highest concentration to be situated
to the east of the Moluccas, directly on the north coast of western
New Guinea. Here the larvae occur right up as far as the Shelf region,
with islands and shallow stretches of sea lying between the individual
EEL SPECIES 12 5
• AfW}vil/a ".OO/oS(! ".00/0$0
111111 ""9vJlia hlormorata
• An9villa o!>$~vra
~ An9villa bor"flMSis
..0<> 150<>
Fig. 53 Distribution of some of the tropical Indo-Pacific eel species (for original
reference, see Table 5).
~."
,..
<0" 1\--. Ang<Jilfa !>icolor biCOlar
. . Angvfl/o bicolor pacifica

• AfW}villa hI_ga.ta,""
Fig. 54 Distribution of some of the tropical Indo-Pacific eel speeies (modified from
Ege, 1939).
126 THE EEL
populations of larvae. It seems reasonable, therefore, to propose the

existence of several, independent spawning grounds. The only way of
sorting the larvae systematically is to divide them into long- and
short-finned individuals. The result of this division shows that the
long-finned eels appear to occur in larger numbers near the coast than
do the short-finned species. In addition, short-finned eels are to be
found as far north as the area just south of Taiwan. The most northerly
station to record long-finned larvae was situated immediately south of
the Philippines.
In the present case it seems that A. hieolor pacifica is the most
abundant species; most of the short-finned larvae (of which there were
129 from this area) were probably of this species. Anguilla obscura,
the other type of short-finned eel in the region, occurs only in small
numbers, as is indicated by Jespersen's (1942) taxonomic studies. The
limited inland distribution of this species in the western part of its
general distribution, which is restricted to a few records from northern
New Guinea (Fig. 53), supports Jespersen's findings. The discovery of
a few A . obscura larvae in the stomachs of tuna fishes (Kacsuwonus
pelamis) confirms the presence of A. obscura in this region (Matsui
et al., 1970).
The long-finned species to be considered, judging from their inland
distribution, are A. marmorata, A. borneensis and A. inten·oris. Together,
these species make up the largest proportion of all long-finned larvae
that have been caught. They can be distinguished from the other
larvae by the number of vertebrae they possess (Table 5). The remaining
larvae must have been composed of A. celebesensis and A. ancestralis.
According to Jespersen (1942), A. marmorata is the most common
species in this area and therefore a predominant proportion of the
long-finned larvae is probably of this species. In inland waterways,
A. celebesensis is also distributed over the whole larval area, so this
species too must have contributed to the larval material, a supposition
apparently confirmed by morphological features. The remaining species
have only a very restricted distribution over the total area under
consideration (Fig. 48), so that there were probably very few specimens
represented among the larval material examined.
Favourable hydrological conditions of the sort which appear necessary
for spawning in the eel, are also found in the region north-west of New
Guinea where the high concentration of larvae occurred (Jespersen,
1942). At a depth of 300 m the temperature was 15 °C and the salt
concentration was higher than in the surrounding water masses.
In both the areas thought to be spawning grounds, i.e. near Sumatra
EEL SPECIES [27
and near New Guinea, the currents are not as dearly oriented as they
are in the supposed spawning grounds of temperate zone eels (Fig. 39).
In the area with the highest concentration of larvae near New Guinea
there is, in summer, a whirlpool, which, although confined to a very
small area, is very strong. It appears as if the larvae, especially those
around Sumatra, could be dispersed in several directions. This would be
particularly important for such widespread species as A. mannorala
and A. hieolor. Anguilla marmorala, for example, is found both far
east and far west of the spawning ground near Sumatra. Of course it
cannot be established from our present knowledge oflarval distribution
whether, with the exception of the spawning ground north-west of
New Guinea, these eels, and other species too, have other spawning
grounds. The occurrence of larvae thought to be A. marmorata has
suggested the existence of an additional spawning ground for this
species in the area used by eels from the south-east Pacific (Castle,
1963; see above). The spawning areas described in this section are
probably not frequented by the more southerly species A. megasloma
and A. reinhardli. Their spawning grounds are almost certainly restric-
ted to an area in the South Equatorial current (see p. 92), as has been
indicated by larval finds from this region (Castle, 1963).
On the basis of the vertical scatter of the larvae it is possible to say a
little about the depth of the spawning place. Figure 55 shows the
distribution of larvae at different depths and is based on figures from
,
, ~.
D'
•
, I
, I
,
0
1
~ 100-150 200 -2W lOO -l50 400 -4W ~5W 000 -6W 700
l~nq!h of \0,", linol .. )
Fig. 55 Depth distribution oflarvae of short-finned (A) and long-finned eels (B) north-
west of New Guinea. Ordinate : number of larvae caught per Io-hour period.
Abscissa: length of tow line paid out. Half the value of the length of the line
indicates the depth of water fished (after Jespersen, 1942; reproduced from
Dana Report, No. 22),
128 THE EEL
the area with the highest larval concentration north-west of New

Guinea. Judging from this diagram, most larvae of the short-finned
eel occur at a depth of between 100 and 200 m, and the larvae oflong-
finned species occur at a depth of about 200 m (provided that one can
safely calculate depth on the basis of the length of line used on the
fishing net). The highest larval concentration in the area around
Sumatra is at a depth of between 120 and 300 m. This applies to both
older and younger larvae (Le. those less than 20 mm long). Most of
the larvae less than 20 mm long were probably confined to water
between 200 and 250 m deep. One can thus assume that the spawning
beds themselves are to be found at about this depth.
The depth at which the larvae were found also varied with the time
of day. At midnight, larvae appeared to be at slightly higher levels
than they were in the morning and evening. Larvae which are less than
20 mm in length apparently do not show such depth changes, which is
understandable in view of the fact that young at this developmental
stage are less active Gespersen, 1942).
The Danish research did not provide much information on spawning
periods. Sampling of glass eels indicates that ascent of the rivers, and
thus the spawning periods too, have few seasonal restrictions (Ege, 1939;
Jespersen, 1942), e.g. during a period of only two months when larvae
were collected in the area around Sumatra, three different-size groups
of A . bieolor bie% r were found. However, research in India (pantulu,
1956) on A . nebulosa nebulosa, which probably also spawns near Sumatra,
suggests that the period of ascent is greatly restricted by seasonal
changes; as in Europe and North America, the ascent of glass eels is
confined to the winter half of the year. Since temperature barriers are
unlikely during the period when the larvae are drifting towards the
continent, it is difficult to see how the observed seasonal rhythm can
come into being. However, it is possible that this rhythm is associated
with the dry and rainy seasons. The migration of silver eels towards
the sea is determined by the flow of water in the rivers, so that
rhythmical spawning times could be conditional on changes in water
level. Near Calcutta, for example, there is a summer and a winter
rainy season which might result in a bi-arumal rhythm. But that this
should affect the glass eels' ascent seems unlikely. The idea that both
glass and silver eel migration, as well as the corresponding spawning
period, might be dependent on water levels is also questionable because
the weather in northern and southern India is not uniform, and
presumably a regular spawning and migratory rhythm could not exist.
A much more likely influence, also determined by equatorial factors,
EEL SPECIES [29
TabJe 8 Average lengths of tropical glass eels at

deve!opmentaJ stage VI A (see Table 3).
Data were collected at various dates
and places (after Ege. 1939)
Species Length j" mm
A. ctlebesensis ,8
A . anceltTalis 49-54
A. nebulosa n ebulosa 49- 54
A. marmara/a 49-5 0
A. reinhaTdti SO-58
A. bie% r meo/or 52-55
A .obswra 49
but which is much more regular, is the currents in the northern areas
of the Indian Ocean. In winter the currents are westerly, i.e. they flow
from Sumatra to the coasts of India (Fig. 39); in summer, however,
they flow in the opposite direction. Thus in summer neither the
leptocephali nor the glass eels could manage to travel a distance of at
least 2000 km from Sumatra to India against a current flowing at one
mile an hour in places (Morskoi Atlas). Detailed research is still needed
to solve these questions.
The tropical eels reach lengths of about 50 mm in the glass eel stage
(Table 8). Measured against the lengths of the two Atlantic eel species
(Fig. 40), they appear to go through a relatively short period of develop-
ment. If one analyses the figures on growth rate in the larvae of A.
hieolor hieolor (Jespersen, 1942), it seems that the developmental
period from egg to larval Stage I (Table 2) lasts three months. In this
species, for example, only Ii months is necessary for the average length
after hatching to increase from 39 to 50 mm. Development into the
glass eel stage cannot take more than four months. In view of the fact
that some of the spawning grounds lie so near the coast it is likely that
this time-span has been over- rather than underestimated.
2.3 Zoogeographical relationships

The various species of Anguilla are relatively closely related to one
another and yet they are spread over the whole globe; research into their
common ancestry raises many questions of a geological, zoogeographi-
cal and genetic narure. Let it be said from the very beginning that, to the
present day, nothing definite is known about a common place of origin.
Therefore these various questions will only be touched upon lightly
in this book.
THE EEL
There has never been any doubt that, for example, the two species
of Atlantic eel are closely related; there has even been a theory that the
European and American eels are members of the same species (see p. 99).
A study of haemoglobin structure showed, for example, that that of the
two Atlantic species is formed in the same way, while that of the Japanese
eel is different (Sick et al., 1962).
The first step in untangling their phylogeny is to try and establish
the common ancestry of these two species, which are clearly so closely
related, and to explain the cause of their unusually marked geographical
separation as adults but their geographically contiguous spawning
grounds.
Von Ubisch (1924) and later other authors (Ekman, 1932) suggested
a link between eel migration and Wegner's continental drift theory.
This theory postulated that the continents of America on the one hand,
and of Europe and Africa on the other, were at one time a single land
mass. In the Cretaceous period this continental block split apart, and
during the Tertiary an ever-widening trench appeared which finally
became the Atlantic Ocean as we know it today. The spawning grounds
of the eel species of that period were presumed to be in the same place
as the present-day Sargasso Sea, i.e. in the Atlantis of the Tertiary
period, a sea (or lake) which, at that time of course, was much less
extensive than the present Atlantic. While the eel's spawning ground,
and maybe its associated hydrological conditions, drifted off in a
westerly direction, part of the eel's home ground, i.e. Europe and
Africa, remained in or drifted towards the east. Thus these eels had to
make journeys of increasing length in order to get to their spawning
grounds.
During the time when doubts were accumulating about Wegner's
theory, the corresponding hypothesis on eel migration gradually lost
its feasibility. Bertin (1956), who quotes the geologist Termerier's
view on Wegner's theory, has the following to say: 'It is a beautiful
dream, the dream of a great poet. But if one tries to embrace it one sees
in his arms only vapour and smoke. It attracts, it interests, it amuses
the mind but the solidity is missing. ' Currently, the basic concepts
of Wegner's theory are enjoying a renaissance. The underlying geolo-
gical aspects have changed somewhat and are now finnly based, but
these features cannot be dealt with in detail here (see Tarling & Tarling,
1971). It is now thought that the continent of America, including the
sea bed as far as the mid-Atlantic ridge, drifted away from Europe and
Africa, the mid-Atlantic ridge being the line along which the two con-
tinental masses were once connected to each other. This theory is
EEL SPECIES 13 1
supported by evidence from anomalies in the palaeo-magnetism of

the sea bed near the mid-Atlantic ridge, and by other evidence as well
(Bullard, 1969; Menard, 1969; McKenzie, 1970).
Thus, it does seem likely that the two types of Atlantic eel did have
a common ancestor, which spawned in the much smaller proto-Atlantic.
The question now arises as to the relationship between these species
and those of the Indo-Pacific. These latter must have evolved at some
earlier time. Central America has not always acted as a barrier between
the Atlantic and the Pacific but it is highly improbable that the eels
migrated over Central America (see below); it seems more likely that
there is some connection between the Atlantic and Indo-Pacific species
to the east. The present-day Mediterranean is the remains of an earlier,
larger sea which existed before America split away from Europe and
Africa. This proto-Mediterranean (Tethys Sea) divided a northern
from a southern continent and was in existence during the Mesozoic
and at the beginning of the Tertiary period. It acted as a link between
the present Atlantic regions and Indo-Malaysian waters. The fauna
of the western Pacific and the Atlantic have retained certain common
features from the period of this Mesozoic sea, features which are still
to be seen today, not only among eels and other fishes (see, for e.g.
Greenwood, 1976) but in other animal species as well. The existence
of eels from the Cretaceous to the Miocene period is indicated by
deposits from Tethys. Geological events caused the eel population
of this time to split up, producing a division similar to that between
the eels of the east and west Atlantic.
The Indian Ocean probably evolved in a similar way to the Atlantic
Ocean. Originally India, together with Australia and the waters to
the south of India, formed their own northward-drifting plateau and
were connected, to the south-west, with Africa (Bullard, 1969). This
provides a good explanation for the separation of Indo-Pacific and
African species; from a taxonomic view point, though, this separation
has not yet resulted in complete differentiation of the forms inhabiting
the African region, although some endemic species have evolved in
the Indo-Pacific region (see p. 110).
For the reasons already mentioned, a certain degree of scepticism
exists with regard to the idea that the Indo-Malaysian region should
be considered as the nursery ground for all species of Anguilla, in-
cluding the Atlantic eels which were thought to have migrated eastwards
from the Pacific into the Atlantic (Ekman, 1932). This view point
has been put forward on many occasions to account for the abundance
of species in the Indo-Pacific and their scarcity in the Atlantic. A region
[32 THE EEL
in which many different species occurred was, in the past, considered

to be the centre of origin for a particular lineage. Geological events
and palaeontological data already described, however, suggest that
speciation in Anguilla occurred after the separation of the land masses
with which the taxa are now associated. As a result of unfavourable
conditions in the Atlantic there was less speciation there, both in the
eels and in other marine animals. The hypothesis on the origin of eels
in the Indo-Pacific has not, however, been totally discarded; para-
sitological-zoogeographical data are still used to support this theory
(Manter, 1955; see p. 356).
3
Post-Larval Ecology and
Behaviour
3.1 The glass eel and elvers: their migration on the

Continental Shelf and into freshwaters
3.1.1 Migration in the sea

The term 'glass eel' here refers to all developmental stages from the end
of metamorphosis in the leptocephalus to the beginning of pigmentation,
i.e. Stage Vb (see p. IS). Following this stage the young eels are called
'elvers', Geographically this account begins at the Continental Shelf,
the region where the account of larval distribution in Chapter 2 was
interrupted.
According to Schmidt (1909a), the Continental Shelf represents a
clear dividing line between larvae and those animals which have
undergone metamorphosis. At the margin of the Continental bloc
there is a predominance of animals which have attained the external
appearance of the adult eel. With regard to pigmentation. however. the
young eels once again deviate from the adult form.
No one has yet been able to explain what causes this sharp division
directly above the Continental Slope in the region of the 1000 m depth
contour. Hydrological factors must be instrumental in preventing the
young eels from prematurely entering regions which are unsuitable.
Clearly eels in the pre-metamorphic stages are carried along by the
Gulf Stream; their locomotory mechanisms are very different from
those of the glass eel (Dean. 1912). The larva wastes little energy in
133
134 THE EEL
swimming and moves without producing any turbulence in the water

(Spacek, 1930); the glass eel, on the other hand, swims in a snake-like
fashion. The larva's narrow, willow-leaf-like form is thought to aid its
relatively passive propulsion during the planktonic phases of life,
whereas the glass eel and the adult are more suited to a benthic existence
(Wunder, 1<)63 & 1964)·
These basic cbanges in locomotion could perhaps be interpreted in
the following way. Movements in the larva are made merely to bring
about a cbange in position, regardless of the direction of progress.
Each body section follows the one in front, rather like the carriages of a
train moving in succession along a curved track. During this process the
head of the larva constantly changes position, thereby enhancing the
chances of it encountering prey. The glass eel, on the other hand, can
make goal-oriented, forward movements and can actively swim against
the current (see below). If larvae were carried into the tidal current of
the coastal regions, they would be swept passively backwards and
forwards and it would be impossible for them to migrate into the inland
waterways (but see below). By its own efforts, the glass eel is, however,
able to migrate actively and thus enter coastal and later inland
waters.
In view of their locomotory characteristics, it still remains to be
explained how the larvae are able, by means of their own activity, to
resist being carried over the 1000 m line and onto the shallow shelf
area. Deelder (1970), however, has taken up a suggeslion by Westenburg
(1952) thaI fishes can perceive differences in depth by using certain
physical factors as cues. For example, fishes swim along the coast at a
certain constant depth by perceiving oscillation resonances which come
from the sea bed. These are produced by waves at the water surface and
their frequency, after being reflected from the sea bed, is increased by a
factor of four times the depth of the water. It is possible that the larvae
respond to these frequencies but they must also be capable of a certain
amount of swimming activity in order to prevent their being drawn into
shallower waters by the current. Observations on freshly caught
leptocephalus larvae showed that the animals are in no way as inactive
or passive in directional movements as was implied above (Deelder &
Tesch, unpublished).
Figure 56 indicates at what time of the year glass eels are likely to
appear around the coasts of north west Europe. It is clear from this
figure that glass eels are already beginning to appear in September on
the coasts of France (and Spain), while they are not found until February
in the mouth of the Channel and near the Heligoland Bight. The eels
POST-LARVAL ECOLOGY AND BEHAVIOUR 135
" _ _ -- - )s,oPt _
~
/
/(1
/ i(J,
MetomOtpho ..S ' :
I 0
~ \
" Stpt. , Oct
" !5., , <i:"''>m!
,- ",
"" ,
Fig. 56 Probable times of arrival for glass eels on the north-western coasts of Europe,
according to data from various authors. Single or double lines beneath Roman
numerals indicate, respectively, the month of arrival according to Bowman
( 1913) and Schmidt (1909a). Data for January and February ( I and II) from
Creutzberg (I96t ) or, in the coastal regions of Germany, from Tesch ( 197t ).
take longer to penetrate the more easterly parts of the sea than to reach
places which are in direct contact with the Gulf Stream. In addition to
distance, temperature is probably also an important factor in controlling
this timing; in the north eastern areas it is too low to allow the glass
eel to penetrate this far so early. Contrary to previous evidence (van
Heusden, 1943; Creutzberg, 1961) the invasion takes place earlier
than was originally thought; the eels do not first reach the Heligoland
Bight in March or April but arrive there sometime during February
(Tesch, 1971). At the beginning of April glass eels were seen in the
Skagerrak around an underwater lamp near the surface or were caught
in nets (Lindquist, 1972). In the Baltic, too, eels appear earlier than
one might expect in view of the greater distance they have to travel.
Herold ( 1933) recorded glass eels in the coastal regions of the cl!ntral
Baltic at the end of May; from the Baltic coast of Schleswig-Holstein
Herrmann (1957) reports that glass eels are usually found in river
estuaries from the middle of May onwards, and in 1957, they were
THE EEL
J,"
"" Nov Dtc
,B ilboo
ftb. Mar
"
...•,
10
. [
02
----------
Oct . 'w Otc. Jan. Ffb. Mar. Apr. May Jun.
Fig. 57 The length of elvers caught at various times of the year in the Mediterranean
region and western Europe (modified after Heldt & Heldt, 192~).
present by the beginning of April. Normally, glass eels first begin to

appear in the freshwater reaches of the river Ems towards the end of
April (Schmeidler, 1963). If they appear in the North Sea in February,
i.e. two months earlier, then they must also be present in the Baltic
two months before the observed times of ascent.
Glass eels appear along the coasts of the Mediterranean at about the
same time as they appear on the Atlantic coasts. The question arises
here whether the larvae need one more year to reach the Mediterranean
than they need to reach the coasts of western Europe (Harden Jones,
1968). Schmidt (1912) found fully grown larvae and larvae undergoing
metamorphosis in the central region of the Mediterranean in spring,
i.e. at about the same time as similar larvae are found west of the British
Isles beyond the 1000 m depth contour. Thus, it seems, even before
metamorphosis is complete, these animals find themselves near the
coasts, so that, with the higher temperatures obtaining there, ascent into
freshwater can begin in autumn. A review of the literature by Heldt
and Heldt ( I929b) shows that, along the west coasts of Italy, near
Livorno, glass eels are found as early as October (Fig. 57). In the Nile
and in the inland waters of Tunisia they have been sighted in January.
Their arrival in the coastal regions can be estimated as occurring one
or two months earlier; in other words, in the Mediterranean too, glass
eels can be expected to appear outside the coastal areas between August
and September.
A developmental period in young eels involving basic physiological
changes (changes equalling those in the transition from the larval to the
glass eel phase) is that in which there is a change-over from salt- to
freshwater life. This transition involves basic c!langes in behaviour
and the beginning of pigmentation (see p. 14). The movements of glass
eels up to this period. are still largely obscure. One point is certain
though, the eels are now capable of active movement in a particular
direction.
On the basis of data summarized in Fig. 56, Creutzberg (1961 )
comes to the conclusion that glass eels in the North Sea are carried along
from north to south at a speed of 7 km a day. Since north-easterly
currents of a similar strength have been observed in the Channel,
Creutzberg asswnes that the coastal journey of the glass eel is largely a
passive one. However, since glass eels are capable of active swimming,
Deelder (1952, 1970) casts doubt on these data; he believes that the
eels must travel faster than this.
According to Schmidt (l909a), metamorphosis is completed and
Stage V reached in September, over the 1000 m depth line. The glass
eels appear near the coasts of Belgiwn and Holland in January, and
near the coasts of Germany in February (Tesch, 1971). Thus, they have
to cover a distance of about 1000 km in order to reach the Dutch coasts,
and for this they need a period of about four months. This would corres-
pond to covering a distance of more than 8 km a day. To reach the
Heligoland Bight requires travelling 1200 km within five months and
necessitates a speed of about 8 km a day. This analysis shows the need
for a greater rate of travel than 7 km a day, but does nOl give sufficient
grounds for casting doubt on Creutzberg's (1961) hypothesis. Neverthe-
less, there is another argument in favour of active migration in glass eels:
how is it that eels in these developmental stages are able to reach various
coasts in the Mediterranean and the Baltic so early? Highly variable
currents occur off the different coasts, and in many cases passive
migration must be ruled out.
Finally, there remains the possibility that the eels are carried along
by tidal currents, though progress would be very slow near the 1000 m
depth contour. However, nearer the coast. such a means of travel does
appear to take place (Deelder, 1952, 1958, I960b; Creutzberg, 1961).
Catches both near the surface and deeper down showed that, at high
tide during the night, glass eels occurred at all water levels, but that at
Timt of d<ly
Fig. S8 Glass eel catches from plankton nets in the Texel current in February/March
1957. These data indicate the concentration of animals near the surface and in
deeper waters, at high and low tide, and during the day and night (modified
after Creutzberg, l¢il ).
low tide only a few isolated individuals were to be found (Fig. 58).
During the day, practically no glass eels were caught in plankton nets
at low tide, but at high tide considerable numbers were caught, though
only in deeper water. This observation, which has been made many
times by a large number of people. leaves one with no option but to
conclude that glass eels take advantage of tidal movements to make their
way inshore. Since they are not caught in plankton nets at low tide,
one can assume that they remain on the bottom, where they are protected
from very strong currents; despite the backflow of water from the tide,
glass eels may even make their way further towards the shore by actively
swimming against the current. Lastly, Fig. 58 shows that glass eels
exhibit negative phototaxis and therefore, during the day, they remain
in deeper waters where the light intensity is lowest.
3.1.2 The movement into freshwater

Creutzberg ( 1961 ) has demonstrated experimentally that glass eels
show a preference for natural, inland, surface water but are not attracted
to freshwater that has been passed through a charcoal filter. Natural
odours are removed from water that has been treated in this way, thus
causing it to lose its attractiveness to the eels (see p. 77). When eels
encounter seawater which is mixed with natural freshwater from rivers
and lakes, they become rheotactic. According to Creutzberg (1961),
this causes the eels to swim against the ebb current (i.e. towards the
source of inland water), whereas at high tide, with its increasing salt-
water influence, the eels yield to the current and allow themselves to be
carried along with it. On the basis of data obtained in the upper tidal
area, Tesch ( 1965) suggests that these changes in behaviour pattern
during the tidal cycle might possibly be partiy the result of some internal
rhytlun, because all the factors which might be responsible for releasing
behavioural changes (such as the influence of freshwater, changes in
turbulence, and overcast conditions) do not always coincide with high
or low tide. These factors may well act as supplementary pacemakers
and provide the necessary synchronization.
As more recent investigations have shown (Schulz, 1975), a distinc-
tion must be made between the effect of freshwater as a medium provid-
ing stimulation on the one hand, and its properties as an environmental
medium on the other. Glass eels also prefer natural freshwater when
given the choice between it and tapwater. It is essential that the eels are
free to leave environmental media of differing salinity (IS parts per
thousand and 36 parts per thousand), i.e. they must have the time and
the opportUnity to move between the various media in order to be able
to make a choice between them. The glass eel's preference for fresh-
water is ascribed to the lowered concentration of the animal's internal
medium; body fluid concentration is higher in young yellow eels
(Parry, 19(6) and they preferred brakish water with a salinity of IS parts
140 THE EEL
per thousand (originating from the Lower Elbe drainage) (Schulz, 1975).
The actual movement into inland waters has been studied closely in
an area where there are no physical obstacles to prevent the glass eels
from being carried, in the way described, in the direction of the inland
waters. Such conditions are to be found in the river Elbe which, with
its funnel-shaped estuary, shows no clear-cut transition between
gradually decreasing salt concentration and freshwater. A long-tenn
investigation was carried out in this funnel-shaped area ; the number of
eels caught in a young fish trawl at various points in time was used as a
measure of the concentration of grass eels (Tesch, 1971). The results of
these catches show that in March and April the concentration of glass
eels per haul increases from 1-6 in the sea to 15--60 in the estuary and
then showed a sharp drop in most cases. The greatest and most sudden
declines are to be seen where the salt concentration in the river is still
low (1-2 parts per thousand), i.e. in places which are not far from the
barrier of freshwater. Here, the lower the temperature, the more
sudden was the decline in glass eel concentration.
The glass eel invasion has been observed at a place on the Dutch
coast where there is a much more sudden transition from salt or brackish
water to freshwater than is the case on the Elbe. At the last dike on the
Ijsselmeer, freshwater is let into the Wattenmeer through sluice
gates. Thus there is a very sudden transition from sea to freshwater
conditions. The glass eels arrive here when the temperatures are still
very low, often as low as 4'5 °C (Deelder, 1952, 1970); if after their
arrival there is a sudden drop in temperature, they naturally experience
temperatures even lower than 4'5 0c. Before the glass eels penetrate into
freshwater, they must first undergo a certain physiologiCal adjustment.
To this end the glass eels remain for a time in the s-altwater outside the
sluice gates. Aquarium experiments have shown that these eels avoid
freshwater during this period ; unlike eels which had been captured
while they were ascending into freshwater, they did not swim into an
inflow of freshwater entering an aquarium.
The glass eels outside the sluice gates of the Ijsselmeer later begin to
gather together into shoals. This behaviour indicates the beginning of
their readiness to ascend into freshwater. Various studies have shown
that this phase begins at temperatures around 6-8 °C (Deelder, 1952;
Creutzberg, 1961 f Tesch, 1971 ). At this time too,-glass eels show a
certain degree of negative phototaxis. This tendency was revealed by
yields from lift nets near the sluice at Den Dever where there is also
an abrupt transition to freshwater (Fig. 59). Here, glass eels have only
been caught when they were in open waters, i.e. when they are fairly
2S. 111.- 7.IV.
a.lv -21. IV
12 IV. - S.V.
6.V -19.V
Fig. S9 Glass eel catches (as a percentage of the total calch) at various times of the day
over the period 1938- 49 from Den Oever sluice on the closing dike of the
Ijsselmeer (after Deelder, 1952, 1970).
active. According to Fig. 59 greatest activity is seen during the hours of

darkness; it begins as the sun is setting, increases until just after mid-
night and then starts to decline again at dawn . These changes in activity,
which appear related to darkness, have been verified experimentally
by van Heusden ( 1943, from Deelder, 1970).
Such negative phototaxis has also been observed on the Ems near
Herorwn, about 70 km above the freshwater-marine boundary
(Meyer & Ktihl, 1952!53). There, with the use of a weir, the glass eel
ascent has been exploited commercially to obtain eels for stocking
purposes. Over a period of many years, observations have shown that
the eels also make use of flood-tides in freshwaters in order to get further
upstream. Hourly checks on the numbers of eels caught indicate that
in May the migration begins at about 2 1.00 hrs when the flood begins
THE EEL
to flow, reaches its maximum at high water, and is over one or two
hours later; the glass eels then disappear from the surface waters
where, until that time, it is possible to see a wide band of eels measuring
2-5 m in width on its nocturnal migration up river (Kuhl, 1955). The
largest catches are recorded when high water is at midnight. If low
water is around midnight, e.g. from 21.00 to 03.00 hrs, no eels are seen
ascending the river, and commercial fishing ceases. From these observa-
tions it is possible to predict the time of the main migration.
A I4-day rhythm emerges from the probable connection between
darkness and high water; this results in maximum surface ascents at the
change of the moon. It should also be noted here that maximum ascents
also coincide with the spring tides. In addition to the factors already
mentioned, the connection between ascents and the stronger currents
of the spring tide should also be taken into consideration.
Temperature is an important factor when attempting to estimate the
beginning of glass eel ascent. As already mentioned, glass eels start
to cross the boundary between salt- and freshwater when the tempera-
ture reaches 6-8 °C; they can be expected in the freshwaters of the
river Ems at temperatures of 9-1 I °C (Schmeidler, 1963). Ascending
Japanese eels, A. japonica, are first caught regularly in rivers when
temperatures no longer drop below 8-10 °C (Matsui, 1952).
Observations in the estuary area of the river Bann (in Northern
Ireland), where glass eels are caught commercially and transported
to Lough Neagh, have provided a certain degree of support for tidally
determined periods of maximum ascent (Menzies, 1936; Lowe, 1951,
1952). Here, and in the British rivers Severn and Leven, however, the
eels are reported to ascend both by day and by night, although commer-
cial fishing used only to take place at night because of labour shortages
(Lowe, 1951; Lubbert, 1910). In addition, reports from the river
Severn in south-west England indicate that eels only migrate near the
water surface at low tide; indeed, they swim along very dose to the
banks (see also Fischer & Lubbert, 1908). This does not, of course,
exclude the possibility that eels might move in the flood current amidst
the inflowing water. In fact, data from Fischer & Lubbert ( 1908) point
to just such an assumption; these authors report that the two-foot
high tidal wave in the rivet Severn near Epney surges upriver with
considerable force, but only lasts an hour, and then only during the
spring tides. The glass eels are caught during this period, i.e. once a
fortnight, for three days before and three days after the highest spring
tides. Fishing with dip nets takes place while the glass eels are 'swarming
upstream against the current' - i.e. at low tide which lasts more than
eleven hours there. If no glass eels are caught at neap tide this probably
means that the flood current, which normally reaches Epney, must have
died out before then. The flood current carries large numbers of glass
eels a long way upstream. When low tide begins, they continue their
own active migration along the banks.
Tidal dependence has also been seen in A.japonica (Fig. 60). In the
river studied, glass eel ascent was coupled with the period directly
following sunset. However, the glass eels only become active if high
water coincides with this period. In the river Ems the daily ascent
maximum occurs late in the day and later in the year j while migration
at high tide seems to be over in the Ems, it is often only just reaching its
peak in the Japanese river and may even have two or three hours more
to go. Not only in Japan but also in the European rivers Ems and Severn,
and, to a certain extent in the river Bann, the largest ascents take place
during the spring tide. The causes of these differences are probably
determined by different hydrological and geomorphological conditions
in the rivers, as well as by the distance penetrated inland.
The latter factor gives an indication of how the migration is progress-
ing and of possible differences in behaviour. The progress of the migra-
tion can also be measured by the degree of pigmentation seen in the
elvers. After certain physiological changes have taken place the glass
eel begins to show signs of pigmentation (see p. 15) as the temperatures
increase (Strubberg, 1913). Unpigmented eels have been found outside
the Dutch sluices between the Wattenmeer and the inland waters, and
in the brackish water of the Lower Elbe. They were active mostly at
night or remained in the lower depths. In the Elbe above Hamburg the
author (unpublished) has found elvers which were mostly semi-
pigmented. Five individuals out of a sample of 16 had reached pigmenta-
tion Stage VI A II~. The other stages, ranging from VI A 114 to VI A
IV 2, were represented in smaller numbers (see Strubberg, 1913; see
also p. 16). These eels, which were caught almost 100 km upriver in
the Elbe, were much more advanced developmentally than those seen
in the estuary of the Elbe. Other types of behaviour are thus also to be
expected (for details on migration under the surface and possible
rhythmical appearances, see Tesch, 1965 j Leich, 1929 and LOwe, 1930).
The upstream migration of glass eels by day in a visible band-like
shoal along the banks has been observed in many areas (Tesch, 1965),
not only in rivers along the coasts of the Atlantic, near the North Sea and
in the Mediterranean, but also on the Baltic coastline (Herold, 1933).
On the Atlantic coasts of France the band may be as much as 4 m wide
and may extend to a depth of 5 m (Wurtz-Arlet, 1961) . In the rivers of
144 THE EEL
" 16.2. MlojIwatfl' 19.10
•,
"
20
II II 21
"
10
14.2 ~WQt ... l1.lS
"
, ,
21 18
" 20 21 22
"
10
~
,•~
" " 20 21 II
"• I•
U . H;';' wet... 18.S0
,
;;
" •
" H.
High wattr 21.05 ~
10
" " lO 21 II
2-'. HiQII wallr 18.10
20
,i
•
" " "

10
A " " 20 21 U 2:5
I~
,..
FIg. 60 The times of glass eel ascent up a Japanese river as rdated to high water and
dusk (modified after Matsui, 1952).
Germany, however, the band is seldom more than a few centimetres or

decimetres wide and only measures about 5 em in depth (Leich, 1929;
LOwe, 1930). Deelder (1970) believes that distance from the bank is
not maintained by visual cues because these migrations also take place
by night. Rather, it seems that the proximity of the bank is detected
through the lateral line sense organs.
Surface migration is seen far inland; in Holland it can still be observed
120 km from the coast, in areas where tidal influences reach a long way
upriver (Deelder, 1970) , and on the Elbe it is known as far inland as
Lauenburg, i.e. 150 km above the Elbe estuary. There, however,
daytime ascent is limited to the uppermost 30 or 40 km, i.e. to the
area above the influence of the flood tide (Tesch, 1965).
As a result of both scientific and economic interests (see p. 150), the
extent of glass eel invasion has been recorded in a number of places
over a period of several years. Particularly comprehensive statistics
are available from areas where glass eels have been used as food and
for re-stocking purposes. With the help of data from Chiappi (1934),
Bertin (1956) has used such ascent statistics from the coast of Italy
near Rome to construct a six-year cycle of invasion maxima and minima.
While Chiappi favours the hypothesis that meteorological factors on the
mainland of Italy are responsible for this periodicity, Bertin (1956),
championed by Gandolfi-Hornyold, supports a theory of oceanic,
meteorological and hydrological influences. In Fig. 61 glass eel statistics
from different points of ascent are compared. It can be seen that there
is a certain degree of similarity in the ascent pattern of glass eels in the
river Ems (in Germany) and the river Bann (in Ireland). Not far from
the Ems, at the last Dutch dike, the figures show trends in direct
contrast with those from the Ems and the Bann. For many years the
number of glass eels along the coast of Denmark has also been seen to
differ from all the other areas observed in the North Sea and the
Atlantic ; data from Chiappi ( 1934) also suggest that the situation in
Denmark differs from that along the coast of Italy. In connection with
this, Herrmann's (1957) observations seem worth mentioning. He
points out that in 1957, on the Baltic coast of Germany, the glass eel
invasion was well above average, and compares this with the minimal
ascent along the coasts of the southern North Sea area. Herrmann
suggests that this could be brought about by a stronger than average
northerly drift in the Atlantic resulting in more eels being carried to the
northern part of the North Sea and thus to the entrances of the Baltic as
well. In 1957 the glass eel ascent was also above average in Denmark.
From this point of view, then, perhaps more attention should be paid
THE EEL
100 &I.., HlH~tc,,"011 tho ...."t of Otft....r1I

r,... ....t ......lionfrono tbo 11·,....._1
• • • • .
19l5
'"'' 1940 , ~,
1950
mGlG1I HI onch 11ft, Romt

Un nIIMlonlr
0 Glau HI catchln lilt AI .... knn
I NotU\fr,. 11'1111li0ii1 lin .. ~l ionI1
aU,. ..illloMl
~1G1S"'C<ltthilltM .;Vtr Eml
I Glauttl eotc!o MG' D.,. Otvlr

1I1001alloll) lintllauHIIoIII)
50
i
i
!
,i
. .
1925 1930 IUS ,~ ,
1950 "55
''''' 1965
Fig, 61 Glass eel ascent at various places in Europe : Denmark, Maccaresa ncar
Rome, the river Bann in Nonhem Ireland, the river Ems in nonh-west
Germany, Den Dever in Holland (compiled from data by Chiappi, 1934 ;
Jensen, 1961; Wiehr, 1967; Decider, 1970),
in Denmark to correlations between the nature of the west wind, glass

eel invasions and later eel yields (see p, 246).
While pigmentation continues to increase gradually, the length of
the metamorphosing elvers continues to decrease until a certain point is
reached (Fig. 57). Aquarium experiments by the author (unpublished)
have shown that eels, which were at Stage V B on 4th April and which,
for the most part, had reached Stage VI A 1v t by 31st July, underwent a
reduction in length of o's em (from 7'5 em to 7'0 em); after the end of
June, Le. after Stage VI A w :, there was almost no decrease in length,
This reduction in length took place regardless of whether or not the
600
500 ,
, , ,
, , ,
20' ,
,
I ,, , I , I , , ".
" VB VIAl VI All I
P~mentQt oo
VI Alii
stages
VI A IV I
Fig, 61 Weight change in glass eels from Lake TUnis, correlated with increasing
pigmentation according to Strubberg's (1913) stages of pigmentation (modi-
fied after Heldt & Heldt, 1929a).
animals received food, and neither did the degree of illumination appear
to have any critical effect on length reduction or increase in pigmenta-
tion. Pigmentation, however, was retarded slightly by keeping the eels
in darkness or by providing a bed of sand under which they could hide.
Temperature, on the other hand, is important in controlling pigmenta-
tion and in length reduction (and thus, later growth) (Strubberg, 1913).
Research into the processes of body reduction (measured in grammes)
and pigmentation in non-captive eels confirms findings from laboratory
experiments (Fig. 62). ]n Lake Tunis weight reduction also ceases at
Stage V] A 1lI 2 ; it is then followed by a marked increase in length.
]f one compares the average lengths of elvers in different regions
(Fig. 57), it is clear that the mean value of samples from the Atlantic
(Severn, Bilbao) and the North Sea is higher than that of samples from
the Mediterranean at almost all times during the period of ascent.
A comparison of weights reveals similar, though not such marked
differences (Heldt & Heldt, 1929b). ]n addition there appear to be
differences in length and weight between eels from the western and
eastern regions of the Mediterranean. Eels from the Balearic ]slands,
THE EEL
Italy and Tunis are considerably longer and heavier than those from
the Nile (see also Lubben, 1930). It is possible that during their journey
through the Mediterranean, where the temperatures are higher, a
greater reduction in length takes place during and after metamorphosis
than occurs in glass eels moving along the Atlantic coasts. Within the
Mediterranean the reduction is more marked the further the eels
migrate or are carried eastwards. Differences in length similar to those in
the European eel have also been seen in the American eel (Vladykov,
1966). In North America the average length of glass eels increased by
6 mm from south to north. Again the question arises whether higher
temperatures in the south bring about an earlier and larger reduction in
length than do the lower temperatures in northern regions. In order to
investigate the question one would have to compare length changes in
eels from various regions of North America at the same time of year, in
the same way as this has been done in Fig. 57.
The first intake of food by metamorphosing eels occurs much sooner
than the cessation of length and weight reductions might lead one to
expect. Almost without exception Eichelbaum (1924) found that
'glassy' eels still had empty intestines; intestinal contents were found in
about a third of those animals with 'light grey' pigmentation. The
majority of the more strongly pigmented eels studied were found to
have food in their stomachs. Studies on elvers during their ascent of the
Geesthacht weir on the river Elbe above Hamburg (on 17th May 1966)
showed that until Stage VI A 1112 (light grey) not one of the eels had food
in its gut (Tesch, unpublished). It was only in the latter stages that the
fishes appeared to have eaten anything. Five Stage VI A IV 1 eels all had
some food in the gut but only one of these specimens contained food. in
any significant quantity. Results of an aquarium study where eels were
provided with newly hatched larvae of the brine shrimp Artemia
salina showed that food. was already being taken shortly after the end of
Stage V b (glassy) (Table 9), despite the fact that a reduction in length
continued up to Stage VI A 1112 - as was mentioned above. In the early
stages (VI AI ) the amount of food consumed is very small and only
begins to increase with further development.
I t is interesting to note that soon after the first intake of food, the
glass eel will eat practically any living animal that is small enough to be
ingested. Experiments with captive elvers have shown that they will eat
the benthic Tum/ex as well as such planktonic organisms as Artemia
and Daphnia. On the coasts of the North Sea and the Baltic the elvers'
prey consists, predominantly, of Mysis, ldothea, copepods and chirono-
mid larvae, although polychaetes, oligochaetes, amphipods, Asellus
Table, Stages of glass eels (after S trubberg, 19 13) which took food during an aquarium experiment in 1966 and t967. T he eels were fed ..
0
on newly hatched A nemia larvae. Temperature 1SoC. Figures to the left indicate the number of animals studied at each develop· ~
mental stage, and figures to t he right (italicized) indicate the number of eels with stomach contents (after Tesch. unpublished)
...,
...>
Sfages
<
'">
Do" VB VI A , VI A VI A VI A VI A VIA III, VI A III> VI A JJJ, VI A VI A W, VI A VIA VI " T otal %
'" "- .. - . - ,", ,". ,"- ...m
10 ....67 , 0 6 3 3' 7 '3
'4.4. 67 '0 4 0
11.4.67 4
"3 ,J • ,7 0 , 0,
'7
0 0 , 0 6
•• "53 "...00
1·5·67 37
'7.5. 67 •
•0 , ,• ,33 ,, , , ,,
6 3 J
'.
"17 17 100
19.5.67 0 , 0 , 0 3 3 3 0 0 0 0 '7 , "..,>
19·6.67 , 0 3 7 16 II " Z
, 0• 0 0 0 0
13.7.67
• ,, , , ,, 4
,• , '3 4 3'
3 1.7.67 0 7 7 , 0 33 ,8 to
."
" " " m
T~I , 0 47 .J '3 7 6 8 7 9 17 10 '7 7 4 , 3 179 78 44 :c
0 " •" , •
,. ,0 80 " 53 • 7' 47 4' ,0 33 >
1·4·- 8.6.1'}66 40 19 29 19 29 8 J , 8 <
49 9 9 ' 7 17 3 ,,
'""• 0 "
", " ".,
45 '3
" " " '" ' 00 ' 00
'. '00 ' 00
'. ' 00
" ,
' 00 ' 00 ' 00 ' 00 '00 0
.' "'"
-
;ll
150 THE EEL
and a selection of insect larvae are also taken, but in smaller quantities
(Eichelbawn, 1924); in earlier stages detritus was also fOl1I1d in the gut.
Thus, even with its very first intake of food, the eel displays omnivorous
tendencies.
3.1.3 Human intervention

In various parts of Europe and Japan the massive glass eel invasion
has been put to commercial use. It must be mentioned here that only
recently have glass eels been caught for re-stocking purposes. Originally,
in areas with a particularly large surplus of eels, e,g. on the coasts of
the Bay of Biscay and in the river Severn in south-west England, glass
eels were used as a food stuff for both human and animal consumption.
Nowadays, in Spain and in Latin American states, glass eels are
treated as an inexpensive speciality, e.g. they are served in soup with
generous amounts of garlic. In the fishing regions of France glass eels
are prepared as a sort of cake.
Previously, however, the use of glass eels as food was negligible and
'the majority was used as fodder, particularly for pigs, or for manure'
(Wurtz-Arlet, 1961 ). In 1924 12-15 lOones of eel fry were caught daily
on an 8 km stretch of the Loire River and whole farming families would
take part in eel trapping. Indeed, France exported 750 tonnes of eel
fry to Spain. Gandolfi-Hornyold (1936) reports that around 1930 the
glass eel market in St. Sebastian, on the Franco-Spanjsh border,
annually sold 237 tonnes of glass eels, 90% of which came from France.
The highest yield for the year was 494 tonnes in 1922/ 1923, of which
250 IOnnes were caught in February alone. In Spain glass eels were used
in the manufacture of preserves and the production of glue. In light of
such figures as those mentioned above, the yields from the river Ems
are very modest (Fig. 61 ); at most, they amounted to about 6 tonnes
(1960, 1963) (Wiehe, 1967; see also Meyer, 1951; Schmeidler, 1957,
1963) and were often no more than a IOnne (Fig. 61 ). The quantity of
eels imported from France by the traditional eel fry distributors, the
'Deutscher Fischereiverband', has increased to over 10 tonnes. Smaller
quantities were imported from Denmark and England and were taken
to Hamburg for distribution. The annual yield of the former German
eel fry station in Epney on the river Severn amounted to 2 tonnes. Bet-
ween 1938 and 1943 about 10 toones of glass eels were caught annually
on the river Bannin Northern Ireland (see p. 151 ) (Fig. 61 ) (Lowe, 1951).
In 1965 and 1966, 13 IOnnes of eels were distributed from Hamburg
and transported to many countries in Europe; the majority went to
Fig. 63 Weighing glass eels ready for transport on gauze-covered frames (Photograph:
Witters).
central Germany with its relatively large number of lakes. In addition

to this, Europe had other customers, in particular Poland and the Soviet
Union. In such countries as these, with large expanses of inland waters,
the demand is naturally very high. Nevertheless, the quantity of glass
eels sent from England to the Soviet Union since 1956 is small in
comparison; it amounts to 2'3 tonnes (Orlov, 1966) and before 1939 it
was only 100 kg annually (Kochnenko, 1958).
Most of the fishing along the banks of rivers is carried out with dip
nets, stow nets, or spade-like hand nets (Fischer & Liibbert, 1908;
Wurtz-Arlet, 1961). In France, flares were often used as well, and
the catch per fisherman per night was between 20 and 30 kg. Nowadays,
motor boats with nets suspended from them are used in glass eel fishing
on the Loire (Wurtz-Arlet, 1961); the total catch per trip is 100 kg.
On the river Bann in Northern Ireland, glass eels are caught in an
automatic trap (Fig. 64). This trap is similar to a device used to catch
pigmented young eels on the river Elde, a tributary of the Elbe near
Domitz (Gollub, 1959). A similar device has recently been introduced
on the river Ems near Herbrum for catching the stream of eels swimming
along the left bank. The ascent channel, however, is not lined with
straw or brushwood as is common in eel ladders in Germany, but
with man-made bristles. In some areas ascent channels for elvers
are also being equipped in a similar way (see p. 164).
A trap used in Japan for catching glass eels and elvers resembles
the fyke nets (described on p. 277) used for catching eels mature enough
for marketing; it was given a trial for catching the American eel on
15 2 THE EEL
t wterltvtt
fft\~5tl;~~#:
:.~ - Eel COIltQillf!"
~ ..
w ttr l_1
Fig. 64 Schematic representation of a glass eel trap on the lowest weir of the river
Bann near Colerain (Northern Ireland) (from T esch, 1967c, after Menzies,
1936).
the St. John River, Florida, but had linle success. There, a lift net
is more suitable; this is an elver trap, also based on the fyke net princi-
ple, and commonly used in the State of Maine, USA. The funnels of
this box-like elver trap consist of vertical walls and open downstream,
as do the funnels in the fyke net-like trap mentioned earlier. The lift
net is erected in shallow waters near river banks along which the stream
of glass eels swims when migrating upriver (Sheldon, 1974).
As early as 1908 a method was being used to transport eels which is
still used widely nowadays (Fischer & Liibbert, 1908). The eels are
carried 'dry', a technique which has proved to be physiologically
advantageous (see p. 23). In 'dry' transport, the glass eels are first spread
out on gauze-covered frames, each frame holding about 2 kg of eels
(Fig. 63). The frames are then packed in a box, ten frames being piled
one on top of the other. The uppermost frame is covered with ice so
that the glass eels remain cool and damp for the whole journey. Using
this method, glass eels can survive trips by lorry over the whole of
Europe. If the eels are to be transported by train only five frames are put
in each box. For air transport, lighweight polystyrene containers are
used; in 1973, 292 tonnes of glass eels were flown from France to Japan
(Forest, 1974). In addition to 'dry' carriage, large numbers of eels are
transported in special types of lorry which are equipped with ventilated
fish containers. This type of transport is particularly popular when
large numbers of eels are involved, e.g. for carrying eels from the west
coast of France, or from the river Ems for distribution in Hamburg.
There is now a good deal of opposition to transporting glass eels from
France as early as February or March and taking them to central and
POST-LARVAL ECOLOGY AND BEHAVIOUR IS3
eastern European countries where wintery conditions are still prevalent
at this time (Koops, 1968; Gedymin & Gottwald, 1968). Losses of up
to 10% occurred during transporlation to Poland, probably because
of the great length of the journey; experiments have shown that the drop
in temperature during transport was certainly not the decisive factor.
However, it is doubtful whether glass eels are able to survive for any
length of time in the wintery waters, particularly in eastern Europe.
For example, glass eels placed in running water at temperatures near
freezing are more likely to drift downstream than to journey on upstream
against the current. Therefore, recipients of glass eels in Poland have
been advised to keep the animals for at least three weeks before setting
them free. Appropriate experiments have shown that it is possible to do
this without further losses.
The distribution of glass eels in Japan gives rise to even greater
problems than it does in Europe. The huge numbers of eels needed
for pond culture has resulted in the glass eel becoming the basic element
in the Japanese eel industry. It is easier to understand the need for
young eels when one remembers that, despite its size, Japan produces
more eels for consumption than does the whole of Europe (Table 2S ).
In addition, the fry must be taken from a population of eels that is
much less widespread than the European eel - i.e. the demand for
A. japonica must be met from a considerably smaller initial supply.
The prices paid for glass eels are correspondingly high. While I kg of
glass eels cost 30DM in Germany in 1972 (soDM in 1976), the same
quantity cost much more in Japan, £12 to £18 sterling in 1973 (Forrest,
1974). Thus, excitement runs high among the Japanese people when
the glass eels begin to ascend, and the amount of time and effort put
into fishing is correspondingly great. In order to meet requirements the
Japanese have even gone as far afield as France to purchase some of the
latter's surplus supply of A. anguilla glass eels for transport to Japan.
The elver ascent, as well as the lives of many other fishes, has been
adversely affected by the by-products of civilization. A cautionary
example is provided by the river Weser with its many weirs which
restrict the ascent of glass eels as early as the tidal region. Wurtz-Arlet
( 1961) reports that in France, since the introduction of dikes in the
Seine, glass eels no longer ascend the river in large numbers. In the
Somme, which produced 2000000 elvers before 1914 and which
provided some SOO 000 in 1926, the elver fishery has been non-existent
for the last 20 years. Nevertheless, supplies from the west coast of
France appear to be so limitless that, for the moment, there is no
shortage of glass eels for stocking programmes.
154 THE EEL
Sewage probably only has an adverse effect on elver ascent if it is

present in high concentrations. I t was found that sewage near the estuary
of the river Schwentine flowing into the Baltic did not prevent glass eels
from penetrating into the river (Neubaur, 1933).
3.2 Migration of pigmented young eels
3.2.1 Dependence on environmental factors

'Young eels', postelvers or 'snigs' may he fully pigmented by the time
they are 7-8 em long, and are more widely known than the unpigmented
glass eels. The 'young eel' period of development is said to have begun
when the eel reaches the pigmented Stage VI B in its ontogeny (see
p. IS). Infonnation from fishermen on the Elbe above Lauenburg
indicates that a surface migration like that described in the previous
section is no longer to be seen in this part of the river, i.e. 150 km from
the estuary. Young eels only occasionally are seen near the surface.
In fact, many eels have already withdrawn from the surface levels long
before this region of the river is reached. If the right temperatures
prevail, it is clear that elvers will continue to develop, even if they are
held back for some reason or other in coastal areas (Suubberg, 1913).
The eel's rheotactic migratory tendencies are retained for some time;
studies on the age and size of such detained eels indicate that the
migratory urge may continue for several years.
Research on over 8000 eels from the weir at Geesthacht, which is
still within the tidal region of the Elhe, has shown (Fig. 66; Penaz &
Tesch, 1970) that only small numbers of eels migrating upriver are
more than 25-30 em in length (see also Mann, 1961, 1963; Tesch,
1967b). Isolated examples of eels up to 40 em in length also occur.
Research at four weirs situated as far as 200 km above the estuary of the
Weser indicate that eels longer than 35 em are unconunon (Tesch,
1966). Opuszynski (1965) reports similar findings after studies at
Masuren, in the drainage basin of the mid-Baltic. Here, the migratory
eels were, at most, 35 em long, and the majority was no longer than
25 em. In research on age and length in eels, Gemzl$e (1906),
who caught young eels in various places in Denmark, also showed
that most young eels are less than 25 em long. Using electtofishing
techniques, Larsen ( 1972) caught young eels in two streams on the
Danish island of Zealand and found that 78-86% were between 9 and
24 cm long.
Observations on the river Elbe (Tesch, 1965) demonstrate that the
POST-LARVAL ECOLOGY AND BEHAVIO U R [55
, ,
4meirtSobo'/,
"
nonll(ll "",I
~ Anguil/Q ongvil/o
• l.o~tro
Septombtr ..
,,
Fig. 6S Daily ascent figures for pigmented young eels and lampreys (Lampetra
f/uviatilis) at the eel pass on the Geesthacht weir (river Elbe), with details of
water temperature, high tide levels, lunar phases and alterations in the weir's
outflow. No control checks were made on the days indicated with a cross
(after Tesch, 1967d).
elver is able to migrate at least 150 kIn upstream before it is fully pig-
mented. Once pigmentation is complete the eel probably manages to
travel considerably further in the same year. Observations on A . nebu-
[osa (which occurs in India) indicate that in one year from the glass eel
stage, young eels are capable of migrating 650 kIn up the river Godavari
(Ibrahim, 1961 ). This is an amazing feat, especially when one takes into
consideration the fact that during this time the eels also increase in
length by 5-15 em. Of course, the distance travelled is considerably
less if migration is hindered by obstructions such as weirs which lack
156 THE EEL
I:
Electrofishi ng duril'Mj doyligM
~l
", -
67 19"
.=9B
o , , iIDID ~Q , "" ,n ., .,
."1 ML " "~
30510 152025 30'5 """ 0
I
Day-ti"" o~nt
17_8 .\Q6~ S.7.1967 10
, ,
lI,OO -tSOO 10.45-21.00
n .. 151 n"'724
o ,5 1015202530
" ,n ~"':''''':'l+'l':'"",~"C7""~:::r'''''-",",-,;n-,,,
5 10 1520Z530J54(J4530 55
0
•
20
~l"
Dey-ti'" oscent
18.S.1964 0.7.1907
"
O.800~lS,40 08.45-16.30
0 "'52 "=627
,"
:1 ,
• " " " " , 20
" " "
.,
Fig. 66 Length frequency distribution of young eels caught by day and night in the
eel pass on the Geesthacht weir (river Elbe), compared with the length
frequency distribution of young eels caught using electrofishing techniques
below the weir (Tesch, unpublished).
proper facilities for aiding the eel's ascent. For example, on the fifth
weir of the river Weser, about 200 km upriver, the smallest eels were
only 15 em long (Tesch, 1966). At the lowest weir, only 80 km upriver,
even glass eels were caught; of course, more larger eels were caught
inunediately above the weir than below it. If one calculates the time
necessary for the eel to grow to a length of 15 em, it appears that these
animals take at least two years to reach the fifth dam (Peiiilz & Tesch,
1970). In this time, then, they cover little more than 100 kIn. However, it
is also possible that only larger eels are capable of overcoming this
stretch of river which has so many dams, and that some of them may
have been able to cover the same distance much more quickly.
In comparison with other species it seems that the upstream migration
of young eels is a slow process, as is indicated by the fact that individuals,
which have already spent two or more years in freshwater, can still be
found in the lower reaches of a river. In other words, young eels do not
show such intensive or 'single-minded' migration as do, for example,
salmon, lampreys and many other anadromous species.
Experimental techniques have made it possible to compare the
migratory behaviour of young eels with that of adult lampreys ( Lampetra
fluviatilis ) which swim upriver with some determination; the results of
this experiment are reproduced in Fig. 65. The means of ascent was
provided by an eel pass (complete with an eel trap) built very close to one
of the weir sections. If this section of the weir was shut off, the strong
flow of water near the opening of the eel ladder ceased, the only flow of
water remaining being that through the eel ladder itself. When this
happened, the ascent of lampreys stopped almost completely. The eels,
however, continued to migrate upriver. This finding must be interpreted
in the following way: like many species offish (Fries & Tesch, Ig65b),
lampreys swim upriver along the margin of the main current, using it as
an orientation guide. Thus they manage to reach higher regions of the
river without detour or hesitation. Pigmented eels, on the other hand,
do not make use of the immediate edge of the main current. In other
words, they simply continue swimming even if the current is reduced
or ceases completely. As a result, they often find themselves in back-
waters and only relocate the main current, and with it the path upstream,
after some delay.
F. Schiemenz (1950, 1952) has shown from his investigations that
glass eels display strong rheotactic responses. However, he adds the
following remarks: 'If the speed of the current continues to decrease,
after a certain point orientation by means of the current ceases; at even
lower speeds or in still waters, the eel swims around haphazardly until it
relocates the current, after which it follows the line of the current again.
Thus, if the current becomes too weak, the eel will seek out a current
which corresponds to the intensity of its migratory drive.' The glass
eel migrates at a speed of 5 em/sec (10-15 em/sec according to Deelder,
1970). Thus, the central current in the Elbe is too strong, both for
glass eels and for the slightly older stages as well (the current is 100 cm!
sec and more in some parts). This means that young eels muSt swim
through the more sluggish regions in order to make their way upriver,
and, as already mentioned, this often results in them entering back-
waters. The current through an eel pass and the slight flow of water
through it can, however, be detected from a relatively great distance
by the eel and can elicit rheotactic responses.
The pigmented young eel does not orient itself directly by the line
158 THE EEL
Table •• The number (per hour) of young eels ascending the eel pass on the right
bank of the Elbe at Geesthacht, and the proportion of individuals which
are thought to have entered the Elbe as glass eels in the same year (after
T esch, unpublished), The figu res in brackets indicate the proportion of
older young eels (excluding glass eels)
Proportion of
Date T ime the glosJ etl R emarks
du ring
daylighl "'
night
gmera/ion of
1964 (a.! %J
22 .00- 10.)0
, '3 •
10.)0- 14·)0
•
15.00- 19. 15 3 ••
.",
)1.5.- 2 .6. 19 · I S- 2O ·5 0
20·50-08·15
7
'0 0
08 . IS-IO.)0
10 .)0--12.00 , 0
0
1l.00- 1)·55 0
,,,,
14·6.- 16.6. 22 .00-15·00
15.1)0-20.)0 ,6, (1 4)
74 ( II )
"
9'
20.)0-08.)0 '70 ( 15) 9'
,", 0)
"
25.6.-26.6. 08 .)0- 18.)0
18.)0-08.)0 " 3 (, ) '7
,",
6·7·-8·7· ' 9·00 -Q9.00
16.00- 17·45
IO.O<l--09.00
, " "0
0
09. 10- 14.00 0
,", " ,,
17·8.-18.8. 1 1 .00-18.00 ( 18) '7
18.00-08.00 '0 ( 10)
08.00- 15·40 6 (6)
06.00-20.00 ( 17)
" (22) ,3
20.00-08.00
"
.",
26.8.-28.8. 12.00-15·00
15.00-- 18.30
18·30-08.00 "
9 (7)
(9)
'9 (29)
'3
",
09.00- 12.00
12.00--15·15 ,
'3 ( 12)
(" "3
,,..
21.9.- 22.9. 10·30- 14·30
14·30-- 18.00
0 P roportion of
the I'}66 glass
18.00-08.30 '0 eel generation
08·30--12.00 < , c. 6%
22.00--10·30
'" Proportion of
'"
10·45-14·45 the 1967 glass
4.7.-6.7. 15 ·00-21.00 '067 eel generation
' 967 21. 15-04·00 ,8, was c. 50%
08·45- 11.15 76, (based on
11.30-- 16·30
'" population
studi~ using
el~trofishing
techniques)
of the bank to the degree shown by the younger stages with little or no
pigmentation; the latter, when migrating, form a light coloured band
moving along the bank. The young eel swims at a slightly greater depth
and is but rarely seen at the surface. To a certain extent its migratory
behaviour seems to be a compromise between travelling near the fast-
flowing current, and orienting by the bank where often it is difficult to
detect any current at all.
The migratory activity of pigmented young depends to a large
extent on the temperature (Mann, 1963; Tesch, 1967d; Larsen, 1972).
Young eels become active at temperatures between 8 and 9 °C, i.e. at a
similar temperature to glass eels. However, it appears that the migratory
activity of young eels is conSiderably more sensitive to changes in
temperature than is that of glass eels which, once they have begun their
ascent, are only adversely affected by decreases in temperature below
the minimum. Figure 65 shows the degree to which young eel activity
depends on temperature in comparison to activity in the lamprey
( Lamperra jiuvjatilis) , as measured by the number of individuals
arriving at the Geesthacht weir on the Elbe. Upriver spawning migration
in lampreys appears to be restricted to a narrow time span and to be
dependent on the seasons or on the periodicity of the moon, while
ascent in eels over the same period of study (18 days) rises and falls
irregularly (see also Mann, 1961, 1963). The eel's activity does not
appear to depend on the levels of the tidal waters which, in this part of
the Elbe, are strongly overshadowed by the downflow of water; indeed,
it may possibly be dependent on the amount of water that is flowing
downstream (Mann, 1961), but technical peculiarities in the trapping
methods used may have resulted in chance correlations. Migratory
activity in eels also declines with decreasing temperatures in autumn and
stops almost completely if temperatures fall below 10 0c. The strong
temperature dependence of migratory activity in young eels has also
been demonstrated experimentally (Wehrmann, 1968). Magnets were
implanted in the eels and were used in conjunction with a magnetic
field detector placed near the individual being studied (Lillelund, 1967);
simultaneous increases in water temperature (from 20 to 25 °C) and
in current (from 0 to 6·5 ern/sec) raised activity to thirteen times its
original level.
According to Mann (1961) and Rosengarten (1954), ascent takes
place mostly at night. However, as is indicated in Table 10, there is no
great difference between night and day. In fact, if in any year the
proportion of eels which arrived as glass eels is large, then ascent during
daylight hours may predominate. If the temperature drops sharply at
160 THE EEL
night, migratory activity may fall below that of the previous day, as was
observed, for example, by the author on the Elbe at the beginning of July
and the middle of August. Figure 66 shows the percentages of YOWlg
eels of different lengths caught during ascent in the eel pass at the
Geesthacht weir on the Elhe. These figures indicate that the proportion
of larger eels is relatively greater at night than it is by day. Bearing this in
mind, it is possible that the greater weight of eels caught at night
(reported by Mann, 1961) is attributable, for the most part, to the
capture of larger individuals rather than to the capture of a greater
number of eels. Thus, these studies suggest that as the young eel gets
larger it changes from a totally diurnal animal into a nocturnal one.
It has been shown experimentally that elvers begin to show activity
if the amount of light normally available is reduced from 120 lux to
9 lux. If the amount of light is increased again, from 9 to 18 lux, this
activity will cease (Wehrmann, 1968). Similar results have been obtained
using slightly larger individuals of the species A. rostrata (Bohun
& Winn, 1966). In order to provide experimental confirmation of
observations made in the field it is important to study animals at various
stages of development, i.e. differentially pigmented glass eels and
pigmented young eels of different ages.
Investigations into the length frequency of young eels in the Hamburg
harbour area of the Elbe have shown that during the summer smaller
eels (between II and 20 em) outnumber larger individuals (between 21
and 30 em) by two or three to one (Ladiges, 1936). However, in Sep-
tember this ratio changes, and equal numbers of both groups are caught.
A relative increase in the number of larger young during autumn
has been confirmed by more recent investigations (penilz & Tesch,
1970). The same trend has been observed further up the Elbe, near
Damitz (Fig. 67) (see also Larsen, 1972). Thus the view that the much
smaller young have already disappeared upriver in autumn, cannot be
upheld; if this were the case they would appear in the upper stretches
of the river. It is much more likely that in autumn, when the days are
shorter and temperatures are falling, the activity of the smaller eels
shows a greater decline than does that of the larger individuals. A similar
tendency is also seen in spring; in most years the ascending young are,
on average, larger in May and the beginning of June than are those
caught in swnmer (Penaz & Tesch, 1970).
Salinity is another environmental factor which could influence
activity in the lower reaches of rivers. As research on salinity preferences
among young yellow eels has shown, experimental animals from the
Lower Elbe regIOns prefer brackish water of 18 parts per thousand
PO ST-LARVAL ECOLOGY AND BEHA VIOUR 161
17.-216. 24.-30.6. t-S.l 9.-1H 16.-22.7, 21-29.7. ~1l-6.a. 7.-14.8. 15.-2Q.8. 2h!Sa 9a.-~9 6.-12.9 13,-19.9
~
}OO
'"
200
WHklycatchts Ntweeo the respKtov~ lunar
--- --No,af per k~
pho$I"'i
-
2<, ~II
220
200
'"', ". '",

'" - _11Q_ J~I li9_ -~-
~16_
'/"
1~_
1 4~ !~- -~
''20" / '
'", I,-
•, "
'00
00
'/
•0
\
20
, '-. V
Fig. 67 The relationship between lunar phases and the quantity (in weight) of
ascending young eels caught in an eel trap on the Elde sluice near Domitz (on
the Elbe) (after Gollub, 1959; Deutsche Fischerei-Zeitung 1959, p. 88,
Neumann Verlag, Radebeul).
salinity when they are given a choice between saltwater (36 parts per
thousand) and freshwater (Schulz, 1975). It is thus conceivable that
young eels which have completed metamorphosis to the yellow eel
stage but have not quite passed through the brackish water regions, do
not develop a further drive for continuing their upriver migration
into freshwater. This could be one reason why the extensive brackish
regions of large rivers are so densely populated by yellow eels
(see p. 56 and 256).
In addition to an increase in negative phototaxis, young eels appear
to have another characteristic which brings about a further period of
activity. Studies by Gollub (1959) at an eel trap on the Elde sluice
near Damitz revealed a period of activity which clearly ran parallel
with the phases of the moon. During the three months' period of investi-
gation, maximum catches were recorded in the last quarter of the moon
or at the new moon (Fig. 67). An increased level of activity was also
seen in free ranging animals (Mohr, 1971).
Mann (1963) was unable to observe such periodicity in fishes collected
80 kIn funher upriver from the weir at Geesthacht. He notes, however,
that: 'The only striking observation was that the maximum catches
coincided with the waning moon (28·8. to 30·8.). From this one might
conclude that the stimulating effect of an increase in temperature was
reinforced during this period.' Mann's other data on young eels migrat-
162 THE EEL
ing upriver (1961, 1963) do not point to any correlation with lunar
phases. Personal observations (Tesch, unpublished) at the same place
also do not allow any definite conclusions to be drawn.
The experimental studies mentioned above (Wehrmann, 19(8) give
support to ecological data; they point to a lunar rhythm in movement.
However, other earlier aquarium experiments (Schoeniger, 1951) did
not, unfortunately produce such evidence. Nevertheless, as silver eels
have shown activity correlated with the phases of the moon (see below,
p. 224), a lunar rhythm of activity in young eels would not be surprising,
and the question arises whether in earlier developmental stages there
may not also be a correlation with lunar periodicity as well as with tidal
rhythms (see p. 142).
3.2.2 Human involvement in the ascent of young eels

As was the case with the glass eel invasion (see p. ISO), the ascent of
pigmented young is also turned to commercial advantage by providing
new stock for fish ponds and other inland waters. The most important
young eel fishery is in the Lower Elbe region. Here the so-called 'eel fry'
are caught in eel baskets and stow nets, i.e. with virtually the same gear
used to catch eels for markets (see p. 273). In this region the proportions
of eels of different lengths are similar to those shown in Fig. 66, although
the groups of shorter eels are not represented. Eels less than 15 em in
length are not caught in the mesh of the fishing gear used; thus the
transportation of young eels is restricted to animals between IS and
30 em in length.
Frequent discussions on the value of these eels as material for re-
stocking have arisen, particularly with regard to their sexual
development and anticipated growth (Schnakenbeck, 1954; Wundsch,
1954; Rahn, 1955b, 1957a; see also p. 53). Judging from recent investiga-
tions in the Lower Elbe region (Peiu1.z & Tesch, 1970), earlier fears that
most of the eels there are males and are thus likely to migrate earlier,
appear to be unfounded (see p. 54). Furthermore, as was suggested
above, eels of this size still have rheotactic tendencies, and thus, when
they are put into waters further inland, they will generally show a
tendency to continue migrating upriver, instead of returning towards
the river esruary, as one might expect of an 'almost silver eel' (see p. 220).
Other means of catching young eels are to place eel traps in hydraulic
installations. These are often used in combination with eel ladders
(described below). The principle behind them is similar to that of the
glass eel trap on the river Bann in Northern Ireland (Fig. 64) : Fig. 68
POST-LARVAL ECOLOGY AND BE H AV I OUR
Eel tr"1lP
!
,
,
£el loddtr ape!\l0\I
,
!
, ,,
/
/: ~
~
Fig. 68 Position of an eel ladder with a box trap for elvers, on the Elde sluice near
Damitz (on the Elbe). Solid lines represent the path of the current, broken
Jines the preferred route taken by the glass eels (after Gollub, I959; Deutsche
Fischerei.Zeitung t959, p. 87 Neumann Verlag, RadebeuJ).
Fig. 69 Part of the fish pass on the Geesthacht weir on the left bank of the Elbe,
together with the adjacent eel pass. The eel pass is lined with brushwood and
covered with wire netting in order to protect the eels from predatory birds
and animals, induding the mitten crab (from Tesch, 1967b).
THE EEL
shows the lay-out of an eel trap built in this way at the outflow of the
Elde sluice at Domitz on the Elbe (see Fig. 67 for records of yields).
It seems important to ensure that the exit of the eel ladder, which leads
to the eel trap, does not open into excessively fast flowing water (see
p. 157)·
Eel populations can of course be improved by facilitating the ascent
of young eels. Their ascent is greatly impeded by structures such as
weirs and, to a lesser extent, sluices, unless some sort of eel pass is
incorporated.
The principle behind eel ladders is to make use of certain particular
characteristics of eels, i.e. not only their ability to swim but also their
ability to climb (characteristics shared by other species which are
snake-like in shape, e.g. the lamprey). Young eels, with their long,
relatively light bodies are even capable of climbing almost vertical
walls, provided that the surface is not too smooth and that the substrate
is slightly moist, The most amazing forays are made over rocks that are
slightly wet (see Meister, 1970). On the basis of srudies made on
A. d£efJenbach£ and A. australis it is thought that eels not more than
8'7 em in length and 0' 53 g in weight are physically the most capable
of overcoming vertical obstacles (Woods, 1964).
Because of these characteristics of young eels, devices to aid ascent
are easy to provide and there is no need to go to the expense of a typical
fish pass. Many fish passes are. in fact, unsuitable for young eels because
the water flow in them is too fast. As a result. additional eel ladders
have been installed next to the usual fish pass in many weirs on the
Mosel. The same alterations have been carried out on the fish pass in the
weir at Geesthacht, though here an additional eel pass is not
really necessary because the currents are very slow (Fig. 69).
Using electrofishing techniques, the author emptied the separate
compartments shown in the picrure and found that on each occasion,
young eels. including moderately pigmented elvers, were present in
every one. Current speeds of 1' 1-1'4 mlsec were measured at the junc-
tions between the compartments; the fishes must be able to cross each
of these critical points (about 20-30 em in length) if they are to reach
the next higher level. Timing of maximum swimming speeds has
shown that eels 7-10 em long swim at between 0,6 and 0'9 mlsec, while
eels 10-15 em long and larger, swim at a speed of 1'5 m/sec (Sorensen,
1951); these speeds could be maintained over a distance of 120 em.
Current speeds as great as 2 mlsec have been recorded at the critical
point in the fish pass on the Mosel at Koblenz (Rosengarten, 1954),
Nevertheless, eels manage to swim through this fish pass, but it should
be noted that they swim very close to the side walls so as to take advantage
of areas with the least current flow. Also, the stretch with the maximwn
current was only 10 em long. Young eels took twice as long to ascend this
fish pass than did cyprinids. This behaviour of young eels agrees with
the findings described above from the Elbe. Further indication of how
young eels are able to overcome obstacles, despite their relatively slow
swimming speeds, is given by observations from the lake regions of
Finland (Jarvi, 1909). Here, over a distance of at least 300 km, the
eels succeeded in ascending 13 waterfalls and rapids ranging from
1'5 to 9'5 m in height and, in one instance, managed a height of 22 m.
About 100 eels an hour pass through the Koblenz fish pass at night.
In comparison with the nwnbers of eels which negotiated the eel pass
at Geesthacht on the Elbe (Table 10), this is a large number. Under
optimal conditions the average number of young eels ascending at
Geesthacht is 500 an hour and, at the maximum, 1000 an hour. How-
ever, such figures were only seldom reached and even 170 eels an hour
was considered to be very good. Despite detailed studies by the author,
it has been impossible to establish how these figures for the eel pass
alone compare with the number of eels using the fish pass on the Elbe
at Geesthacht; techniques such as letting the waler out of the fish pass,
electrofishing, marking the young eels with neutral red (Fries, 19(5),
tetracycline and nicks cut into the fins, were of no avail. Clearly the
compartments of the fish pass are so large that an operation such as
electrofishing or emptying the water from each comparnnen(, is too
time consuming.
Although fish passes do facilitate ascent for young eels, eel ladders
should be erected especially to aid their passage. Eel ladders are
considerably cheaper to build than are fish passes.
Schiemenz (1940) gives much information on the construction of eel
ladders. The sort of devices in common use nowadays are shown in
Fig. 70. The method of construction depends, to a great extent, on the
structure of the weir and the nature of the locality. The basic principle
of an eel ladder is that it should form a link between the upper and lower
water levels such that water flows slowly down it. The flow of water is
retarded by the presence of brushwood, straw or synthetic bristles
(the latter have only recently been introduced) (Kothke, 1964b;
Schiemenz & KUhne, 1964). Eels can either swim against the reduced
flow of water or crawl upwards through the brushwood etc. Figure 71
shows a contraption which was made from brooms with synthetic
bristles; the advantage of such bristles is that they do not disintegrate
as Quickly as brushwood. According to Jens (19<)9) the number of glass
166 THE EEL
"
,
N
,
Fig. 70 Different examples of eel ladders. ( I) Sheltering holes at the base of a vertical
wall. Stones and brushwood in, and in front of, the openings help reduce the
speed of the water. (2) Tubing (at least 30 em in diameter) leading diagonally
down from the upper water level to the lower water level, lined with wood
shavings and brushwood. Bristles taken from the type of trap shown in
Fig. 71 make an even more effective lining. (3) Right: Holes (2 cm in dia-
meter) in the wall of the weir; they are positioned at 5-10 em intervals.
Matting is hung in front of these holes on the downstream side. Left: holes,
positioned as above but with a triangular box placed in front of them; this
wooden box is filled with brushwood and wood shavings. (4) If no holes are
available in the wall of the weir, a matting of twigs (two layers of plastic-
covered chicken-wire filled with wood shavings and brushwood) is placed on
the edge of the drop and anchored in place. (5) As a substitute, sacks can be
sewn together and hung over the edge of the wall, to one side of the main
current (Wilke, 1975).
eels passing upriver is increased fourfold when brushes and not fashine
are used. A structure similar to the one on the Elbe was built for the eel
pass on the weir at Bremen-Hemelingen (on the river Weser) (see also
Trager, 1954). As has already been mentioned, an eel ladder with
brushes has now been installed near Herbrum to aid the ascent of
glass eels along the left bank of the river Ems.
In Mecklenburg a larger number of smaller eel ladders has been
erected (Gollub, 1955, 1959). Control catches show that their installa-
tion produced good results. As an example one may cite an eel ladder
leading into Lake Rudower (a lake of 180 ha in the Elbe and L6cknitz
Fig. 71 Brislie linings which were first used in 1964 to replace brushwood in one of
the two tubes of the eel ladder at the Geesthacht weir on the Elbe (Photograph:
Tesch ).
168 THE EEL
drainage). The number of eels which go up this ladder in a week would

have fe-stocked the lake for a whole year) producing eel yields of
16 kg/ha. With the introduction of a ladder into Lake Dobbertin,
4000 young eels entered the lake which has extensive connections with
the Baltic. Sixty-six per cent of these eels would have been sufficient to
produce an annual yield of 6 kg/ha from the 290 ha lake. It is important
to remember that Lake Dobbertin is one of the lakes most distant from
the sea in the drainage of the river Wamow and that the majority of
young eels have dispersed over the 10400 ha drainage basin before
reaching the lake. From 75 000 to 225 000 young eels annually pass
through the Elde sluice near Domitz (Gollub, 19(3). Such numbers
would provide a quarter of the eels needed to re-stock the waters above
the sluice. There has, however, been a drop recently in the nwnber of
ascending eels, a probable consequence of the Geesthacht weir on the
Elbe which has been built since Gollub's investigation. At another
spot where young eels migrate upriver from the Baltic (Mildenitz
near Sternberg), 16000-80000 individuals have been recorded
annually; by this stage the young have already covered a considerable
part of their journey inland from the Baltic.
It is difficult to calculate just how many eels pass upriver at a key
point in the Gennan area, i.e. at the Geesthacht weir on the river Elbe.
The figures in Table 10 suggest that, on average, at least 100 young
eels ascend per hour, so the daily average must be about 2500. This
would mean the passage of 375 000 young eels during the five months'
annual ascent period. It is reasonable to asswne that similar nwnbers
use the fish pass and the eel pass on the other bank of the Elbe. In
addition, the side canal of the sluice is used by migrating eels. In good.
years swanns of glass eels have been seen here in the morning and are
let through the sluice. In other words, similar nwnbers of eels are
thought to pass through the side canal as use the eel pass on the right
bank. Thus a total of I million glass and pigmented eels can be expected
annually in the Elbe. According to some reports (Schiemenz, 1930),
between 5 and 10 million glass eels migrate annually up the large North
Sea rivers of Gennany. Statistics compiled by the German Fishing
Union indicate that in the last few years slighdy more than 5 million
eels swim up the Ems annually (see also Wiehr, 1967). It thus appears
that there has been a considerable reduction in the number of eels
ascending at Geesthacht, or that the reduction may already have
occurred in the Lower Elbe regions (the harbour at Hamburg). The
million eels that probably pass through the weir would be sufficient
to produce an average yield of 5 kgjha from a water body wi th a surface
area of 30 000 ha (see p. 265). However, the drainage basin in question
is four times that size and a re-stocking programme is therefore necessary
in this region if yields of 5 kg/ha are required.
The darruning of the St. Lawrence River in 1958 by the Moses-
Saunders hydroelectric power plant must have prevented glass eels
from entering the drainage area above it, and could thus have created a
situation like that on the Elbe. However, figures from the Canadian
fisheries for the years 1959-70 do not indicate any decrease in yields
(Hurley, 1973) and there is no evidence of an increase in the ratio of older
eels as a result of a decrease in the nwnbers of younger eels. On the
contrary, younger age groups were more plentiful than older age
groups. For example, the proportion of eels in the 90-100 em length
group decreased from 17'6% in 1964 to 2'6% in 1971 , while the 60-
70 em length group showed an increase from 14'8% to 38'2% over the
same period of time (see p. 249). Probably the young eels manage
to by-pass the barrier fonned by the Moses-Saunders plant, possibly
by using the shipping locks, as do eels which ascend the weir at Geest-
hacht on the Elbe. Thus, it appears that the flexibility of migratory
behaviour in young eels is such that these animals even manage to
overcome quite considerable diversions and hydrological changes
encountered in the course of the journeys.
3.3 The 'yellow eel' stage to the 'silver eel' stage

In general the tenn 'yellow eel' is used to describe all individuals which
have attained a certain size. Externally, yellow eels hardly differ from
pigmented migrating young eels (see p. 18) except, of course, that they
are larger. This section deals principally with eels which have completed
their migratory phase and are over 30 em long. Animals which have
reached this stage are referred to as being in the 'feeding stage'. They
are relatively sedentary and migrate only when meteorological, hydro-
logical and seasonal factors in their environment make a change of
surroundings necessary. Such changes are, however, of very limited
range. Younger developmental stages are sometimes included in the
category 'yellow eels' particularly when dealing with feeding (see also
p. [48) and growth.
3.3.[ Feeding
The period of activity and food intake
Just as the eels' activity (see p. 159) and the fishennan's catch (see
Figs. 88 & 89; Thurow, 1959; Wenner & Musick, 1975) decrease in
THE EEL
1 1 , - - - - - - - - - -- ---------,
.
""
10 - II,vtr Wtn
.... - .... Ri_ Owyfo,h
~\
I \
9 ...... « NOfth'Xo Coent I \ 90
SchlnwHj'HoIstfill J....... }.,...... .
I \
I
I
I
I
I
I
I
---. -.
I
I
I
I
I
I
I
I
2 \ lO
/"...
\
\
\
J
.-~ "
O,~IL"~~-~\~~~-~-rc-~c-~-o~~~C-+
OK. Kor "PI" Hoy Jun. Jul. .1.1>\1. Stflt Oct
Jc~ F~. Nov
o
Fig. 71. Seasonal differences in diet. Average extent to which the eel's stomachs were
filled, based on figures from twO rive:rs in Wales, the: Wen and the: Dwyfach
(from Sinha & Jones, t967b). Percentage of eels with food in the: digestive
tract, based on figures from the: brackish waters of the west coast of Schleswig-
Holstein (from Daniel, 1968).
winter, there is a similar reduction in food intake during this season.

Figure 72 gives some idea of how markedly food intake increases in
March, April and May. The increased activity of the eels searching for
food during the spring also accounts for the higher catches recorded
at that time (see p. 249). Eels clearly eat much more in the swnmer than
they do in the winter months, regardless of whether they are living in
coastal regions or in streams. What is more, higher food oonswnption
in surruner is much more pronounced in the eurythermal and near
subtropical eels than it is in other eurythermal species of fishes living in
the temperate zones.
In addition to seasonal differences in food intake there are also
variations correlated with the time of day. Aquariwn experiments by
the author have shown that eels eat considerably more at night than
during the day. This has also been observed by other workers. A large
percentage of New Zealand eels caught in the early evening have empty
gutS j traps set during the night caught animals with food in the gut
(Cairns, 1942). When fishing for A. dieffenbachi in New Zealand it has
also been shown that baited eel traps are only effective at night (Burnet,
I95Zb). The same results are obtained when baited eel baskets and lines
are used in fishing for the European eel (see p. 273). In a study of the
Indian eel, A. nebulosa nebulosa (Pantulu, 1956), all the eels used for
stomach analysis were caught in the early hours of the morning, thereby
establishing the fact that this species also feeds at night. Nevertheless,
eels do not become totally inactive when in natural daylight. If they
become accustomed to particular feeding times in an aquarium, they
will take food during the day. Anglers, and fishermen too, will confirm
that they have a certain amount of success using bait to catch eels during
daylight hours.
Investigations into the activity of American eels between 20 and
30 cm long have shown that the daily activity pattern is under the
exclusive comrol of a light-dark rhythm (Bohun & Winn, 1966). As is
the case with young eels (see p. 159), the period of greatest activity
coincides with the period of darkness. Activity in yellow eels is probably
even more closely connected with darkness than it is in younger eels,
which, however, showed increasing inactivity in light as they increased
in size.
The question now arises whether swimming activity coincides with
feeding activity. According to the studies just mentioned this in fact
appears to be the case, but research in Poland suggests that these
results should be accepted with some reservation (Opuszynski &
Leszczynski, 1967). Studies on feeding in young eels between 10 and
40 cm in length from four lakes and one river (which connected two
lakes) produced different results. On average 40-80% of the lake eels
had food in the stomach (and in these fishes the stomach was fairly
full), whereas only about 10% of eels from the river had eaten anything
at all and then only in relatively small quantities. The authors explain
this anomaly by the fact that river eels were acting rheotactically, a state
which presumably results in a marked reduction offood intake. Further-
more, only animals that are easy to catch were found in the stomachs
(e.g. Ephemeroptera and Trichoptera), while chironomids were found
in the lower part of the digestive tract. Presumably the eels caught the
chironomids in the lake they had left just before entering the river.
Seasonal variations in diet

Since eels are predominantly benthic in habit their food is derived
mainly from the zoobenthos. Depending on the nature of the water
these animals may be present in varying numbers during the year. Such
seasonal changes are reflected in the eel's diet. The organisms found
in the intestines of eels are shown in Table I I, which is arranged
THE EEL
according to the time of year. The figures show that in the English
streams studied, many more fish were eaten in the spring months and
the beginning of the summer than during late summer, autumn and
winter. This is related to the fact that most of the fishes eaten were glass
eels, 126 of them in fact. The other fish prey consisted of 15 lampreys,
10 salmonids, 4 sticklebacks and 4 flatfish. All the salmonids fOWld in the
eels' stomachs were recorded during the summer months; in this and
other streams, salmonids comprised but a small part of the eels' prey.
Mussels and snails were only eaten in one of the two streams studied,
and then mostly during the summer. Ephemeroptera nymphs, which
constitute some of the eels' most important prey animals, were also
eaten mainly in the summer, as were the Trichoptera larvae, worms,
and food taken from the 'vater surface. Nevertheless, it must be empha-
sized that insect larvae were important prey animals throughout the
year, with dipteran larvae particularly important in the winter months.
Of course, it is difficult to indicate the eel's preferred food during
winter because, as Fig. 72 shows, most had empty stomachs.
Studies carried out at various times of the year in the harbour region
of the Elbe (Hamburg) failed to reveal any particular seasonal differen-
Table 11 The proportion of eels (as a percentage) which fed on various prey animals
at different times of the year in two rivers in Wales (adapted from Sinha &
Jones, 1¢7b)
Du.' Feb.! Aprill JUflll Aug.! Oct.1

Jan. Mar. M.y July Sep. Nw.
Ritler Win
, ,S
,
fishes
Plecoptera nymphs
, "7 '7
7
Ephemeroptera nymphs
,7 '7 '3 7
,8
Trithoptera larvae 3
"
'0
"
30
"7
,"
Dipteun larvae S ·s
Coleoptera, adults & larvae .0
Crustatea
Annelids , , S , 3
.0
Surface food 7
Ritler Dwyfach
fishes 3 '7 S 3
Molluscs 3 '3
,
S
Pleooptera nymphs
Ephemeroptera nymphs ,, 53
60 3' 9
3
3
Trichoptera larvae
Dipteran larvae , 47
,0
,0
9
9
7 3
,,,
Coleoptera, adults & larvae S
Crustacea 3
Annelids
Surface food
'3 ,S
"
ces in prey or in prey frequencies (Ladiges, 1936). Here, the eel's

main prey animal is Tubifex, which normally does not show any seasonal
fluctuations in number. Variations were not detected in the other
important prey animals either, in particular in the number of Cladocera
eaten. Mitten crabs (Erioeheir sinensis) were also eaten in almost all
months; their particularly frequent occurrence in eel stomachs in
September was presumably due to chance. An important prey animal,
especially in spring, is the earthworm, which inhabits newly flooded
areas. Similar observations have been made by Micheler ( 1967) on eels
from Chiemsee, which in addition ate land snails during the spring.
Fishes (e.g. roach, Rutilus rutilus and perch, Perea jluviatilis), on the
other hand, were eaten less frequently during the spring, but increased
to 96% of the total diet in August. Eels from Lake Balaton in Hungary
showed marked seasonal differences in diet (Biro, 1974) . Asellus
aquaticus predominated from March to June, after which mud dwelling
amphipods (Corophium curvispinum ) and other amphipods or Lymno-
mysis benedeni became the most frequent prey animals.
The seasonal diet of tropical eel species is very different from that of
eels in temperate zones. Tropical eels do not cease eating nor reduce
their food intake during the winter, but nevertheless considerable
variations in diet do occur, as has been shown in the case of the Indian
eel Anguilla bengalensis (Pantulu, 1956). The marked predominance of
crab larvae in August is particularly striking; they comprise 79% of the
prey eaten. These animals occur in vast numbers at this time of year,
with the result that even fishes (Hilsa ilisha, Setipinna phasa and Pama
pama ), which in some other months form the eels' main diet, now make
only a small contribution. The frequencies of prawns show similar
peaks from May to July, comprising 42-84% of all prey animals, so
that once again fishes and other organisms occur less frequently in the
stomach contents, especially in May. Adult crabs have several maxima
throughout the year, making the fish part of the eels' diet appear insigni-
ficant in January and February. I t must be emphasized that throughout
the whole year, in particular in January and February, glass eels or
young eels were present in the same waters as the yellow eels. However,
in contrast to the findings from streams in England (see above), in
only two cases had glass eels been eaten by yellow eels.
Choice of food and body size

As with most species of fishes, the eel prefers prey animals which are
not too big and which can easily be attacked. The eel's mouth is not
particularly large, so it can only take in moderate sized food items. If the
174 THE EEL
Table 12 The percentage of eels of different sizes which had fed on various prey
organisms on the rocky North Sea littoral of Heligoland, between September
and October 1965. Fishing method used: fyke nets (after Tesch,
unpublished)
Length range of eels 30-35 em 35-40 em 40-50 cm

Number of eels studied '4
Prey organisms:
" '7
Fishes (predominantly Gobiidae)

Mussels and Snails (Pl"edominamly " '3
"
Liuon'mI {iHorea and Hydrobia sp.)
Decapoda (predominantly Caron!.!! maenll$) ",6 '3
5'
'5
8,
Other Crustacea (Amphipoda, lsopoda,
Mysidacea)
Hydrozoa (predominantly Serm/aria sp.)
3' 4'
"
Bryozoa (Flwrra jQliacea)
4
"4
Polychaetes (predominantly Nereis sp.) 68 63 7
Algae
" 4
eel has the opportunity to eat its prey piecemeal, it prefers to do so,
rather than trying to swallow large prey animals whole. This habit is
dearly seen in an aquarium when an eel is fed chopped meat or fish.
The eel bites into the food and makes quick, jerky movements to the
back or to the side until a piece of a manageable size has been torn off.
In some circumstances the eel twists itself with great skill and speed
several times round its own axis in order to tear off pieces of food. Eels
do this, for instance, when they attack Crustacea. Fishermen from the
river Elbe report that they often find tightly dosed mitten crabs in their
eel baskets (see p. 273) and that the dorsal shell of these animals has
been forced open and the viscera eaten out (Ladiges, 1936). Crab legs,
which had been torn off with the gill lamellae still attached, were also
found in eels' stomachs, indicating that the eel actually attacked the
mitten crabs and did not just pick up limbs which had been shed by the
animals.
The eel is only able to indulge in its predilection for crabs (Decapoda)
when it has reached a certain body size. Table 12 shows the diet of
Heligoland eels caught in the vicinity of the island in water at least
3 m deep. The stomach contents of eels over 40 em long contained an
overwhelming proportion of crabs. Bristle worms predominated in eels
less than 35 em long. A year later the author carried out a study on eels
from the same area and found that many of them had empty stomachs.
In the few stomachs containing food, decapod crabs predominated
(over 50% of the total) both in eels over and, rather surprisingly, under
30 cm in length. However, most of the crab fragments found in the
stomach were from the edible crab (Cancer pagurus) , which had been
" ,.
~ Insect latvat(mainly $;mvlivm, § Crustacea(mainly
Tricllopttra, Odonata, PlltO(ltota Potol/!Ol> ,~,)
and E~n tfl>m>pt.t(t)
I I Verti'bt(ttf'
L-...J (ma,nly f ishnl
Fig. 73 The percentage of different prey animals found in SOUlh African eels of
various sizes (after Jubb, 1961 ).
used as bait in the eel baskets. Thus, in this case, the smaller eels had
been able to utilize a food which would normally be difficult to catch.
Fishes are another type of prey which is relatively difficult to catch
(see Table 12); they are, however, caught in increasing numbers by
larger eels, though not as extensively as are crabs. The fishes found were
mostly gudgeons (Gobio gobio ) which, because they are less than 50 em
long, are also available to smaller eels.
A study of several species of South African eels, presents a similar
pictUre (Fig. 73). While eels 10-20 cm in length had eaten nothing
but insect larvae, the diet of individuals over 20 cm long also included
Crustacea, in particular freshwater crabs of the genus Potamon, and
fishes . The proportion of fishes and crabs increased in eels up to
jP-60 em in length. Crabs comprised the largest proportion of the diet
in eels between 60 and 70 cm long.
In the English streams mentioned above, eels with fish remains in the
stomach also belonged to the larger size group, with an average length
of 31 cm (Sinha & Jones, 1967b). Individuals with insect larvae and
worms in the digestive tract were, on average, only 21-30 cm long.
Eels which had eaten molluscs, and to some extent Crustacea as well
(Gammarus sp. and Asellus sp. ) were the only groups to show a greater
average length. Research on Lough Gill in Northern Ireland (Moriarty,
1973b) has shown that, among groups of eels of various lengths, the
following percentages of individuals had eaten fishes: 7% in the
THE EEL
30-40 em group, 19% in the4Q-50 em group and 71 % in the 50-72 em

group. The percentages of eels which had eaten Trichoptera larvae
were 60%, 44% and 14% for the size groups respectively. The smaller
eels also preferred Gammarus to fishes. Similar observations, namely
that as the size of the eels increased so did the proportion of fishes in
their diet, were also made in the lakes of the Lough Corrib network in
southern Ireland (Moriarty, 1972). Prey animals that become less
frequent as the size of the eels increased were Trichoptera and Gamma-
rus, as well as chironomids, ephemeropterans and Asellus; medium-
sized eels showed a preference for gastropods.
Eels ( A. rostrata ) from Bermuda which are fish eaters have an
average length of 42 em, whereas individuals with mosqui to larvae in
the stomach are only 23 em long (Boetius & Boetius, 1967h). The body
size of American eels from the mainland also shows a marked correlation
with food types (Ogden, 1970). The stomach content of eels less than
40 cm long, which were caught by electrofishing in streams, consisted
mainly of aquatic insect larvae, while larger eels had fed predominantly
on fishes and the river crayfish.
Unfortunately, research on the European and American eels has
concentrated overwhelmingly on relatively small individuals, whereas
in New Zealand most of the eels studied have been considerably larger.
A local species in New Zealand, A. dieffenhachi, normally grows to
significantly larger average lengths than do most other species of eel.
A study of the proportion of trout in the diet of this species shows that
by the time an individual has attained a length of one metre, it is already
almost exclusively a piscivorous predator (Burnet, 1952b ; see also
Table 13). Further studies by Burnet (1952b) and earlier studies by
Cairns (1942b) (Table 13) have shown that both A. die/fenbachi and
A. australis schmidti, the other species of New Zealand eel (which does
not grow so large), both feed on benthic invertebrates, especially when
the eels have not yet reached a very large size.
Some studies have also been made on the stomach contents of larger
Anguilla anguilla from the periphery of that species' European area of
distribution. These eels were mostly between 50 and 80 em in length
and also showed feeding patterns typical of piscivorous predators
(Volf & Smisek, 1955; Drapkin, 1964; Sedlar & Krl:marik, 1967 ;
Morovic, 1970).
Larger eels are known to eat certain species of snails in addition to
fishes and crayfish. According to investigations in a Polish lake (Opuszy-
nski & Leszczyilski, 1967) eels measuring less than 20 em do not eat
any snails at all, the largest proportion of the diet in eels of this size
POST-LARVAL ECO LOGY AND BEHAVIOUR 177
Table 13 The percentage proportion of New Zealand eels, A . dieffenbachi and

A . australis schmidti feeding on various prey animals; eels grouped in size
classes. (Calculated on the basis of data from Cairns, 1942b)
<: 40 em 40-75 em > 75 em
A. dieffenbachi A. dieffenbachi A. australis A. dieffenbachi A. australis
Number of eels studied

with stomach coments
'"
2178 75' ,,' 7'
Prey animals:
,"
"""
Fishes '3
" "'9
MolluSC1 '3
Crustacea 3'
" "7
Ephemeroptera
" 39
,
T richoptera
Orthoptera " 3
3
, "7 47
Hemiptera 7
Diptera
Coleoptera ", "" ,
Odonata "
Arachnida <,
Various insect larvae
Oligochaetes
Assorted prey animals " ,
'7 37 , "'3
3
being made up by chironomid midge larvae. Molluscs first appeared

in the diet of eels between 20 and 30 em long, although in insignificant
amounts (only 14%) but were found in the stomachs of 64 % of indi-
viduals between 30 and 40 em long, and in 84% of individuals more than
40 em long. Among eels between 20 and 40 em long the greatest propor-
tion of molluscs was of the species Birhynia tentaculata, which grows
to a height of 12 nun. Eels over 40 em preferred larger snails, particularly
Radix limosa.
Choice 'of food and head width

Detailed investigations correlating body proportions with food eaten
have thrown some light on the influence of head shape on the type of
food eaten. Earlier studies were unable to establish any significant
difference in diet between the broad- and narrow-headed fonns
(Schiemeoz, 1904, 1910).
The distance between the nostrils, the interpupillary distance, and
head width, expressed as ratios of total length, differed by 12-20%,
10% and 22 % respectively in the broad- and narrow-headed eels
studied (Torlitz, 1922); 13 out of 16 eels which fell into the 'broad-
headed' category had fish remains in the stomach and intestine, but
only three out of a total of 19 'narrow-headed' eels from the same inland
178 THE EEL
Table 14 The percentage of broad- and narrow-headed eels from Norwegian coastal
waters containing different groups of prey animals (after Sivertsen, 1938)
Broad- InuT- Narrow-
headed mediate headed
Number of ~ls studied 76 93 94

Fishes [Gobiidae (inc. Aphia), Pholis, Ga.f/eromus,
,6
Zoarcu, Crrnolabrus, LabrusJ
Snails and Mussels ( Riuoa, Na.fsa, Li/lOrina,
3'
"
6
Myeilus, Modiolus )
" 9
"
Crustacea (Carcinus, Por/unus, amphi pods, isopods) 63 ,6
Worms (A renicola, Aphrodite, Maldanidae,
Lycoridae, PTiapulidae) 6 9
"
waters had eaten fishes. This difference appears convincing if one
considers the slightly larger total length of the 'broad-headed' eels
(70 em in contrast to 64- cm for the 'narrow-headed' eels) to be irrelevant.
However, Micheler (1967) found relatively slight differences in the
proportion of fish in the diet of 'broad-' and 'narrow-headed' eels:
42 % of 'narrow-headed' and 58% of 'broad-headed' eels contained
fishes.
Similar results have been obtained in coastal waters (Table 14).
Here, considerably more fishes were eaten by 'broad-headed' than by
'narrow-headed' individuals. In contrast, wonns were found more
frequently in the stomachs of 'narrow-headed' than 'broad-headed'
eels. In other words, the differences are similar to those found between
smaller and larger eels (Table 12)_ There were no differences in the
preference shown by the two groups for Crustacea.
Both these studies were concerned mainly with eels over 50 cm in
length. However, since smaller eels can also be classified as broad- or
narrow-headed, the question whether differences in feeding- occur
among the smaller eels as well, is of particular interest. Investigations
have been carried out on smaller eels from different lakes in Poland
(Opuszynski & Leszczynski, 1967). A comparison of the diets of eels
from Lake Gardno showed that broad-headed eels had eaten far more
snails than had narrow-headed eels. [Micheler (1967) observed similar
tendencies among larger eels as well.] The latter fonns had eonswned
a considerably larger proportion of insect larvae.
When discussing food selection among eels of varying sizes, it was
established that the proportion of molluscs eaten increases with the
size of the eel. This was also the case in Lake Gardno, a finding that can
be taken to confinn the idea that eating snails and mussels requires a
stronger jaw apparatus; such a diet is available to larger eels and to
broad-headed eels, both of which have a correspondingly stronger head
musculature than do narrow-headed individuals (Teriitz, 1922).
Since eels need stronger head muscles in order to eat molluscs, it is
easier to explain how, for example, they are able to crack open snail
shells with their mouths and then eject most of the shell (Drescher,
18 97).
A comparison of diet and head shape in eels from fresh- and saltwater
reveals the following: narrow-headed eels in freshwater eat mainly
insect larvae, while their counterparts in saltwater concentrate mostly
on wonns (polychaetes). Of course, freshwater eels of the narrow-
headed variety also eat certain species of worm (e.g. Tubifex), when, as a
result of pollution, no insect larvae are available (Ladiges, 1936).
The dependence of diet on food availability

As we have seen already, there are only a few bottom-living organisms
which are not eaten by eels. The spectrwn of foods consumed by these
animals ranges from fish, through mussels, Crustacea, insect larvae,
surface-dwelling insects and worms, to plants. The vegetable compo-
nents of the eel's diet are probably pieces of plant that the eel had
swallowed because the animal prey was attached to them. The same
applies also to Hydrozoa and Bryozoa found in eel guts. It is well known
that eels are not limited to aquatic animals alone, but will also eat
anything edible that falls or is washed into the water (Table 13), such
as air-borne insects and terrestrial worms (Heincke, 1894; Cairns,
1942b; Sinha & Jones, 1¢7b; Daniel, 1968; Ziepke, 1974). It is often
questioned whether, with the exception of fresh bait, eels will eat dead
animals. Most authors believe that eels will not do so, a reaction fre-
quently observed when eels are offered food in aquaria. Eels will not
touch food that has been in the water overnight. Fish bait on hooks and
in eel pots must be renewed every day or else the catch will show a
definite decline (see p. 276, and Schiemenz, 1910).
The eel's amazing spectrum of food reflects its great trophic opportu-
nism. This characteristic is particularly striking when one compares
neighbouring stretches of coastline with different types of bottom,
e.g. Wismar Bay which, in its central region, has a luxuriant growth of
water plants, and the Wohlenburg Wiek, which has a predominantly
slimy bottom with no plants present in the areas where eel fyke nets
are set (see Table 15 ; Drescher, 1897); 15 out of 20 eels from Wismar
Bay had eaten fishes, particularly sticklebacks and sea gudgeon; four
eels had eaten isopods. In contrast, the diet of 42 eels from Wohlenburg
Wiek consisted mainly of mussels; Macoma baltica was found in 88% of
the eels examined and the diet of 60% of the eels consisted exclusively
180 THE EEL
of Macoma baltica. Mussels were also among the other prey eaten, e.g.
Scrobicularia sp., Mytilus edulis and Mya arenaria. Lastly, a few eels
had eaten sand shrimps {Crangon crangon }. Fishes and snails were
absent altogether; in contrast to the Wismar Bay habitat, these animals
only occurred in very small numbers near traps in the Wohlenburg
Wiek.
Very striking differences in the diet of freshwater eels are revealed
when still and flowing waters are compared. Such a study has been
made by the Polish authors Opuszynski & Leszczyilski ( 1967), who
compared a river with certain lakes. Their results showed that the
majority of eels in rivers eat Trichoptera and Ephemeroptera, while
lake eels prefer snails and, above all, chironomids. In the Erne system
of Northern Ireland, Moriarty (I973b) found similar characteristic
differences between the stomach contents of eels from rivers and those
from lakes: in addition to fishes and sand lice ( Asel/us sp. ) the former
contained trichopterans and the latter chitonomids. This preponderance
of Trichoptera and Ephemeroptera in eel stomachs has also been
observed in streams in England. In addition, these English eels con-
tained a large proportion of larval Diptera (Table II), among which
chironomid and Simulium larvae were well represented. The chirono-
mids, however, were found mainly in the smaller eels and particularly
in those from running waters.
The two species of New Zealand eel, which feed mostly in rivers
and streams, prey mainly on Trichoptera and Ephemeroptera, when
they have not eaten fishes or Crustacea (Table 13; Burnet, 1952b). In
streams, however, a considerable pan of the diet also consisted of
snails, in particular Potamopygrus sp. In contrast, a lake study yielded
guts containing large numbers of mosquito larvae and, in flooded areas,
worms as well (Cairns, 1942b).
Different areas within one lake can yield fishes with different stomach
contents. In Lake Balaton, Hungary, eels from regions near the bank
ate mostly sand lice ( Asel/us aqualicus ) or other Crustacea such as
Corophiwn curvispinum and amphipods. In the benthos of open waters,
midge larvae were numerically predominant, and these, with fishes
such as various cyprinid and percid species, constituted the greater
volume of the eel's diet (Biro, 1974).
Certain differences in diet are found in the two species of New
Zealand eel, but it is very debatable whether these should be attributed
to the choice of food available. Cairns (I942b) comments that, despite
the common occurrence of Ephemeroptera larvae in the upper regions
of the coastal rivers, none was found in the stomach contents of A.
australis schmidti living there, which had fed mainly on molluscs and
Crustacea. The differences between the two species (see Table 13)
can just as easily bt: attributed to different feeding preferences among
eels from certain types of water.
Choice of food
Finally, the question arises whether the eel indiscriminately eats any
suitable prey animal that happens to come its way. A study on male
eels from brackish water deals with this matter in particular (Daniel,
1968). It was observed that many eels in the estuary area of the river
Eider frequently ate a large number of animals of the same species
(i.e. monophagy); other authors have observed similar phenomena in
freshwaters (Droscher, 1897; Ladiges, 1936; Cairns, 1942b; Pantulu,
1956). In the river Eider almost 60% of 145 'middle-sized' eels and
82% of 196 'small' eels were monophagous. Studies on A. dieffenbachi
in New Zealand have shown that if up to 10 animals were consumed at
one time, there was a tendency for them to be of several different
species (i.e. polyphagy). However, when 10-30 prey animals were found
in the stomach, there was a trend towards monophagy, a trend which
increased with increasing numbers of prey until, when 1000 animals
were found in the stomach, only one species had been eaten (Burnet,
1952b). Such observations were not made in the author's own studies
on A. anguilla from the rocky littoral of Heligoland (Table 12). There,
the only organism which was eaten in large numbers was the amphipod
Jassa pulchella. But, even when over 400 of these were found in the
stomach of a single eel, two other organisms were also identified.
Only 3 Out of 9 eels with more than 10 Jassa pulchella in the stomach
had eaten this organism exclusively. Most of the other eels examined
had at least two types of organism in the gut. Thus monophagy is not
always clearly expressed, especially where the range of food is as great
as it is in Heligoland.
The question of variation (or lack of it) in the diet plays a certain role
in production biology, above all when there is competition for food
(see below). The question whether the amount of food conswned by the
eel is proportional to the amount that is present in the benthos, has
been investigated in New Zealand streams (Cairns, 1942b; Burnet,
1952b). Results showed a certain correspondence between the quantity
of prey animals in the benthos and the composition of eel stomach
contents. This applied in particular to Ephemeroptera. Preference
was, however, shown for other animals, e.g. Trichoptera and Crustacea;
in other words, the proportion of these animals in eel stomachs was
IS. THE EEL
Table lS Diet of cds from different environments (unless otherwise indicated. the species i!
A. anguilla). T he various organisms are listed inorder offrequency (without percentages)
Where specifically indicated, the percentage of the total net weight or volume of all th(
prey is given, otherwise the percentage of eels which had fed on the organism in questio[
is given. Other localities have been described in Tables II and 13. and detailed informa·
tion is given in Tables 12 and 14 and in Fig. 73
SI!A WATER
Norwegian coast (SivCTtsen, 1938) Nonh Sea, rocky littoral near

H eligoland (Tesch, unpub lished)
Net weigh
Crabs (Carcinw, PortullUS) )8°{. Cr.Ibs (Carcinw) 4'~
Fishes (Gobius, PltcliJ, Gastuo$UItS, Aphia, Bristle ....mml (Nuns 5p.) 35 °~
Zoarus, ClenoIlWrw, Lobrus) ll °'o Sand hoppers (Jrma pu/chella ) 8 °~
"
Shrimps, mud dwelling amphipods Fishes (Gobiidaej I eel)
(GammaruJ Corophium) ' 9% Bryozoa < t"!.
Worms (Arenico/a, Aphrodiu, Maldanidae, Algae < I"!.
Lyooridae, P ria pulidae) IS % Mussels and snails < t"!.
BRA CKIS H WATER
Baltic Coast, open sca bay, Baltic coast, open sea bay, Chesapeake Bay ( Arlluilla rostrara ) ;
few plams (Droscher, 1897) many planlS (Droscher, 1897) (Wenner & Musick, 1975)
Mussels (Mcuoma baltica) Fishes (Pungiriw pungiriw) Crabs ( CallintclU S{lpidw)
M ussels ( Scrobicu!aria, Mytilw, Fishes (Gasluo$uwaculeatw) M ussels (Mya armaria, Mulinia [a!tralis,
Mya ) Fishes (Gobiw nigu) Maroma sp)
Midge larvae (Chironomw) Sea lice (isopoda) Bristle worms (Polychaetcs)
Prawns (Cra~on crangon) Bivalves ( Cardium, Theodoxw ) Crustacea (Amphipoda, Isopoda)
Fro" Fishes ( A/DIll pseudolwrmguJ )
Inse,"
FRES H WATER
Lake, Mecklenburg Lake, England Lake, Ireland Dike waten, North

(DrOscher, 1897) (Frosl, 1946) (Moriarty, I969) Sea coast (Daniel,
1968)
N,
weigh
Moll uscs ( Limnaea, Snails and Mussels Midge larvae and pupae Earthworms 40'0/,
Theodoxus ) ( Va/tiara, Limnata ) 86% (Chironomidae) Fishes (Pungitiw
pungitiuI) 37"/,
Tricho ptcra Insect larvae Other insect larvae Fish eggs
(Gasterosteidae) Il o/c
Fishes (Pwlgitius (Trichoptera, Sialir, A sellus 5p. Insect larvae 3°{
pu~tjus) Ephemeroptera, Fishes ( Puca Leeches 2°,
Midie larvae Olironomidae) 49°" jluviatilis ) Shrimps (Gammarw) I O{
( ChirIJnOmUS) Shrimps (Gammaridae) 21°" SOli!, ( Radix 01Jala ) , 'J;{
Dragonfty larvae Wonns (Oligochaeta) 3 °~ Piantremains to/.
Fishes ( Puea) I o~ Beetle ( Agabw Iybius ) < 1%
Argyrontta (?) < 1%
P OS T - LAR VAL ECO L OG Y AN D BEH AVIO UR
RIVER ESTUARIES, with a panial brackish water influence
Elbe tidal region near Hamburg Harbour River near Calcutta ( A . ntbuwsa )
(Ladiges, 1936) (Pantulu, 19S6) Volum,
Tubiftx s.5'J~ Fishes (Hilsa, Suipinna, PanuJ, a
""
Water fleas (Cladoc::era) 10 °0 few glass eels)
Midge larvae (Chironomidae) ,', C~'"
,',
",
Bait prawns ( MysiJ ) Prawns (Peneidae)
Millen crabs ", Insects
Earthworms ", Megalopa larvae of the cra b
Mussels and Snails ", ( Varunasp.)
,,3
I..«d>n ", Plants (micro- and macrophytes)
Worms (Annelida)
Shrimps (GomnuJrus) I ~o
Prawns ( uandn) < t o,"
North Sea Coast, dike waters Bermuda ( A. rOJlrala ) Kattegat lagoon-like waters
(Daniel, I968) (Boetius & Boetius, 1967b) (Muus, t967)
Net weight
Prawns ( Crangon crangon) :z6% Fishes ( Fundulus bermudat ) Fishes (GasleroSltus and PungiNu
Fish spawn (Gasrnosrew acultalw) 19°0 Midge pupae (Chironomids) Crabs (small Carcinw mamtl$)
Crabs (Caranus mamtl$) 11°0 Snails ( Physa) Mussel remains ( Mya )
Ea rthworms 11°"
Bristleworms ( Ntreis ) ,',
Fishes ( GaslnoSfnlJ acuJtalw ) ,',
Snails ( Radix ovala ) ,',
Midge larvae (Chironomidae) ,',
Fishes ( PutIKi1iw pungiliw ) 1%
Stream, England Streams, North America Lake, South Germany

(T homas, 1962) (Godfrey, 19S7) (Michcler, 1967)
( A . rOflrala)
""
Ephemeroptera 70°'0 Ephemeroptera Bony fishes
Trichoptera 45 ~0 Other insect larvae Plant fragmenlS
Midge larvae (D iptera) 35°0- (plt'COptera, Chironomidae Diptera '3
''..
PlteOptera 30° o Simulidae, Odonara, etc.) GastropOds '3
Fishes ( Phorin w, Lmnpetra) :ZSo° Fishes (RhinidllhyJ sp., Annelids
Leeches (Hirudinea) :ZS°'o Gasterostcidae. Conidae. Algae
MUS5els & Snails 20% Salmon spawn) Detrilus
Trichoptera "
Water lice, Shrimps & Sand hoppers
( A sellw & Gammarus) 10% Tabanid la['Vae " 7
Alderfly larvae ( Sialis ) S%
Beetles S%
higher than in the benthos. These studies also showed that eels, like
other species offish, scorned beetles (Coleoptera) as a food source. Some
relationship between stomach content in eels and the bottom-dwelling
animal popUlations has been observed in American streams (Ogden,
1970). The stomach contents of eels from a lake in southern Ireland
also reflected the composition of the benthic fauna (Moriarty, 1972),
i.e. chironomids, Ephemeroptera, corixids, Gammarus and the snail
THE EEL
Bithynia. However, the stomachs contained a much larger number of

Asellus sp. than was to be expected from samples taken in the benthos.
A water bug (No tonecta sp.) was not eaten by any eels. In contrast to
findings from other studies (Table 15; Droscher, 1897; Frost, 1946),
the snail Bithynia also appeared to be totally unattractive to eels.
The various elements in the eel's diet are analysed in Table 15,
grouped according to hydrological conditions and their importance in
the feeding regime. From this table, and the results cited above, the
following hierarchy emerges with respect to the significance of different
organisms in the eels' nutrition (but not necessarily the eel's food
preferences) :
(I) Fishes, in particular small, bottom-dwelling species such as the
stickleback (in 9 cases), gobies (Gobiidae) (in 3 cases), the Miller's
thumb (Cottidae) (see also Frost, 1946), gudgeon (Gobio gobio ),
loaches (Cobitidae), eels of the same species, and lampreys (see also
Frost, 1946; Perlmutter, 1951; Thomas, 1962). SaImonids (the
object of many studies) appear with disproportionate frequency as eel
food (Cairns, I942b; Frost, 1952; Burnet, 1952b; Godfrey, 1957;
Thomas, 1962; Sinha & Jones, 1967b; see also Kothke, 1968).
(2) Snails and mussels.
(3) Large Crustacea (decapods) such as crabs, prawns and such
edible Crustacea as the brook crayfish (e.g. see Walter, 1910 ; Svard-
son, 1969; Fisch u. Fang, 7, 246,1966; Runge, 1965) and the mitten
crab (Jacob, 1970).
(4) Small Crustacea, in particular amphipods (see also Tesch & Fries,
1963; Fries & Tesch, 1965a, for details on Gammarus tigrinus),
Isopoda, prawns used as bait (Mysidaceae), water fleas (Daphnia)
(Ladiges, 1936), and Megalopa larvae (Pantulu, 1956).
(5) Worms (see also Opuszyilski & Leszczynski, 1967), in particular
tubificids, earthworms and, in one unusual example, the spiny-
headed worm ( Echiurus echiurus) (Kiihl, 1965).
(6) Insect larvae.
3.3.2 Age and growth
Methodological problems
The following section does not set out to give a detailed review of
methods used in studying age in eels. Specialized publications are
available which deal with the methodology of such research in fishes
(e.g. Chugunova, 1963; Bagenal & Tesch, 1977); by and large these
methods can also be applied to the eel. However, the genus Anguilla
does provide a few special problems which must be mentioned here.
The eel is unusual in that its life cycle consists of two quite separate
phases: the oceanic larval period and the continental period, often
incorrectly referred to as the 'freshwater phase'.
The larval period is of different duration amongst the different
species: in the European eel, for example, it is thought to be 2~ years,
and in the American eel, 6 months to It years (see p. 93). Studies of
otoliths indicate a larval period of two years in the New Zealand eel
(Cairns, 1942a), and one of about 18 months in the Indian species
A. nebulosa nebulosa (Pantulu & Singh, 1962).
Age comparison between species is made more difficult by the
varying lengths of their larval life. For this reason, and because the
interpretation of the larval otolith is an unreliable procedure, it is best
to calculate age as dating from the glass eel stage, a system used by most
authors. However, the methods used for counting the years from the
glass eel stage onwards are by no means standardized (Thurow, 1959;
Sinha & Jones, 1 967c). In many cases, data from various authors
concerning eels of different age groups cannot, strictly speaking, be
used for comparative purposes. Therefore, as far as possible results
obtained by one author only will be compared in the following sections;
differences which arise from differing interpretations of annual rings,
will also be dealt with.
Techniques used in the preparation of otoliths for age and growth
studies are discussed by Moller-Christensen (1964), Moriarty (1973a
and 1977) and by Deelder (1975). The otoliths of eels kept in ponds
appear to provide a very accurate indication of the number of years
that have in fact passed (Smith, 1968).
By recapturing marked eels (Fig. 74), it was possible to confirm the
relatively slow growth rate indicated by otolith studies (Peiulz & Tesch,
1970) made on eels along the German coast of the North Sea (see also
Burnet, 1969b). These showed that, at a length of between 40 and
50 em, eels grow 2-3 em a year. Marked specimens also showed a 2 em
length increment on recapture (Fig. 74). Studies on the otoliths of
eels between 78 and 91 em long from Lake Ontario indicated an annual
increase in length of 3· I em; marking and recapture experiments using
eels between 73 and 87 em showed an annual increase in length of
3·3 em - which again canfinns the otolith method for ascertaining
growth (Wenner & Musick, 1974).
A further complication in research on age and growth in eels arises
from the fact that scales first appear when the fish is already relatively
186 THE EEL
I::~"
I' j~
---===-========1
-, r I , I , , , I I , , I ,
~ ~4 8 12 10 W 24 ~ U » ~ « ~ 52
TlrM t lop.ed olter mCI,kinQ(in ",",,~ I
Fig. 74 Growth rate in He1igoJand eels which had been marked internally using
stainless steel tags. Vertical lines represent the extent of variation in length
increase, as obtained from laboratory measurements after the eels had been
re-captured. The figures next to the lines indicate the number of eels re-
captured. Specimens caught I -8 weeks and 52 weeks after marking were
32-45 em 10ng and 47-51 em long respectively (after Tesch, unpublished).
large (see p. 12), and there are interspecific variations in the size at which
this occurs (pantulu & Singh, 1962). Therefore, the normally clearly
defined annual markings on the scales are of very little use if one does
not know the age of the eel at the time of first scale formationj some
differential growth within this period of the eel's life may affect the
actual age at which this occurs.
Age at first fonnation of scales is best determined by comparing the
figures calculated from the scales with those from the otoliths. In
research on the central European eel, most authors have arrived at a
deficit of three years in the number of annual rings (Ehrenbaum &
Marukawa, 1914; Nordquist & Aim, 1920; Rasmussen, 1952j Rahn,
19553). Gemzc;k (1906) found a deficit of two years in the number of
rings on the scales of eels from Denmark, Opuszynski (1965) a deficit
of two to three years in eels from Masuren, and Marcus (1919) a
deficit of four years in eels from England and Germany. In A. rostrata
in Canadian waters the deficit is three years (Smith & Saunders, 1955).
Data from Cairns (1942a) indicate an annual ring deficit of almost
five years in species from New Zealand. In the Indian eel A. bicolor,
the scales usually develop six months after the glass eel stage, and
thus there is no need to correct the calculated age derived from annual
ring counts in this fast-growing species (Pantulu, 1956). Finally, as
Rasmussen (1952) points out on the basis of data derived from several
authors, the difference between the number of annual rings in the
otoliths and those in the scales increases with the increasing age of
the eel (up to five years in specimens over eight years of age); also the
number of annual markings on different scales from a particular region
of the body may vary by as much as ± 2 years in one individual.
For these reasons, scales only appear useful as a first approximation
in estimating age, but, despite this, frequent use has. been made of such
data in recent studies (e.g. see Voronin & Rusetskaya, 1971; Kozlova,
1973; Noskova, 1973).
In no way can scales be used to back-calculate the length of an eel
during its earlier years of life. For such calculations one is forced to
rely on otoliths. Although objections have been raised (Rahn, 1955a),
there is nothing wrong with the use of otoliths for back-calculations,
as biometric studies have shown (Matsui, 1952; Pantulu & Singh, 1962;
Penaz & Tesch, 1970). Other hard parts, e.g. vertebrae, can also be
used as a means of determining age (Ehrenbaum & Marukawa, 1914).
However, none rival the otoliths in providing such a complete reflection
of the fish's earlier life history, including the larval period.
The ring pattern of the otoliths should be seen as an indication of
periodicity in the eel's life whether the species lives in tropical or
temperate zones. Since it is not known how far factors leading to ring
formation in tropical species correspond with those producing annual
markings in eels from temperate regions, interspecific comparisons may
not be fully valid.
In the examples that follow, total body length is used as a measure
of growth. Weight varies greatly with nutritional condition.
Comparisons and means of converting data from different species are
to be found in a separate section (see p. 199).
Differences in growth between males and females, and the age and length
at which migration begins
As has been discussed in the section on sexual development, the majority
of eels which come from overpopulated biotopes and which show a
slow growth rate during the first years of their continental life, tend
to develop into males (see p. 56 and Fig. 75). It therefore seems likely
that, within a given biotope, males will show a slower rate of growth
than females. This hypothesis is confirmed when the results from
several investigations made by various authors are combined (Fig. 76;
see also Ask et al., 1971 ). From the age of four onwards, females are as
much as 10 cm longer than males in all the regions studied in Europe
and Japan.
The only means of establishing in which year of life or at what body
size this difference first occurs in natural waters, is to back-calculate
the length attained at earlier ages (Penaz & Tesch, 1970). On the basis
of such calculations it appears that until the eels are at least 20 em long,
males are still largely undifferentiable externally. Eels from the coastal
regions of the North Sea and from most other areas too, are about three
188 THE EEL
j\~>~>: ;
f
I Ffrawc:lft1ptf(1l1
,-
I
10
,
J Rhyd-hitc.4tt"perml
, .JRhyd-hir
~
I'--r~~~--r'- O
o I II III IV V VI OIIlIU IVVVI
AQtgroups
Fig. 75 Total length and length increase in male eels from (WO different streams in
Wales. Ffraw: population density, about I eel per m", proportion of males,
50% . Rhyd-hir: about 4 eels per m", proportion of males, about 90% (data
from Sinha & Jones, 1966, 1967C).
years old when they reach this length. Since the indications of sex-
correlated size differences are generally still very slight at this age, the
difference must increase rapidly from then onwards. Differences of
up to 10 em have been found in many cases) particularly in eels from
southern Europe. The females of the tropical species A. nebulosa also
show a faster growth rate than the males of this species (Pantulu &
Singh, 1962).
It appears that this secondary sexual characteristic follows a different
pattern of development in the New Zealand eel, A. dieffenbachi (Cairns,
19423). Investigations have shown that the males of this species have a
better growth rate than the females.
As regards the maximum size attained, A. dieffenbachi has a special
position among those species for which data on length are available.
Burnet (1969a) studied the average lengths of migrating silver eels in
POST-LARVAL ECOLOGY AND BEHAVIOUR
., ,.,"" "" Sd1l11OlIto,

"
~"
Rt>yd__ Htli9olor4 Sta, Ustdotl
lI'I!igcIarId , HoIlohd ~
"
"
25 ~~111
l , . " ""
Elbo
H!Uary,
Co<
30
25
........-..-.-.~ ............ ,..............J""'"'T-r"'O ,..............,............., 20

"IWVmWVIII~v~mwvm~V~IIIWV~"IWV~IIIWV
.
Wyk/FOhr,
KIt! Bay, North Sta
...
)
Saltie
' ~.
nearer....
""" vv .,
" "
Ang/Jdla
25 japonfC(I
(JOPOnl
...,...
20
,..............,..............J""'"'T-r"'O,""·""~,.....,.....,""i.
V~W~WV~W~V~~V~~~~WVVWWhWV~Wg~IIIIIWV~W
Fig. 76 Size differences related to age in male and female eels from various waten in
Europe and Japan (Matsui, 1952): the North Sea and the Elbe estuary
(Penaz & Tesch, 1970), the lower stretches of the Elbe (Ehrenbaum &
Marukawa, 1914), the Weser, the Severn (south-west England). the Trave
estuary and Lake Schmollen (Marcus, 1919). streams in Wales (Sinha &
Jones, 1¢7c), the Dutch Wadden Sea a.J. Tesch, 1928), Kid Bay (Thurow,
1959), the lagoons of Valencia and Carmargue (Gandolfi-Hornyold, 1921,
1930). Most authors have treated sexually undifferentiated specimens as
males. Vertical hatching indicates length differences between males and
females.
the Waimakairi stream in New Zealand. Whereas male A. australis

reached a length of 43'4 em and, in this respect, resemble the Atlantic
species and A. japonica, males of A. dieffenbachi reach a length of
59' I em. The females of A. dieffenbachi are also correspondingly larger
at the silver eel stage, anaining lengths as great as 120 em (see also
Burnet, 1969a); the maximum length recorded for A. dieffenbachi
is 200 em. Such lengths are hardly ever attained by Atlantic eels.
The lengths of European silver eels recorded over a period. of several
years in different regions and waterways have been combined in
Table 16. These figures indicate that the average length of male eels
190 THE EEL
lies between the relatively narrow limits of 35 and 41 em, and thus does
not differ much from the average length of the short-finned New
Zealand eel A. australis; even the average length of male Japanese
silver eels (approximately 42 em) is only slightly higher than this
(Matsui, 1952). In northern Europe the animals appear to be longer
than those from central and southern Europe.
The range of variation about the mean lies between ± 3 and 7 em, so
that the smallest male silver eel is 29 em long. In Sardinia there are
reports of a male silver eel only 24 em long (Gandolfi-Homyold, 1934).
The largest individuals were 46 em long. The degree of variation is
greater in the New Zealand eels (Burnet, 19693) and in A. japonica;
in the latter species males up [0 57 em have been known (Matsui, 1952).
In European female silver eels the mean lengths likewise only vary
within very narrow limits. Mean values range from 54 to 61 em in
European waters (see also Apollova, 1969). In contrast to A. dieffenbachi
(see above), A . australis has an average length a few centimetres above
this range (Burnet, l¢9a & b). Anguillajaponica appears [0 resemble
the European eel as regards the larger size of females relative to males
(Matsui, 1952). The mean length of the American eel, A. rostrata,
during its spawning migration is 69 cm (Gray & Andrews, 1971); for
A. mossambica see p. 205.
According to the figures in Table 16, the lower limit for length in
adult female eels appears to be about 38 em. In southern Europe and
Africa, small female eels were found in particularly large numbers
(Gandolfi-Hornyold, 1934), e.g. females measuring 37 em have been
recorded in the lagoon of Comacchio. The upper limits for length in
females vary quite considerably; Table 16 shows lengths of up to
100 em. Eels measuring as much as 130 em have been caught in a lake
in Switzerland. According to Bauch (1954), female European eels may
be as long as 150 em. The maximum lengths of the South African eel
species, whose females, according to reliable sources, can weigh as
much as 18-23 kg, appear to be 180 em or more (Jubb, 19<)1). As noted
above, female A. dieffenbachi can reach a length of 200 em.
Finally, it should be pointed out that the average length of silver
eels from one stretch of water can vary from year to year, though this,
in fact, only occurs to a slight degree. Observations have shown that
the average length of females from Lake Esrurn in Denmark shows a
tendency to decrease (Table 16). The Danish author Rasmussen
(1952), attributes this decrease to an intensified re-stocking programme,
which must have produced a considerable increase in population densiry
and with it, greater competition for food. At the same time there is
Table 16 Average total lengths of emigrating ma le and fema le silver eels in different European waters
Malts Females
Water body Y ear /.mg" Range Nu""" Ag. /.mg" Rang. Nu""" Ag. Rmtarks Author
(=) (= ) (= ) (= )
.
0
~
Lake Esrum, Denmark 1937-8 57"8 49- 6 7 8, 11 "5 Rasmu$$C1l, 1951 ,
Lake Esrum, Denmark 1940-2 58 "7 52-70 9' 11 "4 Rasmussen, 1951 .....
Lake Esrum, Denmark 1943-5 56"5 46-79 8, 11"1 Rasmussen, 1952 >
Lake Esrum, Denmark 1946-8 55"3 49-73 8, 11 "8 Rasmussen, 1952
Lake Esrum, Denmark 1949-50 55"7 49-79 8, 11"4 Rasmussen, 1952 <
'"
Lake Windermere, UK 1940-50 ",-0 60-8 12"5 Frost, 1945, 1961
>
9 47-97 '4 0 ..
Cunsey48e<:k, UK 1939-44 C,40"O 37-46 c" 57'0 46-88 dd + 99 Frost, 1945, 1961
River Bann, " No" of eels = t 122 n
Nonhern Ireland 38 "5 33'5-44"0 Frosl, 1950 0
'944 " 9 7-'
River Bann, Rep , Ministry of
Nonhem Ireland '9O, 4' "3 3S-C" 45 ,,6 54"4 45-79 153' Agr" Belfast, 1966
..'"
0
River Bann, Rep" Ministry of
Northern Ireland ,,,.. 4 1"0 33-c,46 '57 55"3 46-78 8" Agr" Belfast, 1966
">"'
Ijsselmeer, Holland 195 6 36"0 29-43 9 Deelder, 1957b Z
Ijsselmeer, H olland 1920 35 "0 31-38 Tesch, 1928
River Dieze, Holland 8
'''''8660 Tesch, 1928 to
'9'" "
Danish CoaSt C, 41 "0 36-48 7 55"0 45- 8 S 8 Presumably at least
,"'" " '" one year older GeD"\2.;t, 1906
:c
'">
Lake Vran, Yugoslavia 37"2 33-39 ,0 6-7 99 not represent- Haempc:1 & <
Lagoon of O:Imacchio, ati ve Neresheimer, 1914 0
-
haly 37"2 31 - 44 < 6? 4 annual marks on Gandolfi-
Lagoon of Valencia, ''''' the Jellies Homyold, 1934
Spain 4
c" 37'0 31- 46 6-, c, 55"0 46- 100 44 9 Gandolfi
"'"
"" Homyold, '921
Lake Tunis 1930 ,6~ 33- 44 347 7 38- tO I '49 Females not re4 Gandolfi4
p resentative Hornyold, t930
'"
--
THE EEL
thought to have been an increase in the proportion of male eels in this

lake - which would confinn the author's views on the dependence of
sexual differentiation on population density (see p. 56).
Although the growth rate is not as high in males as it is in females
(Fig. 76» it should be noted that male silver eels are between one and
three years younger than female eels when migration begins (Table 16).
Thus, it is not through a better growth rate alone that females achieve
their greater length - they also spend approximately two years longer
in continental waters. Investigations in natural European waters
(Table 16) show that the 'continental' age ranges from 8 to 12 years
in females, and from 6 to 9 years in males. The age range within the
sexes could be determined by regional climatic variations. In northern
and central European waters, male eels are, in all cases) slightly older
than those from southern Europe, on average about Ii years older.
The difference between females from different areas is probably even
greater, though a comparison is not possible on the basis of data from
Table 16. In contrast to age, the average lengths do not differ to the
same extent from north to south. For this reason it must be assumed
that it is not so much age, but length which detennines whether meta-
morphosis into the silver eel stage occurs. This assumption is supported
by studies on eels in Lake Windermere (Frost, 1961) and by earlier
investigations by other authors. The reason why female eels often
considerably exceed the critical length (50-60 em) for metamorphosis
into the spawning migration stage, is that when these eels are between
50 and 60 em long (or less), they grow at a faster than average rate.
In the north-west European Lake Windermere just mentioned, the
age criterion for this stage is thought to be nine years. Thus, if female
eels between 50 and 60 em are younger than nine years or are growing
relatively fast, they will attain final lengths considerably in excess of
60 em. Female Canadian silver eels ( A. rostrata ) are, on average,
12 years old (Gray & Andrews, 1971 ).
Examination of the otoliths or scales of migrating eels) or of sexually
mature eels which have been artificially prevented from migrating
or continuing their migration) suggests that the apparent age of these
eels may be misleading. For example, eels introduced into Lake Como
in Switzerland 45 years previously had a maximum of 10 annual
rings on their scales (Gandolfi-Hornyold, 1932a). Eels which had
lived for over 30 years in a clay pit near Brandenburg and which had
reached an advanced stage of maturity, had developed no more than
6 to 8 annual rings on their scales (Wundsch, 1953). The fact that some
of the eels from Lake Como did not weigh very much and had very
irregular scales indicates that these animals were exceptionally old.
Both cases must be regarded as most unusual. The eels had reached the
silver stage but had been denied the opportunity of migrating.
Presumably further growth could not take place, thus preventing the
usual formation of marks on the scales.
Earlier literature (Walter, 1910) also indicates that, if migration is
prevented, eels may live to quite an advanced age. One specimen was
kept for 55 years in a well in Denmark. Another eel is said to have been
reared in the smallest of containers by a French family; it lived 37 years
and attained a length of about 1'3 m.
Head width and growth

As data on feeding habits have shown, the organisms eaten by broad-
headed individuals differ from those eaten by narrow-headed eels.
In general, broad-headed eels eat animals that are larger and more
difficult to caprure ; the greater strength and size of their jaws enables
them to do this. It is thus likely that the two types of eel will show
differential growth rates. Table 17 sets out the results of a study where
the age of broad- and narrow-headed individuals was determined
separately by one worker. According to this table, broad-headed eels
show a much higher growth rate than do narrow-headed individuals.
The length advantage of the former over the latter increases from
5 em, after 6 years on the continent, to 10 em in eels between 8 and
10 years of age . Another author (Bauch, 1954), who is experienced in
age determination in eels, indicates an even greater increment in
broad-headed individuals, i.e. 2-4 em for every IO em increase in length.
Other authors (e.g. Wundsch, 1916; Ehrenbaum, 1930) also attribute
better growth rates to broad-headed eels (see also Micheler, 1967).
Such differences are only relative, that is, they occur within a given
population. Eels from Heligoland serve as examples of extreme broad-
headedness (Tesch, unpublished); head width was equal to 36% of
head length or 1 :21 in total body length. The latter ratio is approximate-
ly I :40 in narrow-headed eels (Walter, 1910j Torlitz, 1922). Relatively
good growth rates are only seen in Heligoland eels which are less than
Table 17 Average lengths (em) of broad- and narrow-headed female eels from Lake
Sakrow near Potsdam, arranged according to age groups (after Rahn, 1955a)
Age groups V VI VII VIIl IX X
Narrow-headed ec:Js 43'7 46'9 50'0 53'0 59'0 6"0

(No.) (6) (26) (35) (20) ( 10) (6)
Broad-headed eels 53"0 57'8 62'3 69, 7°'1
(No.) (8) (3) (19) (7) (8)
194 THE EEL
30 em long - after which length they do not grow as well as other eel
populations along the coasts of the North Sea (Peiuiz & Tesch, 1970).
These data were derived from eels measuring between 33 and 63 em.
D£fferences in growth due to environmental factors

The main factors influencing individual growth rates in all members
of a population are population density, temperature and the quantity
of food available per unit of area.
Observations from two rivers in Wales, the Ffraw and Rhyd-hir
(Fig. 75), show how population density can influence growth rate
(Sinha & Jones, 1967C). In the first river, female eels in particular
showed a much better growth rate, until their eighth 'continental'
year, than did the eels from the second river. The difference in length
between eels of the same age was as much as 4 em. However, after the
ninth year, and in eels over 45 em long, this difference had almost
disappeared. Appropriate conversion of Sinha & Jones' data shows
that, in the river containing eels with a slow growth rate, population
density was equivalent to four eels per square metre, and in the river
with the better growth rate, it was equivalent to one eel per square
metre.
Where larger eels are concerned, growth rate is not so adversely
affected, even in high density populations. This is due to the fact that
such individuals feed differently, and males over a length of 40 cm
leave on their spawning migration, with the result that there is a sharp
decrease in competition for food once this length has been reached.
]n addition, the proportion of females decreases at high population
densities. Since females made up 50% of the eel population in the
Welsh river with a good growth rate, but only 10% in the river with a
poor growth rate, population density, and thus competition between
eels over 40 em long, was reduced much more in the latter habitat.
Competition in the 'good' river must have been relatively intense among
the larger female eels, which then became members of a population
that had only, in effect, been reduced in size by a half.
Moriarty (1973b) found differences in growth rate among eels from
different water bodies. A poor growth rate and small average lengths
were found in Lough Gill and Lough Conn in Northern ]reland; there,
Io-years-old animals were about 40 em long. ]n the Erne system, on
the other hand, eels of the same age were over 50 em long. In fact, the
average length for all eels caught in the river Erne system was over
10 em higher. On the basis of catch size per unit effort, there did not
appear to be any differences in population density (Moriarty, 1973b).
Moriarty attributes the small average lengths of eels from Lough Gill
and Lough Conn to over-fishing. However, it is striking that these
two lakes are not more than 10 and 25 km respectively from the sea.
The lakes of the Erne system, on the other hand. are all more than 65 kIn
from the sea, with the majority at a distance of considerably more than
100 kIn. One might therefore assume that a greater population density
in the waters near the sea is responsible for the poor growth rate in the
eels living there. In a comparative study of eel populations in Ireland,
Moriarty ( 1974) mentions the various food elements and their propor-
tional occurrence in the eels' diets. In lakes where eels feed mostly
on fishes, growth rate was poor. In contrast, if few fishes were eaten
and the diet consisted mainly of chironomids, the eels showed a good
growth rate. This information immediately suggests possible causes
for these differences. I t is well known that lakes with a rich population
of benthic midge larvae (Chironomus plumosus ) are very productive;
in such waters, large quantities of dipteran larvae and other invertebrate
prey are available to eels. In waters where there is a dearth of insects,
or where there is strong competition for food, eels will begin to eat
fishes. When this occurs, slow growth rate can be expected.
Data on size increases in eels introduced into a new habitat show
that, in natural waters, reduced growth is largely the consequence of
competition resulting from excessive population densities. Eels,
introduced as glass eels, into Lake Worth in Austria weighed between
500 and 800 g after a period of four years (Einsele, 1961 ); on the basis
of such figures one would expect these fishes to be between 65 and
75 em long. According to Fig. 76, however, none of the female eels in
other natural waters was more than 37 cm long by the end of the fourth
year of life.
Glass eels which had been introduced into a lake in the upper regions
of the Volga reached a length of 13'9 cm after the first period of growth,
and 25 cm after the second (Nikanorov & Smotrjaev, 1962). These
figures are more in accord with those shown in Fig. 76 and Table 18.
In addition it should be noted that, with 70 to 75 glass eels per hectare,
stocking was relatively intensive (see p. 265), so that a certain amount of
competition existed. What is more, the area was so far north that, on
the basis of relatively low temperatures alone, one would not expect
particularly good growth rates.
Glass eels which were introduced into Russian dams grew to a length
of 13' 1 cm in the first year and 19'2 cm in the second year after introduc-
tion (Kochnenko & Borovik, 1958). In this case stocking was at a density
of 200 glass eels per hectare. At the same time Kochnenko & Borovik
THE EEL
put 500 glass eels into ponds in the same area ; these eels reached lengths
of 22"2 em and 39"9 em in the first and second years of growth respective-
ly, i.e. they showed a much better growth rate. It appears that ponds
(see p. 330), which reach higher temperatures in the summer and which
present less competition from other species of fish, offer better condi-
tions for growth than do the relatively deep dams. What is more, growth
rates (as compared with data in Table 18) appeared to be relatively
good in the older age groups of those eels which were introduced into
Belorussian lakes. Kostyuchenko & Prischepov (1972) have a large
amount of data showing that between the fifth and eleventh year in
freshwater these eels attained the following lengths: 56,63,69,74,79,
81 and 85 em.
Koops (1g67a) reports that in Lake Balaton in Hungary, stocked eels,
each weighing 7-14 g and measuring 20 em, grew to between 47 and
50 em after one year and had reached go em after four years" The density
involved in this stocking was close toone eel per hectare, i.e. competition
was very low indeed. In addition, the temperature of this relatively
warm water mass may have contributed to this unusually marked
increase in size. In other words, conditions for growth were initially
very favourable.
In general it is very difficult to establish from these studies whether
temperature has a decisive influence on growth rate. If one compares
data from the Atlantic drainage area (see Fig. 75) with those from the
Mediterranean area, it is evident that, in the north, all female eels in
the age groups V and VI (AG V and VI ) are less than 40 em long,
whereas in the region of the Rhone estuary females in AG V are 42 em
long, and in the lagoon at Comacchio females in AG VI are also 42 em
long (Haempel& Neresheimer, 1914). There are also data from Valencia
and Comacchio on eels in AG VII. In Valencia females of this age
group were 47 em long, and in Comacchio they were 46 em long. In the
north, however, all females of AG VII were less than 45 em long.
According to Bellini, female Italian eels in AG IV even attained lengths
of between 55 and 65'5 em (from Haempel & Neresheimer, 1914).
Data from Czechoslovakia (Sedhlr & Dobrota, 1966), where the
waters are relatively warm and only thinly populated, indicate quite
a high growth rate. There, the eels were 48, 57 and 63 em long in age
groups V, VI and VII respectively.
Experiments have confirmed that in an aquarium eels kept at
favourable temperatures show considerably higher rates of growth
than do eels in the wild. Glass eels which were kept in an aquarium at a
temperature of 23 °C and fed on pieces of marine fish, attained lengths
of 35"8 em and more over a growth period that lasted from April to
November (Meske, 1968, 1969). A glass eel kept in a 60 litre container
at a temperature of 23 °C reached a length of 37 em in I4 months
(Dreist, 1968). Another aquarium owner reports that within a year a
glass eel had increased to a length of 40 cm while on a diet containing
90% Tetramin (an artificial food stuff) (Fisch u. Fang, 10, 167, 1969).
The author (Tesch, unpublished) has also observed that glass eels
kept in fairly favourable conditions at room temperature can be brought
to a length of 30 em within six or seven months by feeding them initially
on brine shrimps (Artemia salina ) , and then on Tubljex worms and
fresh mussels. It seems, then, that in northern and central European
waters temperatures are such that the eel's growth potential is in no
way fulfilled.
It is obvious that growth rate is greatly affected by the quantity of
food available. Food quantity varies with population density and is
thus directly dependent on it. For the yellow eel, this situation is
accentuated; as a loner, the yellow eel is in strong competition with its
contemporaries for both food and territory. When several growing
individuals are kept together in one aquarium it is possible to observe
their fights for dominance. Over a period of time one individual comes
to dominate the rest, and then begins to grow much faster than the
others. The animals at the bottom of the hierarchy waste away and die.
This sharp division in growth is the reason why. in pond culture,
eels often have to be kept in groups according to their sizes (see p. 337).
Interspecific differences in growth

Studies of growth rate have been carried out on only a few species
besides A. anguilla. Comparative data are set out in Fig. 18. From these
it can be seen that also within the species A. rostrata there are consider-
able differences in growth rate. Particularly striking is the good growth
rate of eels from the Bermuda Islands. This may be due to the relatively
high temperatures in Bermuda compared with those in the other
regions studied. Growth rates in eels from the Canadian lakes also
appear relatively high (Smith & Saunders, 1955), but they are based
on studies of the scales and are therefore unreliable.
The large differences in growth rate between eel populations of
one species make it difficult to undertake interspecific comparisons,
except of course, when one author has studied different species under
similar environmental conditions. This, in fact, appears to be the
case where the two species of New Zealand eel are concerned. In these
species a size difference develops after the fourth year of life
198 THE EEL
Table 18 The growth of different non-European species of eel compared with growth
in the European eel [the values for which were obtained from rabies compiled
on the basis of results from various authors (after Penaz & Tesch, 1970)]
Species Watt!' body and geograph- S" Agegroup (length in em)

,cal po~irion a I II III IV
A. rostrata Bill's Lake, Canada Jo + 92 25'8 29'7 3H

A. ranrara Crecy We, Canada 00 + 99 21·6 2)'7 31 '4
A. ronrata Potters Lake, Canada 00 + 99 2)-8 )O'j 34"5
A. rostrata Ottawa River, Canada o-o + n 19'7 21'2 25' S
A. rostrata Lake Ontario n 31 "0
A. rourora St. Lawrence River n
A. rostrata New Jersey streams, USA &0 + 99 c. 17
""
A. rostrata Bennuda c,23 3l 43 '7
A.japomca Japan 27"9 31-6
A. australis
schmidti New Zealand &9+ 99 "7 7'8 ,,' 17'5
""
,,'
A. dieffenbachi New Zealand &9+ 29 ,,8 8'7 10·8 16'1 20"4
A. dieffenbachi New Zealand <19+ 22 6'0 11'0 16'0 19'0
A . ntbulosa
,,'
nebulosa
A . anguilla
Calcutta
Average value ""
&2+00 + 92 ,'.
1)"0 21'4
15'5
26'9
23'2
32'6
26'4
and ultimately increases to a difference of 10 ern between eels of a

similar age group. This difference is all the more surprising because
A. dieffenbachi, which grows to a larger size, also has a slower growth
rate. Recent investigations (Burnet, 1969b) do not contradict earlier
findings (Cairns, 1942a). Further research is needed in order to establish
to what extent these differences are environmentally or genetically
determined.
Apart from the examples cited above, different species of eel tend
to show very similar patterns of growth, and even the tropical species
A. nebulosa nebulosa fits in well with the general picture formed by
the other species. Comparative experiments carried out under
controlled conditions are necessary before definite differences can
be established.
Theoretical pattern of growth

Figure 77 represents the mathematically calculated course of growth
in males and females from two different eel populations. The most
suitable formula for male A. nebulosa nebulosa from the estuary area
of the river Hooghly was found to be that of Bertalantfy. These cal-
culations indicated a maximum length of 68'3 ern in this species, a
figure which corresponds very well with data on the lengths of male
silver eels of other species. It is possible that A. nebulosa grows to
slightly greater lengths than do the males of other species (see p. 190).
Agegroup (lmg lh in em) A UlhoT

V VI Vil VIIl IX X Xl Xil XIlJ
39'3 43'9 4S'O 5)'0 58'2 Smith & Saunders, 1955

34'2 36'0 43'0 47'2 48·S Smith & Saunders, 1955
'41 '1 47'7 54"7 56 '5 60'7 Smith & Saunders, 1955
...
29'0 27'6 33·8 38-6 33'0 Hurley, 1972
5°'9 49'S
58'8
53'2
61 '9
53"2
64·8
65'S
68·6
,0
,
7S' 5 78'7
74'6
83' S Hurley, 1972
77" 1 Larouche It 01., 1974
" "
6, Ogden, 1970
,0" " 34 37 4S
Boetius & Boetius, 1C}67b
40 '3 45'2 52'9 56·, 65·8 MatSui,195 2
3 1'2
25'2
24'0
39'5
3 1' 5
47·8 C, 57
39'0 47
"" 6,73 " "
73
Cairns, 19411
Cairns, 1942a
McFarlane, after Cairns, 19411
37"4
3°'0
43'5
33'7 38'6 43"9 jl ·8 54"8 ,,'0 7°'7
Pantul u & Sinsh, 1962
72'1 Penn & T esch, 1970
The Bertalanffy formula was not suitable for calculating the course
of growth in female European eels from the Lower Elbe and Alster
regions (data from Ehrenbaum & Marukawa, 1914). Hohendorf (1966)
was unable to calculate a maximwn length using these growth figures.
As Fig. 77 shows, the growth curve for the Elbe eel does not flatten
out with advancing age; on the contrary, it continues to increase,
Thus, the Ford-Walform function seems more suitable for describing
the course of growth in this species. It is unlikely, however, that female
eels do not show any decrease in annual growth rates nor a gradual
approach to the maximwn calculated length. Larger eels, which were
not available at the time, must be studied in order to demonstrate
such phenomena,
Length and weight

Weight is particularly variable in such an elongated fish as the eel.
Good feeding conditions or a well filled intestine can cause the weight
of an eel to be twice that of another specimen of the same species and
same length. _For this reason, comparisons of growth rate are always
made on the basis of length, Nevertheless, comparing weights can
be very interesting and it is important to be able to provide conversion
factors for use in commercial fishing projects.
A large amount of data on eel weights from various investigations
have been combined in Table 19. More data on weight in A. rostrata
200 THE EEL
Table 19 Weights of eels from diff~r~nt populations and species in r~lation to l~ngth
and, in part, to sex and h~ad form; A, anguilla unless specified
,•
0-,
7 0-,
0-7
9
,-,
0-'
,-, ' -0
"
" ,-.
,-, ,- ,
' -7
"'3
,.
'7 3-3
"
"
3'
' -3 , 3-'
,6
,.,"
IS') 6-3
"
7
'7 6-, 9 6-,
"'9 ' -3 "" 7-'

'-7 II 'S
" 9-3
" 12'1
11 ,8
" " " fl ' l IS'1
"'3 1, '1
"
'3 '4 11 '3 19'2
" 18'5
11'S
"'-9
18'4 I S'S
17'9 "'3 '3
15 '4
11'4
14'0
16,6
" ,,-, 25'0

" 20' 1 10' 1
,,-,
" lS '6 16,6
"3'
25 '4 2) '2 22 '2 2( '8 33'9
'7 30'8 21 '2 25'S
"
26,8
"
'9
27'8
,0
33 29'2
31 '8 31" S6 '3
30
"
37 '2 34'9 38 '3 36'0 33'2
3' 41 'S 42 '6 434 0'0 37'S

3' 46 '4 48'4 45'S 42'9
33 46'6 36'8 62'1 52'1 45'7 48 '2 45'6 72 '9
" "
34 60-6 So'S 17 54'S 51'S
"
51'S 51'S 65'S 55'6
36
37 79'7
85'S
.,-,
66~ 62 ' (
7 2'4 6S')
82'7 76'S
3'
39
40
60-,
" '"
"9
'"
&2 ,8
90'
POST-LARVAL ECOLOG Y AND BEHAVIO U R 201
0-, ,-,
,.,
'-7
37
.-, 3
•,-,
3-7
7-g
g-, ,-,
,-g 6-,
7-7 ,-0
10'6 ' -7
11 '2 10'3
17'2 15'3 1)"6 11 ,6
15"9 14'7
20'S
20'6 24'0 14'0
27 '8 27'S 29'7 23'0 20'0
3°'7 30'S 23 '0
36 '7 36 '3 31 '4 28'0
4S'S 37'S 31'S
,g
39'2 5°'7
5°'7 42 '1
53'7 45'6
62 '8 5 1'3
,,-0
46'0
S3'6
,6'2 61'3 6]'9 54'9

77'1 66-6 66 68' 4 62 '7 6" 0
79 '2 71'S 66-. 81 '0 59'0
8S ,8 7806 " -3
,.
86 88'1 76'7 9S'o 59'0
'" 78'0
go-'
"'. ,.-0
,,6
". ".
"7
"" ' 37
'04
202 THE EEL
Table 19 continued
,. ,,6 ,,6 ."

"0
"""
'"
.,.
,,6
",647
.06
' 49
.,0 ." .,0
."
."
.,.
'49
.6,
,8 '" '77
'7'
.66
49 .8, .8, .8,
,0 '0,
,. .,6 ",
"""
'44
,80 '" '47
,66
",6 '"
,.,'",,8
'9'
",8 ,6,
"
60 377 ,60
6. ,80
..
6,
6, 437 ""
,8,
,,8
6, ,,6
66 ,,0
6, ,8.
,6,
""
...
70 7'9
88,
"8,80 1146
1434
'" ' !l 6 )
"
"0 1884
'"" 0
-
- • -. 1 ~
- ...
~ :;;-~
~
~ ~ ~ ~~
:g-•
-'-t:;: 1~
~ ~
00 ~ '"0 ~ 00 -- :
---,.
.~
.'
~~
..... "'0 .... 00 00 ~ ::!. ~
. ~PiP
~ ;, -~
-a't>:'& >:,& 00 .; ~ ~ ~ t;
l! : !:i
- -
.. "'; .. ~ .. "'0 .. .... ~'O~
~
.~
~
.~>: ~:f ~ .;
e .. o.o <> ..t:
~
,
'. ,
~
' :.til ~ ~.::!.::~~i

~
~. - i -It a
~
"
.
.~ ~ "'O~
, . a ., ,~
~ - ..o::~"'O~
l~ ~£, ~to'o,1t>'o, .0.., ! ~
J! -, ~ ~ l~l;
.2e ,~ ~ ~
~-.
aea~
:<: .. ..... .t:. a::
.N .e<h .§ <h E~ _~ :<:.. ~e .. .-:a::
.~
...... E'"
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.... ..., .... 'Ii ..0::'..0::.", 0. '" 0> ' t
:. ~
~E
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.
.. 0. .. :.1
'.t~ '.;i·E .~~ .~~ .2~ .2-1 .5 ~ lE~ ~i
"~
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."
~ .E 0> '5 .2,~ . 2,~
'G. l "
~ ~ ~~ ~~ l· . . ",
~ ~
~ - a:: ...... a:: ..... a:: ..... l:t: ..... ~ ~ ..J~"I:! '- ~ ~ "I:! ...... ""I:! ...... " ..... ~ ~
'"
,60
..", ,",'"'".. '"'" '"
'47
,
'53 ,,0
,,0 ,,0
", ," ,,0 ",
'" ".
'"'" '"
'94
,gg >0, ' 99 '77 ,6,
,So
'"
' 70
'" >0,
'"
"" '"'44 ".
'" ,So '"
'"
,.. '" ,,0
'" "0 '" '"
," ". ,,0
," ,,0 ,..
33'
'" ,0> ,60 '"
'0'/ '90
'"
'"34' '34
,60
J37
'"
33' '"
,So
34' ,00
'",86
'79 ," '"
,>0
,68 m ',>0
"
'" '"
370 330
'" '"'" ,,0 "0 47J ...
44'
'"
,,0 400
,0,
,60
"" ," ,00 '", So m

'" '"'"
",
'"
,"
", '" 1036
,,, 12 79 1475
"00
,,60
1587
,,"
,,60
1530
204 THE EEL
can be obtained from the length-weight regression line: eels from

Lake Ontario, individuals 48 to 79 em long log 10 w = 2'9810&10 /- 5'59;
individuals 83 to 108 em long log10 W = 3'77 log10 /- 7'88 (Hurley,
1972); eels from two different water bodies in Newfoundland,
individuals 16 to 84 em long log w = - 3'1797 + 3'270610g 1,; indivi-
duals 29-57 em long log w = -2'8955 + 3'0812 log I, (Gray & Andrews,
1971) ; eels from the St. Lawrence River, individuals 56 to 110 em
long w = 4'461. IO - s. P'OI (Larouche el al., 1974). These data show
how greatly weights differ among populations from various
water bodies. Large differences even occur from year to year or from
one season to another (see also Marcus, 1919). Therefore, it is often
difficult to judge whether differences between two or more populations
are not perhaps being obscured by annual or seasonal variations. Other
problems include sex and age differences, differences in head width,
and the fullness of the gut.
Two srudies, each at different times of the year, indicated seasonal
variations in weight (Marcus, 1919). Eels caught in April were lighter
than those caught in September. Good feeding in September compensa-
ted for a reduction in weight which occurred in winter as a result of
unfavourable conditions. In comparison with eels from the lower
stretches of other rivers, such as the Elbe and the Severn, eels from
the river Star were lighter than average.
Tropical eels also show certain seasonal variations in weight. For
example, in A. nebulosa the condition factor, k, was 0'13 and o· 14 in May
and October respectively. In the months preceding and following May
and October the value of k was 0'01-0'02 units higher. Seasonal
differences here are presumably due to the effects of the monsoon.
The weights of eels in Lough Neagh in Northern Ireland (Table 19)
show quite remarkable differences from year to year. In 1965 the eels
were heavier than in 1966. The sheer quantity of data on which these
results were obtained, prevents one from doubting the results.
It is particularly significant that differences in weight are found
between male and female eels; comparisons between eels from different
waters where data are only available on males in one instance and on
females in the other, are not valid. Interesting weight differences
between the sexes can be seen in data derived chiefly from northern
Gennany (Table 19). Here, and in two of the three rivers srudied in
England (Sinha & Jones, 1967c), males, panicularly those in the upper
length groups, were heavier than females. The observed differences
are easily explained if one.remembers that males which become sexually
mature at a length of about 30 em, have a maximum accumulation of
99 A. anguilla
00
.',
,,,
,,, .
,,•
,,
,•
,,,
,
,, .
,,
20
, .
,
,,
1 2 3 4 S 6 7 8 9 10 11 Agtin~ts
Fig. 77 Mathematically determined growth curves. Bertalanffy's formula L , = 68·3

l-e -O· 14 UI {I- o·m 4)was used in the case of male and sexually undifferentiated
A. nebu/osa from the river Hooghly near Calcutta. Crosses represent
observed values (after Pantulu & Singh, 1962). The Ford/Walford
formula L ,+ 1 = 3.19 + l ·o84 · L , was used for female and sexually un-
differentiated A. anguilla from the Lower Elbe and Alster. Circles represent
observed values (from Hohendorf, 1966).
reserves, and are thus bound to be heavier than the females when they
reach the stage at which the spawning migration begins. Figure 78
shows, among other species, the dependence of weight on length in
A. mossambica. The curve becomes steeper after the 100 em mark.
Females of this species reach the silver eel stage when they are 100 em
long; they also become relatively heavier (Jubb, 1961),
Mention should be made of the connection between head width and
weight. Research on eels from Lake Sakrow near Potsdam (Table 19)
has made it possible to compare these two factors. Results from this
area show that, contrary to earlier assumptions, broad-headed eels
are lighter than narrow-headed individuals (see also Torlitz, 1922). An
explanation for this difference is that, with advancing age, the relative
206 THE EEL
w ~ ~ ~ ~ ro ro 00 00 ~ 00 ~ 1~ ~ 1~
m •
,
, Afl?nlla marmorafa
Anguilla,' 9
/abJafa/
,
8
,I 8
I
,
I
,
I
• I
•
,
I
I
I
I
Angul1f6
• ,
.I mossombica
I
, I
,
.I
I
I
I
,
,, / /
/ ,'
2 , , 2
-- '" " .....-' : ... " AnguHla anguilla
--.-;~.;;,~;-:-.::.~.-:.-.
-... :::~~.-' .. -
~ W ~ ~ ~ ro m 00 90 ~ ~ ~ 1m lW IW
ll ~thlcm)
Fig. ,8 The relationship between length and weight in mree species of African eel
(after Jubb. 1961), compared with the European eel (compiled from data on
A. anguilla in Table 19).
growth rate of the eel's girth' ... increases independently of the length.
It thus follows that a slow growing, older eel will be heavier than a
fast growing, i.e. younger eel of the same length' (Rahn, 1955a). And, as
results have shown, broad-headed eels grow better than do narrow-
headed individuals (see p. 193). However, Micheler (1967) could find
no differences in weight between eels with different head shapes.
This evidence indicates that rapid increases in length are not neces-
sarily correlated with rapid increases in weight. Further proof of this is
provided by Table 20, where the weights of eels of the same length
but different ages are compared. The figures show that older eels are
heavier than younger individuals of the same length. Of course, it is
Table 20 Comparison of weight in eels of the same length but of different ages
(without regard to head form) (after Rahn, 1955a)
Lmgth in em Aglgroup
V VI VII VIlI IX X XI
53 'w '40 '35 '60 '70 3"

,8 3'0 335 3'0 "0
6. 3" 36,
'" 4~ 4"
PO ST-LARVAL ECOLOGY AND BEHAVIOUR 207
W · [00
Table %I Length-weight coefficient k~ t' of eels
(A. anguilla ), which ascended into the Elbe
reservoir through the eel pass near Geesthacht in
August [964 (Tesch, unpublished), and of eels
(A. nebulosa nebulosa) from the Hooghly River,
Calcutta (Pantulu, 1956)
w = weight, I = length in em
Number Gemhachl Lerv: th Hooghly

ralWe River
k-'f)o./ue k-vo./ue
, 0"17
0"11
6"0- 6"9
7"0- 7"9
0 "14
3 0"10 &"0- &"9

l 0'", 9"0- 9"9 0" 14
• 0" 10
0"10
10'0-10"9
11"0-1l"9
O"IS
3 0"12 12"0-12"9 O"IS

3 0" 11 13"0-13' 9 0 ' 16
• 0"11
'.
14 "0-14"9
" O"U
0"11
15"0-15"9
16"0- 16"9
0"16
0" 17
0"12 17"0-17"9 0" 17
"9 0 "" '&"0-1&"9 0" 16
" 0"11 19·0-19"9 0"17
" 0"11 20·0-20"9 0" 17
", 0" 12
0"14
21"0-21 "9
22"0-22"9
0 "16
0"17
,3 0"12
0"1&
23"0-23"9
24"0-24"9
0"17
0"17
O"IS 25"0-2 5"9 0 ' 1&
26"0-26"9 0" 18
0 "11 27"0-27"9
impossible to tell whether head width or age is the primary factor here.
It is clear that older eels must be relatively heavier than younger fishes,
because the fonner are nearer to the silver eel stage than are the latter
and, as already mentioned, silver eels are heavier than yellow eels
(see above). Even during the adolescent growth of the yellow eel
stage, relative weight is already beginning to show considerable in-
creases with increasing length, This has been demonstrated both
in the European eel and in one species of Indian eel (Table 21 ).
Bearing these facts in mind, it seems almost impossible to demonstrate
interpopulational differences in weight which are not subject to in-
fluences of the sort described above. Comparisons of the different
results in Table 19 do, however, reveal a further striking difference:
eels from coastal areas are usually lighter than eels from inland waters.
Eels from English rivers, from Lough Neagh in Northern Ireland,
from Lake Sakrow and from an aquarium all have high average weights.
208 THE EEL
In contrast, lower than average weights are recorded for eels from
all coastal and near coastal waters, e.g. the North Sea, Fohr Island,
the Baltic, Kurisches Haft', predominandy coastal waters (Marcus,
1919) and possibly also the Cannargue. However, ·it is difficult to
detennine the extent to which factors such as head width are involved
here - as already mentioned, Heligoiand eels are an example of extreme
broad-headedness. The question arises whether eels from salt or
brackish waters are relatively lighter as a result of osmotic factors
(see p. 42).
Table 19 and Fig. 78 provide a comparison of eels of different
species. The most reliable difference appears to be that found by
Cairns (1942) between A. australis and A. dieffenbachi in New Zealand.
Research by Woods (1964) confinns that members of the fonner
species are lighter than those of the latter species. Woods found a
weight-length ratio of w = 8'03 1\3.18. 1 0- 4 for A. australis and w =
S·7Slt 3' 38 .IO-4 for A. dieffenbachi. According to these fonnulae A. dieffen-
bachi is only heavier than A. australis when it is over 10 cm long ;
below this length it is lighter. When compared with other species
of eel, A. dieffenbachi also appears to lie well above the average weight
(Table 19). Quite considerable differences have been found between
the three species of eel from South Africa (Fig. 78), though it should
be noted that the data on weights in A. nebulosa lamata were collected
by a different author (Frost, 1954). The two species studied by Jubb
(15)61 ) appear relatively heavier than A.n. labiata and A . anguilla. The
difference between A. marnwrata and A.n. lamata over 120 em long
amounts to 2-4 kg, i.e. 30-40%. This difference is so high that it
could be considered species-specific. The larger individuals of A.
"!annorata and A. mossambica studied by Jubb differ considerably
in weight, so that the question of genetically detennined, differential
weight increases may arise here as well. Unlike A . nebulosa labiata,
which, according to Fig. 78, is relatively light, A. nebulosa nebulosa
[according to Pantulu (1956)J is well above average. At a length of
70 em, A. nebulosa nebulosa weighs 1748 g, i.e. twice as much as, for
example, A. anguilla; this also suggests a genetically detennined
difference.
3.3.3 Location, habitat and behaviour patterns

This section deals with various results and isolated observations
which could not be taken into consideration in the preceding sections,
nor in the specialized chapters on sensory and behavioural physiology,
fishing techniques and yields. It deals primarily with features that

are peculiar to anguillid eels as compared with other species of fish,
but it also touches on behaviour patterns, well known in themselves,
which exemplify the eel's special characteristics particularly clearly.
Unfortunately it is impossible to mention every one of the many
interesting observations in detail, and only a selection are dealt with here.
Refuge in tubes and hollows, and contact with members of the

same species
A most striking feature of the yellow eel is the lengths to which it will
go in order to avoid light, and to remain in physical contact with its
immediate surroundings (see p. 70). Negative phototaxis becomes
even more marked with metamorphosis from the glass eel into the
yellow eel stage. A tube placed in an aquarium is bound to be used
by eels as a hide-out during the daylight hours. In the evening,
particularly when it begins to get dark, the eels leave the tubes and
swim around actively (Mohr, 1970). The eel's tactile tendencies are
emphasized by the fact that, during the day, several eels may squeeze
into the same tube. Such behaviour is seen even when several tubes
are available simultaneously. Thus, many tubes may be standing empty
while several eels, tightly packed together, can be seen looking out
of a single tube.
The eel shows similar behaviour in the wild, i.e. it swims around
at night, but during the day it hides (Fig. 79). During fishing operations
in lakes in Schleswig-Holstein it was even reported that 'eels construct
tubes in the mud, which is a metre thick in places. These tubes, which
are diagonal to the surface, fonning angles of about 45°, resemble
a fox's den or a rabbit's burrow at the open end.' The eels look out
of these tubes with about two-thirds of the head protruding from the
entrance (Krause, 19(1 ). It is, however, doubtful whether the tubes
had actually been constructed by the eels. It is possible that funnel
shaped openings had developed as a result of the eel's head movements
and would collapse just as soon as the eel left its hiding place.
Observations from another source suggest, nevertheless, that the
eel is capable of constructing its own burrow. During dike excavations
in the Lower Ems area, a middle-sized eel was seen in a ditch subject
to a 1'5 m tidal change in water level. The eel was caught and, when
digging continued, a metre long tube was found which led diagonally
downwards, ending in a spherical hollow. Here, more eels, which
could no longer escape, were found. Several of these tubes were dis-
covered, and at least 15 eels were recovered. It appears that other
210 THE EEL
Flg. 79 Eels in an edible mussel (Mytilus edulis) field in the Baltic (Photograph:
Krause).
species of animal, wn;dl might have built these tubes and then deserted
them, do not occur in this region (Schnare, 1960; see also Rudsinke,
1960).
The following report, which has a certain fantastic ring to
it, describes how eels creep far into cavities and may even contribute
to their formation (Benecke, from Walter, 1910) : 'Between 1846 and
1847 a large weir was built on the river Brake near MUhlhof, just above
Rinel, in order to provide water for a large complex of fields. A sloping
platform of planks, about 100 paces in length, was constructed below
the weir. Its purpose was to prevent powerful waves of water from
washing over the ground and river banks when the sluice gates were
opened. This boarding consisted of two layers, an upper layer of three-
inch planks, and a lower layer of two-inch planks. The Mtihlhof weir
was so high (13m) that the ascending eel fry were completely prevented
from reaching the upper stretch of the river Brake and the adjoining
lakes. The number of eels caught above the weir - which was previously
very high - was gradually reduced to almost nothing. The construction
of the weir and the sloping plane was completed in 1847. In 1852 the
upper layer of planks began to lift away from the lower layer at various,
randomly distributed places, with the result that the upper layer had
to be removed for extensive repairs. The cause of the buckling was

immediately apparent. Thousands of little eels, each no wider than
a finger, very pale in colour through lack of light, and mostly more
or less squashed flat, filled the space between the two layers of planks;
their concerted force had caused the upper layer to weaken. As fry,
these eels had pushed their way between the layers; they had found
enough to eat and had simply continued to grow until their increased
volume caused the roof of their prison to burst open.'
Walter (1910) ciles further observations to illustrate the eel's tendency
to hide itself in the ground. The following example is important in that
it shows how the eel colonizes the tidal areas of the coast: a large nwnber
of people got together on a beach and dug away the mud from the
silt, ... 'and revealed huge nwnbers of very lively eels which were
lying, only a foot below the surface, on the firm sandy layer below'. It is
also a well known fact that eels bury themselves in the mud when ponds
are emptied; this habit causes considerable difficulties for fish fanners
(see pp. 332, 337). While studying the winter quarters of the lobster
in the estuary area of a river, Thomas (1968) also found American
eels hiding in the mud. I t is unlikely that these eels were trying to avoid
drought; they were probably sheltering from the unpleasant winter
weather.
Digging into the ground is also a form of defence reaction seen in
eels when water levels begin to drop. If the eels remain on the surface
they are exposing themselves to predators and possible dehydration.
Some animals use the tidal current to escape from areas that are drying
out; others, small eels in particular, remain behind, as observations
have shown.
The tendency for eels to cluster closely together is clearly not restric-
ted to tubes and tunnels. It is reported that a group of 15 to 30 A.
rostrata massed together in the outflow of a lake in Nova Scotia, forming
a cluster of about half a metre in diameter. They Jay, without moving,
at a depth of 1'5 m, but disentangled themselves at once when eel
spears were thrown into the water. Drifting eels have been seen in
similar clusters, forming a ball estimated to be two metres in diameter
(Medcof, 1966). Such 'gatherings' of eels are not limited to yellow eels
alone j silver eels show similar behaviour. There are also reports of
European silver eels forming clusters (Nilsson, 1860).
Water depth
There seems little point in asking the question 'At what depth does the
eel prefer to live in salt- and freshwaters?' One comes across eels of all
212 THE EEL
sizes at all depths. When using electrofishing tedmiques in shallow

streams, one frequently catches large individuals in the very shallow
regions. However, in general the majority of smaller eels are found in
shallow waters and larger animals occur at greater depths. This is the
case particularly in areas where the water level remains constant.
Population studies in coastal waters to the north of the Isle of Fiinen
revealed that eels over 20 em in length were commonest in weedy
regions where the water was between one and three metres deep.
Animals of 10 and 20 em length were found in the shallower waters
(Mulls, 1967). Amateur fishermen recommend fishing along the peri-
phery of a bed of water weed, at depths between one and three metres
(LoebeU, 1966). As anglers are interested in catching the larger eels,
these regions can be regarded as the preferred habitat of such animals.
The author's own experience has shown that electrofishing in shallow
water very close to the shore, in lakes with good eel populations, will
result in catches consisting mostly of small eels which have hardly
reached the size of 'migrating stage' individuals (see p. 191).
The old notion that eels, particularly in the sea, only occur in coastal
regions and in relatively shallow waters (Walter, 1910), has not been
confirmed. Earlier it was assumed that eels were limited to the Wadden
Sea. North Sea trawler men are still of the opinion that previously eels
did not occur beyond the littoral regions. In Heligoland too, eels were
thought to be restricted to the regions at the base of the cliffs. This
belief probably arose from the fact that in earlier days eels were seldom
caught in trawl nets, and haddock long lines, baited with sand wonns,
placed in the open seas, also never attracted eels (Walter, 1910). It
would be interesting to know whether there have been any quantitative
changes in the distribution of eels, i.e. whether more eels have settled
in the deeper waters of the Heligoiand Bay than were present there
before about 1960. Environmental influences such as sewage, the
construction of weirs and the introduction of young eels from totally
different regions could have played a part in such a change. There is,
however, no doubt that depths of 20 m and more are suitable as penna-
nent habitats for eels. Confinnation of this statement is provided by
the size of the catches from trawl nets used in deep lakes (see p. 304).
It is, however, important that deep waters contain enough oxygen
for the eel. In lakes this is not always the case. Nevertheless, eels still
frequent such regions, as is shown by the fact that dead eels have been
found in traps placed on the bottom of small bodies of still water. In
addition, the deeper regions of lakes are often unsuitable for eels because
the temperatures there are generally considerably lower than those in
the upper regions. Thus, the deeper waters, which offer protection
from light, are by no means always the optimal habitat for the warmth
loving eeL In the North Sea, temperature is unimportant at depths
of 20 to 30 m. Here the water masses move about almost continuously,
so that vertical temperature differences of the sort found in lakes do
not develop. For this reason too, deep channels in rivers do not deter
the eel.
As long as temperature and oxygen requirements are met, there are
hardly any stretches of water which are not suitable for the eel since
almost all waterways provide the eel with the opportunity to dig itself
into the ground or to hide in hollows among stones or rocks. Of course
eels of different sizes show differing preferences in their choice of
substrate. Anguilla marmorata, for example, is to be found mainly in
the sand or under stones until it has reached a length of 30 em. Larger
individuals prefer hollows (Nishi & Imai, 1969). Plant stands, in parti-
cular the edge of weed beds, are an ideal hiding place for eels, as are
tree srumps, roots and anything else that has fallen into the water and
which can provide shelter.
Survival in air
As we know from numerous physiological studies, neither salt- nor
freshwater pose any limits on the distribution of eels (see pp. 24, 41 ) .
This animal can also survive out of water for extremely long periods of
time (see p. 24), though protection from dehydration is essential, and
life in the air is only a temporary expedient. Observations on the eel's
ability to stay alive in air and to find its way back to water are highly
contradictory. However) there does appear to be good and reliable
evidence suggesting that eels are able to negotiate sand banks at the
beginning of the silver eel migration. 'During the autumnal new moons
in the period from July to October, outlets from lakes near the sea
frequently become silted up as a result of storms, and a wall of sand is
formed. From time to time the Baltic Sea surges inland, flooding this
wall with saltwater. As dusk falls, the eels in the outlets force their way
to the wall of sand and, taking advantage of the remaining flood water,
slithter over the wall and into the sea' (Buckow, 1956). Of course, such
an opportunity to catch eels is not ignored. Bergmann (1970) reports a
similar case; he saw a silver eel three or four metres from the banks of a
small river in the Minelgebirge region of Germany at the end of July;
there had been a heavy thunderstorm that same afternoon. I t appears
that this eel had escaped from a mill pond situated about 70 m away
on a slope. The mill-race was no longer in use and was blocked so that
2[4 THE EEL
the water could only seep through it. The eel therefore had been forced
to migrate overland when the undergrowth was moist enough. Two
other cases, which also occurred inland, are also cited.
Basically) young eels undenake similar goal-oriented 'terrestrial
migrations' when they ascend rivers. However, one is not really justified
in calling these journeys terrestrial migrations because the degree of
moisture is always sufficient to indicate the proximity of water) so that
advance or retreat into areas with more water lies well within the eel's
capabilities.
It is conceivable that the eel is forced onto dry land as a result of
external circumstances. The eel's reaction to the sudden introduction
of sewage into the water is well known. I t escapes onto the bank in order
to avoid the unbearable conditions created in the water . Eels also desert
ponds that have had quicklime or too much liquid manure emptied into
them (Walter, 1910) and will leave an aquarium if conditions such as the
water supply or competition from other eels prove unsuitable. Eels will
escape from aquaria or other containers when their migratory drive
becomes too strong (Boetius, 19(7).
In the Japanese eel pond industry (Fig. 125), and also during experi-
ments in Germany (Seidlitz, 19(9), eels have been lured out of water
at feeding time (see p. 335). In aquaria too, eels will climb out of the
water and up the side of the glass when offered food. Is this ability to
collect food from the land of any importance in natural conditions?
Bergmann (1970) cites eleven reports from people whom he considers
to be reliable wimesses. Three of these reports deal with escapes from
closed areas of water (see above). The remaining instances, in Berg-
mann's opinion, are feeding forays. Although eels are seen repeatedly
at night or in the early morning in pea fields, and particularly on moist
pasture land, Bergmann (1970) rejects the idea that the eels left the
water to feed on the peas. Pea fields as well as meadow land and other
such damp areas should be attractive, because they would harbour
worms, snails and many other kinds of animals. It is said that the eels
find their way back to the water by following their slime trails. In
addition to these data there are many other reports, the validity of which
should be assessed carefully (Allgem. Fisch. ZIg., 27, 341 , 1972; 98, 230,
1973 ; Fisch u. Fang, 14, 694, 1973; IS, 66 and 204-206, 1974).
If eels are removed from water and put on dry land somewhere
nearby, they are unlikely to find their way back to the water. The follow-
ing experiment illustrates this. Twelve eels were caught on hook and
line in a tributary of the Weser. They were placed 4 m from the river
bank in a meadow gully with short grass and were left there for half
a night, after which time none of the eels had moved away. If the place
where the eels were deposited had sloped tow2Ids the river and not been
a depression, most of the eels would probably have managed to find
their way back to the water. Only two or three animals (out of a total
of 10) placed on flat meadow land succeeded in reaching the water,
and these animals were not guided by some instinct for water but were
simply favoured by chance (Rudsinke, 1960) .
Localization and homing behaviour

In connection with its tendency to inhabit tubes and hollows, questions
arise whether the eel lives in this type of shelter permanently, whether it
returns to the same hide-out after its nocturnal forays and whether the
animal remains for any length of time in one place or frequently shifts
its locality. There do not appear to be any studies directly concerned
with this aspect of the eel's life, although modem diving techniques
should provide such observations without any difficulty.
In contrast, numerous studies have been carried out on the extent of
the eel's home range . Data from streams of the Lower Mississippi
show that A. rOSlrala remain in extraordinarily limited areas for long
periods of time (Gunning & Shoop, 1962 ; Gunning, 1963) . For periods
ranging from a few weeks to a year, marked yellow A. roslrata living
in a stream 3 to 6 m wide did not move more than 60 m away from the
place where they were originally located. Of the 23 animals studied,
1 I were re-captured within 30 m of the place where they were first
trapped, and four were less than 15m away. Two animals, which were
caught a year later, were found no more than 50 m from their original
localities. In a stream measuring 8 to 12 m in width, some of the points
of recapture were slightly further from the original trapping area. One
eel, 91 em long, did not show any change in location during a three
months' period of observation (which took place in autumn). Various
collections made during the year showed that two other animals did not
move more than 100 and 150 m respectively, and on most occasions were
found much closer to the original locality .
Marking experiments have shown that, even in large rivers such as
the Elbe, eels are more sedentary than was originally thought (Mann,
1965). Out of a total of 47 eels recaptured, 16 were caught where they
had first been freed, 21 had moved between 10 and 60 m, and the rest
had moved further away. Thus, the home territory of eels in a river
such as the Elbe is much more extensive than in a stream. Other studies
have shown that eels tend to travel upstream for limited distances
during flood tides and to swim in the opposite direction when the
216 THE EEL
water level falls (Koops, 1962; Tesch, 1966; Aker & Koops, 1973).
Flood waters considerably extend the size of the river bed and eels
find ample food in the flooded areas. When the floods recede the eels
are forced to leave these areas if they are to avoid being stranded.
(If they do get trapped, they simply dig themselves into the moist
ground - see above.) The very nature of these unstable water bodies
enforces a certain degree of movement in eel populations.
When living and feeding conditions are more or less constant,
however, the eel has a very restricted home territory, even when there
is colonizable territory surrounding the animal on all sides. In 1968
an experiment was carried out in a rectangular, circa 3 x 4 km, area
with Heligoland at its centre. From a total of 350 eels marked internally,
17 individuals were recaptured; not one had left the rocky area round
Heligoland; 5 of these eels were not recaptured until a whole year had
passed, clearly indicating the constancy of the home area even over
fairly lengthy periods (Tesch, unpublished). These findings confirm
some earlier results (Tesch, 1967a, 1970): a similar constancy was
found in the estuary area of the Elbe, near Cuxhaven, and in an irrigation
lock on the North Friesland island of Fohr. Only I specimen out of a
total of 73 recaptured in the three areas mentioned above, had left its
original home. The region investigated on the island of Fohr covered an
area measuring about 100 x 500 m at the maximum, that near
Cuxhaven was relatively larger (about 30 km in length), while the region
around Heligoland Bay was bigger still (Aker & Koops, 1973).
If eels do change habitats, or even their home range, then this change
tends to take place in the transitional periods between sununer and
winter. Here we are dealing with a change-over from summer to
winter quarters, and vice versa, which is also seen in other species of
fish (Schiemenz, 1960b). Surface ice, cooling of the water, and ground-
ice formation produce envirorunental conditions which fish, and the
eel in particular, tend to avoid. The eel's sensitivity to hard winters
has been discussed in a previous section (see p. 21 I). In rivers, brackish
water and tidal areas eels seek out quiet side creeks and canals where
the water is deep enough to buffer the effects of winter, or they find
gullies in the main river where thermal stratification of the water
produces conditions on the river bed that are at least tolerable. In the
river Hunte, marked eels were recaptured at depths between I and
1'5 m, in the same places in the shallow side creeks as they had been
caught the previous summer, exactly one year earlier. In winter these
eels were found at depths of 2-2'5 m, and some were recaptured 5 km
from the place where they had been caught in the summer (Lubben &
Tesch, 1966). Reports on both eel species in New Zealand indicate that
during the winter these animals live deep down in the sand and mud,
places where in swruner they are never found (Cairns, 1942a).
Large-scale tagging progranunes in river estuaries and the Heligoland
Bight have confinned that during the swruner months eels do not
undertake any migrations at all. In September and October, however,
'large shifts in the populations' took place. From September onwards,
eels caught along the coast near Biisum were relocated in the coastal
regions of Heligoland or in inland waters. It thus appears that the
Wadden Sea area is unsuitable for overwintering. However, eels which
spend the swruner in the coastal regions ofHeligoland, also remain there
during the winter (Aker & Koops, 1970). Environmental factors in this
area seemingly cause few problems for the eels.
According to observations made by fishermen, the American eel
(A. rosrrara ) undertakes similar migrations in the spring and autumn
(Smith & Saunders, 1955 ; Medcof, 19(9). Reports of such locality
changes are particularly frequent in the coastal regions, but also come
from inland waters. In autumn, when the first frosts begin, yellow eels
migrate from the coast into freshwater; this movement takes place
over a period of about three weeks, and lasts for a similar length of
time in the spring when, in May, the animals return to brackish waters.
Fishermen from the Elbe estuary have reported very similar observa-
tions on A. anguilla, and eels used by the author near Wyk on the island
ofFohr for research into homing (Tesch, 1967a, 1970) were caught on
the basis of such spring migrations towards the sea. In the autumn,
these eels show similar activity, but moved from the sea into freshwater.
This autumnal migration is not, however, always in this direction ; it can
also be from fresh- to saltwater. The same applies to the spring migra-
tion. During marking experiments, Aleer & Koops (1973) observed
that in the Eider, a North Sea coastal river in Schleswig-Holstein, the
autumnal migration of yellow eels was directed downstream in the
middle reaches of the river, and upstream in the coastal regions . This
suggests that both populations of eels were migrating towards a common
area in which to spend the winter.
In both autumn and spring the two species of New Zealand eel
showed increased activity levels (Fig. 113A) during the yellow eel stage
and in running waters. The question arises whether the increased
activity seen in silver eels, not only in autumn but also in spring (Boetius,
1967), does not apply to yellow eels as well (see p. 221 ) .
In addition to the constancy with which the yellow eel retains the
same territory each summer and winter, this animal shows a further
218 THE EEL
H,lRUNGEH
C RllfQH PQ,n\
00 Pointsafni-CIlpturf
Q _ _~ RoutlrN)/llOt~",alpo;nt
of=pturHO rtitoSi pomt
<0'
E ,.
Fig. 80 Homing ability in eels transported to North Sea coastal regions. Above: In
1968, 1 [0 eels were caught near Den Dever, at the last dike on the Ijsselmeer
(Holland); the eels were marked and then released to the north of the island
of Ameland. The lines join the release point to the places where 26 of the 110
eels were recaptured (from Deelder & Tesch, 1970). Below : In 1967,636 eels
were taken from the rocky littoral of Heligoland. Of these 175 were trans-
ported to Wyk (Ftihr), and 461 taken to the Elbe estuary near Cuxhaven
(see the two thicker lines), where they were marked and released. Five of the
former and eleven of the latter were recaptured (Tesch, 1967a).
unusual characteristic which has yet to be seen so dearly in any other
species of fish, and which is reminiscent of the qualities shown by
homing pigeons. Yellow eels which have been transported great
distances from their home waters are capable of finding their way back.
This homing ability has been seen in all transportation experiments,
not only in river areas (Mann, 1965; Tesch, 19(6) but also on the coast
and in the open sea (Tesch, 1967a, 1968b, 1970; Deelder & Tesch,
1970). Results shown in Fig. 80, as well as those from a series of other
transportation experiments, indicate that most eels are still able to find
their way back from distances of up to 100 km from home. Beyond this
distance the percentage of successful returns becomes much smaller.
However, isolated individuals have been known to return from distances
of over 200 km. Impairment of the eel's sense of smell did not appear to
reduce homing ability. Nevertheless, odours could playa part in guiding
eels when they homed after displacement from one estuary to another
(Tesch, 1970; Vladykov, 1971). Since conventional marking techniques
could provide no satisfactory solution to the question of orientation
during homing. the author made a series of tracking experiments using
yellow eels provided with ultrasonic transmitters (Tesch, 1974a,
1975a; Fig. 84). The eight experimental animals were from north
Denmark, the Elbe estuary and the Baltic coast near Gdynia. The eels
were set free in the south-eastern part of the North Sea, and were tracked
for up to 17 hours; all the experimental animals, regardless of their place
of origin, swam in a south-easterly to easterly direction (mean course
126 0 ). In other words, only some of the eels swam in the direction of
their home ground. Laboratory experiments have shown that both silver
eels and yellow eels are capable of following a fixed compass direction
despite the removal of all hydrographical direction stimuli (Tesch &
Lelek, 1973a,b). This choice of direction can be influenced by altering
the earth's magnetic field (Tesch, 1974b; see also bibliography in
Tesch, 1975b). The orientation of the yellow eels may be explained as
follows: after being set free in the south-eastern North Sea, the yellow
eels. guided by the earth's magnetic field, take a south-easterly compass
220 THE EEL
course and thus arrive on the coast. Once they are there, those animals
which have not arrived near their home area, orientate themselves
along the coastline. Only about 50% of Danish eels would be likely to
find their way back home, because the other 50% would have to swim in
a westerly direction once they arrived on the coast. Many instances of
eels marked in the conventional way and recaptured on a south-easterly
course from the release point after transportation over distances of
200 km and more (DeeIder & Tesch, 1970) can be explained in this
way. The ability [0 get back to home grounds in the north-west (e.g.
Heligoland; Tesch, 19673) can also be explained. The eels in the
laboratory experiment showed a bi-modal choice of direction. If, in the
sea, eels swim in a south-easterly direction, they must likewise be able
to orient in a north-westerly direction; when eels from the east coast of
England were set free in the western North Sea, they swam in a north-
westerly direction (Tesch, unpublished). The speed and depth at
which some of the yellow eels travelled approximated to that of migrating
silver eels (Tesch, 19743, 1975a). Yellow eels appear to travel near the
surface of the sea.
3.4 Silver eel migration

The migratory movements of silver eels are much more clearly defined
than are the locality changes seen in yellow eels. Because of increased
commercial catches, dates of migration and the routes taken by the
eels migrating from inland waters to the coast are, by and large, well
known and easily plotted. Movement within the continent and on the
Continental Shelf region, which is more easily recorded than routes
taken in the open sea, is discussed in the following sections, the oceanic
part of migration on page 96. The two parts of the journey, however,
are not to be thought of as separate or distinct.
3.4. I Seasonal variation

As mentioned already) the fishing industry has for long known at what
time of the year the silver eel migration takes place. However, it is not
clear from available statistics what proportion of the catch is silver eels
and what proportion is yellow eels (Fig. 89). The monthly yield records
shown in Figs. 8 I and 90 provide a clearer indication of the dates of
the main sdver eel migration.
In the Dutch canals the greatest number of migrating eels is caught
during August, while in a river in Northern Ireland and along the
central coast of the Baltic, most migrating eels are caught in October
and almost as many in September; in the Ijsselmeer a peak is reached in
September. One would have expected eels from the mid-Baltic to
begin migrating earlier than eels from Northern Ireland and the
Ijsselrneer, since eels from Ireland are at least 1500 km nearer their
spawning ground than are those from the isle of Rugen (see also Walter,
1910). Thus it is very difficult to make any definite correlations between
an earlier or later start to migration and the latitudinal or longitudinal
location of particular populations (see p. 251 ).
In contrast, comparisons between inland waters and those nearer
the sea provide a more clear-cut picture. From Fig. 81 it will be seen
that eels in the Dutch canals begin to migrate earlier than do eels from
the Ijsselmeer, which lies closer to the sea. In the Baltic coastal region
near the Isle of Rugen it has been observed that, in the Greifswald
Bodden, which lies near the mainland, silver eel migration begins
two or three weeks earlier than in SaBnirz, and four weeks earlier than
in the area of Lolune which is even nearer the open sea. Furthermore,
increased catches indicating the onset of migration, reach a peak between
September and October along the coasts of Rugen but do not do so
until October or November in the sound (Meyer, 1938). Tagging
experiments (Martinkowitz, 1960, 1961), which show that migration
begins in mid-August in the region round Rugen, fit in with these data
and with information on the direction of migration .... 'There is
usually a temporal displacement in catch maxima from the south coast
of Rugen to the Tromper Wiek. On average, peak catches occur during
September in the Greifswald Bodden and during October in the
Tromper Wiek.' Apparent confirmation of this temporal shift in
silver eel catches from inland waters towards the sea is provided by
Fig. 89 (discussed more fully on page 251), though the data are
contaminated by the inclusion of yellow eels.
Migratory activity is also thought to occur in spring (Boetius, 1967).
There is a natural explanation for this in yellow eels, as has been
discussed above (see p. 217). Several possible explanations exist for
this activity among silver eels which are seldom caught in spring
by professional fishermen. The first activity peak in autumn, when
the eels start to leave their home waters, signifies the beginning of
migration. The second peak, in spring, may be a sign of spawning
activity in the Sargasso Sea which occurs some time in March. Why,
then, are laboratory eels inactive between the bouts of activity shown
in autumn and spring, while eels in the wild are migrating during this
period? There is a second possible explanation: animals which are
222 THE EEL
prevented, either naturally or artificially, from migrating in autwnn,

become inactive in winter so that they can resume migration in the
spring (Frost, 1950). Since large female silver eels predominate in
the spring migrations in the Elbe, for example, it is quite probable
that these animals are stragglers from the autumn migration. Parti-
cularly large eels were also caught frequently among the last migrating
eels in autunm (Table 22).
It would appear that migration periods do not coincide fully in
males and females. This difference first became apparent with the
discovery of a shift in the proportions of different sized eels on the
east coast of the isle of Rugen during the course of the autwnn migration
(Fig. 102). According to Fig. 102, mostly small eels are caught in
pound nets at the beginning of the migratory period. In the following
months the percentage of larger eels increases and reaches 90% of
the catch in November (see Meyer, 1938; Nolte, 1938). The fishermen
- - - Outch conals
- -- Ijsstlmotr
- - Rivtr B~nn
._._ ••. Eost o;o(Ist of Rtic.ltn
40
Apr. May. Jun. Jul. Aug . Sept. Oct. NO'.'. Dec.
Fig. 81 Silver eel migratory period, based on catches from fisheries at four different
places in Europe. Average eel basket catches from canals in Holland over the
period 1947-52 (Deeider, 1954); average eel basket catches from the
ljsselmeer for the years 1950 and 1951 (DeCider, 1970); catches from pound
nets on the east coast of Rugen averaged over the period 1928-36 (Meyer,
1938); ten-year average of bag net catches in the outflow (river 8ann) of
Lough Neagh, Northern Ireland (Frost, 1950).
believe that the large late arrivals have come a long way and originate
from waters furthest inland. It is well known that large female eels are
found mostly in inland waters while the smaller males occur nearer
the coastline. These observations suggest, therefore, that male eels
do not begin migrating earlier, but rather that they have a shorter
distance to cover before reaching the traps. In some Dutch waters
too, the average size of the eels caught increases during the migratory
period (Deelder, 1970). The percentage of large eels was nil in August,
I1 % in September, 26 % in October, 47 % in November and 60%
in December.
Data summarized in Table 22 also indicate that the proportion
of smaller eels in the Elbe continues to increase throughout the sununer
months but then to decrease in the autumn so that, by the end of
September, it is unlikely that males are represented in the catch.
Eels in New Zealand show marked seasonally defined patterns of
migration (Fig. 113). Naturally, activity geared towards reaching
the spawning grounds is centred around the late sununer and autumn
of the southern hemisphere. Monthly records show that males of the
two New Zealand species, A. dieffenbach£ and' A. australis, appear in
traps before the females. Figure 113 also enables one to draw
inferences concerning the temporal pattern of migration in these two
species, viz. A. dieffenbach£ begins migrating first and is followed by
A . australis. An earlier study, however, showed this sequence to occur
in the reverse order (Hobbs, 1949), so the differences may be of an
ecological origin.
In some Dutch waters there is a gradual decline in the average
length of male silver eels over the seasonal period, e.g. the average
length in July is 36'0 em, in August 35'5 em, in September 34'0 em
and in October 33'0 em (Deelder, 1970). The reason for this may be
that, as a result of a lower level of physical fitness, the smaller males
Table 22 Proportion (%) of silver and half-silver eels of different lengths caught in
the middle Elbe ar various times oCthe year 1958 (after LUhmann & Mann,
1961)
Lenglh group Dale of caplure

(= ) 11.). JO·5· 10.6. 41· 1J·7· 19.8. 24·9·
40-50 , ·8 73 10·6 16'9 '·0 10'2 0

51-60 , ·8 4"9 59'6 45'8 JJ"1 18 '6 12'5
61-70 23'8 37' 1 29'8 27'7 J5"1 4°'9 50'0
71- 80 42'9 10'5 0 '·0 19'6 10'2 29'2
> 8, 23'8 3' 0 3·' , 8 0 8·3
224 THE EEL
are delayed during migration and therefore pass particular points

at a later date than do the larger males.
3-4-2 Variations in the time of day, and the influence of light

It is well known that silver eels are caught mostly at nightj they are
in evidence from an hour after sunset until one and a half hours before
sunrise (Walter, 1910). On the southern coast of the Baltic it has been
noticed that when conditions are bad migration only takes place in the
evening, between sunset and midnight. In fact, during the period of
peak catches the majority of active eels is observed within a much
narrower period of time, namely after s~set and before moonrise.
Since peak catches only occur during certain phases of the moon
(see below), this active period is limited to between one and three hours
(Brautigam, Ig6Ia, hj see also Frost, 1950 and Lowe, 1952). A similar
situation is seen in A. rostrata (Medcof, 1966; Winn er: al., 1975;
Hain, 1975).
These studies indicate that, since the periods of activity are dependent
on the phases of the moon, moonlight probably plays a significant role.
The question arises whether, on lighter nights the eel prefers to move
at greater depths, if these are available. Eels are known to avoid artificial
light coming from the land (Rumphorst, 1930), and this fact is used
to guide the animals into fyke nets (see p. 322). The following observa-
tion was made on the coast of Rtigen: 'In pound net systems made up
of several chambers the seaward facing chambers in deeper waters
caught more eels during the full moon than did chambers in shallower
waters; when the moon was not so bright, the opposite was often found
to be the case. Of course, the wind also played a significant role ....•
(Nolte, 1938). In these conditions the observed increase in the number
of silver eels caught when the water was very turbid (Maar, 1947;
Frost, 1950) appears to be the result of decreased light penetration
rather than the consequence of any direct effect from the factors
causing the turbidity.
Edel (197Sb) has carried out experiments to analyse the effect of
light and shelter availability on activity in silver eels. He found that
total average activity per day increased when the eels were unable to
take cover behind objects in the water, with the result that the normal
increase in nocturnal activity was barely perceptible.
343 Correlation with lunar phases

It is known that eels are caught most frequently during the last quarter
of the moon, i.e. when the halfmoon is on the wane (Fig. 82). In the
• • o •I •I 30
%
Ijss~l_
2
0~101~202530
I.
."00"
N •.of
;1\
. \
."H •1,of•
60
./ .\
, . . Y .\.':.
,~'"
'"
"
: c..;-~-
' ,,"'1,:',._.:+-i---,;:-'--T.'---!'i:;----,:
010152025 30
., .,
'"
200
1.5t of Ru~tn
~
000 000
" .I"',·.. '
° ,I " I "I
00 20 25
• • 6 • •
I
Fig. 8% Average frequency of silver eels caught at various places in Europe during the
lunar month. Figures for the coast of Rugen were cruculated by Jens ( 1953)
on the basis of data from Meyer (I938); Rhine Gens. 1953 ); Ijsselmeer and
the Dutch canals (Deelder. 1954. I970).
previous section it was suggested that moonlight probably has some
influence on the daily activity rhythm. Statistical analyses of eel catches
from the coasts of the isle of RUgen, however, have raised doubts as to
whether light alone is responsible for the monthly lunar rhythm in eel
226 THE EEL
migration (Meyer, 1938). Active behaviour was even seen in silver eels
in closed tanks in the Lake District of England (Lowe, 1952). Later
research on escape activity in eels in a sealed-off experimental area
showed that, during the critical period of the waning halfmoon, 6 I %
of the silver eels tried to escape. During new mOOD, waxing and full
mOOD, however, the percentage of eels trying to escape was only 9% ,
13% and 18% respectively (Boetius, 1967). The curve in Fig. 82
representing catches in the Upper Rhine can be divided into lunar
phases with overcast skies and lunar phases with very little cloud
cover (Jens, 1953). In very doudy conditions no reduction was seen
in the maximum yields that are observed during the waning halfmoon.
Laboratory experiments by Haio (1975) have also been unable to
rule out the possibility that, in addition to light, other factors COlUlected
with daily rhythms and monthly lunar cycles might not also playa part.
Further analysis of the lunar month catch curve has shown that
it can be regarded as an ellipse (Jens, 1953). This may well be an indica-
tion of the planetary nature of the intensity curve. An interesting
correlation is revealed if one compares the catch curve from Riigen
(Fig. 82) with the number of meteorites recorded by radar near Stanford
in Onawa (Lang, 1972). The two curves are almost identical and
correspond to the catch maximum obtained during the last quarter
of the moon. However, a causal cormection can be ruled out here.
The various publications cited above suggest very strongly that
moonlight alone is not the sole factor determining the apparent lunar
periodicity of silver eel activity.
3-4.4 Hydrological and meteorological factors
Low pressure areas

There is one weather phenomenon which could not have a direct
influence on the eel and which cannot act through hydrological condi-
tions either. These are microseismic oscillations caused by low pressure
areas over the sea and which are spread out from these areas over
large distances by the sea bed (Deelder, 1954; see also Maar, 1947).
Analysis of 1000 fishing days revealed that microseismic oscillations,
with a periodicity of three seconds, preceded a large increase in the
catch. All other weather phenomena such as air pressure, wind direction
and velocity, temperature, rainfall and cloud cover could not be so
clearly connected with the increased number of eels caught. Studies
on the New Zealand eel have also indicated that microseismic oscilla-
tions might possibly be of importance (see below).
Water level and currents in n·vers
The most critical, weather-dependent, hydrological factor is, without
doubt, the current. If rainfall increases or snow and ice begin to melt,
there is an increase in water flow; water levels and currents also increase.
The number of eels caught during downstream migration is greater
at high water levels than at low water. This phenomenon has been
studied in the outlet areas of lakes both in England and Ireland (Frost,
1950; Lowe, 1952). In the river BaM in Northern Ireland the greatest
nwnber of eels moved downriver during periods of high water. In
the river Elbe as well, catches increased considerably when water
flow increased (Fig. 107). Table 32 shows that, while the daily average
catch was 10 kg per net over the year, 40 to 50 kg were caught during
high water levels (Luhmann & Mann, 1961 ).
It should be pointed out that yellow eels were also included in the
catch records for the Elbe. In fact the picture is often so confused by
other factors that reliable conclusions can only be reached after special
analysis of the data, e.g. in the Upper Rhine, catches were separated
according to rising and falling water levels during the lunar month
(Fig. 83). Lunar influence was dominant in both cases. However,
total catches during rising water levels reached a much higher level,
so that there can be little doubt that increases in water flow assist
eel movement.
Increased rainfall also brings about higher silver eel catches among
New Zealand eel species (Burnet, I 969a). These observations were
made in streams, and thus one can be certain that additional rain can
be equated with increased water flow. In rivers. on the other hand.
there is a greater time lag between rainfall and its effect on water level.
The author is not certain whether improved catches are caused by
increased water volume or whether they are the direct result of falling
barometric pressure. Thus the question of an influence through
microseismic oscillations arises here too (see above).
In general, authors tend to represent increases in catches as dependent
on rises in water level. This is almost certainly inaccurate. It is difficult
to imagine (except in some special cases) how water level could in-
fluence the nwnber of migrating eels. In the Dutch Polders, which
are usually kept at the same water level, silver eel catches vary despite
the constant water level. However, they do increase in relation to the
amount of water that is let through the sluices (Deelder, 1970).
The question arises whether it is increased quantities of water or
stronger currents that lead [0 larger numbers of eels being caught.
In the Elbe (see above), it was found that the nwnber of eels may
...,.,,
228 THE EEL
___ Alf~lIi"9 wattf Ls

" 0
- At ri,,1'IIj .Ottr 1' lItls
" 0
"'"
• 0
, 0
" ,
so
I \
I I
I
" , - -.I I '(

'-
0
?, , ~
o 2Q l5
"
Fig. 83 A comparison of daily stow net catches in the Upper Rhine at rising and falling
water levels, arranged according to the lunar month model (after l ens, 1953).
increase as much as fivefold during high water (Fig. 107). Doubtless

this corresponds to the increased quantity of flowing water. Similar
relationships are seen in the river Bann in Northern Ireland. However,
the current itself only increases at particular points, and then only
by It to 2 times its original strength. Thus, in river areas the greater
influence must be ascribed to the increased volume of water.
However, the height of the water cannot be totally without effect.
In some conditions it definitely has a decisive influence. For example,
when pools are flooded or when larger) static water bodies overflow
as a result of increased rainfall, the eels are given a chance to escape
into neighbouring streams and rivers. It is possible that in the past,
when rivers were not so extensively dammed and regulated, this factor
was more influential than it is today.
Tidal currents
Although current has little influence on the number of silver eels
swimming downstream past a particular point, it does have a
pronounced effect on the direction and speed of migration. Under
20'""
,J
certain conditions this influence does nOt stop when the eel reaches
the sea. In the North Sea river estuaries migrating eels are carried
passively seawards with the ebb tide and are known because of this
habit as 'drifters'. Their pattern of behaviour does not seem to differ
greatly from that seen during the river migration period, the only
difference being that they cease to drift at high tide. Drifting eels are
only caught at low tide in the North Sea river estuaries (see p. 285).
In contrast, yellow eels are caught at both low and high tide, depending
on the time of year.
Some idea of the movements of silver eels in the tidal stream of
the North Sea has been provided by tracking experiments using animals
fitted with ultrasonic transmitters (Tesch, 1972, 19743). The results
of these observations are combined and compared, in Fig. 84, with
data from yellow eels which were taken from the eastern coast of the
230 THE EEL
North Sea or from the Baltic, and then transported to the North Sea.
The courses taken by the silver eels over the ground show a bias towards
a northerly or north-westerly direction. However, the course is by
no means a straight one. If the current is in the opposite direction
to that in which the eels are swimming, the animals appear to make
very little progress. In some cases they may even go in the opposite
direction. If, however, the effects of the tidal current are discounted
and only the swimming movements are represented, the direction of
migration does appear more constant (see the small triangles in Fig. 84).
In general, silver eels swim through the tidal current, without changing
their speed or direction of movement, irrespective of the time of day.
In other words, silver eels do not use the tidal current in the same
way as do glass eels during their shoreward migration; the latter are
thought to drop to the bottom during low tide (see p. 139). Silver eels
have not been found on the sea bed, either in the studies juSt described
or in those by Stasko and Rommel (1974), who tracked silver eels fitted
with depth detecting ultrasonic transmitters in an estuarine area in
New Brunswick.
Wind and currents

In addition to tidal currents, other currents which also influence the
eel's movements, must be taken into consideration. In the Baitic, for
example, the tidal current is of practically no significance, whereas
wind induced currents playa very important part. Rumphorst (1930), in
agreement with Nordquist, states the following: 'On the whole, on-shore
winds are advantageous; they bring the eel nearer to land and thus into
the pound nets. Off-shore winds are usually unfavourable; they keep
the eels away from the coast.' Later publications (Meyer, 1938; Nolte,
1938) report that catches from the east coast of RUgen (Figs. 85 & 101 )
increase when an easterly wind is blowing. On the north coast of RUgen
the catches increase with northerly to easterly winds. In the Greifswald
Bodden, which is open to the east, eelS also become much more abundant
when an east wind is blowing. In most of these places west winds are
unfavourable to the fishery.
One might assume that a west wind on the west coast, e.g. near the
island of Hiddensee, would result in increased catches. However, this is
not the case (Rumphorst, 1930) :
'A very favourable "eel wind" for Hiddensee, in particular for the northern
half of Hiddensee, is an off-shore wind, in fact a sustained easterly and
north-easterly wind, may be even a north wind. Under such conditions
some of the eels travel with the strong coastal currents along the north
coast of Rugen near Arkona; they are temporarily diverted in a southerly
direction round the northern tip of Hiddensee and thus arrive in the
Hiddensee pound nets.'
Observations confirming the fact that relatively large numbers of
silver eels are caught in traps during an on-shore wind have been
reported from Lough Neagh in Northern Ireland, Lake Windermere
in England (Frost, 1950) and the Ijsselmeer in Holland (Deelder, 1970).
On the east shore of the Ijsselmeer, for example, 83'5 % of the total
silver eel catch was landed during strong west winds, while only 50% was
caught during moderate east winds. The author too, has observed that
particularly large numbers of eels are caught on the north shore of
Lake Miiggel near Berlin when southerly or south-westerly winds are
blowing.
With the exception of the island of Hiddensee, all these observations
seem to indicate that certain wind induced currents bring eels nearer
the coast. Attempts have been made to explain the situation on the east
coast of the Isle of Rugen (Nolte, 1938), namely, that on-shore winds
cause the surface water masses to flow towards the shoreline, and this
in turn produces a deeper counter-current which forces the eel to swim
along the bottom against the current, i.e. in the direction of the shore.
This explanation is based on the assumption that an eel swims along the
bottom, or at least in the lower water levels. However, even when this
theory was proposed, there was reliable evidence to make one doubt
the basic assumptions involved. In fact, at the present time several
observations have been made which show that the silver eel also swims
in the surface layers (see below). On the coast of Rugen. it has been
shown that tagged eels can be caught in nets at any depth, both near the
bottom and 2 metres from the water surface. Thus at least some of the
eels are carried towards the coast by surface currents. It may even be
that the majority are carried this way, otherwise it would be difficult
to explain why the highest catches are recorded during on-shore winds.
There is another possible explanation: strong on-shore winds cause
the water to become more turbid. Under the protection of this turbidity
the eel penetrates further than usual into shallower waters before its
negative phototactic responses force it to escape along the coast. How-
ever, with an on-shore wind, the eel does not arrive nearer the west
coast of Hiddensee because its westerly direction of movement (see
p. 237) usually keeps it away from this coast. It is possible that this
active movement to the west first becomes pronounced when the eel
reaches seawater (Tesch, 1974b). In this way, one could explain all the
different wind-dependent catches in inland waters. Here, the rather
23 2 THE EEL
more passive type of movement in mid-water levels predominates,

as stow net fishing in the river Elbe has shown (Kothke, personal
communication).
On the question of the dependence of eel migration on currents
in the Baltic, Westerberg (1975) has carried out tracking experiments
in the sea and claims to have found instances of dependence between
the current and the direction in which the eel swims. The rise and fall
of the water is thought to playa part, and orientation in the currents
could be on the basis of magnetico-hydrodynamic factors. These
possibilities have been discussed elsewhere by the author (Tesch,
1975b). Particularly low current speeds in the Baltic make such a means
of orientation unlikely.
Temperature and depth

According to fishermen in the Elbe estuary, when late summer extends
for a long time into auttlmn, silver eel migration is delayed and the
number of migrating eels may even be fewer than usual. This suggests
that certain minimum temperatures are necessary to initiate migration.
In a tracking experiment in September, three eels were equipped with
ultrasonic transmitters and their movements were recorded in the
Baltic (Tesch, 1976). One of these specimens showed behaviour similar
to that of yellow eels (Fig. 84); it swam in a south-easterly direction.
At this time the surface temperature was over 16 °c' The two other
eels, which were tracked at a later date when the temperature was about
15 °C, behaved in a normal way, i.e. they swam in a westerly direction.
The annual pattern of silver eel migration suggests (see p. 220) that
there is probably also a certain lower temperature limit. Migration
shows a steep decline in November. Only in exceptional cases does
migration continue into January, e.g. in the river Bann, after marked
delays in seasonal rainfall in Northern Ireland (Frost, 1950). Reports
from the same area show that, with the onset of frost, migration ceased.
Tracking experiments carried out in November on silver eels also
indicate that active migration will not take place below a certain mini-
mum temperature limit. As reported by Tesch (1972), an eel which
was set free near Heligoland, in water at a temperature of 9 °C, still
showed a tendency to swim in a westerly direction. Another eel, which
was set free two days later in the Elbe estuary near Brunsbiittel, allowed
itself to drift with the current, not only at low tide, but at high tide as
well, i.e. it allowed itself to be carried back again (Tesch, 1972). At this
point, which lay further inland, the brackish water had already cooled
down to 6 0c, I t was probably the excessively low temperature which
caused the eel to act so passively.
It is possible that temperature is also responsible for delaying the

onset of migration in deeper inland waters which take longer to cool
down. Migration in eels from a canal was delayed by forcing animals
beginning their migration to pass through a lakt:, over 10 m deep. Below
this lake the eels had to continue on through a shallow canal. The
majority of the eels were still here two month~. after they had left the
canal above the lake (Deelder, 1957a).
Low temperatures may also prevent the ee l from migrating along
the sea bed. Data on temperature from an ultrasonic transmitter
attached to a silver eel in the Baltic indicate that at night the eel swam
at depths between 0 and 15m or rested on th e sea bed CWesterberg,
1975). During the day the eel swam at much i:reater depths. Another
eel, which was fitted with an ultrasonic transmitter indicating water
pressure, was tracked by the author (Tesch, 15'76) in the same region;
this eel swam near the bottom for only very short periods and, over a
two day period in which it covered a distance of 100 km, it showed a
preference for an average depth of 17'7 m (the average depth of the
water was 41 .8 m). As will be described below: silver eels in the Baltic
approach the coast repeatedly. Presumably the lower temperature in
deeper waters prevents the eel from taking a route along the coast where
the water is too deep. This gives fishermen the (:hance to take advantage
of the passing stream of eels by using fyke n.!ts in relatively shallow
water (see p. 277; for further details on depth (.f migration, see also the
preceding section and last section of this chapu: r).
Salinity
Once the eel is physiologically adapted for migration, its osmoregulatory
mechanisms will also have changed (see pp. 24.. 42). It is quite possible
that the silver eel will not only tolerate increased salt concentration
(as will the yellow eel), but that it will in fact show a preference for
it. Logically one would expect the silver eel to ~:how behaviour contrary
to that shown by the glass eel, which seeks out freshwater. The glass
eel, however, does not seek out low concentrations of salt, rather it is
attracted by the smell of natural freshwater (st::e p. 77). Does the silver
eel show corresponding behaviour towards seawater, or does it actively
seek out osmotic differences, as is thought to be the case in the salmon
(McInerney, 1964)?
Experimental studies have shown that silver eels become positively
rheotactic when placed in seawater; in freshwater, on the other hand,
they are negatively rheotactic (Hain, 1975). Thus, in freshwater rivers,
silver eels swim downstream, whereas in saltwater they actively turn
against the current - in contrast to yellow eels which show no rheotaxis
234 THE EEL
in seawater. Silver eels with an impaired sense of smell did not show
such rheotactic responses in seawater. It can therefore be assumed that
silver eels detect seawater through their olfactory senses just as glass
eels can detect freshwater. An example which has already been quoted
(see p. 213) shows that this ability to detect saltwater instantly is
essential for migration from freshwater into the sea. In cases where
seawater spills over the sand bars of coastal lakes, eels will try to escape
from the lakes in the direction of the incoming seawater, and often
manage (0 reach the sea after exhausting overland journeys. Fishermen
have also confinned that eels will swim against the tide of inflowing
seawater (Deelder, 1970).
The activating effect of seawater on silver eels becomes particularly
significant when one recalls that as soon as the eel enters seawater a
certain directional tendency becomes apparent (Tesch & Lelek,
1973a,bj Tesch, 1974b). As field experiments have shown (Tesch,
19743), silver eels in the south-western North Sea take a north-
westerly compass course (Fig. 84). This constant direction is an appro-
priate one for finding the northern exit of the North Sea. However,
it is only appropriate when the eel has reached the river estuary and the
open sea. Before this point, i.e. in freshwater, the tendency to swim
north is not apparent, as laboratory experiments have shown (Tesch,
I974b). Stow net fishermen from the Lower Elbe have observed that
the northerly migration appears as soon as the silver eels reach the
brackish water of the Elbe, which flows in a westerly direction before
emptying into the North Sea. Here, silver eels are only caught on the
northern bank, an obvious consequence of the eels' tendency to move
in a northerly direction as salt concentrations increase. This choice
of direction, which is activated by the salt concentration, is probably
dependent on geomagnetic factors (Tesch, 1974b).
3.4.5 In subterranean waters

A further special characteristic of the eel should be mentioned here,
namely that not only young eels and yellow eels, but also spawning
eels travel for long distances underground during their spawning
migration. In view of the eel's many other peculiar qualities, this is
perhaps not surprising, although it is a behaviour pattern which is only
rarely seen in other species of fish.
Evidence of subterranean migration has been found in the Karst
regions of northern Yugoslavia, where a large number of rivers flow
long distances underground before emptying into the Adriatic. One
of these rivers is the Timavo j eels are caught in its estuary at the small
town of the same name. Two hundred yellow eels and three hundred
silver eels were marked with small cuts on the fins and were set free
at three places 28 to 42 km above the estuary. Twenty-nine eels were
recaptured between 40 and 188 days later; all were in the migratory
stage. It should be noted that the river flows 200 to 300 m below ground
and is only connected with the surface at a few places. Eels from the
river Timavo are light in colour, a well known phenomenon among
cave dwelling animals (Sella, 1929). The observed migrations are
perhaps not so surprising when one recalls that eels in epigean waters
show a tendency to retire into hollows and cavities (see p. 2(9).
The American eel ( A. rostrata ) may also cover large distances
underground. In the Santa Fe River, Florida, one collection produced
12 eels from below but only one eel from above a 'natural bridge' some
3 to 4 km in length (Hellier, 1967)·
3.4.6 In the sea, with special reference to the Baltic

The Baltic is particularly suitable as an experimental area for studying
silver eel migration in the sea. Extensive silver eel fisheries, matched
Fig. 8S Well-known spawning migration routes in the Baltic, based on the results of
marking experiments by several authors (Nordquist, 1904 ; Trybom &
Schneider, 1908 ; Maar, 1947 ; ManinkQwitz, 1960, 196 1) (O riginal).
236 THE EEL
Table 23 Silver eel catches in the sea, from trawl nets (in the Nonh Sea, mostly from the herrin~
fishery); based on data from various authors
Fishing area Place of tapture Deprh Dalt Species lAng., Weight
(m) (=) (g)
North Sea NW Dogger Bank A. anguilla .00

",.,.
11.10.24
S by W of SUgen ,So 18.11.24 c·500
'"
Kattegat SSE of Skagen 18.11.24 C·500
Swedish west coast D ec. 1924 " " ",
near Vioga Dec. 1923 c. so c.250
North Sea O CI. 1912 7'
Noss Head Dec. 1923 " 7S
Todhead, south of Aberdeen Nov. 1924 67
Ung""
West of Dogger Bank, " Apr. 1924 " so'S '97
S4 So'N,0030'E
Q
7S Oct. 1924 46'S '70
North Dogger Bank,
SSo37'N, 0°40'E
NW Dogger Bank,
66 Oct. 1932 6,
'"
6,
SSoIO'N, 10 30'E
SE of D evil's H ole,
60 Oct. 1932
'"
"
S6°:z6'N,oo:z6'W Oct. 1931 6, ,6,
" Dogger Bank -Autumn 1958
Western North
Atlantic Block Island, USA A. rostrala
SE Chesapeake Bay 15-20 5·12.67 51-58
"" "
" SE Cape Cod 7· 11.69
."
" " SE Cape Cod 7. 1 1.69 " 31
" Assateage Is. 61-66
Japanese Sea near Awashima Is. 0- 2 50 20.1.64 A. japonica 54'
near Niigata 0-120 8-4·55 338
NE of Sado Is. 0-35 0 30.11.55 73 760
N of Awashima Is.
off Agano estuary ~ .,
0-250 5·10·55
16.12·57 "6,
8,
74
1140
Sado Straits 4·1.61
.
~,,,
6,7'
0-210 20.12.61
Shallow 4.4. 62
" ,60
Awashima Is. 24·2·64 ,8 m
" 0-2 10 29·12·64 g, 938
Pacific New Zealand, east of
Cape Farewell
'" 10·5·71 A. ditffenb(lChi '00 '700
by no other area (see p. 281), make it possible [0 recapture marked eels

along almost the entire coastline, and thus marking experiments to
study spawning migratory periods have been underway since the
beginning of this century.
Individual experiments, of which there are a large number, will not
be discussed in detail here; Fig. 85 provides a summary. Judging from
this diagram, by far the greatest number of silver eels is recaptured on
the north coast of the Baltic. This is true for both those animals set free
in the eastern region, in the gulfs of Finland, and those originating from
the south and the north coasts. Eels marked on one of the Baltic coasts
Rt mDrks Re/treltus
Sex organs no! panicularly well developed Fischerbou 17, 120- 121 , 1925
" """ "" " "" "
"
Widdiofovaries ' "' cm " "
"
Widdi of ovaries ' "2 em; diam" ofoocytes 0"13 mm
Widdi of ovaries 1"2 em; diam" of oocytes 0"23 mm
" "
"
.
.,
Lengdi ofptttoni fin 3"0 em Schnakenbeek, ! 932 ; sec also Schnakenbed:, 193<
1"9 em ; widdi of ovaries 0"9 em " .. "
3"1 em; 1"9em .. " "
2"4 cm; " " " !"7cm
" "
" " 2"7 em; " 1"3em
"
LUhman & Mann, 19S8;
5 specimens Maritimes, 1969(Winter) ; 3-S
3 " , diameter of oocytes 0"28 mm Wenner, 1973
Oocyte diameler 0" 17 mm
Males
"
6 specimens, oocyte diam"0"36 mm ; 80nad wei8ht 30-37 g
"
Oocyte diameler 0"03 mm; gonad weight 308
"
Honma,l966
0"28 mmj " ' 3g
0" 29mm; 22g
Males; gonad weigh! 0"48 g
Oocyte diameter 0" 20 mm ; gonad weight 31 g
"
"
" 0"33 mm j 34 g
" " 0"29mm j 199
0"28mm ;
Oocyte diame!er 0"29 mm ; gonad weight 51 & Horuna, 1966

Lena:di of pectoral fin 6"6 em ; widdi of ovaries 5"7 em;
diameter of oocytes: maximum o"S mm ; gonad
weight 302 g Todd, 1973
crossed the Baltic and were recaptured on the coast of Sweden. Eels
which originated from the area around Riigen swam west or north-
west to arrive on the nearest coastlines, those of Denmark or Sweden.
Eels released from Sweden were, without exception it seems, recaptured.
on the same coast or were trapped. by Danish fishennen in the most
northerly exit of the Baltic.
From marking experiments on the coasts ofRiigen, it looks as though
the usual westerly migration (see p. 229) tends towards the north-west
rather than the south-west, otherwise more eels would have been
recaptured in the western regions of the Baltic. Thus it appears that
THE EEL
Table 24 Daily distances travelled by silver eels; data obtained by using conventional
marking techniques
Locality Oiltame trawlled (k.m) Speed Remarkl A uthor
(k.m/day)
B<lltic ><= '3 Trybom & Schneider, I<}OS

B<lltic > '30 T rybom & Schneider, 1<}O8
Baltic "
55- 6 3 Maximum speeds Maar, 1947
Baltic Average speeds Maar, 1947
Baltic Rugen to Swedish coast "
'5 Goal orientated
(95 bn) migration in
October M artinkovntz, 1960, 1961
Baltic Riigen to Danish ooast <0 Goal orien tated
migration in
October Martinkowitz, 1960, 1961
Baltic Riigen to Swedish ooast 5 Less goa l oriented
migration in
,
( US km)
September Martinkowitz, 1960, 11)61
Baltic Riigen to Danish roast Migration in
( 151 km) September MartinkOwitz, 1960, 1961
Eastern
North Sea '5 Liihmann & Mann, 1958
the main migratory course is, at least at that point in time, not yet
directed towards the Sargasso Sea (Deelder, 1970). If it were, the course
would have to be 2400 • In passing it might be noted that eels which start
migrating from the exit of the Mediterranean would never reach the
Sargasso on such a course.
Results of marking experiments are not available from other regions.
For the North Sea area, for example, one still has to rely completely
on conjecture. This is simply due to the fact that eels, especially those
in the spawning migration stage, are seldom caught north of Heligoland.
Only a very inadequate picture of migration routes can be deduced
from the few isolated specimens that have come to light (Table 23).
On the basis of available information it is impossible to tell whether
eels pass through the Channel or leave the North Sea on a northerly
course. There are two instances of eels being marked and recaptured.
The two specimens caught in the Elbe, one in the autumn of 1957, the
other in the autumn of 1958, were marked and set free there; each was
recaptured within the same year, one near Hvidesande on the Ring-
k~bing Fiord ICX)() m from the coast, and the other in the Lim Fiord
near Svanholmsl</b (Lillunann & Mann, 1958). These two examples
indicate that the migration route is very far to the north. North Sea
animals equipped with ultrasonic transmitters also swam in a north-
westerly direction (Tesch, 1972, 1974a, 1976). If one compares the
direction taken by these eels with that taken by the eels marked in
Rugen, one gets the impression that the migration route in the northern
areas of Europe really does show a north-westerly tendency.
3-4.7 Behaviour and speed

As has already been mentioned (p. 232), it appears that silver eels drift
downstream in the middle depths of rivers. If stow nets are set directly
on the river bed, they catch fewer eels than if they are placed nearly
in mid-water. Silver eels are not necessarily lone migrators. As is the
case with yellow eels, they have been seen to drift in groups (see p. 211 ).
As yet, nothing definite is known about the depths at which silver
eels migrate in the Baltic (see p. 233). Various sources report that, at
least during the night, eels can be found immediately beneath the
surface. Using floodlights to illuminate the surface in the Great Belt,
it was established that some silver eels migrate very slowly just below
the water surface. This surface migration ceased with the appearance
of the moon (Fischerbote, 21, 316-318, 1929).
The American eel shows similar patterns of behaviour (Maritimes,
1969, Winter, 3-5). During the period of the new moon in November
a female eel was seen in the coastal region of Florida. In addition, a large
number of eels was caught 5 to 10 miles off the coast of North Carolina,
and one specimen was caught at night (with the help of floodlighting)
beyond the 180 m depth contour off Rhode Island. Other anguilliform
fishes show similar behaviour during spawning migrations, e.g. the
shallow water worm-eel, Ahlia egmoncis (Cohen & Dean, 1970).
It is possible to obtain direct observations on the depth at which
eels swim by equipping migrating silver eels with ultrasonic trans-
mitters that register either water pressure or temperature. Five different
tracking experiments with eels in a bay between 20 and 60 rn deep, and
with a salt concentration over 30 parts per thousand showed that the
animals made frequent changes in depth and from time to time made
excursions to the sea bed (Stasko & Rommel, 1974). At the
beginning of one of these experiments six changes in depth were
registered per hour; this figure dropped to 3'5 towards the end of the
period. Preferred depth did not appear to differ between day and night,
and when a constant depth was recorded, it was between 0 and 5 m below
the surface and was maintained for 25 % of the total observation period.
Similar behaviour was shown by a European eel tracked in the shelf
region near Biscay for four hours one night over depths of more than
50 m (Tesch, 1976). This eel ventured into water over 32 m deep five
times in the first hour, two to three times in the next two hours and
none during the last hour of the study. The fact that eels of both the
species studied made frequent deep excursions at the beginning of
240 THE EEL
each experiment, suggests that the animals may have been affected
by experimental conditions.
Similar, but less extensive diving patterns wefe seen in the Baltic
(Westerberg, 1975; Tesch, 1976). It is possible that one Swedish eel,
whose depth could be inferred from temperature recordings, was
disturbed by the presence of the research vessel directly above it; this
animal remained on the sea bed by day, generally without moving
(Westerberg, 1975). In contrast, the author followed one silver eel for
47 hours; this animal remained active the whole time and its behaviour
pattern did not change between day and night. The eel's preferred
depth was 18 m but this varied greatly, apparently in relation to the
total water depth. The nocturnally active Swedish eel tended to remain
near the surface. On the basis of casual observations and the experiments
cited above one can conclude that silver eels in various shelf regions of
the North Atlantic often move in shallow waters during the night.
It is possible that there are behavioural differences in regions with low
salt concentrations and high turbidity and those with a normal
concentration and low turbidity.
As regards distances travelled, some eels cover considerable distances
in one day. Not only is the individual's swimming capacity of importance
here, but the combined effects of swimming speed and current must also
be taken into account. In the Elbe, tagged silver eels cover at least
30 km a day (Luhmann & Mann, 1958). In the sea, however, the speed
is slower (Table 24). It should also be remembered that in the Baltic
favourable currents are infrequent. There, during the most active period
of migration, progress is usually about 15 km a day. At the beginning of
the migratory period, when eels are less active, only 5 to 8 km are
covered in a day. In the North Sea distances travelled are usually
about 25 km daily. These figures for the North Sea are backed by
observations on yellow eels which travelled at least 25 km under favour-
able conditions in order to reach their home ground. In exceptional
cases daily travel of three times this distance appear to be possible
(Tesch, 1967a; Deelder & Tesch, 1970), and there are isolated records
of similar distances covered by silver eels in the Baltic. Tracking experi-
ments in the North Sea and in the Baltic have provided the opportunity
for direct investigation into travelling speed in silver eels (Fig. 84;
Tesch, 1972, 19743). In the North Sea 48 km a day were covered, and
similar figures were obtained from the Baltic. Such speeds would be
sufficient to complete the 6000 to 7000 km journey to the Sargasso Sea
by spawning time in March (see p. 93). If eels have five months (Novem-
ber to March inclusive) to complete the journey, they must cover
between 40 and 50 km a day.
4
Fishing Yields
4. I The development of eel fisheries

We can be sure that eel fishing was practised in prehistoric times.
Numerous archaeological finds indicate that fishes, induding eels, were
part of the diet of our ancestors, but unfortunately we have learned
little about the methods used to catch the eels or about the numbers
caught.
In Denmark, fishery statistics go back for many years; information
on the quantity of eels caught since 1885 are available for a large pro-
portion of aU Danish waters. Since the figures for 1885 are not signi-
ficantly below those of the last few years, it seems likely that, for a
long time before official records were kept, eels must have been caught
and sold in appreciable quantities. The largest Danish fyke net fishery
started in the Belts; its annual yield between 1902 and 1916 for
example, was from 400-1400 tonnes. Between 1885 and 1895, the same
fishery, together with rod and line fishing, was already bringing in
as many as 333-585 tonnes a year (Jacobsen & Johansen, 1922). Between
1940 and 1959 the silver eel fishery in the Belts, which is also based on
eel baskets, was giving yields of 500-1000 tOMes a year (Jensen, 1961).
It thus appears that catches generally have remained more or less
constant since 1885.
Figure 86 shows annual catches from the North Sea fisheries of
Norway, Derunark, Holland and Germany over a considerable number
of years. This diagram also fails to indicate any significant increase in
241
242 THE EEL
Table lS Eel yields from the most important eel (Anguilla spp.) producing countries
in the world (in 1000 t) (from Yearbook Fish. Stat. FAD, anti Bull. statist.
Cons . perm. Explor. Mer)
Country 19J8 1948 '949 1950 1951 195 1 1953 1954 1955 1956 1957
Japan
Denmark
Italy
,-, ,-, ,-, ' -s
10'4
... "
,-, , -8 , -8 3'7
"
,-8 , -8
Holland
Sweden
" "
,-, " " "
,-, ,-, '-7 '-s ,-,
New Zealand
GDR ,-,
Gennany FRG 0- ' 0-, 0-, 0-, 0-, o-s 0-, o-s 0-, 0-,
Poland 0' o-S 0-7 0-' 0-,
F~~ '-0 ' -0
Spain 0-, 0-' 0-' 0-' 0' 0- '
Northern Ireland 0-' 0-, 0-8 0-'
Canada ' -0 ,.,
USA 0-' 0-,
Morocco
,8 ,8 ,8
World Iotal
" " "
>0 >0
" " "
catches during the period reviewed, although there are changes brought
about by such factors as the ending of the successful Ijsselmeer fishery
in Holland. The sometimes marked variations in catch sizes are brought
about by economic trends, the effects of war or of biological factors.
Information on the large I talian eel fisheries in the lagoons of northern
Adria can be traced back even further (Bellini, from Walter, 1910).
Between 1798 and 1824 a yearly average of 962 tonnes of eels was
caught in the lagoons of Comacchio. From 1825 to 1870 the catch sank
to 493 tonnes, but then increased, remaining at an annual average of
858 tonnes over the next 10 years, only to drop again to 391 tonnes
until the beginning of the twentieth century. At the end of the eighteenth
century catches in Comacchio alone were equivalent to one third of the
total present-day yield for the whole of Italy. The lagoon fishery of
Comacchio was already flourishing in the fourteenth century. According
to Walter (1910), the rent alone at this time was 266 600 Marks, so that
it is unlikely that yields in the fourteenth century were much below
those of the nineteenth century. It thus appears that eel fishing is a
long established branch of the European economy.
Considerably less information is available on the quantity of eels
caught over the rest of the Continent or, for that matter, over the whole
world. Statistics from the International Council for the Exploration of
the Sea and from the United Nations' Food and Agricultural Organiza-
tion (FAO) provide a limited and rather incomplete review from 1938
FISHING YIELDS 243
9' 1 8'4 9'0 11'5 10'7 12·6 ,6'2 187 19·8 22·8 26'7 26'5 19'4 16·8 16'5 17']
3·3 4'0 4'7 3' 3-' 4'0 3"3 3':Z 3" 3'S 4'3 3'7 3"4 3':Z 3"3 ]'6
:Z'I ]'0 :Z'7
" ,-,
,-,
3' 3'S :Z'2 3'0 3':Z 3 '2 3':Z 3'4 3':Z 3'3 3' 2 3'2
:z·8
1"8
3'S
:z '8
3' 1
1·8 "
,-,
1'9 :Z'4
1'9 1"8
:z., 2·8 ]'1
I', :z·o 1·6
:Z"7
1,8
:z·8
"7
1'5
I ':Z
1' 2
1'4
1' 1
1'2
"1
1' 1
0' 1 0'1 0'] 0"4 0'9 1"5 2' 1 1'3
1·6 1'4 1' 1 1'2 I'] I ':Z 1"0 1' 1 0·8 0'9 0"9
0'4 o·S 0"4 0'5 0'4 0' 4 o'S 0·6 0"6 0'5 0'5 0'5 0'4 0'4
I ':Z 0 '7 1"0 1'0 1"1 0'9 1'0 1' 1 1"1 1'1 1'0 0'9 0'9 0·8
0'7 0'9 1'3 1'4 I 'S "7 I '] 2'0 2'7 "9 4 '2 4 '9 2 '7 3'9
o'S o'S 0'5 1'9 :Z' 3 1'7 1'7 1·6 I 'S 1'5 1'2 1 '2 1'5 1'2
0'5 0'7 0·8 0'7 0·6 0·8 1'0 0·6 0·6 0·6 0"8 0·8 0' 7 0·8
0' 4 0'3 0'] 0' 4 0·8 0,8 0', 0·8 0'9 1'1 1'1 J ':Z 1'1 0'7
0·6 0·6 0'4 0' 4 0"5 0"8 0,6 0'7 0·8 0'9 1'0 1' 1 0'9 1'1
0 '3 0'] 0"3 0"3 0'] 0'] 0'3 0'3 0'3 0 '3 0'0
:ZS 27 28 ]1 ]0 ]1 ]5 ]9 41 45 51 51 45 45 39 40
onwards (Table 25). These figures suggest a certain increase in world

yields from about 1960. The period between 1938 and 1960 reflects a
general depression, which is certainly attributable to the effects of the
Second World War. In Poland, for example, fishing on any significant
scale was only able to develop after 1945, and therefore it was many
years before statistics for Poland even began to appear. However,
countries such as Denmark and Sweden, which were unaffected by
world conditions, have more or less constant yields for the whole of
this period. A detailed analysis of Table 25 indicates that, by and large,
Japan is the only country to show genuine increases in its catches.
In 1938, and from 1958 to 1962, catches remained at 10 000 tonnes,
whereas in 1964 and 1965 they rose to 16200 and 18 700 tonnes respec-
tively. Closer inspection of the statistics shows, however, that lakes and
rivers in Japan have not yielded larger quantities of eels in the last few
years than they did before. The increase in total yield must therefore be
attributed solely to the artificial culture of eels in ponds. Figures which
distinguish between yields from pond culture and catches from lakes and
rivers have been reproduced from the Yearbook of Fishery Statistics,
FAO_
19]8 1958 1961 1964 1967 1968 1969 1970 1971 1972 1973
Lake$ and River ].:z 2·8 ]'4 2"8 ]'2 ]'1 3'2 2"7 2·6 2 '4 2" thousand I
Pond Culture
244 THE EEL
In general, figures taken from several countries indicate that yields

from natural waters, regardless of the fishing techniques used, have not
increased to any great degree but that there is considerable potential for
yield increases by the use of efficient pond culture methods.
4.2 Annual and regional fluctuations in yield

4.2. I Variations over the whole North Sea region
Since the density of eel populations in, for example, the whole of
Europe, depends, presumably, on a single spawning of larvae, it can be
assumed that fluctuations in their number will have a similar effect
on eel catches. In addition, climatic changes could cause eel catches
to rise and fall to the same extent over large areas. The North Sea is
an example of a fishing ground which is relatively unifonn bydro-
graphically and which is utilized by a number of different countries.
Since economic conditions in these countries do not always coincide, it
is unlikely that similar variations in their total eel catches are the result
of common economic factors. Figure 86 enables one to compare North
Sea catches for more than 50 years. It shows that, where the catch
pattern for the previous year is available, in 36 out of a total of 44
comparable years, the catches for at least three of the four countries
rose and fell simultaneously, and in only eight years did the four
national catches show little agreement, or dissimilar trends.
4.2.2 The effect of differing proportions of males and

females on variations in catch size along the
southern coast of the Baltic and in inland waters
Table 26 enables one to compare the yields from various coastal regions
in the southern Baltic. With the exception of the West Gennan Baltic
regions, it is not possible to compare yields until post-war development
was completed, i.e. until 1954. In all regions the year 1963 stands out
from all others because of the unusually high catches. 1963 was followed
by a general depression lasting two years, after which the catches
returned to their fonner levels. Statistics derived from fisheries research
carried out at Rostock in 1964 and 1966 include data on the different
sizes of eels caught in the central Baltic. These figures show that the
peak catches in 1963 were clearly brought about by an increase in the
numbers of small eels (Size III ). A further analysis of this phenomenon
indicates that the majority of these small eels were very probably male
silver eels on their seaward migration.
FISHING YIELDS 245
Germany
Fig. 86 Eel yields from the: N orth Se:a coasts offour countries adjoining the: N orth Sea
(data from Bull. statist. Com . perm. Explor. M er.).
Since the proportion of male silver eels along the entire south coast
of the Baltic was panicularly high in 1963 but showed a decline in
1964/65, it seems reasonable to assume that this was a reflection of a
particularly abundant year class which had migrated to the area as
glass eels about seven years previously (jensen, 1(61).
Nine years after this presumed invasion took place there was also
an increase in the number offemale silver eels leaving the area. In 1964
too, i.e. eight years after the arrival of the glass eels, a limited increase
was also seen (Fisheries Research, 1964). Furthennore, one would
expect increased catches between 1963 and 1967 in inland waters as
well. Table 27 shows the relevant figures for Schleswig-Holstein and
the German Democratic Republic (DDR). Whereas in Schleswig-
246 THE EEL
Table 26 Eel yields from the coastal waters of the central and western Baltic (in
tonnes)
Year Total cQCljra/ region Fishery control area Year
FOR (from GDR (fro»! Folk & Stralsund* Kurisehes Haff

Bull. Jt(uiJf. Lauterbach, r9fis (jmm Roy, (Gajgalas, 1966)
Cons. pam. & Fisch. Forsch., 1961)
Explor. Mer.} 1964-1<}(6)
14"4 1947
'7"0 1948
1949 3S3 1949
,,6
20"9
195 0 27'1 1950
1951 '57 64'9 32·8 195 1
7S,g
1952
'" 45"3 1952
,,.
1953 '9S 107"3 132"' 1953
1954 "7 107'4 295"2 1954
,,'
1955 166'0 268·6 1955
1956 132"' :z8:q 1956
1957 '53 907 145'6 272"' 1957
,,6
1958
'"'49 126"5 264"3 1958
,"" ,""
1959 '77 1)4"5 273"7 1959
,6,
,,.,
,,., '73
'SO
9'"
110·6 )07"7
245'2 ,,.,
,,.,
,,.,
,,.,
'44
34'
93'
11 6 1
240'4
368'4 ,,.,
,,..
,,., '7S
'0'
97'
645
35 8 ',
,,.,
- induded in data for (he coastal region of the G D R
Holstein yields were more or less constant from 1961 onwards, in the
DDR catches showed a decrease in 1963 and 1964. The year 1966
brought almost the highest yield of the whole series. Schleswig-Holstein
does not fit into the present picture because it has a relatively high
proportion of males, probably the result of its proximity to the coast.
In the DDR, on the other hand, there is a preponderance of females,
due to the fairly great distance from the coast, so that the expected
increase in catches three years after the occurrence of an above average
number of males did in fact take place.
4.2.3 The relationship between the west wind and glass eel
invasions
Danish expens (e.g. Jensen, 1961) believe that increases in the size of
glass eel invasions along the coasts of Europe are caused by suonger
west winds during the spring. The winds increase the surface currents
and thus further the progress of the drifting glass eels. For this reason,
silver eel yields in areas with a preponderance of males have been
FISHING YIELDS 247
Table 1,7 Eel deliveries (in tonnes) to the Kiel marine fish market
from the inland waters of Schleswig-Holstein
(Herrmann, 1967 and Hoffmeister, 1967) and from the
inland waters of the GDR (Muller, 1961; Menzel &
Zobel in the annual reports on inland fishery production
in the GDR, DBch, Fisch, Ztg" 1962-1967 )
Year SchluViig-Holstein GDR

1954 ,60
1955
195 6 31 '9 ''99"
1957
1958
39'0
43-5 ,8,
"'"
1959 47'9 '54
,,.,
'960 66·, 43'
,,.,
HO
'"
'"
1962 54'4
,,., 58'1
53"3
467
4l'
1965
,,66 54'0
54'3 '"
54'
.,
1905
" " "
•• 11BlImrneter
y~
diff, F-S
ovrnl<J'
"
1}~12
'" " " 50
Fig. 87 Yields from the Danish silver eel fisheries in Kattegat and the Belts from 1912
to 1959, and the differences in barometric pressure between Fant) and Skagen
seven years previously, i.e. 1905-1952 (after Jensen, 1961 ),
compared with differences in barometric pressure between Skagen and

the isle of Fano (see Fig. 87). It is known that air currents over the
earth's surface move more or less parallel with the isobars. If a large
difference in barometric pressure occurs between Skagen (to the north)
and the isle of Fane (to the south), then it is highly probable that the
isobars will run in an east-west direction at both places and, depending
THE EEL
on the size of the pressure differences, westerly winds will blow. This
pressure difference has, therefore, been used here as an indication of
the strength of the west wind drift; parallel trends were shown to exist
between catches and differences in barometric pressure. Since glass
eels are also caught in Denmark (see p. 145), additional analyses were
made to see whether the quantities of glass eels caught 7 to 10 years
earlier correlated with the silver eel catches. The results were positive,
suggesting that it is no longer difficult to predict the size of catches
7 to 10 years in advance.
4.2.4 The influence of seasonal temperature levels

I t might seem very obvious that one could link summer temperatures
with eel catches, particularly those of yellow eels. Such correlations
have been made by Danish scientists since 1922 (Jacobsen & Johansen,
from Jensen, 1961). While there is an indisputable correlation between
these two parameters, there also appears to be a connection between
winter temperatures and eel catches. In cold winters, for example, eels
suffer considerable mortality (Johansen, from Jensen, 15)61). This was
confirmed by fishermen from Heligoland, who pointed out that after
the cold winter or 15)62/1963, for example, there were very many
deaths (see also p. 24). Eel deaths were also reported from inland
waters during the cold winters of 1962/1963 and 1969/1970 (Rahn,
1963; Tesch, 1964; Mattern, 1971).
As already mentioned, the winter of 1962/1963 was extremely cold,
with noticeable mortality among other organisms too (see pUblications
in Helgol. Wiss. Meeresunters. , 10, No. 1-4, 1964). This must have had
some effect on eel yields in central and northern Europe. However,
contrary to expectation, large numbers of eels were caught on the
southern coast of the Baltic in 1963 (Table 26). In the inland waters of
Germany relative changes in catch size were variable. Along the North
Sea coast of West Germany (Fig. 86) the catches for 1963, and for
1964, were similar to those in the inland waters of the DDR, i.e. below
average. However, in contrast to West Germany, the North Sea fisheries
of Denmark, Holland and Norway showed increased yields in 1963
(Fig. 86). On the basis of these statistics, it is debatable whether one
can conclude that cold winters, particularly those when temperatures
do not fall to extremely low levels, have a significant effect on eel
populations and thus on catches the following summer.
4.2.5 Catch variations in Canada
The Canadian eel fishery has shown considerable fluctuations in yields
over a period. of 45 years (Eales, 1968). The maximum, 1200 tonnes, and
FISHING YIELDS 249
the minimum, 350 tonnes, were reported in 1933 and 1948 respectively.
Catches have been increasing since 1948 and reached 800 tonnes in
1965. The reasons for these marked changes are not at all apparent,
though presumably they are associated with economic factors or
fishing techniques.
Ever since the more valuable and popular species offish have become
less frequent in the markets, the eel has become an important element in
the fisheries of Lake Ontario (Hurley, 1973). In the last few years eels
have comprised 8% of the total catches and 14% of the financial returns
of the fisheries on Lake Ontario. From 1959 to 1970 the total Canadian
eel catches fluctuated between 49 (1962) and log tonnes (1964). Since
the St. Lawrence River was blocked by the construction of the Moses-
Saunders power station in 1958, declines in yields were to be expected
because of the markedly adverse effect such an obstruction would have
on the ascent of eels. Despite this, however, catches between 1959 and
1970 did not show any such decrease although there was a noticeable
decline in the average length of the eels caught. This decrease in size is
not the result of a decline in the number of eels ascending the river,
but is a consequence of the high fishing intensity.
4.3 Variation in catch size throughout the year

As is already apparent from the preceding sections, the quantity of eels
caught shows considerable seasonal variation. In addition, there are
//;
, '
; ~l
•' ,g.
, ,
i~
.
,
,
,, ,,
.
,
.j ,,
,
,.- "" '
,,
, ' ,f
:,
,,
,, ,,
,,
,
,,
, .,
Jgn_ Feb. Mgr. Apr. Mgy Ju... Jul. AUCj. Sept. Oct. NOlI. Dk
Fig. 88 The mean monthly eel catch expressed as a percentage of the mean annual eel
catch in the North Sea and the Baltic from 1961 to 1965 (based on data from
Land u. Forstwirtscha/t, Fischerei, Fangergebnisse der Hochsee· u. Kwten-
mlherer!.
250 THE EEL
variations between different waters. Among the factors contributing to

this variability"are the unrelated peaks in activity seen at various stages
in the eel's life cycle, stages which appear in different waters with
varying frequency. The seasonally determined activity rhytlun, and
thus the monthly catch distribution, is, in many ways, determined by
the yellow eels, i.e. by those fishes which remain in continental waters
for about 7 to 10 years. Waters through which eels pass on their seaward
spawning migration show a quite different picture. Many waters
containing yellow eel populations arc, however, influenced to a greater
or lesser extent by the migratory peaks of silver eels.
Yellow eel activity is, in the first instance, dependent on temperature.
This fact is illustrated by a comparison of the 1953 and 1955 monthly
German catch record in the western Baltic with prevailing water
temperatures (Thurow, 1959). The peak catches coincide with the
highest temperatures and thus occur at the height of summer. Yields
drop to practically nothing in December, and only begin to increase
appreciably in April. A similar pattern of catch distribution is seen in
Fig. 88, which compares monthly catches in the Baltic and in the North
Sea. The varying proportion of silver eels causes a fairly marked
displacement of the catch maximum to the autumn.
"
~g/h
- C.~tl'<ll reQcllH )
Ilivllf' OIkr
lQkI Con.tlltlCl_ 'I.
.. ....... Lowtf rrQchn Rl'Iifll " · ___ •
• .!\ "
./
, o
•
, ,
,
~ ....
0 , v VII VIII IX ,
FJg. 89 The
" '" " " " '"
percentage monthly spread of total annual eel catches in Lake
Constance (Bodensee) from 1964 to 1966 (from Land u. Forstwirtsdw./t,
Fischerei, Fangergebnisse der Bodensujischern), and the eel stow net catches on
the Rhine (after H ass, 1964). Monthly yields in kg/ha in the middle (near
Seelow) and lower reaches (near G artz) of the river Oder from 1958- 1963
(after Albrecht, 1964).
,.
FISHING YIELDS 25 1
000
.. ,~
------ ---.' .... .

90
ao - -
, --
-
.-
,..
,r- ••• • '._
70
00 ..
,..------I
SO -
'"
JO
20
00
..,
ns 30:5 20.6 lH
'"
Obwr.ation datts
24.9
Fig. 90 The proportion of yellow and silver eels in a fishery on me river Elbe above
Hamburg during 1958. Dotted line: silver eels. Discontinuous line: half-silver
and silver eels. Continuous line : yellow eels (after Luhmann & Mann, 1961).
Furmer observations in 1967 gave a similar result (Mann, 1968).
Catch distribution in rivers (Fig. 89) resembles that in coastal waters

with smaller proportions of migrating eels. The Oder and the Rhine
show the highest catches during the sununer months. Peak catches in
Lake Constance, however, are greatly displaced, i.e. they occur in
spring. Since this lake lies in the upper drainage area of the Rhine, its
silver eels would have to begin migrating considerably earlier if they
are to reach the river estuary in autumn, hence the unusually early
maximum catch in Lake Constance is most likely attributable to the
need for an early start to migration in the upper stretches of the river.
Any other explanation is highly improbable; Lake Constance is a deep
water mass in the Lower Alps, and thus takes a long time to warm up
in spring, a situation that is unlikely to produce increased activity in
yellow eels at this time.
Catch distribution in the middle and lower stretd:..es of the river
Oder indicate a similar tendency. A relatively early maximum occurs
in the middle reaches in June but is not seen in the lower stretches.
Monthly records of catches from eel traps situated in Mecklenburg
lakes each showed a maximum catch in spring and autumn (Table 33).
Russian observations in Lithuania also indicate that there, too, silver
eel migration is already in evidence in the spring (Matschenis, 1965).
However, no definite conclusions can be drawn because catch statistics
THE EEL
do not clearly differentiate between silver and yellow eels (Fig. 90;
for silver eel catches see p. 220 and Fig. 81).
Increased total catches are a well known phenomenon along the coast
in spring (see p. 217). On the whole, however, these increases are not to
be attributed to an increase in silver eel migration, but are probably the
result of movement among yellow eels leaving their winter quarters.
This movement is similar to the change in habitat which takes place in
autumn during the search for winter quarters, only it is in the opposite
direction.
4.4 Population density and catch sizes from an

economic and ecological viewpoint
4.4. I Regional differences

The occurrence of a large or small number of eels under natural condi-
tions is primarily a function of the inunigration of young fishes, which
in turn depends on the length of the uninterrupted waterway providing
access to the sea. This is one of the most striking features of tables
dealing with eel populations and catches (Tables 28-31 ). Differing
local conditions make direct comparisons difficult. The large coastal
rivers of West Germany, for example, cannot be compared with the
waters of Nova Scotia and New Brunswick. While the ascending eels
are still clearly detectable for hundreds of kilometres in the large
German rivers, distances of only about 10 km appear to playa role in
the small brooks and streams of Canada (Smith & Saunders, 1955). No
obvious differences seem to occur in the relatively small bodies of
water in New Zealand (Burnet, 1952a). Natural conditions have, of
course, been largely obscured by the march of civilization and
industrialization. For example, a lake in close proximity to the sea may
have only relatively small populations of eels because it has become
blocked by dam and dike construction.
It is also striking that the biomass and yields from running waters in
New Zealand are greater than those for streams and rivers in other
continents. Streams studied in New Zealand contain 66 to 530 kg of
eels per hectare (Table 28), whereas in Lower Saxony the figure is
only 1'5 to 100 kg/ha, and 43 kg/ha is considered well above average
(Tesch, 1967b) ; a stream in Denmark contained 75 kg/ha. There are,
unfortunately, no other comparative data for Europe. In New Zealand,
catches of between 33 and 1520 kg/ha are recorded, while mean values
can reach over 100 kgJha (Burnet, 1952b). In Germany, the corres-
FISHING YIELDS 253
ponding values are between 3 and 50 kg/ha, with average catches of
little more than 10 kg/ha.
lt thus appears that in New Zealand eel production is higher than in
the temperate zone of northern continents. There may be various
reasons for this: ( I ) either or both of the two eel species (A. dieffenbachi
and A. australis ) which occur in New Zealand grow faster and are
therefore more productive than is A. anguilla; (2) temperature levels
are higher in New Zealand than in central and northern Europe,
e.g. winter temperatures do not fall below 6 °C in the Horokiwi stream
(Allen, 1951); (3) the eel populations in New Zealand are larger because
commercial fishing is not very intensive; (4) in New Zealand waters,
competition from the few other species of fish, with the exception of
the introduced salmonids, is not very great.
There are no comparative studies on possible differences in growth-
rate between the various eel species. We only know that, on average,
A. dieffenbachi in New Zealand grows to a much larger size than does
A. anguilla in central Europe (see p. 188). The high temperatures in New
Zealand are probably the main cause of the higher than average catches
there. If this is so, one would expect large eel populations in streams near
the coasts of southern Europe, and this does appear to be the case. One
angler in Spain commented to the author on the large numbers of eels
in Spanish streams, and noted, in panicular, the large size of some of
these fishes. A magazine article (Fisch u. Fang, 1966, p. 276) also
mentions the abundance of eels in a river near Barcelona.
In support of the assumption that higher temperatures are the main
ultimate cause of high eel biomass in New Zealand, is the fact that in
Europe too, those standing waters with the highest eel yields are found
in the south (Tables 29 and 31), namely the lagoons of Comacchio in
Italy. A water body in northern Europe which is fished as intensively as
this classical eel-producing area in southern Europe, is Lough Neagh
in Northern Ireland (Table 31 ). Lough Neagh is comparable in size
and is stocked intensively with glass eels; practically all the silver eels
leaving this lake are caught, and yellow eels are trapped in large numbers
too. However, Lough Neagh does not produce nearly such high catches,
20 kg/ha, as do the northern Italian lagoons, 30-40 kg/ha, and in some
cases even as much as 90 kgjha (D'Ancona, 1961). Less comparable is
Lake Convent in north Germany, which also used to be cited as a
classical area of high eel catches, although with a yield of 36 kg/ha it
doubtless represents an example of the optimal use of an eel population.
The high yield here is a consequence of the large natural immigration
of young eels from the nearby Baltic, coupled with a relatively high
254 THE EEL
Table 28 Biomass of eel populations in various waters

Water body Geographical position Size of area Year Popula/ion in
studied kg/ha
( i>lha )
E,/ TOIa/
H oroldwi stream New Zealand, Wellington 0·3 Before 1952 530
Wainui-o-mala stream New Zealand, Wellington 09 Before 1952 3>0

D oyleslon stream New Zealand ± 0' 1954 '30 ,63
Hanmer Road stream New Zealand ± 0·3 1956 66 "0
Bill's Lake New Brunswick, near
Passamaquoddy Bay ,., Before 1955 79'3 So,
Brandstrup stream Jutland, River Gudena,
Kattegat VI + IX 1950 75 ,,6
Lake Coleback Maine, NE coast of USA '0 1958 47'S 117'0
",
Potter's Lake New Brunswick, near
Passamaquoddy Bay Before 1955 90 23'0
Trefry's Lake Nova Scotia, SW coast 21'6 Before 1955 '·9 [9"7
Lake T edford Nova Scotia, SW coast 21 -0 Before 1955 ,·6 4°'5
Pool, near
GundS¢magle Vaerebotal, Seeland '0 1959 ' ·9 482'0
Lake Jesse Nova $oolia, SW coast 18 ' 2 Before 1955 0·' 22 "2
Boars Back Lake Nova Scotia, SW coast 22"6 Before 1955 0' '9"3
Naera Stand NordfUnen, D enmark 430 '9S S '5'

Hinau stream New Zealand, North Island O"2? '968/69 ,88
".
H inaki stream New Zealand, North Island O"2? '968/69 '09 88,
"
T en Mile stream Narrowsburg, N ew York '968/69 '73
productivity and very light fishing because of the small size of the lake;
if Lake Convent were as large as Lough Neagh, similar yields would be
most unlikely. Even the lakes in Schleswig-Holstein, which are subject
to intensive stocking progranunes and which, in some cases, have
abundant, natural influxes of young eels, cannot produce maximum
catches much above 20 kgjha (Table 31); in certain instances, however,
yields as high as 40 kg/ha have been recorded (Herrmann, 1967), but,
such figures are from smaller lakes which should be compared with
Lake Convent (note the size relationships of the lakes in Table 31 ).
The question of how New Zealand waters, which are mainly fished
FI SHIN G YIELDS 255
% eels in Other important species Characteristics of the Literature

lowl of fishes water body
popula-
lion
River Trout population of
about half this weight Stream, 4 m in which Burnet,1952a
River Trout? Stream, 8 m in width Burnet,1952a
Riv er Trout, 133 kg/ha Stream, 3 m in width Burnel, ' 959a
River Trout, 54 kg/ha Stream, 3 m in width Burnet, 1959a
Couesius plumbrus (Cyprinidae)
Brook Trout ~3mdeep, ,"6kmfrom Smith & Saunders, 1955
the sea
Stream, I m wide.
Salmon and Trout Electrofishing Larsen, 1955
CalOj/omus commersonnii Max. 9 m deep. High primary Rupp & De Roche, '965
(Catostomidac), Smelt productivity
39·0 Black Bass, £lox niger 92mdeep, 6 km from thesea Smith & Saunders, 1955
14·9 Morone americana '"I m deep, 1·6 km from Smith & Saunders, 1955
(Serranidae) the sea
6.3 Morone americana (Serranidae)
Fundulus diaphanus ~ 2·' m deep, 5 km from the sea Smith & Saunders, '955
0·4 Roach, Perch, Pike Oecasionalconnection with the

River Vaerebo Larsen, 1<)61 a
Marone americana II 2.4 m deep 25 km from Smith & Saunders, 1955
,., (Scrranidae), Bullhead

CalOs/omus commer50nnii
the sea
9 2·7 m deep, 48 km from
(Catostomidae), the sea Smith & Saunders, 1955
Bullhead
Flounder, Eel pout, Lagoon, S '" 2-20 parIS per Muus, 1967
Gobiidae thousand
6, River Trout, Philypnodo./
breviceps Small stream Hopkins, 1971
9' River Trout, Philypnodon
breviceps Small stream Hopkins, '971
SemQ/ilus CllTporalis,
Catostomus commersonnii Small stream Engstrom-H eg& Locb, 1971
for sport, can produce a higher population level than can European
waters, which are fished commercially, must be decided separately
for each case. The reduced level of competition from other species in
New Zealand must play a not insignificant role in this increased pro-
ductivity. Cyprinids, which are the most common species found in
'eel rivers' in north Germany, are totally absent from New Zealand
(except for a few instances where attempts have been made to introduce
them ; Allen, 1957). Thus the only competition is from brown trout
( Salmo trucca) and rainbow trout ( S. gairdneri) which have also been
introduced into New Zealand (see Hobbs, 1948; and below).
256 THE EEL
Table 29 Eel yields in coastal waters and coastal lakes

Waurbody Geographical position Area in Year Yield in kg/ha
hectares
Eels Total
'Valli' Venice & Cornacchio ,0= Before 1961 )0--40

Lake Convent Mecklenburg, nr. 29"0-45"4 58 "7-12 1"3
Bad Doberan 6, 1954- 1<}61 935·6 ~79·3
Ijsselmecr Holland 220000 1954-1962 C.IO
Strelasund South coast central

Baltic 24300 1953-1965 7.0 14"9
Kleines Haft" South- west central

Baltic, Oder estuary '9= 1963-1965 6·, 41"6
Wismer Bay South coast-western
Baltic 12200 1963- 1965 43 8·,
Dar~ Bodden chain South coast-western
Baltic ,,= 1963-1965 3·' 22"1
R Ugen Bodden chain South coast- central
Baltic 15500 1963- 1965 3·0 14"9
River Pecne South coast-central
BaIlie, Oder estuary ' 77 00 1963-1965 '3 39"2
Greifswalder Bodden South coaSt-central
Baltic 53400 1963-1965 ,8 7·,
Frisches H air South coast-central

Baltic 86 100 1936 + 1939 4·:Z ( 1936) 13"4(1936)
Kurisches Haff South coaSt-central

Baltic 16 1 300 1927- 1938 0·84 119 (1936)
K urisches Haff South coast-central
Baltic 16 1 300 1947- 1964
"
4.4.2 The eel in competition with other fishes and with

crustaceans
The large areas of coastal water shown in Table 29, although producing
above average catches, have smaller yields than do the small coastal
lakes. It should be noted here that migrating silver eels pass through
some of these waters and thus the catches do not consist only of fishes
originating from that locality alone. The entry in Table 29 for the
yield from Baltic coastal waters consists of freshwater fishes only, but
marine species make up a considerable portion of the total catch from
these regions, so that the total yields are considerably higher. In 1952,
for example, freshwater fish catches, which included the eel, came to
I I kg/ha, but a total of 76 kg of freshwater and marine fishes were
caught per hectare (Subklew, 1955). Competition due [0 the abundance
FI SHING YIELD S 257
% eels in Other important species ChorlUuri$ll'cs of Literature

tOlol of fishes waur body
yield
Grey Mullet (Mugi{) Lagoons D' Ancona, 196 1

30-65 Pike 13% Coo.stallake, 300 m long;
if, 45 T ench 4% connected by canal to the
Baltic Sea Gollub, 1963
Pike-Perch, Smelt Formerly a sea bay, now
with dikes. Yields used to be
higher
47'0 Roach 29"5% Connecting water between Fisch. Forseh.

Perch ± 20% two sea bays RO$!luk, 1963-1965
Roach 27"2% Estuary wilh low sal!
15"7 Bream 12"8% concentration
Besides the eel, mostly Open sea bay
52"8 marine fis hes
Roach 32"6% Long, strongly freshened lagoon
14'2 Bream 19"5 %
Roach 29"5 %
47"0 Bream 17"4% Long sea bay
Roach 41 "3% Estuary wilh low salt
'·9 Bream 24% concentration
Excluding Herring Fisch" Forsch.
37"7 ( ± 60 kg/ha), ROJIOCk, I963-1965;
Perch, Roach Open, shallow sea bay Subklew, 1955
Bream, Pike-Perch,
Ruffe Lagoon-like, coastal lake Subklew, 1955; Neuhaus,
'96,
Smelt (± 29 kg/ha),
Ruffe 13 kg/ha Lagoon-like, coaslallake Gajgalas, 1955
Bream ± I2 kg/ha Maniukas, 1961
Pike-Perch 3 kg/ha
of species in these waters may have played an important pan in bringing

about such relatively small eel yields.
The question also arises whether larger eel populations could exist
in coastal streams which empty directly into the sea, if there was no
competition with other species of fish in the same waters. This applies
particularly to trout streams as they are known in Derunark (Larsen,
1955, 1961b). With a total average biomass of 246 kg/ha, the figure of
75 kg/ha for eels is not panicularly high (Table 28). The same must be
said of the River Aue tributaries (Tesch, Ig67b), which are situated
near the Elbe estuary. These waters have the highest eel biomass
(100 kg/ha) for the streams studied in Lower Saxony. However, eels
comprised only 35% of the total biomass of 286 kg/ha. Estimates for
20 Danish streams produced a similar picture (Larsen, 196Ib): on
average there were 5100 trout per hectare, but only 850 eels.
In many instances the competing species were trout, which, to a
25 8 THE EEL
Table 30 Eel yields in funning waters

WaitT body Geographical position Area in heaareI Year Yitfd in kllha
&/, Total
Werra and Several years
tributaries Weser-Nonh Sea
Od~ Oder-ccnlral Baltic
In H essen, 281
Frankfurt on Oder
around 1960 4' '4
68- 100
)2'0-59"5
to Gartz, 3198 1<}61-1963 944"0 ; 8,
Beste, Bamitz T ravc-wcstern Baltic Bad Oldesloe, 3 11-38 44- 83
195 8- 1964 ¢23"o 959"3
Trave Travc-western Baltic Above Bad 11-35 77- 1 74
Oldesloe, Il 1958-t964 tj 19"5 9112
Tralle Trave-westem Baltic Below Bad 8- 2 5 55- 129
Oldesloe, 18 1958-t964 ¢ 15"7 993"7
Westr Weser-North Sea Above Nienburg,
' 90 1<}61- 1 963 12"2 27"0
Westr Wescr-North Sea Below Nienburg,
'97 1900-1<}62 7" 12·6
Ell>< Elbe-North Sea Werben,above the '"9-35"0 34"1-126"5
Havel estuary 18<}6-1928 if, n", 977"9
Ell>< Elbe-Nonh Sea Werben, above the 0"4-6"2 /1"7- 109'8
Havel estuary / 9 27- 1937 92"6 925"0
Ell>< Elhe-North Sea Below
Schnackenburg 1956-1963 25-50 "0
t..hn& Several years
tributaries Rhine-North Sea In H essen, 104 around 1960 9 64
Fulda & Several years
tribularies Rhine-North Sea In H essen, 1652 around 1960 8 8,
DiemeJ& Several years
tributaries Weser-North Sea In H essen, 230 around 1960 8 68
M""I Rhine-North Sea Trier 10 K oblenz 1951-1961 7
,,'
Rhine Rhine- North Sea Below Mannhcim,
3500 1956-1961
Em, Ems-North Sea Above the estuary,
,,8 1954- 1955 ,,8 26",
considerable extent, feed on the same organisms as does the eel (Sinha
& Jones, Ig67b). On the basis of population studies in trout streams,
Larsen, (Ig6Ib) comes to the conclusion that the trout is dominant in
small streams but only if the environmental conditions are favourable
for it.
In other cases the eel was the dominant species, particularly in
streams where conditions are, in many ways, unfavourable for trout,
particulldy for its reproduction. Of course large nwnbers of cyprinids
occur in such areas, and as already mentioned, they compete with the
eel to a great extent for food.
The discovery that in typical trout streams the eel does not provide
the expected high proportion of the total fish biomass may be due to
the fact that, unlike trout, the thermophilic eel does not maintain a
sufficiently high metabolic rate for optimal production in these generally
cold waters.
FI SHING YIELD S 259
% eels in total Other important Characteristics of water body Literature

carch species of fishes
T rout Barbel region, tributaries,
,7' Salmonid region Buhse, 1967

40-65"5 Bream, c" 25%
"
Pike, 10% Bream region Albrecht, 1964
2t-50 Lowland streams, 2-5 m wide,
~ 38"8 anglers' catches Herrmann, 1967
13-32 Lowland river, 20 m wide,
'$17"4 angler yields
9-19 Lowland river, 20 m wide,
~ 16"7 angler yields
47 Bream, Pike}
Bream region
Bream, Pike
"
'$ 14"2
Pike, 11 "2%
also Bream, Roach
}
Bream region Pape, 1952
Pike, 12"4% }
~ 10"3 also Bream, Roach
23-45 Pike, Burbot Bream region Mann, 1964
Barbel-cum-Salmonid region Buhse, 1967

"
Barbel-cum-Salmonid region
" T rout
Barbel-cum-Salmonid region
" Trout
Cyprinids, Pike Bag net yields l ens, 1967
Pike, Whitefish Hass, 1964,

Rameil,I967
Pike, Whitefish Bream region, professional and
" sport fishing T esch,19li7b
When conditions for the migration of young eels are optimal, a

further factor influencing eel productivity in running waters is the
nature of the river bed ; eels and trout have similar preferences in
this respect and some competition will result. As Burnet (1952a)
found, streams which offer many hiding places have the highest eel
yields. An ideal stream includes water plants and overhanging banks,
or plants arid trees which hang into the water or which have fallen
into the stream. Such factors provide most effectively for the eel's
negative phototactic tendencies.
The results of studies on the eel's diet (see p. 169) give the impression
that, above all, the eel and the trout are competitors for food. This fact
has received a great deal of attention in the British and American
literature. In Britain (and to a certain extent the British Common-
wealth) the commercial and sporting value of the eel is very much
260 THE EEL
subordinate to that of salmon and trout, so that the emphasis is primarily

on protecting the salmonids from the eels (rather than the other way
round). Judging from the results of many studies (see Sinha & Jones,
I¢7b as well as the literature cited therein), there can be no doubt
that, at least in summer, the three species are strongly competitive.
This fact is often used as the main argwnent against tolerating eels
in salmonid waters.
The possibility that eels might prey on the spawn, fry and young
of salmonid species is often considered of little importance (see in
particular Sinha, 1969). Nevertheless, studies on the New Zealand
eel (Cairns, I942b) and the American eel (Godfrey, 1957), indicate
that a great deal of such predation does take place. The heaviest losses
of salmonid fry as a result of eel predation in Canadian streams occur
in spring and early summer when the salmonid fry are emerging from
the gravel beds. It is at this time that the eels, which have not eaten
all winter, exhibit their first and strongest feeding activity (see p. 169).
Detailed research on the effects of eels on trout populations have
been carried out in New Zealand (Burnet, 1959, 1968). These
demonstrate that one cannot simply say that eels have a clear-cut,
negative effect on the trout. Electrofishing techniques showed that,
in a stream 3 to 4 m wide, the biomass of trout ( Salmo trutta) was 56
kg/ha and that of the eel was 68 kg/ha (64% of the eels were A. australis
and 36% were A . diefJenbacht) . After removal of the eel population,
however, there was no appreciable increase in the trout biomass. In
contrast, in a stream 2 to 6 m wide which had, as a consequence of
abundant hiding places, a well established colony of eels (47 % of
which were A . australis and 53% A. diefJenbacht) , decimation of the eel
population produced a very marked effect. The quantity of eels caught
in the first year in various sections of the stream amounted to between
400 and 1090 kg!ha. Trout catches, on the other hand, were not more
than 16 to 121 kg/ha. Only one year after decimation of the eel popula-
tion there was a large increase in the trout biomass. Four years later,
the trout yield had risen to 134-240 kg/ha. Due to a marked decrease
in mortality during the first year of life, the number of ttout in their
second year was increased by a factor of between 2'7 and 9'9. As a result
of this increase in weight and, above all, numbers, the growth rate of
individual trout showed a sharp decline. In one section of the stream
the trout now took four instead of two years to reach a length of 28 em.
In addition, their nutritional value worsened, and the fish became less
attractive to anglers. In a control section of the stream, where the eel
population had not been decimated, there was no change in the trout
FISHING YIELDS 261
population. There can thus be no doubt as to the validity of the results.

If one compares eel and trout populations in European streams (see
above) with those in New Zealand, in most cases there appears to be no
question of any adverse effect on the trout. Eel populations of the density
seen in streams in New Zealand occur only rarely in Europe, and in
places where this does happen, the majority of the population tends to
consist of small males (see p. 53). As far as one can tell from research on
feeding habits (see p. 173), these small males do not present much of a
threat to young trout.
It is much more difficult to determine the effect of a dense eel popula-
tion in standing waters, and to ascertain whether and to what extent
other species of fish have a positive or negative influence on the eel
popUlation - positive, in that they provide food for the eel, and negative,
in that they might compete for the same food or that they prey upon
the eel itself. This last possibility can also have a positive effect in
cases where the eel population is too large and, unless it is reduced,
suffers as a result of intraspecific competition for food. It is very difficult
to establish what effect the eel has within the animal communities of
standing waters or, conversely, the effect these envirorunents have on
the eel, because the interactions in fish corrununities are more complex
in such places. Broadly speaking a stream represents an extreme
biotope in which the bulk of the population is made up of salmonids
and eels. In most standing waters, however, many other species may be
present, and furthermore, the adaptability of these species is such
that, if need be, they are capable of moving into other biotopes.
Eels almost certainly had an adverse effect on the brook trout popula-
tion of a 154 hectare, oligotrophic lake in the drainage area of the St.
Lawrence River in Canada (O'Connor & Power, 1971). In addition to
sticklebacks and smelt, this lake contained 0'34 kgjha of brook trout
( Salvelinus Jontinalis ) and a considerably larger population of eels,
Other lakes in the same area, which were limnologically similar, were
without eels; in one of these lakes, in addition to char ( Salvelinus
a/pinus ) , 3'1 kgjha of brook trout were caught, and in two others, the
yields were 1·6 kgjha and 4'3 kgjha of brook trout respectively, In
the lake populated by eels, 7 out of 22 eels studied had eaten brook
trout (56% of the total volume of food found in these stomachs consisted
of brook trout). Monality among brook trout in their third year of life
was twice as high as it was in the other lakes; of course this, as well as
the following example, are only paralleled when the balance in isolated
lakes with extreme biocoenoses is disturbed by intruders such as the
eel.
262 THE EEL
~l~ 1898 1899 1900 19 01 1902 1903 1<Xl4 1005
:i-l,llilD QJl Cln

:bnn llJln.-11111111 •
1930 1931 1932 19n 1934 1935 19}6 1937
1950 1951 1952 IOSl 19H 19S5 19')6 1957
Fig. 91 Eel and whitefish (Coregonus) yields (in kg/ha) in a whitefish lake in Schleswig-
Holstein recorded over several periods since 1898 (modified after Herrmann
& Marre, 1961, and Herrmann, 1967).
It is possible to study the eel's influence on population composition

by the massive introduction of this or other species, followed by 100g-
term observations on the yields of all species under equally intensive
fishing. There are several examples of such studies in lakes in Schleswig-
Holstein (Herrmann & Marce, 1961; Herrmann, 1967). The most
convincing effect is seen in yields of Coregonus nasus and eels from
a lake in Schleswig-Holstein (see Fig. 91 ). Figure 91 combines yield
figures which were reproduced in instalments in various publications.
It shows that, in comparison with yields of Coregonus nasus, only small
quantities of eels were caught between 1898 and 1905. However, after
the introduction of eel stocking programmes at the turn of the century,
there was a sharp decrease in Coregonus nasus yields, which dropped
to less than 0'5 kg/ha between 1919 and 1949. Yields during part of
this period (1 930 to 1937) are reproduced in Fig. 91. ' In the mid-
195OS, as a result of intensive stocking and intensive fishing with
monofil gill nets, the Coregonus nasus yield showed a short-lived in-
crease of up to 3 kg/ha; during the same period eel yields were at a
similar level. However, with further intensification of the eel stocking
FISHING YIELDS
prograrrune and an increase in eel yields to between 4'5 and 6 kg/ha,

Coregonus nasus catches have declined so sharply that fishing for this
species is now practically non-existent' (Herrmann, 1967). This period
is illustrated in the lower half of Fig. 91 (1950-64).
In the first instance, such competition is probably attributable
to the predatory effect of the eel on Coregonus nasus. The eel not only
eats the spawn but it also attacks larger members of this species, This
is particularly clear from the fact that Coregonus nasus caught in gill
nets 'when the nets are near the bottom' (Herrmann, 1967),
are frequently eaten by eels. Similar observations have been made
in Lake Ontario, Canada. There, the white fish Coregonus clupeaformis
caught in gill nets were eaten by eels, and the nets completely tangled
(Hurley, [973).
Further examples of competition between eels and other species
of fish in lakes in Schleswig-Holstein are to be found in the references
cited above (Herrmann & Marre, 1961 j Herrmann, 1967). The com-
petition between eel and carp is easily understood since the latter
feeds on benthic animals and thus encroaches directly on the eel's
domain. For this reason, intensive, simultaneous cultivation of the
two species has been avoided in many places. The eel shows a similar
interaction with tench ( Tinea t inea ) . The bream ( Abramis brama )
must also be included among the eel's direct competitors, though
it is not difficult to control this species in such a way as to favour the
eel.
The eel is an undesirable competitor with valuable edible crustaceans
like the lobster ( Homarus vulgaris ) and the crayfish ( Astacus astaeus ) ,
Proof of this competition, however, is only available in the case of the
crayfish (Svardson, 1969, 1972). In southern Sweden, the eel and the
crayfish occur in many waters, with the crayfish commoner in the
eastern part of the country and the eel commoner in the west. In many
lakes these two species are sympatric. In a total of 159 cases, lakes
without eel populations showed a significantly higher yield of crayfish
(2'3 crayfish per hectare as opposed to only 1' 1 crayfish per hectare in
lakes where eels also occurred). In cases where the mean eel yield was
0'22 kg/ha, crayfish hardly occurred at all. In contrast, an average eel
yield of o· I I kg/ha meant that the crayfish could manage to survive in
the same waters. Svardson (1972) cites several examples of an increase
in the crayfish yield when, for some reason or another, there was a
decrease in the size of the eel population. Conversely, an increase in eel
catches meant a decrease in the number of crayfish caught, The best
known example is Lake Vanern; the construction of a canal leading [0
264 THE EEL
Table 31 Eel yields in inland lakes

Water body Geographical pesition Area ill Year Yield in kg/ha
hecrareJ Eels Total
Lough Ncagh Northern Ireland 39= Before 1 C}66 C. 20·0
Pike and Bream

L>k~ Schl~wig- Holslein c. so '954-1964 g'8-20-3 20'9-51"3
~ 1)'0 ~ 35"3
Bream Lakes Schleswig-Holstein 50-100 1959-1964 7"2- 15'6 24"2-126-7
~ 12'0 464"2
Bream Lakes Schleswig-Holstein c. So 1954- 1964 4'9-1)'5 43")-71'0
~ 7"9 ¢ 55"1
Bream Lakes Schleswig-Halslein 100-200 '954-1964 3'0-9"4 16-6-61 -3
¢ 6'0 rJ 32'4
Roach Lake Schleswig-Holstein ,= 1949-1964 )"5-7"7 20-1-46-4
¢ 5"0 ¢ 29' S
Coregonus Lakes Schleswig-Holstein ,= '949-1964 2 "2-6'2 12,8-26'0
¢ 4' 2 ¢ 18·6
Lake Rogglin Mecklenburg, near Schwerin '00 1957-1961 2'o-1S"1
97"9
Lake Steinhude Lower Saxony 3= 1959-19<)2 4·' 20"3
Lake Crecy New Brunswick, Fundy Bay
" 1949- 1963 I"S - S"!
~ 3"7
Waters in the
region round
,.•
Schwerin M~klenburg 25700 1954- 196 1 1"9-4"5 24"0-- 27"6
~ 3"2 ~ 2S"0
Lake DUmmer Lower Saxony ,,00 1¢x>-1962 11 "2
Lake Storkow SE of Berlin 3" 1937-1949 0"3-6"1 17"6-45"0
~ 2"45 ~ 24"83
Lake Bederkesa Norrhern Lower Saxony '3' 1957- 1958
" 9,
Eder reservoir Hessen, SE of Kassel '00 1937- 1958 0"03-0"90 15"0--98"8
~ 0"33 ~ 59"'
Eder reservoir Hessen, SW of Kassel '00 1959- 1965 2"2-4"1 59" 2-8 1"3
~ 2"8 ~ 7 2 "3
Lake Sakrow SE of Berlin '00 1949- 1956 0"6-3"7 7"2-38 "0
t/J I"g t/J 24"0
Waters in 2510 1¢o--1963 I "S-l"O 13"7

West Berlin t/J I"g
,. , ( l<}6l)
"
Lake Gibson New Brunswick, Fundy Ray 1945- 1951
Oder reservoir Southern Han: '3' 195 9 '·3 4·9
the sea rendered possible the through-passage of eels which therefore

populated the lake and from that time onwards there was a rapid
decline in the lake's crayfish population.
There is no evidence to support the hypothesis that the decline in the
lobster population on the rocky littoral regions of Heligoland could
have been brought about by an increase in the number of eels. The
increased number of eels caught in trawl nets in the Heligoland Bight
since 1963 (see p. 303) cannot be taken as evidence for an increase in the
eel population since this fishing technique, based on small-mesh nets,
FISHING YIELDS
% eels in Other important ChIJT<uuristics of W(lur body Liter(lture

total catch species of fishes
Salmonids (e"g" Salmon) Frost, 1950 ; T esch, 1967c;

Coregonids Rep" Foyle Fish Comm.
Average of several lakes
6-" Average of several lakes

; '9
8-26 Average of several lakes
; "
6-46 Average ofscverallakes
Hernnann, 1967
; '9
1I-35 Average of several lakes
; '7
u -44 Average of several lakes
1$ 21"4
Eel yields from stocking alone Gollub,I963
Bream, Pike-perch Pike-perch lake Tesch, 1967b

Salmon ids Low productivity Smith, 1966
Mostly lakes Gollub,I963
Pike, T ench Pike and Tench lake

Large Bream 17"9%
Pike u"4% Bream lake Muller, 1952
"
0" 1-1 "4
Bream, Pike-perch
Roach, Pike-perch, In the Miuelgebirge, both pro-
Tesch, 1967b
1$ 0"6 Perch fessional and sport fishermen Buhse,I967

1"7-5"7 Roach, Pike-perch,
1$ 3"8 Perch
1"8-50"7 Bream 14"3%
¢ 7"5 Pike 11 "6%
Roach 14"3% Bream lake Kahn, 1957b
Bream, Pike, Predominantly Havdlakes; Buchin & Muller, 1967
Pike-perch proressional fishermen only" Smith & Saunders, 1955
, T rout, Roach
Average depth 4 m
Angler yields,
Minelgebirge Tesch,I967b
was not employed before that time. Along the west coast of Sweden,
where eels and lobsters occur together, there are as yet no indications
that the eel has had an adverse effect on the lobster populations
(Svardson, personal conununication).
4-4.3 Improving yields by stocking

As was mentioned earlier, the further a body of water lies from the
coast, the smaller is the natural immigration of eels into it, and therefore
266 THE EEL
the lower are the yields. In Brandenburg and in large areas of Mecklen-
burg catches are already much lower than those in Schleswig-Holstein
(Table 31), where the average is 6'5 kgfha (Herrmann, 19(7), and
while in north Mecklenburg, near Rostock, yields are 7 kg/ha, in the
district round Cottbus, south-east of Berlin, they barely reach the
1 kg/h> mark (Koops, 1967». As> means of compensating for these
differences, stocking programmes were introduced as early as the rum
of the century (see p. 104).
A well-known example of the successful introduction of eels is Lake
Vitm in East Pomerania. Here, stocking has been in progress since 1909;
within a period of 30 years, the yield was increased from 0'7 kg/ha to
g·o kg/ha (Fig. 92). On average, 14 eels were introduced per hectare
per year (Strophal, 1930). Once the progranune had taken effect,
average yields of 4 kg/ha were obtained, so at least 3 kg/ha can be
considered the result of successful stocking. By far the greater propor-
tion of eels used were young eels (snigs) with an average weight of 20 g.
The person in charge of the stocking programme believed that the
introduction of eels at two to three times the scale used would still have
been within reason in order to attain the maximum yield on a pennanent
basis. Mter comparing the number of eels introduced with the number
'"
'00
I
'"
, l~:::J.l-.Q)ll .LL..Uf.J J,
1910 1915 1920 1925 1950 1955 1960
LG kt Yilm Lakt Yllm
Fig. 92 Eel stock and yields in Lake Vilm, 1750 ba (based on data from Stropbal,
1930), from Lake Rogglin, 200 ha (based on data from Gollub, 1963 ; and
Koops, 1967b), and from the Eder reservoir, 800 ha (based on data from
Buhse, 19(7). Outer columns - young eels ; inner columns = glass eels ;
curve 2 yield.
FISHING YIELDS
subsequently caught, Strophal (1930) concluded that 40% were

recovered; he asswnes this figure to be about 20% for the glass eels
stocked. Yields of other species of fish, including pike, roach, perch and
tench, decreased as a result of eel stocking. Of course it is impossible
to tell how far such decreases may have been due to other changes in
the lake. Financially, however, these losses were easily compensated
for by the increased eel yield.
A further example of stocking is Lake Rogglin in northern
Mecklenburg (Table 3[; Fig. 92). Gollub ([963) says that before [949
it was extremely unusual to catch an eel in these waters; in 1959, eels
weighing 500 g were being caught. However, as far as one can tell from
Fig. 92, catches of any recordable size have only been made since 1957.
The majority of the fishes were caught in stationary eel traps.
Although the first effects of eel stocking can be detected after four
years and, in the case of the warmer Lake Platten in Hungary, after
only three years (Koops, 1967a; Biro & Laszlo, 1970) the most marked
effect is first noticeable six to seven years later. In Lake Vilm (Fig. 92),
results were not seen until after 8 to 10 years. In Lake Rogglin, the time
lapse may have been even greater, but as there are no catch records for
the later years, this cannot be checked.
An average of 1 kg of young eels per hectare is necessary to obtain a
mean yield of 8 kg/ha. This stocking rate would have to be higher if
one were to include the number of glass eels introduced. However, any
noticeable result from glass eel stocking cannot be expected to take
effect until after the period covered in Fig. 92. If one takes into account
the size of the young eels, then an annual stocking figure of 30 eels
per hectare is necessary to obtain a yield of 8 kg/ha. These calculations
for stocking intensity and yield correspond with the figures for Lake
Vitm, where the actual rate was, in fact, at only half that intensity.
It thus appears that, in practice, with four to five young eels per hectare
it is possible to obtain an increase in yield of 1 kg/ha.
A less conclusive stocking experiment is the Eder storage dam
project, in which young, mostly glass eels, were introduced into waters
previously free of eels (Table 31; Fig. 92) ; some elvers were also
introduced during the period 1950-52. Data on weights provided by
Buhse (1967) were converted into numbers of eels in order to compile
the graph shown in Fig. 92. (Figures obtained from the eel export agency
in Hamburg show that one kilogramme of glass eels corresponds to 3000
fishes, and the same weight of young eels corresponds to 55 animals.
The last figure was derived from the average proportion of weight to
number of elvers from the Lower Etbe, which figure usually fluctuates
268 THE EEL
between 50 and 60 individuals per kilogranune.) According [0 the

diagram, the first peak yield, between 1959 and 1962, occurred after
intense stocking activity in 1950. In other words, the introduction of
glass eels becomes effective after about 10 years. This agrees with data
obtained by Jensen (1g61), who has shown that female eels spend
about 10 years in continental waters. The catch maximum of Ig61 is
hardly likely to have resulted from the introduction of elvers in 1950/
1952; these young eels probably gave rise to the small maximum of
0'7 kg/ha in 1955. One cannot use the Eder dam experiment to calculate
the quantity of eels needed for stocking because this water body is not
sufficiently isolated from surrounding waterways; the still actively
migrating young eels could easily have found their way out and entered
adjoining rivers. The only way one could have determined the effective-
ness of the stocking prograrrune in this drainage area would have been
to catch all the silver eels as they migrated downriver. Nevertheless, the
results do show that the annual introduction of close on 100 glass eels
per hectare did give rise to a noticeable improvement in yield. Compared
with other examples, and measured against expected yields from
young eels, this project was only a quarter as successful as it might
have been. In other words, in isolated water bodies, yields from stocking
with glass eels would have been four times greater, or, looked at in
terms of individuals stocked, 25 and not 100 glass eels would have
been sufficient to give the observed yield of 3 kg/ha. In completely
isolated water bodies, 13 to 15 young eels are sufficient to produce a
yield of 3 kg/ha, i.e. half the number of young eels are necessary to
achieve the same yield level as is obtained with the use of glass eels or
elvers.
Schaperdaus (1949) also found a similar relationship between the
relative effectiveness of stocking with elvers and glass eels; with young
eels one can expect losses of 40 to 60%, and with glass eels, losses of
80%. As noted above, Strophal (1930) reports similar findings.
When calculating glass eel losses, it is important to make certain
distinctions (Tesch, 1967b). Extensive water networks, which allow
the elvers to distribute themselves over wide areas and long distances,
result in higher losses than do isolated water bodies where the eels
are confined to the area of release. In the relatively small lakes
of Germany, for example, at least 10 glass eels are needed to produce
a yield of I kg/ha, whereas in the 39 000 ha Lough Neagh in Northern
Ireland, 45 glass eels are necessary to bring about the same increase
in yield (Koops, .t¢7a). Twenty-three to one hundred glass eels per
hectare were introduced for the first time into 43 Belorussian lakes
FISHING YIELDS
with a total area of 48 000 ha. The lakes had an unrestricted outflow;
only 4% of the eels were recaptured (Kostyuchenko & Prishchepov,
1972).
In central Europe even less favourable ratios can be expected with
intensive cultivation. For example, if one compares the quantities
of elvers introduced into various lakes in Schleswig-Holstein with
the resulting yields (Hernnann, 1967), it soon becomes clear that
stocking success is nowhere near as high as YJas to be expected from
the German experiments discussed above. In the last few years, an
average of almost I kgJha of elvers has been introduced into lakes in
Schleswig-Holstein. In a few isolated cases, like 50 hectare pike and
tench lakes, as many as 5 kgJha have been used. Despite the use of
such large quantities, there was only an increase of, at the most, 20
kgjha (from 10 kg/ha up to 20 kgjha). Without doubt, the fact the lakes
already had an above average eel population, must have had a consider-
able effect on the results. In addition, since yields of 20 kg/ha are
already approaching the upper limit for average production (see p. 270),
the cost of stocking would be well above average and, once the upper
limit is reached, additional stocking will fail to produce any yield
increase at all. Therefore, elvers introduced at the rate of 5 individuals
per hectare per year only serve to increase catches by I kgjha in waters
where there are few or no eels, as for example, in the three cases dealt
with earlier. Thus the profitability of stocking declines as the density
of the existing eel population increases - a general rule in the economics
of fish stocking.
As regards the effectiveness of stocking with young eels, figures
from Schleswig-Holstein show that 3 to 4 years must elapse before
one can expect to notice any appreciable increase in yields; with glass
eels, 8 to 10 years are necessary. The effects of glass eel stocking on
later yields are also demonstrated in an example from the river Mosel
(Jens, 1967). This author compared the yield curves resulting from
glass eel stocking carried out 2 and 5 years previously. In both cases
agreement was found between the curves, and it is concluded that
stocking may produce effects 5, and possibly even 2 years after the
introduction of the glass eels. However, in view of the examples given
earlier and the results of Danish studies (Jensen, 1961), this is unlikely.
In Scandinavian waters too, the effects of glass eel stocking were seen,
at the earliest, 7 years later. Maximum results were recorded after
9 to 10 years, and as many as 14 to 15 years later some effects were
still detectable (Eh, 1929). In Belorussian lakes, newly introduced
glass eels were first recaptured 5 years later and the majority was
270 THE EEL
caught after 7 to I I years (Kostyuchenko & Prishchepov, 1972). The

introduction of a generation of glass eels, it seems, is profitable to the
fishery about II to 12 years later.
New stock produces results much faster in southern waters than in
central and northern Europe. Data on the first attempts at stocking in
Lake Balaton (70000 ha), Hungary, have shown (Koops, 1(673) that
only one year after the first batch of young eels had been introduced,
eels measuring 47-50 em were caught. But, it appears that here too,
several years elapsed before any economically significant effects were
seen (Biro & Laszlo, 1970). However, the fact that fishing techniques in
Lake Balaton were not completely geared to eel fishing, may have
contributed to the time lag.
4.4.4 Summary of yield and stocking success

Waters near the coast give higher yields than do lakes and rivers situated
further from the sea. However, in bays open to the sea, the quantity of
eels caught per unit of surface area is relatively low, probably as a result
of competition from a large number of other fish species. In central
Europe, catches are between 2 and 10 kg/ha.
In lakes
High yields, particularly those in small coastal lakes, are between
10 and 40 kg/ho.
Average yields are between 3 and 10 kg/ha.
Low yields are less than 3 kg/ha.
In rivers
High yields are between 20 and 50 kg/ha.
Average yields are between 5 and 20 kg/ha.
Low yields are less than 5 kg/ha.
Yields from streams fished mostly by anglers are not comparable
with those from streams fished conunercially. However, if these streams
were to be fished conunercially, the yields would probably be, on
average, higher than those in lakes, as a comparison of eel biomass
in lakes and streams has shown. In New Zealand, eel populations and
yields are higher than they are in Europe and North America. This
is probably attributable to higher temperature levels in New Zealand.
Judging from yields in the lagoons of Italy, waters in southern Europe
have considerably higher yields than do the coastal regions of the
North Sea and the Baltic.
In cold water streams, the only fishes which compete with the eel
for food are the salmonids, and these do not seem to have a markedly
FISHING YIELDS 27 1
adverse effect on eel population sizes. However, in both warmer running

and standing waters, there are a large number of competitive species
which may reduce eel yields. These competitors include, in particular,
cyprinids such as carp and tench, as well as other commercially less
valuable species. Eel populations which are being improved or
maintained through human intervention, tend to have an adverse
effect on the yields of coregonid species.
In central European waters with small eel populations, or no eels
at all, 4 to 5 young eels (snigs) (each weighing between 20 and 50 g)
must be stocked per hectare in order to produce an increase in yield
of I kg/ha. In lakes with larger eel populations, considerably more
elvers are needed. If glass eels are used for stocking purposes, double
the quantity are required. Affluent waterways joining the system
mUSf also be included in the stocking programme.
5
Fishing Methods
Hardly any other species of fish is caught in so many different ways

as is the eel. Fyke-nets, traps and stow nets are used during the eel's
seasonal migrations and daytime wanderings; baited long lines, other
fishing tackle, fyke nets and trawls are used during the eel's foraging
uips; weirs, barrier nets, fences and banks of lights in combination
with stow nets and fyke nets are employed during the eel's spawning
migrations. In freshwaters electrical devices are used to trap the yellow
eel, and trawl nets are used as well. Seines, spears, dip nets or lift nets
are now in less frequent use than in the past.
Thus the eel's intermittent migratory drive, its need for food, its
aversion to light (negative phototaxis), its need for contact (thigmotaxis),
as well as its various habitats and behaviour patterns are all turned
to the fishennan's advantage. The eel is found in deep and shallow
water or in open waters; in some instances it will even leave the water
altogether. It may drift in streams and tidal currents or it may swim
against them (negative or positive rheotaxis). The eel can be found
on sandy, muddy or stony substrates in both freshwater and saltwater;
as far off-shore as the Continental Slope it is possible to catch the eel
almost anywhere; it is over the depths of the ocean that the eel first
becomes elusive. Excluding the capture of larval forms, no fishing
technique has yet been successfully employed to catch eels in the open
ocean.
Because of the multitude of methods used to catch the eel, it is
27 2
FISHING METHODS 273
difficult to give an exhaustive account of this field within the limita-
tions of this book; thus only an outline will be given here. Technical
details have had to be omitted; they can be found in books on fishing
techniques. Although such books do not always deal directly with
eel fishing, they give descriptions of many types of gear of a basically
similar nature, and in far greater detail (e.g. Seligo, 1926; Peters,
1935; Scbnakenbeck, 1940, 1942; v. Brandt & Rutkowski, 1956;
v. Brandt, 1957, 1959, 1964; Breitenstein, 1956, 1960; Jager, 1960;
and various authors in: EIFAC Technical Paper No. 14, FAO, Rome
1971).
5.1 Eel traps
Certainly the most widely known method of capture is the fish trap
based on the principles of the eel basket. Usually this trap is not used
on its own but in combination with other pieces of equipment. The
principle behind all types of eel trap is, by and large, the same. The
main differences are in the methods used to lure or guide the fish into
the trap. The latest method is to use channelling barrages or leaders
(walls of netting) which obstruct the eels' passage and guide the animals
into the trap. Bait is also commonly used to lure eels into the opening
of the trap. In addition, use is made of the eel's need for cover
(thigmotaxis), and its tendency to move away from light. Klust (1969)
has published a review of the types of trap common in Germany, in
particular those used in the larger rivers.
5. I. I. Eel baskets
The earliest form of eel basket is certainly the fish pot shown in Fig. 93.
Its early origins are due to the fact that primitive men were doubtless
able to build eel baskets by using plants as raw materials. The basic
principle behind the two (or more) chambers of the eel pot, whether it
is made from natural or synthetic fibres, or from plastic materials, can
be seen in all more recent developments of the eel basket. Although the
term 'pot' is only really appropriate for this device, the name has also
been used to refer to later designs made from textiles. In the coastal
fisheries of the North Sea and the Baltic traps made from netting are
referred to as 'baskets' (Bobzin, 1966) while the original wickerwork
basket type is in fact referred to as a 'trap net'. In inland waters the
term eel basket appears again when the original design made from
osier switches is being referred to (Briinunerstiidt, 1955); Fig. 94
274 THE EEL
Fig. 93 Sketch of an eel pot with two funnels; in many countries it is woven out of
reeds or supple twigs (Bohzin, I966; Deutsche Fi5cherei-Zeitung 1959. p. 33.
Neumann Verlag, Radebeul).
shows an example of such a design still being produced in 1965 by a

specialist craftsman in Hamburg.
A slightly more elaborate model is a basket made from the Spanish
reed (peddig reed) or rotan palm (Calamus ). It sinks soon after it
is placed in the water, thus ensuring the likelihood of an early catch.
The trap is produced from similar material in Japan; it is also well
known in North America (Eales, 1968) - i.e. it is used the world over.
Wooden barrels, slats and, most recently, wire are used to form the
outer casing.
One might imagine that this almost primitive apparatus would
gradually become obsolete, but this certainly has not happened. Its
manufacture in plastic has given it a new lease of life (Mohr, 1963a, b;
Koilike, 19643) (Fig. 95). Its catch records are similar to those of
wickerwork baskets as long as it is not used in tidal waters where, it
seems, a trap of somewhat different construction is needed. Originally
the plastic model was twice as expensive as the wickerwork basket.
Due to its higher cost the plastic model (described by Mohr, I963b)
Fig. 94 Eel pot from the Lower Elbe region used by the author in experiments near
Heligoland (Photograph: Tesch).
FISHING METHODS 275
Fig. 9S Plastic eel pot. Above: the upper side with a roof~like light shield. Below : the
lower side with twO iron rods acting as stabilizing weights. The b asket is made
from IS individually replaceable parts (Kothke, 19643) (Photograph: Klust).
which is woven and not formed from moulded parts, had to last at
least twice as long as the wickerwork type. The relative price of pots
and baskets made from moulded parts may, however, become more
favourable as time goes on, since continually rising costs of craftsman-
ship will make the wickerwork model less attractive, while the price
of plastic goods will show a proportionate decrease.
The eel basket is worthy of note not only in professional circles but
also among amateur fishermen. Due to its ease of use, the basket can
be set up without difficulty by the layman. It is popular among amateur
fishermen and for experimental and research purposes. I t is used to
its greatest advantage in rivers and tidal waters with dense eel popula-
tions. Its small size and robust construction make it ideal for use in
strong and changing currents.
In the tidal areas of the Elbe as many as 60 eel pots or baskets may
be firmly anchored to a long, securing rope; each basket is fastened
at 5-6 m intervals with a short rope, so that the individual eel baskets
neither touch nor damage one another as a result of the movement
THE EEL
of the current. Bait is placed in the last compartment of the basket,

and the eels are attracted by the diffusion of the bait's odour.
Experiments where the eel's sense of smell has been impaired show
the extent to which the eel depends on the bait and its smell for finding
the basket (Tesch, 1970). Eels were caught, predominantly in baited
baskets, in the estuary of the river Elbe, were marked and then set
free; 22 % of the eels with undamaged olfactory organs were caught
again whereas only 12 % of those animals whose sense of smell had
been impaired experimentally were recaptured. At another place,
where unbaited, netting eel traps (chiefly fyke nets) were used, 12 %
of the undamaged marked eels and I 1% of those animals with an
impaired sense of smell- i.e. only slightly fewer - were caught a
second time. Any professional fisherman who has set up eel baskets
with and without bait next to each other, will confirm these results.
The success of the catch is also considerably affected by the type
of bait used - as can be confirmed by anglers (see p. 310). At different
times the eel reacts only to particular baits. At certain times, for example,
it can only be caught using smelt. Anyone who feeds eels in an aquarium
will confirm this apparent wilfulness in the eel. It always takes a certain
time before the animals become used to alternative types of food.
Fishermen from the Lower Elbe regions give the following explanation
why eels prefer smelt as bait only at certain times: while the smelt is
still small, it does not manage to escape from the eel. During this
period the smelt is still small enough to pass through the mesh of the
wicker baskets. As the smelt gets bigger, it manages to escape from
the eel. With their hunger aroused by the bait, the eels are eager for
the smelt and are thus attracted into the baskets from which the larger
smelt cannot escape. This is thus the beginning of the period for smelt
bait - i.e. when the smelt is slightly larger in size. Other examples
of suitable baits are shrimps (Crangon sp. ) or prawns (Neomysis
vulgaris) , mitten crabs, earthworms, crushed mussels and the common
whelk. ( Buccinum undatum) , fish spawn, shredded lampreys and even
pieces of salt herring (Mohr, 1963a). According to fishermen, only
fresh bait is taken. Bait left in the water longer than 12 hours loses
its effectiveness. This useful finding has also been demonstrated by
the author in experiments in aquaria; food remains left after feeding
are not eaten the next day.
Baskets or pots placed just before weirs and dams can also be set
up without bait. If the eel's passage is temporarily blocked during
upstream migration, it will seek shelter (negative phototaxis,
thigmotaxis). Because of this tendency to seek shelter, eel baskets
FISHING METHODS 277
made out of synthetic material are constructed with a roof-like shade
over the middle section. The use of such baskets or pots in combination
with woven materials as leaders - as described by Walter (1910) and
Seliga (1926) - is less common nowadays and has been superseded
by the use of modem equipment.
Although eel baskets are basically for use in river and tidal areas,
they can, with certain alterations, also be used in lakes (Briinunerstiidt,
19S5). In such situations they are used in combination with artificial
spawning beds for dace, white bream and bleak ( Leuciscus rUlilus,
Blicca bjoerkna, Alburnus alburnus ) , made from conifer twigs (e.g. from
pine trees), or else they are placed in natural spawning grounds. The
eel, a spawn thief, visits such places and, from time to time, takes
refuge in the pots, which must be placed in the immediate vicinity.
Even as early as the spawning time of the pike, the eel can be caught in
the spawning areas. When the spring floods begin to recede, eel baskets
are placed in these areas, in the hollows, ditches and channels where the
eels withdraw as the water level drops. Not only wicker pots (Feddersen,
1893; Fisch u. Fang, 10, 160, 1969) but also netting eel baskets of many
different sorts (e.g. fyke nets) are used in the spawning grounds.
S.1.2 Fyke nets

Following the basic principle of the wickerwork eel pots, eel 'baskets'
made from cord netting have been developed. Initially cotton and
other natural fibres were used; nowadays, however, synthetic thread
and wire are more common (for the advantages of synthetic netting
materials, see v. Brandt, 1961 and Oray, 1968). Figure 96 shows one
type of gear used in many places in inland waters and on the coast,
the fyke net. The first funnel has a relatively wide opening and has a
Fig. 96 Fyke net or winged eel basket (after Walter, 1910).

278 THE EEL
~ .~? .••••
Fig. 97 Paired or double baskets from the Baltic coast of Germany (after v. Brandt,
1959)·
square or U-shaped cross-section, while the last funnel (the codend

funnel) forms a very narrow, slit-like aperture where it opens into the
farthest chamber (Brietenstein, 1956), from which the eel finds it almost
impossible to escape. Three, instead of two funnels may also be installed.
The number of frames (hoops) varies between 5 and 7; their diameter
can be very variable too. The length of the two wings is optional, or
they can be left out altogether and replaced by a single 'wing' in the
middle, a so-called 'leader'. Another basket, which faces the entrance
of the first, can then be placed at the other end of the leader, and further
fykes can be added, as is shown in Fig. 97. Eel fykes with adjacent end
chambers can also be led directly into one another thereby producing a
double fyke net whose entrances open in opposite directions (Bobzin,
19(6). On the coast near Mecklenburg such eel fyke nets are in common
use; they consist of six double fykes strung together with two end
fykes, and measure a total of 42 m. Double fyke nets are 2·30 m long,
the end fykes are 1'7 m and the leaders or guiding sections between them
are 3'6 m in length. Such fyke nets are usually made on circular frames
or hoops (Figs. 96 and 97); however, the first two frames of the
Mecklenburg fyke nets are semi-circular. The linking leaders, and
sometimes the first frames too, are weighted with lead to ensure close
contact with the bonom. Such relatively small fyke nets are easier to
handle, are seaworthy, and may therefore even be used in the strong
tidal currents near Heligoland as well as in rivers. On the coast of the
Baltic Sea it is common for 20-30 fyke net chains of the type just
described to be joined together, producing rows over 1 kilometre in
length. Normally, however, such rows are only 300-500 m long. The
ends of the rows are anchored to one another and are marked clearly
with a buoy. Long rows of fyke nets are also marked at intervals with
small floating flags. During the sununer months these combinations of
fyke nets are used along the coast where almost only yellow eels are
caught.
In lakes, wir.ged fyke nets are likewise in common use. Here, however,
it is more usual to anchor the wings and leaders with stakes. In addition
lanes are cut through the reed swamp. Several winged fyke nets are
stretched out behind one another, each between three stakes; the nets
are arranged in lines within the pathways cut through the swamp.
FISHING METHODS 279
Fig. ,s Combination of eel baskets for use in the wide littoral zone of a lake (after
Walter, 19IO).
The layout becomes more complicated when there is no reed swamp

and the littoral region is very extensive. A combination of central
and non-return valves, wings and leaders (Fig. 98) must then be used;
these are able to trap a large part of the passing population of yellow
and silver eels. Ingenious constructions are also used at the outflows
from lakes for it is often illegal to completely block the outlet. In many
cases this is not possible anyway, because of water traffic. Since, usually,
the eel does not approach the outlet directly from the middle of the lake,
but from the sides, central fyke nets with wings and non-return valves
are laid down in the lake, leading from the two banks of the outlet.
Besides this system offyke nets with three chambers there are a large
number of other possible combinations which are, of course, not used
exclusively to catch eels. The round fyke net has proved a success for eel
280 THE EEL
.•
J
...
, ...
Fla. 99 Pound net used on the west coast of Hiddensee (photograph: T esch).
fishing. It is easy to move, since it is erected with only four poles,

although it does consist of four sections - i.e. a winged chamber, the
wings and the leader, to which are joined three further chambers
FISHING METHODS 281
without wings. The non-return trap is also well known; unlike the model
shown in Fig. 98, which consists of several fyke nets, it comprises only a
single net with extended wings, which, as in the pound nets (in Figs. 99
and 100), fonn a box-like non-return valve, so that the fish is always led
back to the basket (Breitenstein, 1956; jager, 1960).
Enthusiasm for different constructions knows no bounds among eel
fishermen. Practically every fishing site has its own characteristics
which are strongly influenced by water depth, the gradient of the
sides, the type and extent of the bottom, the weather and the current.
Hydraulic structures, water traffic and such like must also be taken
into consideration. As a result, knowledge and experience playa large
part in the installation of this type of fishing equipment, for, although it
is possible to buy such basic parts as fyke nets, wings and leaders,
these may not necessarily be the right size. Thus the fishermen must
be prepared to take up a netting needle to alter or even reconstruct the
basic net.
Finally, the simple braced bag (cylindrical or barrel trap) should be
mentioned. These are seldom, or only occasionally used by professional
eel fishennen in central Europe but are used frequently by amateurs
because they are easy to handle. They consist of three hoops covered
with netting, each with a single entrance at either end. Of course, the
eel mana2es to escaoe frpm such device§,relatiyelv g~ickl¥, The techni-
calities of construction and the importance 0 a built-in non-return
device (funnel) in fyke nets used to catch eels have been demonstrated
by Kipnick (1965). The relationship between the width of the netting
mesh and the size of the eels caught is illustrated by Fig. 117.
5.1.3 Pound nets

Pound nets are very large j one can install them in combination with
winged traps, i.e. fyke nets, or additional structures can be joined on,
such as spacious trapping chambers made out of netting with large
flaps. Antechambers may even be added. Pound nets are used along the
coastline or in large lakes where it is worth covering a relatively wide
area to catch a correspondingly large number of eels. A typical region
where pound nets are used is the Baltic Sea coast where, particularl)i
in the western and central parts, migrating eels are (Q be found in
relatively high concentrations. Finland's annual eel catch does nOl
even reach 50 tonnes whereas Sweden catches 2000 tonnes and Denmar~
4000 tonnes a year (Table 25). On the German coasts over 2000 tonn~
of eels were caught in 1939 (Neuhaus, 1967).
282 THE EEL
Fig. 100 Drying pound nets on Hiddensee (Photograph: Tesch).
During their westward migrations (see p. 235) silver eels often move
up to the coastlines which hem them in along the northern and southern
sides of the Baltic, and where they are therefore easily caught. Many
coastlines, in particular those of Sweden, Denmark and Gennany) are
not as exposed to the prevailing west winds as are, for example, the east
coasts of the North Sea, so that the weather and the sea cause fewer
problems. A favourable factor in the Baltic is the almost complete
absence of tidal currents, with the result that the water level remains
constant. The use of pound nets in the western part of the Baltic has
therefore developed into an important element of the coastal fisheries;
pound nets are used chiefly to catch silver eels.
The eel pound net has evolved from the herring pound net whose
large dimensions suggested that it could also be used to catch large
numbers of eels without posing any technically insuperable problems in
FISHING METHODS
" Vitti
HIOOEN
SHU"" . •.\
Fig. 101 Pound nets used around Rugen. Arrows indicate traps erected in aurumn, i.e.
traps intended mainly for silver eels. Between the island of Rugen on the
one hand, and the islands of Ummanz and Hiddensee on the other, there are a
further 80 eel basket sites not shown here (modified after Roy, 19(1 ).
the inclement weather of the outer coast. In the most important eel
fishing region (the coasts off the Isle of Rugen), the large pound net
was further developed in 1911, after silver eel fishing had properly
begun there in 1895 (Schlieker, 1957). Figure 99 shows pound nets in
use near the island of Hiddensee; its floating guide is kept upright by
spherical floats and is held in position by anchors. This equipment
differs from the fyke nets described previously in that the single
chambers are surrounded by vertical walls of netting and are open at
the top. Figure 100 clearly shows the dimensions of the netting walls
of these pound nets. A leader may stretch from the beach or from a
sandbank for as much as 160 to 1080 m out to sea (Nolte, 1938). The
whole structure may be erected as much as 12m below the surface
THE EEL
(Bobzin, 1965). With very long barrages two or three trapping chambers
are used. It takes 8-10 men to erect one such trap; a group of
.independent fishermen usually get together for the job, and each man
has his own particular fishing grounds. Figure 101 illustrates the position
of the many pound nets round the isle of RUgen.
Whereas in recent years stake weirs have been replaced to an ever
increasing extent by anchored floating leaders, there now also s,eems
to be a trend towards anchoring the floating net walls of the chambers
(Martinkowitz, 1961; Bobzin, 1965). The leader is anchored by its
lower edge and the trapping chambers are fastened to a steel wire
frame that is bent around them. The trap is kept at the surface by
buoys which are designed so that the net walls begin to sink if the
current reaches more than 0'33 m /sec. The first chamber has an upper
'apron' and the second chamber has a 'roof of thin netting; both these
structures are intended to prevent the fish from escaping as a result of
the motion of the sea or of strong currents. Due to their flexibility
pound nets with anchors are more resistent than those with stakes;
moreover, they take about 50% less time to set up and dismantle.
The first time anchored traps were used they were shown to be equal
in catching power, if not superior, to staked baskets (Bobzin, 1965).
Using this type of trap it may even be possible to fish at greater depths
than before (Martinkowitz, 1961).
As on the coasts of Rugen, pound nets have also been used on the
Schleswig-Holstein coasts of the Baltic since the 19505. They were
not, however, developed from the Pomeranian pound traps but from
the Swedish or Danish 'Bundgam' which has been used for many
years to catch herring, cod and flatfish. In 1964 there were already III
traps from the Flensburg fiords to the bay at Neustadt; these gave a
considerable boost to eel catches in this area and led to other fishing
methods becoming less popular.
Size II
~ BttjiMi~Of
""~~ Stptember
~StPttmblr
• • , , • I I I , I
o
" '''''"
Fia. 103 The proportion of large (I) and small (II) silver eels caught between
September and November in Sassnitz (Nolte, 1938),
FISHING METHODS
'Bundgarn' is Danish and means ground basket. Such traps represent,

so to speak, a compromise between a pound net with its vertical netting
chambers open at the top, and a fyke net with its closed frames (Klust,
1965). Fyke nets attached to the main chamber in the direction of the
current are also to be found on pound nets, both in the inland coastal
waters of the area around Rugen and in lakes as well (Schlieker, 1957).
The 'Bundgarne' from Schleswig-Holstein are installed with floating
leaders measuring 150-600 m in length, and extending to a depth of
up to 6 m (Henck, 1965). In 1964 materials for a 'Bundgarn' with a
a barrage measuring 200 x 6 m cost 8000 DM. A trap cost a further
3000 DM if it was made by a net factory and not by the fishennan
himself.
Henck (1965) gives the following description of fishing with a
'Bundgarn': 'The eel, which is swimming along the coast on a mainly
east-west and later north-south course, comes across the leader which
has been laid out diagonally from the coast, and tries, by swimming up
to the surface of the water, to find a way out. Since, due to the constrUc-
tion of the weir, escape is impossible, the eel then swims down deeper
into the water. At the end of the leader the eel, guided by the wings, is
led into the entrance chamber and later into the main chamber. Usually
the eel then comes across the entrances to the two winged fyke nets
(with two funnels). To take the catch from the 'Bundgam' it suffices to
remove just the fyke nets and not, as previously, the whole trapping
chamber!
The annual fishing season for silver eels lasts from the end of August
until the beginning of December ; catches usually reach their peak in
October. At the beginning of the season small eels predominate in the
catch, whereas from the peak through until the end of the season larger
eels are caught. On the whole this observation is confinned by records
from the coasts of Riigen (Fig. 102), although there the highest catches
occur more often in September than in October (Rumphorst, 1930).
In fact, this generalization is more applicable to the Greifswalder
Bodden (which lies further inland) than to the east coast of Rugen
(see p. 221 ). Tagging, and recent fishing statistics, have confirmed the
tendency implied here - i.e. for the eels to travel from south to north
and later, in a north-westerly direction (Martinkowitz, 1961)
(see p. 235).
5.2 Stow nets
In recent decades the use of stow nets, bag nets with fixed mouths
(also known in Britain as coghills) has become more common. This
286 THE EEL
preference for stow nets in European rivers and coastal waters is, in
many ways, due to the fact that the eel is the only remaining species of
any commercial value - i.e. fishes of less commercial value are no
longer in demand in the markets. Basically, as far as construction is
concerned, the stow net is a simplified version of a large fyke net. Stow
nets used in eel fishing even have a funnel device, the typical characteris-
tic of an eel trap. The main difference between the two types of net
is that the eel is drawn passively into the chamber of the stow net by the
current, whereas, on the whole, the eel actively makes its own way
into an eel trap.
The adult eel makes use of currents when it travels over the Continen-
tal Shelf to its spawning grounds, and, unlike the younger non-migrat-
ing eel, does not move mainly along the sea bed but chooses the open
waters where it can utilize the strongest currents. In fact, the eel travels
where, for hydrodynamic reasons, stow nets are also best positioned.
This property of stow nets means that all such nets, regardless of their
individual structure, are valuable eel catching devices.
A technical problem in the use of stow nets is how to keep the net
chambers open in the current. In practice there are three solutions to
this problem, as follows.
5.2.1 Staked stow nets

The net is held open by stakes driven into the ground. This type of
net is used in the Lower Weser and Lower Ems areas and in Holland.
Up to so stow nets are set in a row along the coastline and are directed
against the high or low tide currents. However, they are gradually
displaced by shipping and, in Germany, are only to be found on the
rivers Ems and Stor (pape, 1970j Nolte, 1967).
5.2.2 Anchored stow nets

A large right-angled frame keeps the net open (e.g. the anchored stow
nets from Altenwerder on the Elbe) or an upper and lower beam only
are used (e.g. the eel jschokker' from the Rhine). One can explain the
principles of this type of trap most easily by describing the net separately
from the boat to which it is usually attached (Fig. 103). The boat to
which the stow net is fastened, is held in position by anchors. Anchored
stOw nets are laid in position by relatively large boats; because they are
attached to a boat and to one or more easily manageable anchors,
anchored stow nets are more mobile than staked stow nets. This
FISHING METHODS
".
Fig. 103 Anchored stow net from the middle Elbe. A-B, net anchor; B- B, chain,
10-15 m long; B,-C, wire cable, 10-15 m long; C-O'_4' securing hawsers,
20 m long ; 0,-0, and 0 3-04 horizontal beams, 4-8 m long; 0 ,-° 4 and
0,-°3 , vertical beams 4-5 m long ; bag length, from frame to H, 15-20 m
long; F H , codend, 2 m long, with built-in throat (after Kisker. 1926).
mobility is the chief reason for their use in broad river basins - e.g. in
the tidal areas of the large North Sea rivers as well as on the Upper
and Lower Rhine from Kehl onwards (Kriegsmann, 1966). They will
remain popular as long as pollution from sewage does not put an end
to stow net fishing in general (Felszykiewicz & Wurzel, 1966; Quednau,
1966; Traluns & Denzer, 1967).
In the Lower Elbe and its estuary, in places where the currents are
strong and numerous, it is possible at times during the migrations of
the silver eel to see up to five 'Altenwerder' stow net fishing cuners
very close to one another. The 'Altenwerder' stow net fishers - certainly
the oldest representatives of this branch of fishing in Germany - use
two stow nets set on either side of a motor cuner lying at anchor.
The length of the beams guarantees an opening in the net of about
8'5 x 4 '5 m. Cotton net chambers, 24 m in length, weigh about 50 kg,
and perIon net chambers weigh about 30 kg. In addition, there is the
codend which, for eel fishing, is divided into two sections each with a
built-in funnel. It has a total length of 5'5 m (K6thke & Klust, 1956).
Nowadys, the anchored stow net cutters from the Lower Weser use
some of their stow nets with only an upper and a lower beam, and are
thus similar to the eel 'schokker' nets on the Rhine. Figure 104 shows
an anchored stow net that was introduced for the first time in the Middle
Weser in 1964. It should be noted that the eel 'schokkers' of the Rhine
remain in the same place for the whole of the fishing season and therefore
do not have a motor. The boat is anchored in such a way, however,
that the vessel can be moved after the nets have been brought in.
The booms, which are hung out on one or both sides, together with the
nets that are fastened to them, are 8-12 m long and span a 4-6 m high
net which measures 24 m. When fishing, the codend of the stow net is
stretched over the first hoop of a fyke net (Podsadlowski, 1949 j Klust &
288 THE EEL
Fig. 104 Anchored stow net cutter, with traps on each side, in the middle Weser
below the Landesbergen storage dam (Photograph: T esch).
Schiirfe, 1954; K6thke & KInst, 1956). One can safely asswne that
most anchored stow nets in use today have synthetic netting (Klust,
1954)·
5.2.3 Otterboard stow net

A net chamber with an opening at least 16 m wide and with wings 2S m
in length is fastened to the shore by ODe wing which is itself attached
to a cable. The second wing is secured to an otterboard which is used
in the same way as in trawling but which is larger in size. The otter-
board in nun is fastened from a distance to the shore. The otterboard is
driven into the middle of the river by the force of the current, thus
holding the large net wide open. Using one of the ropes that secures
the otterboard to the land (the steering cable) and a steering device, the
otterboard can be brought near to the bank in a short time and the
river can thus be cleared very quickly (Fig. 105). Slightly larger nets
are not constructed with wooden onerboards, commonly used until
now, but with a sectional board made from metal (Schlieker, 19643, b).
The superiority of the onerboard stow net over the previously widely
used framed stow net on the river Elbe is clearly illustrated in Fig. 106.
On the river Oder conditions were equally favourable for the two
types of stow net. In Ig61 an onerboard stow net near Graben (Berlin)
yielded a catch of 10 tonnes of eels; five anchored stow nets with
FISHING METHODS 289
Cod tnO ""th throcu
~ OirvctKm
of current
Fig. lOS An otterboard stow net as used in the Elbe and the Oder.
frames, positioned in the vicinity, caught a total of only 4'6 tonnes.

In 1961 the latter were replaced by an otterboard stow net. The two
stow nets now recorded catches of 10 tonnes of eels each (Schlieker,
19643-). This conversion to otterboard stow nets resulted in an estimated
increase in production of 700-800% .
It is surprising that the otterboard stow net took such a relatively
long time to be adopted into general use - Hugo Kothke invented it in
1937. It is interesting to speculate whether the otterboard stow net will
also take over from the anchored stow net in the Upper Elbe and Upper
Weser regions (Klust & Scharfe, 1955). There the raison d'etre for the
anchored stow net combined with cutters stems from the continual
need to tum the traps as the tide changes. In addition, the bank often is
too far away to allow for anchorage on land. A further problem arises
THE EEL
55
• •
5~
D
Fig. 106 Eel yields before and after the introduction of otterboard stow nets in the
Elbe near Gorleben. A, average annual catch from 8 to 10 framed stow nets
before regulation of the Elbe; B, lo-year average from the framed stow nets
after regulation of the Elbe; C, 6-year average from otter board stow nets
made out of cotton thread; D, 1954, after the partial introduction of otter-
board stow nets made out of perlon netting, and during unfavourable
water conditions; 1955, after complete change-over [Q perlon nets (after
Kothke & Klust, 1956).
when it is necessary, or possible, to change the fishing site frequently.

Only a completely new invention could possibly suit these conditions.
Figure 89 shows the annual output from the Middle Oder fisheries
where mostly stow nets are in use. June shows the highest catch records.
Just how much the silver eel contributes to these figures is not clear
from the statistics. Some indication of the proportion of silver eels is
given in Fig. 90 where an analysis has been made of the percentages of
yellow eels, silver eels and eels which are in some intermediate stage.
Whereas in May, June and July relatively few silver eels were caught,
from August onwards by far the largest proportion consisted of silver
eels. In contrast, the proportion of yellow eels was high at the beginning
of the season but then dropped back to just a few percent as September
approached. Presentation of data in this form enables one to see that
catches from stow nets in no way consist exclusively or even chiefly of
silver eels, but also include a large proportion of yellow eels. These
findings have been backed up by the results of stow net fishermen from
the Elbe estuary. There, the first large catches in the spring consist
exclusively of yellow eels. The size of the catch then shows a sudden
decrease, and only in September do large numbers of silver eels appear.
It thus seems that in the Rhine and in the Oder too (Fig. 89), it is not
only silver eels that make up a considerable proportion of the catch.
FISHING METHODS 29 1
Fig. 107 Average daily catch from 4-5 Querboard stow nets in the Elbe above
Lauenburg in 1956 ( .. " . ,) and 1959 (- -). Above. temperature and
water flow; below. lunar phases (afler Luhmann & Mann. 1961),
Figure 107 shows the daily catches from an otterboard stow net in
the river Elbe near Gorleben, It indicates that the catch depends very
much on water flow, When the flow of water is low, the daily catch
may be as little as 2 kg, When the water level is high the figures may
rise to almost 40 kg per day. With an average flow, the annual mean
catch varied between 4 and 18 kg per day per net, depending on the
fisheries concerned (Table 32), On occasions when water levels have
Table 32 Ouerboard stow net catches (in kg per nel per day)
in the Elbe below Schnakenburg (after Luhmann &
Mann. 1961 )
W<lttr flow Y t <lr kg/net/day kg/net/day
m"/sec Fishtry A F;sht ry B
877 19S6 1S'22 11'95

904 1957 10' 18 , .",
'" 1958 18' 12 8·88

57 '
499
1959
' ,00 ,."
7·" 3"8S
4'58
risen more than usual outside the fishing season, as in 1957, catches have
not shown a corresponding increase (LUhmann & Mann, 1961).
Of course other types of fish, both adult and young, are also caught
in eel stow nets, though not in very large numbers. In order to protect
292 THE EEL
the populations of YOWlg fish, devices have been thought up to enable

such fishes to escape (Wundsch, 1934; v. Raben, 1937, 1952). In
practice, however, these have not been introduced (Schiemenz, 1962),
probably because they were not efficient enough.
5.3 Permanent eel traps

In streams and smaller rivers, where the question of disrupting river
traffic does not arise, it is fairly conunon for permanent eel traps to be
constructed. Such traps can be used where there is sure to be a fairly
regular flow of water, where the current and volume of water is not too
great, and where pollution and floating debris do not damage nor clog
up such a device too quickly. With permanent eel traps it is necessary
to cordon off the whole or part of the stream or river in order to guide
the eels, which are swimming downstream, into the traps. The same
principles, then, are used here as in ' temporary' eel traps (e.g. stow
nets). With the latter the eel is led through funnel-shaped netting
wings into the net chamber which, in many cases also has its own funnel,
positioned so as to prevent the eel's escape. Similarly, in some stationary
eel traps stow net-like chambers are used whose wings are simply
fonned from huge, securely built fences or barrages. Traps constructed
in this way are used mainly in rivers. In streams one finds, ahnost
exclusively, rigid chamber traps which consist pardy of duckboards.
Eel weirs span rivers and are fitted out with stow net-like chambers
as well as rigid ones. I t is important to distinguish between eel weirs
which are positioned in streams, and those which are used in rivers in
conjunction with turbine plants.
5.3.1 Eel weirs in rivers

The best known example of a large eel weir is that of the river Bann in
Northern Ireland. The conditions there are very favourable. Northern
Ireland lies in the path of the glass eels' vanguard arriving from the
Atlantic. The river Bann is supplied with water largely from Lough
Neagh which is 39000 hectares in area and is the largest lake in the
British Isles; therefore the prerequisites for a large area of fish
production are met (see Table 31). Floating debris and pollution are
infrequent and the outflow varies from 9 to 470 rn 3 /sec (Frost, 1950).
Figure 108 shows a diagram of the massive weir below Lough
Neagh where eight stow nets (known in Britain as coghill nets) can be
brought into use. Gates, to which stow nets have not been attached,
FISHING METHODS 293
can be shut off by vertical wire netting flaps (butterfly gates), so that
when the water flow is low, for example, trapping is limited to just a
few stow nets. In addition, these 'fishing eyes' can be completely
blocked off so that at extremely low water levels, water will flow only
through the stow net gates.
The individual nets are 12 m long, each with an additional 6 m of
codend, which contains a built-in non-return funnel. After the codend
has been taken in, the catch is ferried over to floating containers moored
next to the nets. The nets can also be emptied by the use of tubing
led from the codend straight into the containers.
Half-way between Lough Neagh and the estuary in the Atlantic,
there are other stow net weirs of the old style (Fig. log), like those that
are also common in Canada. They catch a mere fraction of the amount
caught by the new traps situated further upriver. This is not surprising
because the new traps presumably catch all the eels coming from the
lake before they reach the older traps. Fishes caught in the lower,
older weirs obviously originate from the river Bann alone and from a
small lake that lies below the new dam. Apart from this, the older dams
are very susceptible to floods. In addition, half-way between the lake
and the sea there is a further new style eel trapping device called a
'self-fishing weir'. Prominent features of this structure are two huge
funnel-shaped leaders that extend more than 50 m into the river and
Got~!, O?frl Gates,open
'"""
fishing
whtntht
Fig. 108 Diagram of the eel weir at Toome on the dver Bann (Northern Ireland)
(after Frost, J950),
294 THE BEL
Fig. 109 Section of twO old ~tI weirs. one behind the other, near Porma, on the river
Bann (Photograph : Tesch).
have a correspondingly large number of wire netting flaps. These, when

closed, are intended to guide the eels into enormous storage chambers
which, at the same time, act as containers. These very costiy and out-
wardly imposing structures obviously did not come up to expectations,
with the result that the old weirs nearby have remained in use (Tesch,
1967c). The best and most effective structure is thus the newer weir at
Toome which was described earlier.
According to figures for the years 1960-6 (provided by the Ministry
of Agriculture for Northern Ireland), the silver eel fishing industry,
represented largely by this one weir, brought in an annual average of
240 tonnes - i.e. more than 6 kg/ha. If the figures for yellow eels
caught in the lake during the same 7-year period are included, the
average totals 744 tonnes per year. Approximately 32% of this above
average yield is attributable to the eel weir alone, which employs an
extremely small labour force. What is more, 96% of the silver eels were
caught during the months August to November (Frost, 1950), so, in
fact, it would have sufficed to man the weirs for these four months only.
Of the total annual catch 0'4% was caught in June; 1'5% in July;
9'9% in August; 32'4% in September; 45'2 % in October; 8'6% in
November; 0'4% in December; and 1'6% in January. Thus, as in
north Gennany, the main part of the migrating eel population is on
the move in October (see p. 221 and Fig. 88).
The silver eel fishery on the river Bann is certainly an extreme
example of optimal conditions. The question arises here whether the
FISHING METHODS 295
Fig. 110 Canadian eel weir. Measurements are in feet (after Day, 1948).
yield would increase or decrease if the use of fyke nets, long lines and
trawls (previously seines; Frost, 1950) were discontinued. Permanent
eel traps are serious competitors to all other fisheries in the same area,
especially when permanent eel traps are sited above the other fishing
gear. In the area of Lough Neagh the permanent eel weir is at a dis-
advantage, and its productivity depends on the intensity of other
fishing methods in Lough Neagh.
It seems difficult to find favourable situations for the erection of eel
weirs. In 1964 there were 40-50 eel weirs in the Canadian province of
Nova Scotia (Fig. IIo). However. in 1962 such weirs in the
coastal provinces of Canada contributed only 7% of the total
catch. In general, their annual catch barely reaches the 2 tonne mark,
though there is one well known weir in the province of Quebec which
has an annual yield nearing 30 tcnnes (Eales, 1968); the Question
arises. in cases of such inefficiency, whether the cost of installation is
covered by the catch. One of the largest and most efficient fishing
structures in Canada cost about 100 000 dollars to install and the
annual maintenance costs are between 2000 and 5000 dollars.
The fisheries in the lagoons of Venice and Comacchio should not
be left out of this account of stationary eel traps (Fig. I I r ). The net-
work of traps fonning these fisheries are built de novo in the same place
every year by securing bundles of reeds to rows of posts. I t is necessary
to dismantle the network every year so as not to impede the annual
spring migration of glass eels from the sea to the separate 'valli' - i.e.
the various sections of the lagoon. which are separated from one another
by dams. In each 'valle' there are one or more of the eel trap networks
shown in Fig. I I I. A single network is often not sufficient since as
many as 100 toones of eels may enter the traps in a single night. On
average a 'valle' is about 2000 hectares in extent (the maximum is
296 THE EEL
Fig. III Diagramofoneofthe fishing engines in the lagoon at Cornacchlo. a, entrance

to the eel trap; b, outer court; c, connecting channel; d, entrance to the first
chamber; e, first chamber; f, trap; g, entrance into the second chamber;
h, trapping chamber for grey mullet and flat fish, with fencing through
which eels can pass; i, entrance to eel trapping chamber; k, eel trapping
chamber ; 1, holding baskets; m, shipping lock ; n, canal to the sea; 0, sluice;
p and r, two different 'valli' (ponds separated by dams where the fish are
farmed); q, dike; s, canal for lagoon ship traffic (after Samartine, from
Liibbert, 19(8).
Fig. I12 Diagram of an eel trap with a covered container for collecting the eels. The
dotted lines on the right indicate duckboards laid in the direction of the
current. The collecting chamber (on the left, below) also consists largely of
duckboards. Distance between the slats: 20 mm.
17000 ha) (LUbbert, 1908). The annual catch for a 'valle' is between
30 and 40 kgjha; the total area of productivity amoWlts to 50 000 ha.
In the whole lagoon area of the northem Adriatic about 1000 tOlUles of
FISHING METHODS 297
eels are caught annually (D'Ancona, 1961). There is little point in
making a direct comparison between eel weirs and these 'valli'; in the
canals of the fagoons, for example, the current is much less, so that the
weir fences need not be nearly so stable as they are in the rivers. There
are also some modem structures, as for example those in the northern
area of the lagoons near Venice. Eales (1968) has described fashine-like
weir fences in the tidal areas of the coasts of Canada. These are not
installed in the rivers but in the tidal stream, in order to catch the eels
which are moving backwards and forwards. Maximum catches of
15 tonnes are possible.
5.3.2 Eel traps in streams and rivers

Eel traps were often installed where weirs were already in existence,
in particular near mill ponds. A sloping grill was placed where the water
flowed over the weir into the stream below, thus acting like a sieve to
trap the eels (Fig. 112). Many such weirs have been constructed from
wood or masonry specifically for the purpose of catching eels. The
Swedish 'eel box', a type of eel trap without a weir, is a simplified form
which can be placed in smaller streams. They are very common in the
area around Kassel (Buhse, 1967). The lid can usually be locked so as
to prevent theft. Hofmann (1960) has described a modem version of
this type of trap which can also be used in larger rivers near turbines.
'" •
'" '"
'"
, .. ,
"
.L,~9-~TT'-rf,
618910111212345 6 7 a 9 10 11 12 1 2 :5 4 5
Fig. 113 Monthly catches from an eel trap on South Island, Nt:w Zt:aland. Ld't:
A . dieffenbacht.'. Right : A. awtralis. A, yellow eels; B, malt: silver t:els;
C, female silver eels (after Burnet, 1969a).
Table 33 Monthly catches from individual eel traps in the Weser region and the lake regio n of the Baltic coas t of Mecklenburg (after ...
Walter. 19 10) and in the rive rWadang. Masuren (after Seliga. 1926). ( Figures in italics : highest and second highest annual values) '"00
Year February M arch April May June July A ugusr Sepu .... 0<,_ N~'" Toral
Weser region (in kg)

Around 1900 , , ., ,.
kg)
Mecklenburg lake region (in "
188 3 ,,0 " 940 "
75 ' ,6, ,006
1884 '05 :IllS
'" )5' ItII. 370 ,08 ,69 53 19
1885 ,80 rrlO '09 ,60 8,0 '77 47'
1886 1210 "0 IZIS '40 rZ7°
'41'
516 5
1887 ,,. ,6, ,60 ,88 "07
1888
,.,.,. ...",
,6>0 139 1 r8I7 .,60
'" ,69 ,88 8313
1889 ,0 ,60 6" 1167 1147 .06 ,<0 66"
" 90 ,0, 17JJ 69' 97' 69' '97 ,68 '" 5457
189 1 ,0, 75° ,., 1IJ] 148 1 1034 ,., '0 60)5 ...
1892 ,0, 10 49 '34 90 ' ,00 4954
1893 .,6 '",0, ,
. ,6, ,0' 1)12 ....
1894 105 62 ", 1407
'" 604 ,,, '"
,6, 90 57 80
'"''""
1895 10566 '44 11 050 '00' '0 ...'"
,.,. '4'
".". 6,0 8" 6" ,00
'''' 1820 '"
'07 ", '99'
52 16
1897 050 ,8, 74' 875 ,., 49 380 1
Annual mean '"
over 1 5years >74 1119 ,., '66 88, 68, '77 69 ,.06
River Wadang (no. of eels)
..,. 66 lZ60 ,,, 79 ,0
,,0 .6, , '70'
1787
,,,,
"97 47' '"
.,' ", ,."
1898 ,0> ' 90 '"" ,.. m , '34'
"99 05 " ' 70 ,8} ,,0
'" "0 1894
,,.. ,., ."
1'6 ,00 ,,66
'90' , , '"u, ,,8 ""0, '"'49
'91
."" .,60
'34
'" '" >0. '" 1140
}"
'90' .,
"., ••• ",
,<0 ' 09
, 68 ,6 " ".6, 1155
'90' '94 33'
Annual mean '" "
over 8 yeats , 74 '97 ," '47 ..0 "44
", '" ." "
FISHING METHODS 299
Eel traps, especially if they are placed at the outlets of lakes, can
collect huge nwnbers of eels. I t is not uncommon for 500 kg to be
caught in one night (Hofmann, 1960). In one trap on the river Fulda an
average of 2500 kg was trapped annually; however, in a trap nearby,
on the river Werra, only 100 kg were recorded, Figures as low as those
from the Werra were recorded in 1900 for a trap in the Weser area
(Table 33). Klemke (from Buhse, 1967) estimates the average annual
catch for the year 1935 to have been 500 kg for a stationary eel trap in
Kurhessen. In an outlet from a lake near Mecklenburg the annual
average weight of eels caught for the IS years preceding 1900 was
5000 kg (Table 33), A trap installed in a tributary of the Duna (White
Russia) proved to be far superior to the stow net system that had been
used there earlier. The annual catch amounted to 6000 kg (Dubovski;,
1963).
Eel traps can thus yield between 100 and 6000 kg a year and in some
cases even greater amounts [according to Mann & Klust (1952) 3000-
4000 kg of eels were caught each night in an eel trap in a sluice during
the period of the silver eel migration in Denmark's Ringkobing Fiord].
These discrepancies are probably due to the difference in size of the
various water bodies above the traps. Streams which flow out of lakes
and which are thus connected to an area of productivity often many
hundreds of hectares in extent, are clearly superior to others with the
same water flow but with a drainage area of only a few hectares. For
this reason eel traps in lakeland areas of Central Europe are like gold
mines and their catch records are a well guarded trade secret. The
owners of such eel traps are often looked upon with envy. A certain
amount of jealousy is inevitable when newcomers place their traps in
areas used by oth~rs. In such cases only joint stocking of a given area
of water with elvers is a fair solution.
I t is absurd to forbid the construction of eel traps by law (Hofmann,
1960). From an economic point of view eel traps and weirs in fairly
productive areas represent the most efficient method of fishing.
As is the case with fyke nets, stow nets and weirs, the periods when
the fish are caught vary with the type and the region of the river concern-
ed. While substantial catches are recorded in autumn in the area around
the river Weser, the catches in Mecklenburg and Masuren remain high
throughout the six summer months (Table 33). It does not seem that
one can explain these high figures by suggesting that the eels in Mecklen-
burg migrate earlier (e.g. Walter, 1910) - rather it seems that they
are due to yellow eels being caught as well. Similar findings have been
reported from the stow net fisheries on the Elbe (Fig. 90). Catches in
300 THE EEL
New Zealand also included considerable nwnbers of yellow eels of the

two species which occur there (Fig. 113). In particular the long-finned
yellow eel (A. dieffenbachi) was recorde.d over the whole year, with
increases in numbers in early spring and in sununer. Silver eels, on the
other hand, showed only a single marked increase - from summer until
early autumn (see pp. 217 & 220 et seq.).
5.4 Seines and trawls

The most widely known type of fishing net is the trawl. However, in eel
fishing it has, until recently, only been used to a limited extent. After
unsuccessful efforts to find fishing gear that was consistently efficient,
the trawl was introduced in many areas for catching the financially
rewarding eel. In contrast, the seine began to recede more and more
into the background, a tendency which was reflected not only specifically
in eel fishing but generally in all coastal and inland fisheries. The main
reason for this is bound to be the large labour force needed to operate
seine nets.
All the fishing methods discussed so far have been of a passive nature.
Trawls and seine nets, however, involve active movement - i.e. they
are not kept in one particular place, but are actively drawn through the
water. The fish are gathered in and trapped after they have, if necessary,
been aroused or startled into action. For this purpose the netting bag
•
•
,
,,
\, , ,
, ....
F\ .... F
"" .'
" .'
'·'· ... ..r·
\ /
\ls
"
Fig. 114 Seine fishing. A, anchor; b, buoy; B, boat; L, warp; F, wing; S, bag (after
Walter, 19IO).
FISHING METHODS 30 [
is constructed with wings, the ends of which have tow ropes or hawsers
attached to them. In eel fishing the wings and the bag are weighted
down on the underside so that they drag against the bottom thus
trapping the fish there. Seine net fishing is only dealt with very briefly
here.
A type of drag net, a modified seine net, has been specially designed
for eel fishing; it originates in Denmark and is used mainly in Schleswig-
Holstein (Fig. 114). The bag is 12 m in height and length, and the wings
are 52 m long. They are pulled into the boat by hand. In general,
few eels are caught in the ordinary nets that are used for inland fishing,
probably because the size of the mesh is too great. The net pictured
here is said to have trapped 15 tonnes of eels in one summer (Walter,
[910).
The operation of a seine always requires heavy manual labour - that
is, with the exception of the few large seines in use which are operated
by mechanical power. I t is thus obvious why eel seines are gradually
being replaced by fyke nets , Nowadays seine fishing on the Baltic
coast of Schleswig-Holstein is only of secondary importance (Neuhaus,
[9(9).
'Puis' fishing, a technique based on slightly different principles from
that of seining has, on the other hand, increased in this region. The
fishes are not actively surrounded by the netting wall, but are driven
towards a wall of netting by the use of noise - i.e. by hitting the water.
They are not trapped in a bag but are entangled in a trammel net. The
latter is a wall of netting made up of three layers; the mesh of the two
outer walls is between 100 and 120 mm in width and the mesh of the
inner wall is IS mm wide. When the fish pushes against this wall, the
inner wall bulges out towards the other side and the fish is trapped in the
little pocket thus created. The netting wall consists of 4-6 trammels,
each 30 m long, which are used to cordon off a corresponding distance
along the bank. Each fishing session lasts 25-30 minutes and produces
5-10 large eels (each weighing more than 500 g). With one boat and
two fishermen the daily catch may reach 125 kg (Freytag & Mohr, 1973).
It was generally feared that this fishing technique might have adverse
effects on other methods, but research by Mohr (1974) has failed to
substantiate this fear.
Trawl nets have long been used for eel fishing; the small-meshed
net used is called an eel bottom trawl. Before the introduction of motor
vessels the trawl was drawn by two sailing trawlers or by a single boat
sailing before the wind. Nowadays this type of trawl is either operated
by two motor boats and is called a pareja (twin trawl) (Fig. 115), or it is
302 THE EEL
equipped with otterboards which keep the wings apart and is known
as an otter trawl. The use of eel trawls from sailing vessels is common
on the 'Bodden' and 'Haft's' on the Baltic coast of Germany. Otter
trawls used from power boats are fished mainly in the central Baltic,
e.g. on the Oderbank and, since 1964, also in the North Sea south-east
of Heligoland. The gear shown in Fig. 115 can be trawled with two
boats on larger inland lakes, or it can be used at sea with two otterboards,
and fished from a trawler with a motor capable of developing at least
150 horse power; on inland waters a speed of I' 5 knots is sufficient
(Steinberg, 1964). A further type of trawl net which often includes
eels in its catch, is the beam trawl used in North Sea coastal fisheries
as a shrimp trawl. A recent technique is to use trawl nets in combination
with an electric current (see p. 320).
Trawling has only been permitted in the open sea beyond the three-
mile limits. In coastal and inland waters there are many laws against
trawling. In some cases even seining has been restricted. The main
reason why trawling has been prohibited in inland waters is to protect
other methods of eel fishing, such as fyke nets and eel long-lines. For
example, eel long-lines could go unnoticed and be dragged along with
the trawl, fyke nets could be damaged, or the annual catch could be
reduced because two fisheries using different fishing methods would
have to share the yield. In addition, fishing itself could become un-
economical through over-fishing - i,e, the catches may only consist
of small eels which have not had time to grow to a reasonable size
(Deelder, 19<)5). A further objection to trawling is the possibility of it
causing biological damage. This might include not only the destruction
of the vulnerable young of other species if they are caught in the
trawls, but also the uprooting of water plants used by the fishes as
spawning sites. The whole history of trawling in coastal waters is
littered with incidents of protest and the threat and implementation
of legal restrictions, in which the possible destruction of plants and
animals has been used as the main argument against trawling. Neverthe-
less, the desire to protect passive fishing methods such as fyke nets
and long-lines has played a subtle part in the framing of such restrictions.
Trawling is obviously unnecessary and undesirable in areas where it
could have an adverse effect on another, profitable means of fishing,
particularly if, as a result of over-fishing, the optimal fishing intensity
is exceeded. Nowadays such a situation is quite commonplace in most of
the larger inland waters of Europe and also in a large number of coastal
areas. The introduction of traps which require less manpower for their
operation is becoming more frequent in some areas and will probably
FISHING METHODS 303
Bri dle, 10m

,
l1Zmm hemfl) SprtQd~, 0·6m
I ForWQrd ~>ght
7kg ~l>ain
I
Fig. liS Diagram of a pareja (twin trawl), which is used in inland lakes and is pulled
by two motor boats (about 10 m long) with 15-20 horse-power engines
(after Steinberg, 1964).
be even more widespread in the future, but, in off-shore regions,

trawling is the only practicable fishing technique currently available.
The increasing size of the catches brought to the North Sea fishing
port of Btisum shows how attractive trawling for eels has become:
in 1964, 10·6 tonnes of eels were caught; in 1965,29 tonnes; in 1966,
45 tonnes; in 1967,78 tonnes; in 1968,64 tonnes (Neuhaus, 1969).
The catch fluctuated, from 1970 to 1975, between 58 and 98 tonnes; the
average at Btisum was 79 tonnes, and for the whole trawl fishery in the
North Sea it was 92 tonnes (Aker & Koops, 1976). According to the
results of marking experiments (Aker & Koops, 1970) about 40% of the
eel popUlation is caught annually. Fishing in this region can thus be
re'tarded as very intensive.
According to Neuhaus (1969) even the shrimp trawlers from
Schleswig-Holstein are reported to be increasingly interested in
catching eels. From time to time as much as 10 kg of eels per day were
caught in shrimp beam trawls operated in the sandbank regions. Eel
trawls (i.e. trawls specifically designed for eels) in the coastal fishing
grounds recorded catches of 25-75 kg per day in June 1968 - i.e. at the
beginning of the fishing season (Neuhaus, 1969). At the height of the
season the daily catch often reached 100 kg and in some cases was as
much as 150 kg per day. The annual catch records of several vessels
stood at 5 tonnes,and a few even reached the 10 tonne mark (according to
statistics from the marine fish market at Cuxhaven). The largest
catches were recorded in September and October. I t should be empha-
sized that, despite the fact that the fishing season occurs so late in
autumn, catches do not consist mainly of silver eels. This is probably
due to the cycle of wanning and cooling of the North Sea taking place
later in the year.
In 1969 eel trawlers from the traditional fishing grounds off the
304 THE EEL
Oderbank, Central Baltic, landed 250-350 kg per trip, and in one case
1000 kg was reported (Hoffmeister, 1970). These figures are probably
very similar to those recorded in the south-eastern regions of the
North Sea. In the western part of the Baltic, near the Bay of Lubeck,
experiments in 1969 using eel trawls were less successful. According
to Norwegian studies more narrow-headed than broad-headed eels
were caught in trawl nets; the reverse was found in fyke nets (Sivertsen,
193 8).
Eel trawls, which were first introduced in the lakes of North Germany
in 1960, had good results initially but then began to show a decline in
yield. The introduction of the pareja or twin trawl (see Fig. I IS), raised
the size of the catch again. The number of eels caught increased by
100- I 50% and in addition large numbers of fishes of other species
were caught. Eel fishing was most successful at night. Trawling for
105 minutes produced a maximum of 335 eels. Trawlers fishing in the
North Sea seldom catch this number of eels. In combination with a
narrow meshed outer cover to the codend, the eel trawl can also be
used to catch smelt which, in tum, are used as bait in eel long-line
fishing (Steinberg, 1964). A combination of trawls with electrofishing
gear has produced further improvement and increases in the size of
the catches (see p. 320).
5.5 Angling and line fishing

The term 'angling' conjures up pictures of hooks or similar objects
which are baited and left for the eel to bite. Fishing rods are used not
only in fishing for sport but also by professional fishermen. The diffe-
rence between the two is that the professional does not watch over his
rod nor does he only have one rod in use at a time. Each rod must,
as it were, fish on its own, and is set up as part of a whole battery of rods.
Although rods are the least expensive type of equipment and can be
installed with very little financial risk, they may require a relatively
great amount of man-power to operate them and are not suitable for
catching large numbers of fish; however, since the eel realizes a high
price in the markets of Central Europe, it is particularly suited for
this method of fishing.
Various types of hook are used in sport and professional fishing.
The hooks commonly used in sport fishing are illustrated in Fig. 116,
A and C. Because of its characteristic shape, the bait remains secure on
the Pescaro hook. The solid structure of this hook also prevents it from
being bent (Loebell, 1965). The two hooks illustrated in Fig. n6,B are
FISHING METHODS
those most commonly used in eel long-line fishing; in inland waters

the so-called Kirby hook is also common. This has a reduced point which
is most popular in the sizes 2/0, I /O, I and 2. On the coast, smaller hooks
(2, 3, 4 and 5), are more popular, e.g. circular hooks with a long shank,
which are particularly suitable for fishing with bait (v. Brandt, 1959).
Anglers like to use relatively large hooks (no. I /O) so as to avoid catching
very small eels (Bawngartner, 1968). Oster (1961) has the following
to say on this matter: 'When the foraging eel comes across the bait it
cannot contain itself and rushes at the poor little worm, gulps it down
and subsequently chokes. Experience has shown that the size of the hook
hidden in the bait does not play an important role. The main point is
that the hungry animal should get the hook stuck in its mouth. ' It thus
appears that the size of the hook should, to some extent, correspond
to the size of the eel. Since darkly coloured hooks are less noticeable
they are used in preference to lighter coloured ones.
A 'snood' or 'leader' is fastened to the hook, though this device is not
absolutely necessary for eel fishing (see Jugi, 1970). Any type of line
is suitable, but a leader 30 em long is useful in hand angling (Loebell,
1965). With eel long-lines the snood should not be too short: according
to v. Brandt (1959) the best length is 1-1'5 m; LUbben (1968), however,
suggests 60-70 em. To prevent the eel swallowing the snood, the latter
must never be longer than half the distance between the points where the
snood is fastened to the main line.
More or less any type of rod can be used in angling as long as it is
easily manageable. Only when the line is being cast far out is it neces-
sary to have a reel. 'If a reel is necessary then it is important that it
should wind smoothly and that it should have a relatively high gear
• • c
Fig. 116 Different types of eel hook. A, hook without barb; B, hooks with flat, and with
lateral eyelet; C, Pescaro hook, very strong, with sinuous point (Loebell,
Ig6S)·
THE EEL
ratio so that the fish can be brought in quickly once it is hooked. The
most suitable types are the Nottingham reel, which has a cylinder with
a large diameter, or the multireel, which is strongly built and has an
appropriate gear ratio. Many eel anglers refuse to use stationary reels,
or do so unwillingly, because this type of reel prevents a feeling of direct
contact with the fish on the end of the line .... ' (Loebell, 1965). Man-
made monofil fibres are recommended for use as angling lines. Fifty or
60 Nm lines are good as basic equipment because they can be used to
land heavy eels which are able to wrap themselves, with considerable
strength, around water plants, stones and other objects.
Sinkers with a hole bored through them should be used to keep the
hook and line on the bottom or in the path of the current. 'The lead
weight should be placed so that the eel does not drag it along when it bites
at the bait, as this will make the animal suspicious. The sole purpose of
the lead is to keep the bait on the bottom should the current become too
strong. I usually use a lead shot as it is less likely to get hooked up
among the stones on the ground' (Pape, 1966). In calm waters, any
sort of lightweight angling lead is sufficient, but in the sea and strong
currents pyramidal or hexagonal sinkers weighing up to 250 g are the
most suitable because of their relatively large surface of contact with the
bottom (see Loebell, 1965).
If there is a thick layer of mud on the bottom, the lead, leader, hook
and bait may sink into it. In order to prevent this from happening. an
auxiliary line is mounted on the main line with a simple peg. The main
line runs through the loop fonned by the peg. and the lead weight is
fastened to the end of the auxiliary line. A small piece of cork is attached
to the leader and the bait then floats a hand's breadth from the muddy
bottom (Castan, 1967).
Pape (1966) suggests that: 'There is another peg with a spring clip
just in front of the leadweight on the main line. It is meant to counteract
the twisting, writhing movement of the eel and to stop the line from
becoming tangled. Besides, with a spring clip, it is possible to change
the leader quickly and easily in the dark.' Loebell (1965). however,
does not recommend the use of an eyed leader because, although the
leader can be replaced quickly, it has the disadvantage of the loop
being a weak point with a high risk of breaking, and he recommends
tying a slip knot where the leader is attached to the peg.
A float is nO[ always necessary when eel fishing, e.g. in a strong
current, or when fishing from the shore or from a boat. Whenever it is
difficult to keep the line taut - e.g. when casting wide, or when there
are beds of weeds, or unusual currents, or when the water is very
FISHING METHODS
shallow - a sliding float can be tied to the line (Loebell, 1965; Pape,
1<)66).
Bawngarmer (1968) gives this description of angling with a sliding
float at night in the Lake Chiem region: 'Small eels always swallow the
bait straight away, usually without causing the rod to move visibly.
They then become aware of the hook, press themselves against the line
with their tails and, with the first tug, anchor themselves firmly to the
ground. The fisherman's job now begins; he has to ease the eel gradually
away from the ground without snapping the line and to draw the eel in
without it re-anchoring itself. The marauding silver eel (a large eel),
in contrast to his smaller brothers, will make off with his booty. In this
case a hefty tug is in order. The escaping thief, once he has taken a
a firm hold, will then only succeed in planting the hook more firmly
into the front part of its mouth. So far, I have never had to cut free a
heavy eel. The predatory eel can put up an amazing fight: it does not
struggle wildly but tries to shake itself free on the spot. If that has no
effect, the eel will reveal an astounding ability to anchor itself to large,
heavy objects on the ground. The eel grips branches, planks and reeds
with incredible strength and it becomes impossible to free the animal by
physical force alone. One is obliged either to continue pulling both the
eel and the object to which it is anchored, or to wait until the creature
relaxes its hold - which may take hours. A gentle tug every now and then
will establish how far the eel has got with its manoeuvres.'
Czechoslovakian anglers have devised a means of demonstrating
the yield per month of various species of fish, including the eel. If one
rates the number of eels caught during July at 100, then the expected
yields for January to December will be as follows: 0-0-0-0-0-48- 100-
70-18-5-0-0. Eels caught during April and May are not included as
this is the closed season. Catches of any appreciable size are only to be
expected during the warmer months of the year (Fisch u. Fang, 5, 316,
1964). In areas with a dense population of eels, e.g. in the harbour at
Hamburg, the daily catch may amount to 8-10 eels under favourable
conditions. There is one report of 46 eels being caught in a period of
15 hours (Fisch u. Fang, 7, 268,1966).
The drifting eel jug line reflects a step towards a more commercial
approach. It is made out of reeds or sometimes from a bundle of simsen;
other buoyant objects - e.g. corks or tin cans - are also used. A line
with a hook at one end is wound round the object. The length of this
line depends on the depth of the water - i.e. its length should be twice
the depth of the water (v. Brandt, 1957). The eel jug line is allowed to
drift freely in the water. A light wind is ideal because it blows the eel
308 THE EEL
jug around slowly, and since it changes its position so often, it has
more chance of catching eels than does a stationary eel line (see below).
However, this sort of device is only suitable for smaller expanses of
water or bays where the eel long-line is blown by the wind. Richard
(1938) reports that when the eel jug is cast onto calmer areas of water,
it should not be unrolled fully because the bait must not be allowed to
rest on the bottom and thus prevent the eel jug from drifting about.
v. Brandt (1957) says that the end of the line should be lightly fastened
about half a metre (in greater depths probably more) above the hook
so that the line unrolls as soon as the eel bites. It is obvious from the
description given above of the way the eel will approach the bait, bite
at it and then try to escape, that this type of unattended device is
effective even without human intervention. The fixed handline is simi-
lar in principle to the drifting eel jug (v. Brandt & Rutkowski, 1956).
It is hung from the bank or is suspended under water through a hole
in the ice. In other words, it is not mobile like the eel jug.
Eel long-lines (mentioned above) are also left unanchored. The
operation of this sort of gear demands both skill and experience (Buch-
holz, 1948). It is quite common for the crew of a boat (at least two men)
to set up and bait over 1000 hooks. It is said that practised fishermen
can bait and set as many as 2000 hooks in two hours and do this in such
a way that, when then the line is taken up, many of the bait fishes are still
alive on the hook (peters, 1935). This technique for catching eels must
therefore be regarded as an established method in professional fishing.
In practice, the snoods are fastened to the main line at intervals of
2-4 m. The best material for the main line is perJon (Rankovic, 1957);
'PerJon endlos') or better still, plaited perlon, is a good, staple fibre.
The thinner the line, the more successful is the catch. Eel long-lines
in lakes have between 150 and 300 hooks attached to them; in rivers
considerably fewer hooks are used (30-50; Jager, 1960). Lubben (1968)
has even suggested that 2 or 3 hooks are enough and that the line should
be 5 m long. With three such lines this author caught just as many eels
as he did when using a line with 100 hooks. 'After all, one can only catch
as many non-migratory eels as frequent a given area ... '. 'The larger
predatory eel is a very territorial animal and when one fish is caught,
the area thus vacated is not immediately taken over by another fish.
I have always had the most success with eel long-lines on the first night
spent fishing in a given area. On the second and third nights the catches
have gradually decreased.' When using eel long-lines, it is thus
advantageous to change the fishing areas frequently.
The lines must be weighted with sinkers if they are to stay on the
FISHING METHODS 309
bottom. It is particularly necessary to do this during the day, when the
eel does not move from the deeper waters. Eel long-lines are usually
only set towards evening so as to catch the nocturnally active eel and
are lifted in the morning. Such lines may even be equipped with floats,
for at night the eel rises to the upper levels of the water. Floats are
sometimes also necessary when the lines must be prevented from
sinking into soft mud.
Bait fishes must be attached to the hook in such a way that the tip of
the hook only just protrudes.
Long-lines will last longer if they are put into an eel hook holder
or clamp when washed or cleaned. In this way a good catch is ensured.
In some cases a potassium permanganate bath is used for cleaning the
equipment; a pinch of this chemical is added to a bucket of water
(v. Brandt, 1957; Jager, 1960). Special care is needed when putting on
the bait as well as when handling the long-lines. If the smell of oil or
petrol were to be transferred from the boat's motor, it might well fright-
en off the eels which are very sensitive to smell (see p. 75).
In addition to the type of bait used (see below), it is also important
that the line is laid properly. v. Brandt (1957) describes the process as
follows: 'A stone is attached to one end of the line at the beginning
(the stone should also be from the same lake so that it does not differ
in smell). If possible, the line should be dampened (so that it will sink
better) and then cast out loosely in a bow. This is done from the boat
as it is being rowed slowly forwards. The slacker the line the better.
If the line is baited in the boat, it is often better to leave one hook un-
baited than to hold the line in order to put the bait on the hook and
thereby pull the whole line tight. The slacker the line, the less likely is
the eel to notice the resistance and thus swallow the hook. Most fisher-
men prefer to do the baiting in the boat. Only when it is very windy are
the lines baited beforehand on land. The baited hooks are packed in
sand in a flat box or on a board, so that the line will run out smoothly
when laying the snare.'
Another form of line fishing is that in which the line, about 2-3 rrun
thick, is set up over a river (Jager, 1960). The snoods and hooks, which
are attached in the same way as in long-line fishing are checked in the
morning and the evening, the fish that have been caught are removed,
and the hooks are rebaited (see above).
Long-line fishing is wide-spread and is common in areas such as
southern Europe (Rankovic, 1957) and North America (Eales, 1968).
The best catches are recorded in temperate conditions, when the water
has reached the normal summer temperature - i.e. in May and June
310 THE EEL
Table 34 Monthly distribution of the total annual eel catch

in Lake Skutari in Yugoslavia from 1938 to 1955.
Almost all the eels were caught by long lines
(after Rankovic, 1957)
January )"32% July ' ·47%

February 7·99% August '"37%
March '4·77% September 1·18%
April 27·20% October 5'48%
M,y 20·7°% November 8.40%
June 2·38 % December 5·74%
Table 35 Baits recommended for use in different types of angling

Type of bair Type of angling Locality References
. Lugworms (Armico/a), mussels, shrimps

(Crt:l1lgon), sprats, sand ~l (mostly
narrow-headed ~Is using worms
and mussels, broad-headed eels using
sprats, smaller ~ls using shrimps) Long line Coastal waters v. Brandt, '959
Pieces of herring Mackertlline Baltic, Lubeck Bay
at a depth of t5 m Fisch u. Fang,
(pelagic) 8,6,1967
Polychaetes (Nereis sp.) in November,
more effective than lugworms Fisch u. Fang,
( Areniccla) Hand line Flensburger Forde S,30, 1967
Sticklebacks, during their spawning Fisch u. Fang,
migration in spring Long line Hun te, tidal area to, 165, t969
Earthworms, small herrings, cheese Brackish water, Fisch u. Fang,

tinned meat, pieces of sausage Hand line Bremerhaven 8,298,1967
Bleak, gudgeon, roach, weather fish,
pieces of larger fishes or crayfishes
after the skin has been removed;
the following can be used as substitutes:
frogs, shelled land and water snails,
horse I~ches; if nothing else is
available: pieces of meat, worms for
smaller eels, fishes for larger ecls and Drifting eel
broad-headed individuals jug line Inland waten Walter, 1910
Small fishes, e.g. small perch, available
in large numbers; \eeches, frogs,
earthworms Long line Inland waters Seligo, 1926
Bleak, roach, sticklebacks, roffe, smelt,
etc. or earthworms Long line Inland waters Jager, 1960
Strongly smelling cheese and sausage Fisch u. Fang,
containing lar~ amountS of garlic Hand line Inland waters 7,360,1966
Pieces of white fish (cyprinids) cut in Fisch u. Fang,
tapered strips Hand line Inland waters 7,367,1966
Preferred fishes: perch, ruffe, roach,
white bream, rudd, bleak, stickleback,
gudgeon, smelt (sexually mature
fishes are panicularly effective as
bait); earthworms, cockchafer
larvae, crushed pondsnails Long line Lakes v. Brandt, 1957
FISHING METHODS 311
Type of /mit Type of angling Locality Referencts
Bleak and other white fish, gudgeon, if

necessary pondsnails, Tubifex, crabs,
eanhworms Long line Lok~ Buchholz, 1948
Pieces of ed, glass eels and young eels Lake near Fisch u. Fang,
less than 15 cm long Hand line Cuxhaven ' ,300, 1968
CoregO>lUS cut into small pieces Hand line Lake Pion Fisch u. Fang,
10, 109, 1969
Dead, dewmposmg fishes (surprisingly Fisch u. Fang,
enough) Hand line Gravel pit 9, 142, 1968
Smelt cut into pieces, used in Jan uary
during the spawning migration of Hand line Weser, cold water Fisch u. Fang,
this species (ground line) outflow 7,279, 1966
Cheese Hand line Rivers in the Nonh
Sea tidal area and Fisch u. Fang,
southern Gennany 8,356, 1967
Earthworms, mitten crabs and cray
fishes (cut up, and even boiled), for
predatory eels and broad-headed eels,
fishes as bait, cheese, fresh liver and
lung, dead frogs, lugworms Hand line Castan, 1967
Allis shad, smelt, herring, various
species of whitefish, mussels Long line Canadian waters Eales, 1968
(Eales, 1968). Eels are, however, still caught later in the year, up until
autumn. In Lake Skutari, on the Yugoslavian-Albanian border in
southern Europe, where the temperature seldom drops below the
minimum for eel fishing, or only does so for a short time, the best
catches are recorded in March, April and May (Table 34), and the
lowest catches during the warmest months (June, July and August).
In Canada the yield per fishennan per year from long-line fishing is
not quite 2000 kg. There, a long-line with 300 hooks costs between
25 and 30 dollars, and an additional 20 dollars a year is necessary for
maintenance. In Canada, as in Gennany, this method of fishing is used
both in inland waters and along the coast (Eales, 1968).
As has been mentioned already, long-line fishing is commercially
more rewarding than fishing with drifting eel jug lines, since a greater
number of hooks is used. However, as research in Lake Sakrow has
shown, the catch per hook is greater with eel jugs (Rahn, I 967b). Over
a five-year period, the mean catch per hook of a 100 hook long-line
was 0·7 kg, that from a 100 eel jug line was 2·9 kg.
Table 35 indicates that high yields are obtained with a wide range
of baits used in a variety of places; the earthwonn is mentioned most
frequently in this table. The success of this bait is, presumably, not
altogether unconnected with the fact that it is easy to come by and
3[2 THE EEL
"
",
''''"
;-,
"t· . . .,; \/
Worm bGi t / I \ f i.h bait
" .I / \
\
\
\
I! \'
I! \ \
" I ,I \ '.
, I "
I I '.\
'\~
I /
.,
.::"J
\
\
29
" 99
"
Sm<llIotI I. Si lver tel CQicilH
baskets ne<l r ", usil\lj HI baskns
" tht bQ.nk~
,
" : (16 - 16 mlll IIIUh)
'~ /
/ ,\
/,' \ ~
1 ,' ','., E,I bosket,
/ I , (\4111111 m" hl
'I ,' \'
"
in09tn water
/ \,' V::
I ,' \ ;
I ' \
"
"
I' \ "
J :' \ "
I : \ '.,
/
I
.- :
!
\ ""
..
,
.
'.. ------
/
\
,. \.
29
Total length (emI "
Fig. 117 Length-frequency distribution of eels caught in the Kurisches Half using
various fishing methods (modified from Gajgalas, 1969).
therefore is often used. I t appears that lugworms ( Arenicola) , ragworms

(Nereis) and other related species are equally successful. Almost all
kinds offish are good as bait and, in professional fishing, the smelt tops
the list. Mostly predatory eels, and wide-headed eels in particular, are
attracted by such bait. Figure 117 enables one to compare the length
of eels caught by various types of gear. It is dear from this figure that
in the Kurisches Haff most eels measuring between 54 and 59 em are
caught using fish as bait. However, with earthworms as bait, the
greater proportion of eels only measured 40-50 em. In comparison
with other gear these methods result in the smallest eels being caught.
Even fishes which have begun to putrify are effective as bait, although
one is generally reconunended to use fresh bait only. Crustacea, in
panicular shrimps ( Crangon ) and mitten crabs, also make good bait.
Snails and mussels can be used if their shells have been broken; frogs
and leeches are used as bait too. If small animals are unavailable,
foodsruffs containing albumen, such as pieces of meat, liver, lungs and
sausage can be used. In fact, small pieces of cheese are particularly
effective as bait.
There is little doubt that the eel is attracted to the bait by smell
(see pp. 75,276). As has already been reported, there are certain animal
amino acids which are only attractive in combination with one another.
If these amino acids are artificially synthesized they will also attract
eels but not in such large numbers. Possibly a substance could be
synthesized that will attract eels without first having to condition the
animals to respond to it. Of course, this bait would have to smell
strongly enough to override the effect of any other attractive odours in
the water.
So-called 'naring' should not be left out of a description of rod and
line fishing. This method requires a type of bait which has already
shown itself to be most successful in angling - i.e. the earthworm.
Using a coarse, blunt needle, medium to large earthworms are threaded
onto a strong woollen thread until a string of earthworms measuring
1-3 mhas been formed; at least 20 and up to 80 earthworms are needed.
This string of worms is then wound round the hand and the loops thus
tormed are tied together tightly with thread j the two ends of the
woollen thread are also fastened off. This produces a bundle of about
10 loops (see Fig. 118). One then needs a very strong rod between one
and two metres long j a broom handle will do. A line measuring abouI
3 m is tied to the rod (the line can be longer or shorter depending on the
depth of the water) and a lead weight (the 'oaring weight') of at least
Fig. 118 Hank of worms used for naring (after Loebell, 1965).
3 14 THE EEL
100 g is hung on the end of the line. The hank of worms (sometimes
fastened to a spring clip) is tied just below the lead weight. The bait is
now suspended from a boat or from land and allowed to hang a few
centimetres above the bottom. A float is not necessary; a gentle up and
down movement can, in fact, be advantageous. If an eel bites there is
usually a slight but continuous pull or, more often, the line jerks.
Almost immediately the line must be pulled steadily and not too quickly
out of the water. Only when the eel is above the water must the action
quicken. Since the eel has bitten into the woollen skein and is now
banging with its teeth caught in the woollen thread, it can easily fall off
and escape. In order to reduce this risk, a perforated box is often
floated near the line or is suspended from the edge of the boat. The eel
should be landed in this box as quickly as possible. ~ perforated umb-
rella can be used instead of a box. Naring is also practised in France;
the techniques involved differ only slightly from those used in Germany
(Schalensky, 1975).
Naring is most common on the coasts of the North Sea and the
Baltic. These areas have dense eel populations and, with a certain
amount of skill, it is possible to bring in large catches with this teclmique.
It is quite common for several eels to bite at one time. Grunert (1969)
reports that well over 100 eels have been caught with one naring device
in a morning spent on the Lower Weser. Reports from the drainage
area of the Lower Ems show that catches can vary a great deal from
season to season, and were good in May and October. In one evening
a single angler landed 6 kg of large eels suitable for smoking, as well as a
bucketful of smaller eels. Within a period of two weeks several other
anglers each caught some relatively large eels weighing a total of more
than 25 kg. The best catches were made when a west wind was blowing
and the tide was high (Fisch u. Fang, 10, II , 1969). Similar reports of
successful naring have been received from other sources; 30 or more
eels per hour is not uncommon, according to Walter (1910). However,
on the whole, naring is a method of fishing in which small eels make up
the greater part of the catch.
In the Wadden Sea naring is practised in the tidal channels and on the
river delta. The biggest catches (and the largest eels) are made at night,
but, during the day it is possible to achieve a sizeable catch if one is
willing to accept smaller eels. Loebell (1965) even suggests going further
upriver and naring in the trout regions. Unlike night set long- lines,
naring will only catch eels, and thus it is possible to reduce the eel
population without endangering the trout population. Naring is
recommended to fishermen who are opposed to electrofishing (see
FISHING METHODS
p. 318) but who want to regulate the species content of their waters in
a narural way.
5.6 Lift nets, spears and other equipment

Another eel trapping device which should perhaps have been mentioned
in the section on nets, is the lift net. Basically this is one of the most
primitive types of fishing equipment and was not in fact originally
used to catch eels.
A lift net is made from a square piece of netting varying in area
from almost a square metre to several square metres. The netting is
stretched over crossed frames or over a rectangular frame to form a
square, trough-shaped sheet. Lines from the four corners of the net
meet at the middle of the frame, and the whole contrivance, suspended
from this point by another line, is lowered onto the bottom. With large
lift nets a crane-like jib, complete with winch, can be used. After a few
seconds or minutes, or even much longer, the net is pulled up again,
bringing with it any eels which were just swinuning over the netting
or which happened to have lingered there. A lift net is in use near the
openings of the sluice at Brunsbiittel, where a certain degree of success
is ensured by the dense population of eels continually swirruning past.
Two traditional eel catching implements, the eel spear and the rake
(see Fig. 119, below right) are more widespread and even more primitive
than the lift net (Walter, 1910; Buckow, 1955; v. Brandt, 1959; Kienast,
1961; Wundsch, 1962; Eales, 1968). The eel is impaled on the spear or
jammed between the prongs of the rake (Fig. 119). Barbs on the prongs
prevent the eel from forcing its way out again. The objects in the top
row and on the bottom left of Fig. 119 are for sticking into soft ground.
If the river bed is too hard, the device on the right of the middle row
in Fig. 119 is used. The blunt prongs extend beyond the finer prongs
and thus protect them. The two spears in the middle row, on the left,
are made of flexible material and are used on soft ground. The two
strong, outer prongs move apart as the eel's body passes between them
and then move back once the eel is inside the ring - i.e. the eel cannot
slither out again. This device inflicts only a very small wound on the
eel because the centre prong is so fine.
Eel spearing is illegal in most areas of Germany, though, in some
cases, licences may be granted. The use of a rake to catch eels has been
strictly forbidden for a long time; the rake is a hook-like device which
succeeds in wounding a disproportionately large number of eels without
actually catching them (Fig. 119, below right).
"
~ .
Fig 119 Different types of eel spear (Altona Museum ; Photograph: Marschall).
FISHING METHODS
Fishing with eel spears is the oldest form of fishing still in use.
About 10000 years ago, in the middle stone age, men were using spears
carved from the leg bones of red deer, and very similar in shape to the
metal spears used today (Wundsch, 1962). The bone spear heads were
attached to the shaft with animal sinews or possibly even eel skin
was used.
Eel spears are used mainly in winter when the eels remain motionless
on or near the bottom. This is a time when fishermen would otherwise
probably have to be idle. Spears function well in water up to 4 m deep.
Whether practised legally or illegally, this method of fishing is more
widespread than is generally thought.
Along the Mecklenburg coast the daily catch per fisherman is
between 4 and I I kg, the average yield being about 7 kg; a great deal of
expertise is, however, involved. The best time for fishing is when the
weather is overcast. The majority of eels caught, 90% to be more precise,
are of category III variety - i.e. they are small. In addition, a large
proportion of the catch is undersized and thus not suitable for marketing,
which is indeed the main drawback to this method (Kienast, 1961). It is
reported that in Canada a skilled fisherman can, under favourable
conditions, catch 40-50 eels weighing between 200 and 1000 g in two
hours. Here, as in Europe, the use of eel spears is limited mainly to the
coastal regions (Eales, 1968).
In Japan, 'refuge' traps made out of bamboo poles are used to catch
eels (Fig. 120). This teclmique makes use of the eel's dislike of light
(negative phototaxis) and its tendency to take cover under objects
(thigmotaxis). Two or more hollow poles are tied together and are
suspended in the water on a cord. The eels then hide in these poles.
Anyone who has kept eels in an aquarium will have observed similar
Fig. no 'Refuge' trap made out of bamboo (after v. Brandt, 1964).

THE EEL
behaviour if small tubes are placed in the water. This is an easy way of
getting eels out of an aquarium and the same basic principle is used in
eel fishing. In some cases the tubes may be baited in order to make them
even more attractive to the eels. The process of landing the tubes must,
of course, be undertaken with great care so as to prevent the eels from
escaping. Sometimes divers are sent down to block up the tubes before
they are lifted.
'Shelter trap fishing ' in Europe is based on similar principles. Loose
bundles of twigs are bound together, weighted with stones, sunk in the
water and marked with a float. The eels use the shelter traps as hiding
places. Mter the bundle has been taken from the water it is put into a
large dip net so that the eels cannot escape. Walter (1910) describes a
similar way of catching eels in the brackish lakes and ponds of North
Friesland. When the ice and snow are lying thick on the ground and the
eels leave their winter quarters, holes are made in the ice and bundles
of straw are put into the water; the eels then creep into the straw when
they come up to the holes to gulp air.
5.7 The use of electricity

Since the Second World War electricity has become an important aid
to eel fishing. Reference is not being made here to the use of electric
gadgets which provide mechanical power or light to catch eels, but to
the direct effect on the fish of an electric current in the water.
An anode is placed some distance from the cathode in the water, and
a current of 220V passing between the two points strikes any fishes
which happen to be in the vicinity of the anode. Fishes further away from
the anode are less affected by the current and, after their initial shock
reaction, may well manage to escape. If the eel comes within the central
range of the direct current it will swim up to the anode in a state of
galvanotaxis. Shortly afterwards the eel rolls on its side in a state of
galvanonarcosis. The fisherman takes advantage of the eel's inunobility
and catches it with some conventional device. A dip net is usually
sufficient since, in shallow waters, the eels come to the surface when
under the influence of electrotaxis. The combination of electricity and
a dip net is generally the best method for catching yellow eels in streams,
small rivers, the inshore regions of lakes, and in shallow standing
water areas. The anode itself is often constructed like a dip net. In
most cases, however, one or more additional dip nets are needed to
cope with the great number of fishes that often suddenly appear
(Fig. 121 ) .
FISHING METHODS 3[9
A petrol driven electric generator is the most common source of
power, but batteries or a mains connection may also be used. In the
last "few years pulsed direct current has been used instead of direct
current. The former requires less energy and allows the use of lighter
units, an advantage when one is fishing in streams situated far from
roads.
In Germany, as elsewhere, it is necessary to obtain official permission
for fishing with electricity (Schiemeoz, 1962). In many areas proof is also
required that the person in charge is sufficiently qualified to use this
fishing technique. Safety regulations too must be satisfied (Meyer-
Waarden & Halsband, 1975). It is usually possible to find out about
electrofishing equipment and where to obtain it, from official sources
or, better still, from a fisheries institute. One is strongly advised not to
Fig. UI Electrofishing in a trout stream which had to be cleared of eels. The anode,
in the form of a dip net, is submerged in the stream. On the right, and behind
the man with the anode, there are two other men with dip nets catching the
stunned fishes. The other people in the photograph are waiting to take the
fishes in buckets to a larger container (Photograph: Tesch).
32 0 THE EEL
build such gear oneself; conunercial equipment is usually fitted with

the necessary safety devices and thus avoids difficulties with local safety
regulations and also reduces possible danger (0 life and limb.
In deeper waters a dip net is not suitable for catching the
inunobilized fishes and, until recently) electrofishing gear was not used
at depths greater than two metres. However, by combining electrodes
with a trawl oet it has become possible to fish at any depth. When the
current is switched on, an electric field is created between the two
electrodes. This electric field extends from the lower electrode for a
short distance into the ground with the result that eels hidden there
are also caught. Alternating current has been used by the author because
it is physiologically more effective, and because there is no need to drive
the fish towards a particular electrode. With alternating current the
fish is aligned, in galvanotaxis, diagonally to the lines of the electric field,
immediately becomes galvanonarcoticized, and is thus swept into the
forward moving trawl net. In addition to this combination of gear, which
seems ideal for inland waterways (Predel, 1968; Harrop & Predel,
1969), variations in the number and arrangement of electrodes, and in
the choice of current are possible. In the sea, where several such
combinations have been tried out (Meyer-Waarden & Halsband, 1975)
it is necessary, because of the high conductivity of the water, to use
an impulse current in order to save energy. Tests in the brackish water
of the Carmargue (in the Rhone Delta) were successful (Halsband,
1971; Halsband & Rask, 1974). To date electrotrawl nets are not
available for eel fishing at sea but experiments are in progress.
An electric current can be combined with trapping methods other
than dip nets and trawls. In Sweden, salmon swimming upstream were
driven into a trap by using two leaders acting as anodes and another as
the cathode; catches were considerably higher than with purely
mechanical leaders. Similar devices are used in North America; here
both salmon and lampreys are channelled into traps by the scaring
effect of an electric current (Meyer-Waarden & Halsband, 1975).
Similar traps can be erected to catch migrating eels in rivers (Dobler,
1964). However, it must not be forgotten that silver eels, the so-called
'drifting' eels, appear to have reduced physiological sensitivity (Meyer-
Waarden, 1955) and this, of course, can be a disadvantage when electri-
cal barrages are used to deflect and protect eels from turbines (see p. 360).
Using electrofishing methods it is possible, in a single operation,
to catch the greater part of the eel population in streams up to 10 m
wide. In lakes, of course, such a clean sweep is not possible. Table 36
shows the degree to which electrotrapping cuts into the population.
FISHING METHODS 321
Table 36 Eel catches (in kg) using different trapping devices in the roo ha Lake
Sakrow (Potsdam) before [he introduction of electrofishing in r950 (after
Rahn, 1957b)
1949 1950 195 1 1952 1951 1954 1955
Elecrrofishing
Seines
Fyke nets
'3
20"0
45"0
' ·0
341 "5 193" 5
' ·0
,0
161 "5
11 "0
112 "0
4·' ,.,
19"0
22"5 ' ·0 13 "5 4·0 43"S

Long lines and drifting
eel jug lines 28"8 22"S 16"0 12"0 t3"0 63"S 66·,
T otal 57"t 9 1"0 36 S"5 222"5 199"0 184"0 134"5
On introducing this method for the first time in Lake Sakrow (Table
36), many more eels were obtained than were caught by all the other
methods combined. However, the success of electrotrapping did show a
gradual decline in efficiency. Preswnably that part of the eel population
which could be caught by the use of electricity was reduced more
extensively than was the population in mid-lake, which was fished by
eel long-lines and drifting jug lines. A comparison of the average size
of individual eels caught by the various methods shows a difference
between those caught in electrofishing traps and those caught by other
means" The average weights in 1954 and 1955 were as follows: 793 g
from seines; 405 g from fyke nets; 380 g by long-lining; 478 g from
naring, and 245 g from electrofishing (Rahn, 1957b). Eels caught by
other types of gear were, in part, twice the weight of those caught using
electricity. The latter were dearly younger individuals.
A probable reason for this size and age range of eels being caught
by electrofishing lies in the fact that they are caught in regions near
the shore, i"e. in reed swamps and in shallow areas that extend out from
the bank. In general the smaller eels inhabit the shallower waters
(see p. 212). It seems that these eels are not part of the population
which would never otherwise have been caught, but rather that they
are individuals which would have been caught later with other gear.
The example of Lake Sakrow shows that the eel population is fished
more completely when electrofishing is used. In this case the nwnber
of fishing trips could have been reduced and consequently so would
have the cost of fishing per hectare of water surface. How far labour
requirements can be cut down depends, to a large extent, on how much
other fishing techniques can be reduced.
The effectiveness of combining trawl nets with electricity for fishing
in lakes is dearly illustrated by figures from the larger Brandenburg
lakes" For example, after introducing this method in Lake "Ocker, 40%
322 THE EEL
of the total eel catch came from electrotrawl nets (Hattop & Predel,
1969). In addition, an analysis of yields from 12 different lakes revealed
that, on average, about 30 kg of eels (and 20 kg of other fish ) are caught
in each electrotrawl per working hour. This type of fishing is, of course,
only worth-while if the area to he fished is sufficiently large. Before
introducing electrofishing it is important to establish whether other
fishing methods will continue to be economically viable or whether
they will have to be reduced. Experience with electrodip nets in small
streams has shown that most other fishing methods immediately become
uneconomic.
All in all it seems likely that electrofishing will be recorded in scientific
history as having had a revolutionary effect on the eel fisheries.
5.8 Combination with light

The eel's great sensitivity to light is a well established fact. In the
centuries-old silver eel fisheries of North Italy (see p. 295), it is reported
that fires were lit along the banks of canals in order to stop the migrating
silver eels from moving on (Petersen, 1906).
The first systematic studies on the degree to which eels are distracted
by light were carried out by Petersen (1906) in 1904 and 1905. Using
lights suspended over the water at the narrow outflow from a lagoon,
Petersen succeeded completely in stopping the movements of eels into
a fyke net placed behind the lights. Petersen was working on the assump-
tion that silver eels in lakes swim directly below the water surface
when migrating and therefore underwater lights would be unnecessary.
However, his experiments were carried out in water that was barely a
metre in depth, so nothing could be deduced about the effectiveness of
lights placed over deep water.
It is surprising that, until recently, relatively few experiments were
carried out to determine the practical use of light barriers as channelling
devices. Some good experiments were carried out by Lowe (1952)
and Frost (1961) between 1941 and 1944 on a river almost 10 m wide in
the Lake District of England. These experiments, however, only show
that one part of the river can be cordoned off with lights so that the eels
are caught in the other half. The effectiveness of the diversion was
measured by the relative number of eels that entered the dark and the
light sides of a trap. In all these experiments the number in the dark
side was higher and amounted to as much as 72 % of the total catch.
Later experiments in Holland have shown that illuminating the
water surface results in larger catches in eel stow nets. Here it was also
assumed that silver eels migrate mainly in the upper water levels, and
are forced by surface lights into stow nets placed at greater depths.
Observations on the Large Belt in Denmark using flood lights at night
have confirmed the fact that silver eels travel directly below the water
surface (Fischerbote, lI, 316-318, 1929). Although one cannot deny
this technique a certain degree of success, it is of little practical impor-
tance (v. Drirrunelen, 1951; from Brautigam, 1962).
Traps which divert the eels in a horizontal direction before catching
them, have been erected in Northern Ireland in combination with the
system of traps on the river Bann (described on p. 292). It appears
that the eel weir at Toome is so efficient that the addition of lights had
little effect. Only in the last 10 years have barriers of lights been used
experimentally in Germany. These trials showed that, with the help
of suitably placed lights, very large catches could be obtained. It is,
nevertheless, very difficult to prove that increased yields are specifically
the result of light barrages. Extensive experimental investigations are
needed, especially in coastal waters.
The result of an experiment in an inland waterway, the river Peene
near Anklam, are very convincing; when the light source was extin-
guished, the number of eels caught was only 5- 10% of the catch made
when the light was on. The trap was set up in the estuary of the Peene,
at a place where it was previously impossible to erect a system of fyke
nets because these would have impeded river traffic, the main argument
in favour of erecting a light weir rather than a netting weir.
In this relatively small eel trapping system 7 tonnes of silver eels
were caught within a period of three years. The light barrier was
equipped with thirteen 200W bulbs and was made of the simplest
materials; no special or costly devices were needed at all. The cost of
the total system - i.e. the light barrier and the traps - including cost of
upkeep, was 6000 DM.
Experience has shown that if the current exceeds 2' 5 m /sec silver
eels seem able to break through the light barrier. The light should
penetrate to a depth of at least one metre, the fields of light from each
bulb must overlap, and the chain of lights should not exceed 100 m if
it is to remain effective. Also, if the barrage is too long, the light has no
effect on the eels. Presumably the eel gradually becomes accustomed
to the increase in the amount of light and its movement is no longer
inhibited.
The type of trap used in conjunction with lights is a three funnel
fyke net with a non-return valve. The trap should be set up with the
codend deeper than the mouth and pointing in the opposite direction
324 THE EEL
.so S - -- j--------------------
I
/ . ... ···· d
I
10. I
•
so
40 •••.• - ~
"10
•
t~ ~ffi«S1jJMiSt».W.<m.wmW;((WJ.wN'""iiX<YJ)'&J%%i.ZV»""""v)S\)"KWz:y;;:;:::v;;::::w:~
~
" ... ". ,so ... ."

Fig. U2 Plan (above) and bottom profile (below) of a chain of lights (a), netting
barrage (b) at the exit of the Darss and Barth Bodden chain near BarhOft.
Measurements on the left and lower edge are in metres (after Brautigam,
I96Jb).
to that taken by the migrating eels, because, as far as the author can
make out, the flight reflex causes the eel to swim downwards and into
the current (Holke, 1964; Rennert, 1974).
A Polish research worker has made similar observations in a river
only 10 m wide (Swierzowski, 1964). This river was cordoned off with a
barrier of lights just before the point at which a small canal, with a
power station, left the main river. The efficiency of the light barrier
was tested by placing traps in both the river and the side canal. When
the light barrier was not functioning) the catches recorded in the side
canal amounted to 15% of the total. When the lights were switched
on, 58% of the total catch was trapped in the side canal (and 42% on the
river). However, if the current in the river was increased by opening a
sluice gate, the proportion of eels caught in the side canal dropped to
32% . Thus, in this example too, a large proportion of eels was carried
through the light barrier by the strength of the current. In addition,
the effectiveness of the barrier continued to decrease throughout the
night.
Conditions in coastal waters pose many more problems (Brautigam,
196Ia,b, 1962). The use of light barriers in suitable areas may appear
to bring about marked increases in the numbers of eels caught. However,
. 00
IS lQ 2S '0
f -----xpttlllber - - ----<
Fig. 1Z3 Catch records from pound net and light system shown in Fig. 122.
- - '=" catch from the traps with lights; ----- = catch from the double
pound net basket upstream; -'--'-' = catch from the downstream pound
net behind it (after Brautigam, 196rb).
the use of light barriers may be less effective if the areas involved are too
extensive, and there are only a few places where short light barriers
would suffice. Nevertheless, the vast numbers of eels which can be
caught along the coast during the silver eel migration act as a lure to
pioneers in the use of light barriers. In order to avoid disappointment,
certain basic principles must be followed: as far as possible, the fyke
or polUld nets should be fitted with netting leaders, and light barrages
should only be used in shipping lanes and areas where the water is too
deep for netting barrages.
Examples of favourable trapping sites for catching silver eels in the
Baltic on their seaward migration are the exit of the chain of banks
behind the Darss peninsula and the sea connection of the Rugen
Bodden (e.g. the large and small Jasmunder Bodden). Three experi-
mental traps (two of which are situated in these areas) have produced
more or less conclusive proof that banks of lights can improve silver
eel fishing very effectively.
Figure 122 shows the position of the barrage at the exit from the
Bodden chain behind the Darss peninsula. The string of lights is about
80 m long, and the trap is a pound net, similar to the one pictured on
p. 280. Additional pound nets with traditional leaders are placed
upstream and downstream of the combined trapping system. Catches
32 6 THE EEL
from the experimental trap and from the up- and downstream traps
are compared in Fig. 123. Results show that the pound net with the light
bana~ aLw.a.'".:!\. GlUJlJu. 1ll.Clt:./! than. thr. cJnnhlr.. ?QJlosi 01'1. ~JJJatrJi I ~
stream of it. Nonnally the latter would be expected to show a bigger
catch since it has twice the number of baskets. The downstream
trap caught even fewer eels. September 12th was the best day for
fishing: the eel basket with the light barrage yielded 1000 kg, the double
pound net in front of it caught 250 kg, and the trap behind it only
100 kg.
The second system of traps at the exit of the Riigen Bodden chain
has likewise proved successful. However, it was unfavourably situated
and its barrage of lights (250 m in length) was too long, so that the
results were less conclusive (Brautigam, I96Ia,b).
Another device also situated at the exit of the Rugen Bodden has
proved very effective (Brautigam, 1962). A wedge-shaped chain of
lights was erected with its apex facing into the oncoming stream of
migrating eels. The chain of lights was positioned at the deepest point
of the Bodden exit, and net barrages, each connected with a pound net,
were set right up to the bank on either side. That is to say, in contrast
to the system described above, the eels on their seaward migration
were guided away from the current in the middle and were led towards
the banks and thus into the pound nets. These were situated not far
from the banks, with their cod ends pointing towards the central current.
In other words, these traps were positioned like those at the entrance
to the river Peene (described earlier), the only difference being that
here both banks were brought into use. Those eels that managed to
break through this combined system were caught later in a set of
pound nets lying further downstream. This set of traps completely
blocked the exit of the Bodden with a netting barrage spanning the
entire width of the exit. With the lights switched on, only 12% of the
silver eels passed through the combined system and were caught in
the. control system situated downstream to it, whereas when the lights
were off, 65 % of the eels were caught in the control system.
The effect of a light chain on yellow eels was quite different. Over 70%
of yellow eels passed through the lights, regardless of whether the latter
were turned on or off. The effect of the chain on yellow and silver eels
increased slightly when flashing lights were used. However, this
increase was not significant in the case of yellow eels, thus indicating
that there is no point in using chains of lights for catching yellow eels.
The results of experiments with light chains on the river Peene near
Anklam point to a similar conclusion (HOlke, 1964; see p. 323).
FISHING METHODS 327
The 200W lights on the trap near Anklam were 10 m apart, and
floated, as a result of their own buoyancy, between 20 and 40 em above
the spot where they were anchored j they gave an illumination of
1.32 lux over a radius of up to five metres. The chain of lights should
not form an angle of more than 400 to the direction of the eel's migration.
Ninety-six percent of the silver eels were caught with the help of
the light barrage when it formed an angle of 400 j with the angle at 600 -
i.e. when the line of lights formed a less acute angle - the trap caught
only 80% of the eels passing that way. In this particular trap, stake
weirs were shown to be the most suitable net barrages. In certain types
of current, floating weirs are easily pushed under water, with the result
that the eels swim over them. Experience with stake weirs showed that
light barrages should not exceed ISO m in length, and the lights must
not be placed too near the surface of the water; they must also be in
an area free of water weeds, etc. The eel trap must be at least 50 m
from the chain of lights, or else the size of the catch is reduced. The
space between the lights and the traps must be bridged by a net barrage.
In an article on the productivity of light barrages in Bodden waters,
Brautigam says the results of the experiments just described indicate
that an increase of about 50% in the size of the catch is possible. Normal-
ly IS tonnes of eels are caught annually at the exit of the Bodden chain,
behind the Darss peninsula near Zingst. With a light barrage the yield
should increase to 23 tonnes. The additional expenditure on a chain of
lights, electric cables and an adjoining barrage, would be covered by
just one tonne of eels.
The yields for other species of fish will remain unaffected. Since
positively phototactic fishes will congregate behind the light barrages,
it is possible that traps in this position may also show increased yields.
While carrying out these experiments with light barrages the following
positively phototactic fishes were encountered: roach (Rucilus rUlilus ),
perch ( Perea jluvialilis) , pike ( Esox lucius) , pike perch ( Slizosledion
lucioperea ) and herring (Clupea harengus) .
6
Eel Culture and Live Storage
It is widely thought that the eel is unsuitable for fish farming. No doubt
this idea stems in part from the not insignificant number of unsuccess-
ful attempts to establish eel pond culture in Germany. However, even
the carp, one of the most popular European pond fishes, does not enjoy
such a long history of pond cultivation as the eel. It appears that over
2000 years ago glass eels were caught and kept in ponds in Macedonia
to be eaten as a delicacy (Eales, 1968). There is nothing simpler than
obtaining young eels for rearing. For cultivating trout, carp and other
fishes costly systems are needed in the early stages of rearing, whereas
for the eel these stages are by-passed by using glass eels. In some
circumstances certain devices can be used to ensure that the eels
migrate of their own accord into the ponds.
There has been no lack of attempts, both successful and unsuccessful,
to rear eels under controlled conditions. One is reminded of the lagoons
at Cornacchio, which were already well established in the eighteenth
century (see p. 242). However, these Italian waters do not really serve
as an example of controlled rearing. Production in the Adriatic lagoons is
left very much to chance; the only man-made structures are the rearing
basins. The lagoons of northern Italy can thus only be described as a
transitional stage between true pond culture and normal fishery
practices.
The brackish ponds of Arcachon near Bordeaux, which also origina-
ted in the eighteenth century, are of a similar, but somewhat more
EEL CULTURE AND LIVE STORAGE 32 9
advanced type. As in Comacchio, the pond populations are maintained,
at least in part, by utilizing the ascending young eels, and in part by new
stock imported from other waters (Le Dantec, 1953). Feeding experi-
ments were carried out as early as 1889 (Gobin, 1889). There is a
certain amount of opposition to the intensification of the eel industry
in southern France; here, in the Mediterranean lagoons and the Rhone
Delta, with their natural supply of glass eels and relatively favourable
climatic conditions, production has thrived for a long time without
additional stocking. The most recent manuals on eel culture (Usui,
1974; Forrest, 1976) are based almost exclusively on the well-established
eel farming techniques used in Japan and Taiwan.
6.1 Eel farming in Japan
6.1.1 The importance of eel farming

The Japanese were responsible for a breakthrough in intensive eel
rearing when they introduced high population density culture and
feeding. By 1900, iritensive feeding of young eels was established in
Japan (Walter, 1910). The first attempts in this field go back to 1880
(Matsui, 1952), but only in the last few decades have the effects of
intensive rearing in Japan had any detectable effect on the economy
(see p. 243). During this time intensive eel farming has increased so
considerably that the yield from natural waters is now far surpassed by
that from ponds. Eel cultivation in Japan is today more widely practised
than are carp and trout farming. According to figures cited by Brown
(1969 ; see Fritzsche, 1970), in 1965 about 8000 tonnes of carp and
5700 tonnes of trout were reared by Japanese fish farmers, whereas the
figure for eels was 16000 tonnes. The exceptionally wann climate,
especially in the south of the Japanese islands (where, in fact, most of
the eels are reared; Fig. 49), and, equally important, the highly favour-
able marketing conditions, are probably the basis for such healthy
development. Average temperatures in the Japanese eel ponds are
27 or 28 °C in July and August and 7 to 8 °C from December to March,
whereas in Germany the highest monthly average is unlikely to be more
than 20 °C. Eel farming has also become established on the island of
Taiwan (Formosa) to the south of Japan (Hickling, 1970).
Eel culture is still in the experimental stage in Europe (see p. 335)
and Canada (FA 0 Aquaculture Bull., 6,14, 1974), although in parts of
Japan results similar [0 those using A. japonica were obtained when
using European A. anguilla glass eels flown in from France (see p. 152).
330 THE EEL
However, A. anguilla does not seem particularly well suited to various

ecological factors it encounters in Japan (especially the higher tempera-
tures). ]n many areas yields were low as a result of slow growth and
losses due, among other causes, to ec(oparasires (Cueff, 1974; Chen,
197 2 ).
In 1957 the number of farms in Japan rearing eels for human con-
sumption was 764. By 1962 this figure had grown to 1893. The total
pond area, however, only increased from 1009 to 1257 hectares during
this same period, an indication of the high population density type of
culture practised.
6.1.2 Pond systems and yields

Before 1962 pond yield was between 5500 and 7200 kg/ha (Koops,
1966b). It seems, however, that these figures apply mostly to pond
systems using standing waters. By 1964 two-fifths of the eel farms were
using running water and producing a total yield of 26 000 kg/ha; the
remaining ponds yielded only 6000 kg/ha (Brown, 1969, see Fritzsche,
1970), In Taiwan 13600-26300 kg/ha were produced (Hickling,
1970). One can expect a yield of at least 10000 kgjha from German
trout farms, while carp ponds in central Europe only produce about
300 kgjha. In other words, the cultivation of eels in Japan is so intensive
that it can be compared to trout breeding in Europe.
In 1964 many of the eel farms were only 0'3 to 1'5 ha in area and the
majority were less than o·g ha (Brown, 1969, see Fritzsche, 1970). The
average labour force employed by these concerns was 1'3 workers per
farm (Koops, 1966a). Presumably most of them were family businesses.
In intensive cultivation the surface area of the individual ponds
used for fattening eels is considered to be quite large if it is between
800 and 5300 m 2 (Matsui, 1952). In Shizuoka, the most important
production area, this figure is said to average as much as 4500 m 2 in
some instances (lnaba & Uehara, 1962). Elvers are kept in very small
ponds, often in round containers 5 m in diameter and 60 cm deep.
When the eels reach lengths of between 8 and 12 em they are put in
containers measuring 30 to 100 m2, and at lengths of 20 em or more, in
ponds 200 to 300 m 2 in area (Usui, 1974),
In ponds using a running water system (see above), only one part of
the relatively large network of ponds is connected into the through-flow
of water at any given time. Since 1953 the tendency has been to in-
corporate resting pools measuring 50 to 60 m 2 into the pond (Inaba &
Uehara, 1962) ; these pools are kept particularly well supplied with
EEL CULTURE AND LIVE STORAGE 331
FlJ. 12... Covered Japan~ eel pond. Ldt : feeding place with additionaJ protection
from light. Right, in the background: paddle wheels to provide aeration
(photograph: Dr. FlUchter).
fresh, aerated water. During the heat of summer, with the lowered
oxygen tensions at night, temperatures in the resting pools are 4 to 6 °C
below those in the actual fattening ponds, and the oxygen concentration
1s0'9-3'8 mill higher. In addition to the inflow offresh water, there is a
pwnping mechanism to ensure proper water circulation (Fig. 124).
Between 3'2 and 12 tonnes of eels can be kept at night in one of the
resting pools just d escribed.
Eel ponds in Taiwan are stocked with 0"5-0"7 kg of elvers per square
metre and an increase in weight of 1"6-2"6kg/m t is expected in the young
eels (Hickling, 1970; Forrest, 1974). Calculations show that 0"4 kg/m 2
of glass eels in Japan produces 1"2 kg of young eels at least 8 em in
length, 0'5 kg/m2 of eels of this size produces 1"6 kg of young fishes
at least 12 em long, and that 0"4 kg/m2of individuals of this size produces
4 "0 kg of animals suitable for marketing, each weighing 190 g (Usui,
1974). Forrest (1974) reports yields of 3 to 12 kg/m2 of eels ready for
conswnprion.
Most pond systems are made of concrete or bricks; the rearing
basins supplied with running water are smaller than the conventional
ponds. In 1964 only 6"4% of aU pond systems in Japan still had earth
33 2 THE EEL
walls (Brown, 1969, see Fritzsche, 1970). It is particularly important

to construct ponds from concrete, stone Of, at the very least, wood
because eels will burrow into earth or mud (see p. 21 I ) . If one were to
build eel ponds in the same way as carp ponds, it would be impossible
to harvest a very large proportion of the animals when the time came
to empty the pond. Since a large number of predatory bird species will
attack eels, it is advisable to erect a cover over the ponds to protect the
fishes (Matsui, 1952). Covers also help to achieve and maintain the
highest possible water temperatures (Fig. 124).
6.1.3 Rearing techniques

As with trout fanners in Europe, eel cultivators in Japan have shown a
tendency to specialize in either the production of fish for stocking
or for the table. However, this trend only began in 1955. From 1952 to
1956, elver raisers recorded yields of between 12·5 and 34·3 young eels
per square metre.
Eels for stocking are reared in ponds by feeding up glass eels. After a
period of growth, the latter can attain a weight of 9 g (Matsui, 1952)
which corresponds to a length of about 20 em. More recent data indicate
an increase in length of 30 em in the first year (Brown, 1969,
see Fritzsche, 1970).
The rearing of table fishes in the fattening ponds takes place in the
second and sometimes even in the third year. Matsui (1952) found that
the eels had an average weight of 28 g in their second year and of 83 g in
the third. This corresponds to lengths of 25 and 40 em respectively.
More recent reports (Brown, 1969, see Fritzsche, 1970) indicate that
eels reach a suitable marketing weight in their second year and have a
final weight of 150 g. It should be noted that, at every transfer, eels
in Japan are clearly sorted according to size, so that it is difficult to
provide data on average ages. Since the Japanese prefer relatively
small eels, the animals are ready for marketing at a size corresponding
to the beginning of sexual maturity in wild male Japanese eels (see
p. 189)·
Mortality in the first year is calculated at about 34% , but drops to
between 6 and 7% in the second and third years. Since eels do not eat,
or eat very little during the winter, a loss in weight can be expected
during this period. Weight loss over the period November to March is
thought to be between I and 13%, the average being 4.1 % . The smaller
the eels, the greater the weight loss (Matsui, 1952).
6.1.4 Feeding
When using fresh food, e.g. fish, the feeding quotient necessary for
producing table fishes is between 7'5 and 9'0 (Koops, 1966a); it is
slightly higher for eels than for trout (Schaperclaus, 1967). In Taiwan
the feeding quotient is as high as 7'6-19'9 (Hickling, 1970); in Japan, it
is 7 (Usui, 1974; Forrest, 1974). The quotient increases during the
course of the cycle of cultivation. In the first year, i.e. when rearing
young fishes, it is between 5 and 6, in the second year about 8, and in
the third year 10 or more. Therefore the most economical eel rearing
programmes are those based on the shortest possible cultivation cycle.
The daily food ration is between 5 and 15% of the fish's total weight,
depending on the seasonally determined temperature level (Matsui,
1952), and is thus relatively high compared with that for trout
(Schaperclaus, 1967). If the temperature drops below 10 QC, no food is
given; elvers even stop eating at 13 QC. For this reason, artificial heating
is common in spring in Japan, and structures like glass houses are built
over the ponds (Fig. 124). Eels consume 85% of their annual food
ration between July and October.
The use of dried foods, which is becoming increasingly common
nowadays in both eel and trout farming, requires a lower feeding
quotient. Advertisements for the products of one dried food manu-
facturer in Japan advise a feeding quotient between 1-2 and 2·6 (see
below) ; Usui (1974) suggests 1'4. The daily food rations, as based on
the eels' weight, are correspondingly lower. Another manufacturer
claims that, as compared with fresh fish as food, a quarter to a third the
quantity of dry food is needed.
Nowadays the larger eel farms have gone over to the exclusive use of
food pellets (Brown, 1969, from Fritzsche, 1970). The smaller fanns
only use pellets when the price of fresh fish is too high.
Fresh food is generally made up of fish, fish waste and fish meal.
In addition, silk worm cocoons (dried or raw), Mysis and liver are used
in smaller quantities (Koops, 1966a). The fish is cut into small pieces
or strung on wires. During the first two weeks, glass eels are given
pieces of mussel flesh or chopped earthworms, then Tubifex, and
finally, increasing quantities of chopped fish meat as well (Usui, 1974).
Two Japanese firms give details about the dried food which they
manufacture. The basic ingredients are fish meal, liver meal, skimmed
milk powder, alpha starch and yeast. Food manufactured for young
eels differs slightly from that prepared for fattening eels. Both finns
claim a raw protein content for their products of 45 to 52% ; young
334 THE EEL
FIg. I2S Wire·mesh feeding boxes used in Japanese eel ponds. Above: boxes with
wooden frames. Below: in the backgroun~, wire 'baskets' in which the food is
placed. In the foreground one can see how the eels actually worm their way
into the bowl containing food (Photograph: Dr. Fluchter).
eel food contains about 2% more protein. Analysis of the dry eel food
indicated 2-5 to 6% fat, 0- 1 to 0-6% raw cellulose and 15% ash, as well as
numerous vitamins, minerals and trace elements_ Amino acids essential
for growth in the eel are: methionine, tryptophan, threonine and
valine (Nose, 1969)- A test to determine the best composition for eel
foods gave the following breakdown: casein, 54 ; gelatine, IS; dextrin,
8; corn oil, 6; cod liver oil, 3; cellulose powder with vitamins, 6;
mineral mixture, 7 ; L-tryptophan, 0-5; L-cystine, I; water, 200 (Arai
et aI. , 1971). Before it is given to the fishes, the dried food is mixed with
water and a special oil used for this purpose; 10% of oil is used at low
temperatures and 15% at high temperatures_ Takai & Ishii ( 1971)
carried out experiments to establish just how attractive eels find the
artificial foods_ If both natural and dried foods are used together, the
latter should be given first_
Food is presented in boxes covered with a wire mesh, and suspended
just below the water surface (Fig_ 125)_ The size of the mesh varies
from 3 to 15 nun, depending on the size of the eels to be fed_ Wooden
boxes with a feeding channel 10 to 15 nun wide may also be used. If
the food is floating on the surface, the eels will come out of the water
to feed_ The methods just described make it possible to remove uneaten
food from the ponds, which is particularly important even in the
case of elvers_ Feeding usually takes place in the morning, and in some
cases in the evenings as well_
6_ 1-5 Profit and expenditure

In 1964 the average costs for maintaining a running water system
(see above) were 13541 dollars and for standing water ponds 18286
dollars_ The two systems showed average profits of 2113 and 2142
dollars respectively, per fish farm_ A breakdown of production costs
in 1964 for a running water system (and, in parenthesis, that for standing
water ponds) was as follows: eel stock, 24-9% (16-1 %) ; food stuffs,
54-7% (5 1-1%) ; management, 2-5 % (3-6%) ; wages, 4-8 % (11 %) ;
electricity, machinery and fuel, 3-1 % (3-7%) ; maintenance, including
temperature control systems, 2-6% (2-3%); interest, 2-8 % (3-6%);
insurance, rent and taxes, 3-7% (6-5 %); miscellaneous, 0-9% (2-1 %) _
6.2 Experiments on eel farming in Europe

As was mentioned at the beginning of this chapter, attempts to establish
eel farming were made earlier in Europe than in Japan_ However, in
336 THE EEL
Europe very little has come of these efforts, that is, if one excludes the
extensive, almost lake-sized 'farms' in France and Northern Italy
(see pp. 295, 328). Since fairly large numbers of experiments in this
field have shown a promising start, it seems appropriate to give a brief
account of them here.
As early as 1929 an attempt was made in Gennany to improve an eel
population by feeding the animals (Speichert, 1952, from Wundsch,
1953). Although this experiment was not carried out under properly
controlled conditions, but in a sealed-off clay pit, it indicated that the
eels responded well to feeding. After initial set-backs, which are often
difficult to avoid in such experiments, the following took place. When
sizeable pieces of fresh fish were put into the water, large numbers of
eels massed together to eat the meat, leaving only the bones. After a
period of daily feeding the eels gathered with great regularity, apparently
as if trained, at the sound of the food being prepared, i.e. they behaved
in the same way as carp and trout are also known to respond under a
similar routine. One of the pre-requisites for successful fish farming
seemed, therefore, to be fulfilled. However, the extent and uniformity
of the eels' growth increments appeared to be unsatisfactory: after
seven years a few specimens had attained a weight of 170 g, others
weighed only 50 g; the great majority of fishes introduced as elvers
showed hardly any size increase at all. The daily food allowance, as
calculated from the available data, was, however, at most I % of the
weight of the stock fish, i.e. far below the quantities used in Japan
today (see p. 333).
Only recently have controlled experiments been carried out in such a
way that the results can be properly assessed; these results have been
combined in Table 37. According to these data, young eels have been
reared for use in 'fattening units', and fishes of a size suitable for human
consumption have been produced. It is also evident that eels can be
cultivated at the lower temperatures prevalent in central Europe.
Weight increases appear, in part, to be no worse than those achieved in
Japan.
Only a few of these experiments provide convincing evidence
suggesting that eels were produced in economically satisfactory
quantities. One of the more successful eel farms is on a warm water
system in the district of Altenburg. Here, eels weighing 30 g showed an
average increase in weight of 130 g. The initial stocking weight was
increased fivefold, without using an excessively high food quotient.
In 1968 the yield was 2650 kg of eels. The population density ( 150-250
kg/m 2), which is unusually high, was maintained by a substantial
through-flow of water, namely c. 200 rna of warm water per hour.
It should also be noted that dried food was used in conjunction with
fresh food.
Difficulties have been encountered because of the markedly diverse
growth rates of young eels in ponds; frequent sorong of the eels into
size groups is thus unnecessary. This differential growth is probably
due to adverse conditions, especially to low temperatures. However)
even glass eels reared in warm water, and fed, showed differential
growth, with some individuals growing to a large size and others scarcely
showing any weight increment at all (Meske, 1968).
If eels are kept in ponds, it is important that they be provided with
sufficient hiding places. In one case) corrugated asbestos sheets were
used. High density stocking also seems to have a positive influence on
both thigmotaxis and negative phototaxis. Food intake does not increase
until a certain maximum population density is reached. Eels thus
seem idealy suited to crowded living conditions as is also evident from
their tendency to crowd together in natural conditions (see p. 211).
When food is placed on the water surface) it appears necessary to
provide a certain degree of protection from light. For this reason, a
roof 10 m 2 in area, resembling those used in Japan, was built over one of
the Mosel experimental ponds.
As was noted before, one must prevent the eels from taking refuge by
burrowing into the substrate. If this does happen, even harvesting by
eiectrofishing becomes a problem, and therefore a solid bottom and
sides to the pond are essential.
In Europe, as in Japan, the importance of phytoplankton in main-
taining a reasonable oxygen level has been recognized (Nolte, 1975).
One should aim at producing water which is dark green in colour due
to the presence of Scenedesmus sp., Chiorella sp. and Pediastrum sp.,
etc., so that even as late as August, the oxygen concentration in the water
remains above 4 to 5 mgtl at night. Blooms of blue-green algae such as
Microcystis sp., Aphanizomenon sp. and Anabaena sp., however,
lead to severe oxygen depletion and should be controlled by using
copper preparations at the rate of 1-2 kg/ha.
Pond and aquarium experiments in Germany have shown that it is
possible to rear eels on dried food alone (Meyer-Waarden & Koops)
1968; Koops, 1971). In pond experiments on the Mosel (see above) the
feeding quotient when using fresh fish was 7-1 I, whereas it was possible
to keep the quotient at between 1·8 and 3'7 with the use of dried food.
In these experiments the water temperature was 21 °C and the eels'
daily weight increase was I % of the body weight.
33 8 TH E EE L
Ta ble 37 Experimental eel culture

Ty~of Wattl' Si:t of ttls SizeD! tels Total Total Stocking
Location rearing umdiljol1s 011 intra- at mdof !OOg}1l weight density
urnt duuion experimml on intra- almliflj
ducrjon lrpnimenl
Berlin Natural Glass eels l SI summer 7-16

pond, 26- 5'0- 1I "2g, cds/m'
9S m' '4"5-17"9
on; 2nd
summc-r,
18-22 em
Godeben, Ponds, Spring Glass T8g ,kg 75
=~, loom., ~,~ eels, 0"4 g (0'5-5"') «Ism'
E'''' with loamy
soil
Gorleben, Ponds, Spring Young max. ISO g 210 kg Stocking '50
central 100 mt, water eels, 13-6 g weight eels/m'
E'", with loamy doubled
soil
MOdenl River River Young 30·8 g 350 kg S40q ,0

Mo," pond, ~,~ eels, IS g. eds/m'
...
700 m ' 2) -' em
MOdenl River River 93. ' ,.,

M"" pond, water «Is/m'
700 rn'
Milden! 2 river River 11 9 and 220 and 225'5 and 449'9 and 3 and I
Mosel ponds, 700 water 49 g rC'S- 109 g res- 753 kg res- 162 1 kg eel/m'
"d pec;livciy . pectively pcclivcly respce- rcsp.
I l ooo m" tively
Darp, Bailie Brackish 3-S pamper Young 44'4g, 4-33

pond, thousand ~", wi th some eelsfm "
6som' salt. Temp. 46·S g upto l sog
divided by 16'1 °C in
nets Sept.,
8 ~C in Oct.
Darp, Baltic Brackish Young 90 g, with

so
pond,
6som"
eels, g
,00 •
some up to
di vided
by nets
WieJenbaeh, Pond,
near
Munich
110m',
with 90 m"
Water
temps.
too low,
Glass eels,
eels,
O'lS g
1'4 g with
some up
to 11 '3 g
1'9kg
'" 68
eelsfm"
of water average
16.s oCand
IS '3 °Cin
Ju ly and
August
respectively
Rosiu, Channel ""'d Young Category SlOelting 150-150
district of system water eels, 30 g 1, 15% weight kg/m '
Alten burg outlet Category increased
11,70% fivefold
(160 g)
C..atc.:rorv
EEL CULTURE AND LI VE STO R AGE 339
% Duration Ty/U of food Fudillll Daily ration Rnnark s Author

Mor/a/ily of experi- used quotiem as % of
mrn' population
weight
<.7 Freshwater Miiller,I<}64;

months fish and see also
and c. 8 mussels Miiller, 1967
months
resp.
3°% 7 months Predomi- s-rs% Migrat ion drive dis- Koops, 196s;
nandy natural appeared after 2 see also
food months; much Kijthke,l966
natural food
7'8% 3 months Marine and max. 10 % Conditions bad; 15% Meyer-
freshwater eels with cauliflower Waarden&
fishes, tumoun; I l % losses Koops, 1968
shrimps, on introduction as a
mitten crabs resuli of bacterial
disease
13% 6 momhs Fishes fro,n H)-II 10% in Conditions bad ; 24% Koops,
the Mosel warm eels with ClIulifiower 1966a;
(Cyprinidae), weather, tumours on imroduc- see also Jens,
smelt, spleen,
blood
1'6% in
October
tion '96'
<.6 Fresh fish 7-' Almost all eels at the Meyer-
months silver eel stage when Waarden &
experiment finished Koop, t<}68
5 and 6 Mostly fish 3·7 and With both fishes and Koops, 197t
months in the firSt to resp. dried food, the cost of
respec- pond, d ried feeding per kilo-
tively food in the gramme of weight
,,..
second increase was c. 3·50 OM
4 months Freshwater max. Food intake too low, Lauterbach ,
fishes, edible 12 ·5 % probably the result of
mussels poor water supply and
a population density
that was tOO low
2 months 6' A continuation of the

previous experiment ,,..
Lauterbach,
46% 4t months 45 % spleen, 2-3% of 8ohl, I<}68

45 % fish roe, stocking
10% yeast weight
12 months 50% fish, 4-6 K eepi ng elvers Seidlitz, t969

50% dried proved difficult,
food continual sorting
necessary
340 THE EEL
The dried food consisted of 50% crude protein and 17% crude fat.
This finely ground meal was mixed with an equivalent weight of
water, in a concrete mixture, to produce a paste which did not dis-
integrate in the water for some hours (Koops, 1971). Instead of lowering
the sieve-like feeding troughs onto the bottom, as was done in earlier
experiments, a feeding table, based on the Japanese model, was placed
directly on the water surface (Fig. 125); the top of the table was made
from wire mesh 3 cm2 diameter. Protected by the light screens
mentioned earlier) the eels quickly learned to feed from this raised
platform.
When rearing eels for the table, it should be remembered that cultured
fishes attain proportionately greater weights than do wild specimens.
Comparisons with eels from rivers and from the Baltic showed that
pond-grown eels of the same length were 20% heavier (Meyer-Waarden
& Koops, 1968). Elvers taken from coastal waters for rearing in ponds,
therefore, at first show only a slight increase in length, whereas the
weight increase is quite considerable (Koops, 1965, 1966a, 1971).
It is clear from the discussion above that commercial eel rearing
demands a considerable outlay if it is to be successful. Large ponds
with an extensive through-flow of water appear to be more effective
than conventional ponds, but rwming costs are, of course, much
higher. Labour costs are not low either. Several times a year all the eels,
and especially the smaller individuals, must be sorted according to
size. In addition, food has to be prepared, feeding has to take place at
least twice daily, and food remains and sediments must be removed.
If fresh food is used, the installation of cool storage is also necessary.
Eel rearing cannot be considered profitable unless the total outlay is
covered by incoming revenue and there is an attractive profit as well.
These and other factors have led to the disappearance of eel farming
in southern France, despite the relatively good climatic conditions
there (for an account of recent cultivation experiments in southern
France, see Seltz, 1974). Attempts have been made recently in Germany
to utilize warm water effluents from the cooling systems of power
stations (Koops & Kuhlmann, 1976).
6.3 Live storage

It is often necessary when catching and selling eels to keep them
alive for some time, as for instance, when delaying the sale may mean
higher prices later. Some eels have an unpleasant flavour when first
caught and it may be necessary to keep them until this has disappeared .
EEL CULTURE AND LIVE STORAGE 34 1
A well-known example is the so-called phenol flavour. This taste does
not originate from the phenols themselves, but from accompanying
substances, and is produced by detergents. In a 'holding' experiment,
eels were kept for three weeks in water at 16 to 20 °C; this led to a
marked, but still unsatisfactory decrease in the unpleasant flavour
(Mann, 1960). After three months the tail region, which is particularly
susceptible, still tasted slightly unpleasant in a few of the fishes. It is
thought that feeding the eels leads to a more rapid improvement in
flavour.
Feeding has other advantages too, as it counteracts the otherwise
unavoidable weight loss which occurs while the eels are in captivity. In
connection with this, experiments have been carried out in the small
ponds at Gorleben (see Table 37; also Koops, 1965), where 800 kg of
eels, with individuals ranging in weight from 0'4 to 1'5 kg, were kept
in one of the 100 m 2 ponds, at a temperature of 16 °C. Previous
experience had shown that a weight loss of 10 to 20% could be expected
within a period of three to six weeks. In this experiment the eels were
given 16 kg of food daily, and the total weight loss after five to six
weeks was onlyo·s kg. Since the daily food ration amounted to about 2%
of the body weight, this percentage can be regarded as the minimum
quantity of food necessary to maintain body weight. It thus appears
that the daily rations shown in Table 37 are, in some cases, too low to
allow more than the maintenance of existing body weight. Un-
fortunately, it is not known from these experiments to what extent the
phenol flavour disappeared, but it is almost certain that the holding
period was sufficient to make the eels suitable for marketing.
7
Diseases, Parasites and Injuries
Nowadays, as a result of much research, we have a fairly extensive

knowledge of the diseases which occur in fishes of economic importance.
In many cases stocking or intensive pond culture results in eel popu-
(ions of a particularly high density, and for this reason, various serious
diseases are not uncommon in the eel; consequentlyJ general books on
fish diseases tend to deal with eels in some detail (e.g. Schaperclaus,
1954; AmIacher, 1961; Reichenbach-K linke, 1966). This section is
therefore restricted to those diseases which have direct economic
consequences; other diseases can be dealt with more effectively in
tabular bibliographical form. For further information, particularly
about diagnosis and treatment, reference should be made to the works
cited above.
The term 'injury' is used here to refer to mechanically inflicted
damage. Damage affecting the whole eel population, which is more often
ecological than pathological, is described in the sections on ecology and
conunerdal fishing (e.g. see pp. 150, 162,248).
7. I Virus and bacterial diseases

By far the greatest proportion of epidemic-like diseases which have
severally reduced whole eel populations, are the result of virus or
bacterial infections. The majority of diseases of economic importance
are therefore dealt with in this section. While it was possible to study
342
DISEASES, PARASITES AND INJURIES 343
bacteria in eels using well tested methods, research on viruses meant
covering totally new ground. Particular difficulties arose, for example,
when it became imperative to investigate the cause of 'cauliflower'
disease, an infectious ailment which, although it has epidemic-like
characteristics, is clearly not of bacterial origin. A further instance of
viral infection was found in the erythrocytes of the American eel
(Doyle, 1971 ). Using cell cultures of eel kidneys, Wakabayashi &
Egusa (1969) have developed new techniques which are useful in the
study of viruses.
7. 1.1 Cauliflower disease

Investigations suggest that the pathogen responsible for cauliflower
disease should be regarded as a virus. Infonnation on this disease started
to accumulate after the Second World War, though its occurrence
was first reported in 1907 (Nagel, 1907) ; the first scientific investigations
took place in 1950 (Christiansen & Jensen, 1950, Schaperclaus, 1953).
Cauliflower tumours can be considered a typical eel disease although
cod and, according to other reports, bleak (Amlacher, 1961) are among
the species of fish which may also be affected.
The disease has probably only reached an economically significant
level in the eel. Unfortunately its devastating effect is seldom appreciated
because it is not manifest in a dramatic fashion, such as mass deaths.
However, if one takes into account the fact that as many as 24% of
young eels in ponds may suffer from the disease (Table 37; Koops &
Mann, 1969) and not recover, then its true extent becomes clear.
One must, for example, expect complete losses among infected young
eels which are taken from brackish waters and used as stock for ponds.
Infected individuals develop skin growths on one or more parts of
the body surface. In most cases, however, these skin tumours fonn on
the upper or lower jaw (Fig. 126). In appearance they resemble a
cauliflower, hence the tenn used to describe the disease (Schaperclaus,
1953). Initially the swellings are whitish in colour, but later become
light brown, turning even darker in colour with increasing pigmentation.
The swollen mass may become so large that the protrusion from the tip
of the head can exceed the length of the head itself. Affected eels are
often very emaciated. The fat conent of badly affected individuals
between 19 and 29 cm long drops from 1·6-5'0% to 1% or even less
(see Table 43); the average weight decreases by 35 % (Ltitunann &
Mann, 1956, Koops & Mann, 1966). Although there is no evidence of
high mortality among these animals in captivity, affected eels which
344 THE EEL
Fig. 126 Eel with cauliflower disease (Photograph: Marschall).
are caught in traps die more easily than do healthy animals. The disease
is commoner in smaller eels than it is in larger individuals: in pond
reared animals the incidence is 27% in eels between 15 and 19 em
long, 22 % in eels 20-24 em long, 19% in eels 25-29 em long, 14% in
eels 30-34 em long and only 3% in animals over 34 em long (Koops &
Mann, 1969). This can be explained by the fact that affected animals
stop growing, and have a higher mortality rate.
Histological investigations have produced the following picture of
the affected area: a mass of undifferentiated tissue replaces the highly
differentiated epidermis. This change or transformation begins in the
stratum basale and gradually replaces the nonnal structure of the
epidennis (Schmid, 1969). 'The growths are called fibroepithelial
tumours or papilloma, i.e. they are villous formations with protruding,
tissue-like papillae which themselves have small, lateral, secondary,
tertiary, etc. papillae covered in an epithelium-like tissue' (Schliperclaus,
1953)·
As regards the cause of the disease, it has proved impossible to detect
the presence of bacteria, and none could be cultured from infected
tissues, so it hardly seems likely that the pathological changes are
brought about as a result of bacterial action (Christiansen & Jensen,
1950; Schaperclaus, 1953; Schmid, 1969; Schubert, 1969). On the
other hand, virological evidence from cell cultures was positive, i.e. in
experiments using fish cell cultures, the infectious agent was isolated
from the blood of diseased eels and was shown to be present after
several cell passages (Pfitzner, 1969; Schwanz-Pfitzner, 1973). With an
electron microscope it was possible to detect a virus in these cultures,
and electron microscopic studies of the growths appeared, at ' least
partially, to confirm this result. Inoculation of blood from infected eels
into healthy animals had a positive result, but, final confinnation will
only be possible with the infection of eels by the virus from the cell
cultures. It is assumed that here one is dealing with a Viremia, which is
seldom the case in tumour producing viruses (Pfitzner & Schubert,
1970).
It appears that the virus is only active for a short time in water.
Wounded animals are, therefore, particularly, if not exclusively
vulnerable. For this reason it seems that densely popUlated groups, and
especially animals under intensive cultivation, are most likely to become
infected (Pfitzner, 1969). If possible then, only healthy animals should
be used for pond rearing. Natural populations with very high densities
are also at risk.
Another possible factor in the pathogenesis of cauliflower disease is
the effect of waste substances on the eels' skin. Within 24 hours,
fluoride ions and mono-iodine acetate produced a tumour-like multi-
plication of cells in the eel's epidermis (Peters, 1970a, b). Presumably
the chemicals act by restricting oxygen uptake and by increasing the
exchange of glycolytic substances. The effect of these inhibitors is
annuled by inorganic diphosphate and also, presumably, by the
improved energy supply to the cells. Using the same principles, it
was possible to destroy the cauliflower tumours chemically, causing
them to drop off. There thus appears to be a treattnent for this disease,
but further research on the origins of cauliflower tumours is still
needed. The pock-like growth or folds on the skin of glass eels, described
THE EEL
by Bremer (1974), probably have other causes and, in the opinion of the
author, are due to osmotic imbalance.
Since it first appeared in the coastal regions of the Baltic
at the beginning of the century (Schaperclaus, 1953), cauliflower
disease has gradually spread over almost all areas of the Baltic and the
North Sea. Recently it was found that the disease also occurs along the
west coast of Scandinavia (Gullestadt, 1972). Another case was reported
in the Black Sea (Radulescu & Angelescu, 1972), where eels are only
found very occasionally (see p. 104). Cauliflower tumours have yet to be
found in eels from Belgium and the British Isles, and to the west and
south of these countries.
The disease not only occurs in brackish waters but also in freshwaters
where population density is high (e.g. ponds). Between 1957 and 1959
5'6% of eels in the Lower Elbe were affected, and in 1967 the figure had
risen to 12% (Koops & Mann, 1969). These authors did not find any
seasonal fluctuations. Observations on the weir at Geesthacht in 1964
and 1965 (Tesch, unpublished) indicated a certain increase in the
incidence of tumours between spring and autumn.
7.1.2 Fresh- and saltwater eel diseases

There are two serious bacterial diseases which the eel may contract
and, although different, both have very similar external symptoms.
One is limited to freshwater animals and is caused by the bacterium
Aeromonas ( Pseudomonas ) punClala (Schaperdaus, 1930, 1934); it is
generally known as red pestilence. As the name implies, the second
form, the saltwater disease, occurs in the sea or in brackish waters. It is
caused by the bacterium Vibrio anguillarum (Bergmann, 1909;
Schaperclaus, 1927, 1934) and is also known as the red tumour or red
disease.
The symptoms of the two diseases are compared below (Schaperclaus,
1954)·
Freshwater eel disease

(I ) Eels without external symptoms of the disease swim around
slowly; shortly before death they hang, crumpled up, as though
suspended from the surface.
(2) Red spots, bands or flecks spread, with varying intensity, over
the whole body (but especially in the abdominal region), the fins
and the anal region.
(3) White, often bluish patches appear as a result of loss of mucus,
epidermiS and even parts of the dermis.
(4) Raw, exposed areas develop. The musculature is exposed and
surrounded by pieces of red or white necrotic dermal tissue. Large
numbers of such spots are seen particularly on the tail, which may
look almost as though it has been eaten away.
(5) Boil-like swellings, ranging in size from a pea to a walnut,
develop on the head, particularly in spring. These sometimes burst
and heal up, leaving a white scar; large wounds tend to lead to
general infection and death.
(6) Blindness.
(7) Internal haemorrhages, bleeding in the liver and inflammation of
the large intestine.
Saltwater eel disease

( I ) Sudden death without any external indications of disease.
Shortly before death the eels come to the surface and may, from time
to time, show spasmodic convulsions.
(2) Rash-like reddening of the skin, fin margins and anus.
(3) Swelling and reddening of the musculature around the heart
regIon.
(4) Tetantic spasms which produce contraction of the anus.
(s) Red, often tumorous swellings which may be covered with a
transparent growth of skin.
(6) According to Schaperclaus (1927) and Nybelin (1935), the
liver often shows haemorrhagic patches I nun in size, i.e. a slight
reddening.
It is quite clear that the pathological results of these two diseases are
very similar. For those who have experienced dropsy in carp (also
induced by Aeromonas punctata), freshwater eel disease in particular
will be a not unfamiliar phenomenon. In the case of both diseases,
external symptoms may not be seen if the animals begin to die in large
numbers.
Vibrio anguillarum bacteria of type A occur in the gut and liver of
10% of non-diseased animals. The proportion of infected eels rises to
60% before the outbreak of the disease. During the acute phase, in
contrast to the previous stages, the kidneys, heart and blood also
become infected, that is with type B bacterium (Mattheis, 1964a).
Reduction of the haemoglobin content, the haematocrit and the
erythrocyte count could be seen not only in cases with obvious external
symptoms but also in animals where external symptoms were not
detectable (Kreutzmann, 1973).
A red fin disease, also connected with anaemia and an increase in the
348 THE EEL
Table 38 Parasitic infections in eels, caused by fungi, sporozoans, ciliates anC!

crustaceans, and which have damaging effects on the commercial eel
fishery or pond culture
Parasite Host animal Water body No. of affected Size and number Behaviour
animals of dead animals
Saprolegnia A . japonica Eel ponds, 19"4% of Lethargic

parasitica Japan (sec 1442 kg
p. 32 9)
Myxidium A .japonica Eel ponds, 100 kg in each None, 33-38 em Almost

malSui Japan (sec pood long nonnal
P·3 29)
Ichthyophthirius A. a'fllUil/a SlOrage dam at A few 100 spcci-

mulriftH;s Klingenberg, mens, small eels
Saxony, very weighing up to
low water level 250 g
Ichthyophthirius A. anguilla Small ponds in Young eelg

multijiliis Gorleben and
Wielenbach (see
Table 37)
Goezia sp. A . anguilla Lower Elbe U>-30 em

(Nematode
larvae)
Argulus sp. A . anguilla Lagoon ponds, Death resulted Serious dfccts

Venice (see if water on conunercial
p. 295), salinity temperature eel fishing
15-29 parts rose above
per thousand 25 °C
number of immature erythrocytes and leucocytes, has been reported

in the Japanese eel (Enomoto, I969a, b). It is possible that this is in
some way related to the European eel disease. Usui (1974) refers to
Aeromonas liquifaciens as the bacterium causing similar symptoms in
A.japonica. A red disease found in A. anguilla in the Randers Fiord and
in the North Sea near Derunark is known under the same name (Hansen
Nature of the distase Time of infwion Control ami cure of Remarks Author
and danwge andwauT the disease
umperalUTt
Strong fungal growth 14-21 °C Bathing for 20-90 se<.: Some eels Hoshina &
mainly on the head and aV = 16 °C in Malachite green were also Ookubo, 1956
tail of the animal March to May 1:20000 to I: ,0000. infected with
40 min in KMn01 . PseUl:wmonaJ
I: 10 000. 4-40 mm m
formal in I : 1000 to
1:3000
Large numbersofwhite June Duration: Hoshina, 1952

cysts over the whole 23 °C 20 days,
body except the head moo
and gills unsightly
The whole epidermis is Control impossible Timmermann,

destroyed and sloughs unless one uses try- 1939
off paflavin, chloramin,
idithyorapid and
raising the salt con-
centration (Schiiperdaus,
J954; Rekhenbach-
Klinke, 1966 ; Tesch,
1968a)
The whole epidermis is 0·'S-0·t8 g/m' Koops, 1965

destroyed and sloughs Malachite green Bohl, 1968
off
Tubercles, mainly in M,y Dem:cke,

stomach, also intestine, 1935
cutis, musculature
Body surface, with up Infe<.:tion occur- Control in ponds is This parasite D' Ancona,
to 1000 specimens per red at tempera- impossible, otherwise can exist in 1960
eel; even in the gill tures above lysol, potassium salt concen-
cavities of some eels 10-15 °C pennanganate, trations of up
vermicide to 40 parts
'"
(Schaperclaus, 1954;
Reichenbach-Klinke, thousand
'¢O)
& Bonde, 1973). Pseudmnonas anguil/iseptica has been mentioned in

cOIUlection with the Japanese eel (Forrest, 1974).
Between 1930 and 1933 alone, Schaperclaus (1934) studied 28 cases
of eel disease, the majority of which were probably outbreaks of the
freshwater disease. It is likely that the 'infectious septicaemia' des-
cribed recently in eels from the Lower Oder (Chelkowski & Stankiewicz,
35 0 THE EEL
19<)4) can be attributed to one of the two eel diseases described above.
Saltwater eel disease has caused extensive damage. In 1930, 10 to
15% of the eels caught in the Baltic died after one day in captivity.
After eight days the captive eels had all died. Losses in the Saaler Bodden
in 1932 were estimated at 30 tonnes. In 1931, 1000 eels were picked up
along a stretch of shore ontyone kilometre in length on the isle of Vilm.
In 1959, it seems, the disease did not have quite such a devastating
effect; according to the inhabitants of Vilm a few hundred dead eels
were washed ashore in August (Woiter, tg(0). It has been reported that
in two days 300 kg of dead eels were washed up along a 3 km stretch of
shore at Strelasund (Mattheis, 1960). This disease always appears at
the same time over wide stretches of coastline in the southern Baltic
as a result of high temperatures. The following quote gives some idea of
the extensive distribution of the disease: 'It was possible to start one's
journey in the Stettiner Half, to continue it over the Greifswalder
Bodden and the waters of southern Rugen as far as the Breitling near
Wamemiinde, to enter the lower stretches of the river Trave or to travel
up the Kaiser Wilhelm Canal at Holtenau as far as Rendsburg - every-
where one came across armies of large dead eels.' (Meyer, 1933; quoted
from Schaperclaus, 1934)
In the North Sea also the saltwater disease causes a considerable
amount of damage from time to time (Aker, 1970). For example, in
1969 50% of the eels landed at Busum in August and 62 % of those
landed in September were diseased. All these animals were from the
relatively deep waters (more than 50m) southofHeligoland. The disease
seems to have subsided over the last few years: in 1969 and 1970 the
incidence was 60%; in 1973,30-4%; and in autumn 1974 only 0-1%
(Aker, 1975). However, not one case has been reported from the coastal
regions of the Heligoiand Bight. The disease occurs simultaneously
in the fiords of east and west Jutland. For some time eels in the North
Sea have been known to be infected with Vibrio anguil/arum
(Schaperclaus, 1954).
High temperatures, from 19 to 22 °e, are the pre-requisites for
virulent activity in this bacterium_ The onset of low temperatures
checks the disease at once (see also Moroga & Egusa, 19(9). Saltwater
eel disease is also connected with pike disease, which is caused by the
same organism. Pike disease often breaks out shortly beforehand in
the same year (Roy, 1961).
It is impossible to combat the disease in natural waters. Containers
which have come into contact with diseased eels should be disinfected
(by using a strong solution of caustic soda or by letting the containers
DISEASES, PARASITES AND INJURIES 35 1
dry out thoroughly). Cool water reduces the danger that the eels will
die while in captivity; endangered animals should not be kept more
than three days. Antibiotics mixed with the food, given as an injection,
or used as a bath, help combat the disease in aquarium eels which,
according to the author's experience, are also easily infected. Eels do not
appear to suffer unduly if given a five-hour bath in a solution of, for
example, 250 mg/l of tetracycline, and this has a definite dampening
effect on the disease. For the methods used in feeding or injecting
antibiotics, reference should be made to the techniques generally
employed in treating dropsy in carp (e.g. Schaperclaus, 1956).
Gill disease and swelling of the intestine, which occur in the Japanese
eel, are not to be confused with the infections described above.
Chondrococcus columnaris is thought to be the causative agent in gill
disease (Forrest, 1974; Usui, 1974), which is characterized by the
apparent erosion of the gills. This disease is also known as 'bacterial
gill infection'; it led to losses of80 to 90% amongst glass eels uansported
from Italy to Japan (Ghittino, 1972; F AO Aquaculture, 4 (4), 12, 1972).
It should be noted, however, that bacterial infections usually have a more
marked effect on A. japonica than on A. anguilla. A disease of unknown
origin called branchionephritis had a devastating effect on A. japonica;
in 1969/70 it destroyed 2600 tOlU1es of eels of this species (Usui, 1974).
7.2 Infestation by fungi, protozoa and metazoa

Parasites in eels have less effect on the commercial eel fisheries than do
viruses and bacteria. Nevertheless, in isolated instances, they can be
the cause of considerable losses, as is clear from Table 38. One of the
most dangerous parasites, which occurs over and over again and which
causes a great deal of anxiety among owners of ponds and aquaria, is the
ciliate protozoan Ichthyophthirius mull1fil£is. This parasite is not
specific to eels although it is found particularly frequently on them.
This may be the result of the eel's tendency to live in hollows or in
very restricted areas on the bottom, where the exchange of water is
minimal and where several animals often live in very close proximity
(see p. 209).
The parasite I. multifil£is occurs only in freshwaters and semi-
brackish waters. Since eels also live in saltwater, and occur both in
temperate and tropical waters, one should exercise great care in identi-
fying the causative agent of the so-called 'white spot' disease. In sea-
water Cryptocarion irritans and in freshwater aquaria Ichthyophthiriodes
browni could well be responsible (Tesch, IS)68a).
35 2 THE EEL
Of the remaining ectoparasites, an infection by the fungus Sapro/egnia

parasitica is also a frequent cause of illness in eels (Table 38); it is most
likely to occur as a secondary infection, following injury. In 1965 and
1968 this disease is reported to have killed several hwtdred tonnes of
animals in Japanese eel ponds (Usui, 1974). It occurs throughout the
world and is so conunon among other species of fish that a detailed
description would be superfluous here. Under reasonable living
conditions, serious damage is unlikely both in captivity and in the
wild.
Other parasites seldom have a harmful effect, that is, as far as the
European eel is concerned. Table 39 provides a review of the parasitic
fungi, protozoa and metazoa found in the eel. This table is in no way
complete; the list of acantbocephalans could probably be extended
quite considerably. According to Manter (1955) there are, to his know-
ledge, 15 known species which parasitize eels; Table 39, however,
only lists 10 of these species.
Relatively little is known about the parasites of Indo-Pacific eels.
For all that, the number of possible parasites in the eels appears rela-
tively high. Of course, the large number of Anguilla species and their
world-wide distribution does contribute to this effect. However, in
comparison with other species of commercial fish, A. angu£lla has a
quite nonnal level of infestation (Reichenbach-Klinke, 1966), even
though one might expect an above-average level in a species whose life
history is spent partly in salt- and partly in freshwater.
From the zoogeographical and ecological viewpoints, parasitological
research on the eel provides some interesting results. Even before
studying homing ability and habitat constancy in eels (see p. 215), the
author had evidence to suggest that yellow eels around Heligoland
could not have migrated there from the coastal regions, but must have
been living in seawater for some time. The trematodes and nematodes
found in the stomachs of these Heligoland eels were of species that
only occur in marine fishes, e.g. the trematode Sterrhurus musculus and
the nematode Contracaecum aduncum (Table 39; Schaperclaus, 1954).
The discovery in eels of parasites which only occur in marine fishes,
shows that the eels can, as it were, take the parasite from its nonnal
surroundings of a particular salt concentration into another with a
different salinity. This is well illustrated in the Baltic; here, the salt
concentration continues to decrease gradually as one travels further
eastwards. The trematode Podocotyle atomon is found further east in
eels than in those animals which nonnally act as the first intennediate
host, e.g. amphipods, isopods and mysidaceae. The same applies in
DISEASES, PARASITES AND IN J URIES 353
Table 39 Parasites which usually only cause slight damage in eels. A. anguilla is the
host in all cases where the host animal is nOt specified. Examples are from
Reichenbach-Klinke (1966), unless otherwise indicated
Type of Pararift Host, affecud body region, locality, number oj
animals aJJlCud, author
Fungi
Saprolegtlia parasitica Very common (example in Table 38)
Flagellates
Trypmwsol1lQ granuloswn Transferred by leeches, propagation by binary fission
in leeches
Sporozoans
Cryptobia markewitshi Transferred by lee<:hes (Nolte, 1915)
Eimeria angui/l(U Mostly in the gut, A . T'Q:ltrata, river estuary (Hanek &
Molnar, 1914)
Myxidium girardi Produces white cysts in the kidneys (~de, 19(6), in
A. rostrata cySts on the skin and Sills (Ghittino et al., 1974)
or attacks the liver and gut (Hanek & Molnar, 1914)
Myxidium em:helypurygii A .japonica, fins, in ponds, 5-11 white cysts on the dorsal or
anal fin (Hoshina, 1952)
Myxidium ma(Jui (Example in Table 38)
Myxidium tYUll(U Gills, gallbladder, gut, ponds near Munich (Bohl, 1968)
MyxoSOl1lQ ( Lentospqra ) dermatohia A. japonica from the sea, CYStS on the whole body surface
(Hoshina, 1952)
PIiJtoplwra angwillarwn A. japonica, musde tissue (Usui, 1974)
Ciliates
Trithodina anguill(U Gills, southern Bohemia, Lausitl (Lom, 1970 ; Nolte, 1915)
Trithodinella epizoou"CiJ
lththyophthin'us multijiliis Skin, gills (e)Cample in Table )8) (see also Mann, 1961)
Trematodes
Te/ogf.llter opis/lwrthis A . dieJJerlbachi, A. australis, New Zealand, freshwater
(Manter, 1955)
SUgodexamenl a"lwilltu A. dief!mlxuhi, A . australis, New Zealand, freshwater
(Manter, 1955)
Coitoc(UQU1l anaspidis A. dieffenbathi, A . australis, New Zealand, freshwater
(Manter, 1955)
Dactilogyrw IJini A.japonica (Manter, 1955)
Gyrodacrylwangui//(U Gills, Lausitl (Nolte, 1975)
Diplostomum spathacewn Metacercaria, worm star, waters round RUgen (Reimer,
'966)
Hapl(frchis angwillarum A. marmorala, Philippines (Manter, 1955)
Surrhurus grandipqrus Marine (Manter. 1955)
Sterrhurus mweulus Gut, North Sea/Heligoland (Tesch, unpublished; Identi-
fication, Dr. Reimer, personal communication)
Deropristis inflata A . a'Wwilla and A. rOSlrata, waters round RUgen (Manter,
1955; Reimer, 1966)
Buuphalus polynwrphw Freshwater (Manter, 1955)
Azygia longa A. ros/rala, freshwater (Manter, 1955)
Azygia anguill(U A . japonica, freshwater (Manter, (1955)
Crepidostomum '(frnlltum A. rostrata, freshwater (Manter, 1955)
Crepidostomum brevittllarum A . rOJ/rata, freshwater (Manter, 1955)
Cmtrovarium lobo/ts A. rO$trafa, freshwater (Manier, 1955)
Hdicometra jascia/a Marine (Manter, 19S5 )
ucithlXhirium rufOfJiride Marine (Manter, 1955)
Lecithcchirium pavidwn Marine (Manter, 1955)
354 THE EE L
Table 39 Continued
Type of Paranu HOSI, affected body region, locality, number of
animals a/fecu d, aurhar
Derogaus van"cw A . roslrala, marine (Manter, 1955)

'Distomum vittllosum' A. "mrOla, marine (Manter, 1955)
Tubulowsicula pinguis A . rOll'orD, marine (Manter, 1955)
Tubufowsicula anguil/tu A. japl)flica, marine (Manier. 1955)
Brachyphal/us cnmarU$ A . anguilla and A . roJlrata, waters round ROgen (Manter,
1955; Reimer, 1<}66)
Optgasu:r angui//i A. nebulosa nebulosa, marine, India (Manter, 1955)
Htmiurus communis Waters round Rugen (Reimer. 1<}66)
Podocotyle a/omon Gut, waters round Rugen, 27% of eels (Reimer, T<}66)
Podocotyle als50ni A. roJlrata, marine (Manter, 1955)
Plo~ioporus OllguialUS Waters round Rugen (Reimer, 1<}66)
SphlW'Qslotml br~ Go<
CrowcrocotCum skrjabini Lake in Hungary (Murai, 197 1)
Paracardicoloides yamagurii A. reinhard/i, blood v~sels, rivers in Australia (Martin,
1974)
Cestodes
Triaenophorul Iricuspioorus T ape worm, gut, Greifswalder Bodden >5% of eels
(Engelbr~ht, 1958)
Tn'aenophorU!i nodulo$U.f
Bothriocephalus c/aviceps A . anguilla and A. roSlra/a, tap!: worm, gUI, Greifswalder
Bodden, 1·5 % of eels, rivers in Spain and Canada, lake in
Hungary (Gandolfi- Hornyold, 1929 ; Bangham & Venard,
1946; Engelbr~ht, 1958)
Diphyllobothrium lalum Larva (final host Man), musculature
Proleoctphalus macrouphalu$ A. anguilla and A . ros/rara, tap!: worm, gu t, Greifswalder
Bodden 8·2% of eels, rivers and coastal wa ters of Spain,
lake in Hungary, river estuary in Canada (Gandolfi-
Homyold, 1929 ; Baer, 1948 ; Engelbr~bt, 1958 ; Doby &
Jar~ka, 1<}66)
Protecocephalus perclU Valencia in Spain (Gandolfi-Hornyold, t929)
Nematodes
C01/l'(lCaecum squali L arva ( ? GUI)
Conlracaecum aduneum Greifswalder Bodden, North Sea near Heligoland
(Schip!:rclaus, 1954; Engelbrecht, 1958, Tesch,
un publish ed, according 10 diagnOSis by Prof. Dr. Osche,
personal comm unication)
Gut, Greifswalder Bodden 5% , Oderhaff 1) %. Finnish
gulfs )8-75% of eels (Schneider, t904; Levander, 1909;
Engelbr~ht, 1958)
Orr/eppina longiuima A. mossambica, South AfriOl: Buffalo River, eels < 20 cm
long (Jubb, 1<}61 )
Spinitu tu$ inemris
Campanarou,gelia campanorougeuu A . mormorata, gut, South Vietnam (Hoa & Khue, 1(68)
Ich/hyobronana l1Iedini Go<
Phi/ometra a/x1omioolis Body cavity, common in the river Elbc (Mann, 1<}62)
Cont4lllanu5 iacusrrn
Camallanus tnmcatus Lake in Hungary (Murai 197 1)
GlHlzi a sp. (Example : Table )8)
P(.Iraquimperia ttnern"ma Gut, Great Britain, Germany, Spain, USSR (Morav~,
'966)
Type of Parasite Host, affected body region, locality, number of

animals affected, author
Paraquimperia adilUm A. rosrrara, gut, USA : Lake Oneida (Moravec, 1'}66)

Paraquimperia anguillae A . bengalemis, gut, India : Poona and Nagpur (Moravec,
''J66)
Filaria anguiliae A. japonica, eye (ishikava, 1916)
Merabronema salv elini A. ros/rala, stomach, gut, river estuary in Canada (Hanek &
Molnar, 1974)
Haplonema adilUm A. roslra/a, gut, river in Canada (Hanek & Molnar, 1974)
Acanthocephalans
Neaechirwrhynchus nllili Gut. Oderhaff 6% , Finnish gulfs 8-25 % of eels (Schneider,
1904; Levander 1909; Engelbrecht, 1958)
Pseudoechinorhynchw davula
Metechinorhynchus sa/moms
Metechinorhynchus c/(l'IJu/a Rivers, Spain (Gandolfi-Hornyold, 1929)
Acanthocephalus anguil/ae Gut, Greifswalder Bodden 0'7% , Oderhatf 60% of eels
(Engelbrecht, 1958)
Acanllwcephalus ludi Gm, Oderhaff 19% , Finnish gulfs 25-54% , England, 75%
of eels (Schneider, 1904; Levander, 1909; Engelbrecht,
1958; Mishra & Chubb, 1969)
Acanthocepha/us garoi A.japonica
PomphorhynchuJ /aevis Gut, Baltic 61 % Greifswalder Bodden 6% of eels (Pohl,
1956; Manter, 1955; Engelbrecht, 1958)
Corynosoma semerme
Tanaorhamphus ambiguIIS A. rostra/a (Manter, 1955)
Crustacea
Ergasi/w gibbus Gills, egg clusters up to 6 mm in size
Ergasilus cek"is A. rostrafa, gills, river in Canada (Hanek & Molnar, 1974)
Argu/us foliaceus (Example in Table 38)
Lernaea cyprinacea Skin, mouth cavity, A. anguilla, A . japonica (Usui, (974)
the case of Deropristis inflaca, Hemiurus communis, Podocoty le atomon and

Brachyphallus crenatus (Reimer, 1966). Such instances are referred to
as an 'east drift'. They are explained as follows: infection takes place
when the eel is still young, i.e. in a migratory stage, and passing through
areas of higher salt concentration. Migration continues into regions
with a lower salt concentration, i.e. eastwards, thus transporting the
parasites into these regions as well.
There is also the reverse situation, i.e. eels caught in the sea which
have freshwater parasites. Such instances have been reported from
waters round the isle of Rugen (Reimer, 1966), and involve the
trematode Plagioporus sp. and also, it seems, the cestodes
Bothn'ocephalus claviceps and Proceocephalus macrocephalus. A possible
explanation for such cases is that the eels could have been caught in the
356 THE EEL
narrow coastal region where either the salt concentration was very low
or the transition from freshwater to saltwater was very abrupt, so that
even short forays or the cbange of habitat in spring or autunm (see p. 215)
would have made it possible for typically freshwater parasites [0 be
found in fishes living in the sea.
Extrapolations from parasitological studies which suggest that the
Pacific is the centre of origin for all eel species must be viewed critically
(Manter, 1955; see also p. 131). Such conclusions are based on the
following argument: of the 20 trematode species found in the Atlantic
eels, 14 are also found in other species of marine fish and 5 in other
freshwater fishes. Only one marine species, Deropristis injlata,
parasitizes the eel alone. But out of 9 species in the Indo-Pacific,
8 species and 3 genera are specific to the eel; 5 or 6 of these come from
freshwaters and relatively few (3 or 4) from the sea. Therefore, far more
Anguilla-specific trematodes have been able to develop in the Indo-
Pacific than in the Atlantic, thus suggesting the existence of eels in the
Pacific region for a longer period of time.
The idea that Atlantic eels have a greater number of marine
trematodes because their journey through the sea is longer than that
of any Indo-Pacific species, also raises a number of problems. The main
objection to this theory is that most parasites are first acquired after
metamorphosis, i.e. during coastal and freshwater periods of the eel's
life history. Besides, how can the parasite's life cycle be completed if its
intermediate and final hosts are not present in the regions far out at sea
through which the eel larvae must pass on their journeys to the coasts?
Nevertheless, parasitological-zoogeographical investigations result in a
great deal of valuable information, and the opportunity to complete our
knowledge of the biology of the eel is, as yet, far from exhausted.
7.3 Teratological and other defects

In the eel, as with other species of fish, one comes across morphological
changes in various organs the causes of which cannot always be ex-
plained fully. Often, these changes may be the result of a disease which
the eel has survived, or they may be due to the after-effects of injuries.
The eel is a particularly resistant fish and often recovers from injuries
which would be fatal in other species. Thus, defonnities of the body and
organs are seen relatively frequently in eels. In other instances, how-
ever, the abnormality is clearly of a different nature, such as a deve-
lopmental disturbance or an hereditary anomaly.
The following are instances of changes in internal or sensory organs
which may have been brought about by pathological conditions: the
formation of kidney rumours (Plehn, 1924; Scheer, 1934), ovarian
cysts (Eiclher, 1935), cyst-like fibromyoma in the body cavity
(Schiiperclaus, 1954; Mattheis, 1964b), mesenterial fibrosarcoma
(Wolff, 1912; Plehn, 1924; Mattheis, 1964b), inflammation of the
swimbladder (Flemming, 1954), the formation of a stone in the body
cavity (Reichenbach-Klinke, 1956; Tesch, 1958), macrophthalmia or
exophthalmia (Mercier & Poisson, 1927; Vladykov, 1973). Unfortu-
nately, in most of these cases only the pathological-morphological
condition has been described and nothing is said or known about the
cause of the disease.
As far as one can see from the literature cited, kidney ttunours are
fairly common in eels. In many cases the growths in the connective
tissue of the kidneys were so pronounced that they completely dis-
placed the normal kidney tissue. Ovarian cysts are similar in character.
With regard to the inflanunation of the swim bladder, the latter was
abnormally large and, instead of containing gas, was filled with spongy,
white tissue and fluid. The majority of cyst-like sarcomas appear to be
benign rumours and, in most cases, the causes were probably localized
processes, such as injuries and hyper-regeneration. It is interesting to
note that the neoplasms made up as much as 34 to 49% of the body
weight and that the nutritional condition of the affected eel was generally
normal (Mattheis, 1964b).
It is difficult to explain the two instances of stones (calcium phos-
phate) found in the body cavity; one of these stones was 13"5 mm in
diameter. the other 20 mm. I t is possible that the stone had developed
from an ovarian cyst. Reichenbach-Klinke (1956) talks of a 'simple
stone formation on the inner wall of the body cavity'.
Exophthalmia can be caused by various diseases.
One of the best known deformities in the eel is curvature and distor-
tion of the backbone. Schrader (1930) studied II such cases. A further
five instances of similar abnonnalities have been described by other
authors (Scheuring, 1927; Schaperclaus, 1954; Stolk, 1956; Kochenko,
1958; Wunder, 1969). The reason why such a large number of cases
are known in the eel is probably due to the fact that a crooked backbone
is much more noticeable in the eel than it is in a less elongate fish.
As is shown in Fig. I27A, the body also becomes crooked because of
the malfonned backbone, and the individual vertebrae are also deformed
(Fig. I27A). Wunder's (1968) explanation for this type of malformation
is that, as a result of injury) the young eel suffers a retardation in growth
of the skin. The nonnal number of vertebrae develop and the vertebral
THE EEL
colwnn is forced to fit into the available space through the formation
of a large nmnber of curves.
Shortening of the backbone which is much less common, has different
characteristics (Wunder, 1968). In certain regions of the body the
vertebrae are squeezed together and shortened (Fig. 127C), but the
body as a whole remains straight. In the specimen described by
Wunder, the total number of vertebrae, as compared to that in normal
eels, was reduced from 114 to 110. The body appears much deepened
(Fig. 127C) and is reminiscent of the larval body form (Fig. 36). This
type of malformation also reminds one of the compressed vertebrae
and deepened body sometimes seen in pond raised carp. Therefore, in
eels, as in the carp, the cause is generally thought to be attributable to
hereditary factors and to some disturbance in the differentiation of the
vertebral colwnn at a very early stage of ontogeny.
• I I
~ ~M
-=.. . . . . .: .: :- .;: : :. . ... . . ..-_.
Fig. 127 Eels with abnormal spinal columns. A, eel with curvature and distortion of
the spinal column; three vertebrae, left: normal vertebra, centre: vertebra
deformed as a result of pressure from one side and traction on the other,
right: vertebra showing even greater distortion. B, normal eel. C, eel with
shortened spinal column; section ·of the vertebral column (after Wunder,
1968, 1969)·
Reports of colour anomalies are not infrequent (e.g. Pavesi, 1894;
Neubaur, 1924; Rumphorst, 1929; Thumann, 1953; Jones & Pantulu,
1954; Fisch u. Fang, 8, 96, 1967). The author has also come across
several such cases. The anomalies range from almost white animals
(albinism) (Walter, 1910), through yellow (xanthochromatism) and
golden-yellow specimens, to eels which are flecked with black or are
marbled in appearance. Since there is a melanophore or chromatophore
(i.e. pigment) deficiency in eels that are lighter than normal in colour,
one might assume the cause to be some genetic anomaly, possibly
brought about by mutation. However, Thumann (1953) describes four
xanthochromic eels which were kept for a long time in a well-lit aqua-
rium. After 8 to 10 weeks these animals assumed the coloration of
normal eels, and did not return to their previous colour even when kept
in the dark. It thus appears that the xanthochromatism was only
temporary (see also Ehrenbaum, 1930), and it is doubtful whether one
can regard such anomalies in colour as genetically determined. It should
be noted that the term 'yellow eel' is used for a normal growth stage
in eels and that abnormally coloured yellow eels are quite different.
The same applies to the term 'golden eel' (see p. 18) which is sometimes
used in England instead of 'yellow eel'.
Another anomaly, the cause of which has never been established,
is a double liver (see p. 34). It is possible that injury or epigenetic factors
during early development may be partly responsible; the same applies
to malformation of the caudal fin (Gandolfi-Homyold, 1918a).
The very extensive deformities resulting from injuries at relatively
late stages in development range from cases where the cause is unknown
to very obvious wounds. A severe constriction of the stomach and its
transformation into a fluid-filled bubble (Wunder, 1967) are among the
malformations very probably attributable to mechanical wounds. Such
stomach wounds are thought to result from swallowing a fish hook,
which tears the stomach fundus and leads to a functionless protube-
rance or stomach cyst being formed. The cardiac and pyloric sections
fuse to form a shortened, but functional stomach. Steen and Berg, 1966
(see p. 38) report a case of haemoglobin deficiency which was also
probably secondarily caused by a fish hook injury. Such injury may
stimulate hypertrophy of the corpuscles of Stannius (Lopez &
Fontaine, 1967). Eels that have swallowed hooks are, it is sad to say,
frequently caught, and it cannot be over-emphasized that such cases
can and should be reduced to a minimum by using secure knots and
strong leaders. While on an experimental fishing trip using electrofishing
methods in the middle reaches of the river Weser, the author encounter-
THE EEL
Fig. 128 Eel from the central Weser with part of its caudal region missing and a fish
hook still embedded in its mouth; see text (Photograph: T esch).
ed an extremely malfonned animal (Fig. 128). The photograph shows

that the synthetic leader, protruding from the mouth, had produced
a wound in the comer of the mouth which was already healed, and
contained a large pigmented growth.
The eel illustrated had, in addition, suffered a second injury, a
consequence of industrialization: it appears that turbine blades had cut
off a considerable part of its caudal region. It is surprising that this eel
managed to survive. The ratio of its pre-anal to post-anal length
(Fig. 34), nonnally 1 :1'4, was 1 :0'8; in other words it had lost almost
half its post-anal body. Mechanisms involved in somatic regeneration
in cases such as this have been examined in the caudal neurosecretory
system (see p. 63) of a Japanese eel (Katsutoshi, 19653, b).
The reports, investigations and complaints about rurbine injuries
to eels are numerous (e.g. Lundbeck, 1927; Kroezus, 1954; v. Raben,
1955, 1958; Butschek & Hofbauer, 1956; Schiemenz, 1958, 1960a;
Muller, 1962). Obviously, the long-bodied eel, passing through a turbine
in the course of its migrations, suffers most frequently from such
injuries and may even be cut to pieces. Kaplan turbines in Germany
are thought to injure approximately 18% of eels that swim through
them. In southern Sweden investigations into damage from Kaplan
turbines produced figures as high as 40 to 91 % (Svensson, ftom Muller,
1962). The damage depends on the flow of water and the position of the
turbines as well as on the length of the eel. The speed of the blades is
also important. A pennissible speed of 10·8 m/sec has been calculated
(v. Raben, 1955). A sure means of protection can only be obtained
from grids placed over the turbine openings; grids with a 2 em gap
between the bars have become compulsory as a result of various fishing
laws. Electrical installations are now used to tum fish away from the
turbines, a method that has proved more economical than the use of
grids which reduce the flow of water.
DISEASES, PARASITES AND INJURIES
Finally, sewage and other pollutants should be mentioned, the

effects of which result mostly in the death of the fishes. If individuals
are able to survive the actual effect of the pollution - and this is more
likely in eels than in other fishes - then they usually recover relatively
quickly. Eels react to lack of oxygen and to toxic substances in the same
way as do other fishes - they come to the surface of the water. In addi-
tion, eels near land struggle to the bank and try to save themselves by
leaving the water. In some cases they survive if, as may happen in a
river, the pollution is transitory.
Permanent damage from pollution seldom occurs, unless the
pollutants produce long-term metabolic effects. A noteworthy example
of this is the reduced water absorption from the gut and altered brancial
enzyme content produced by DDT (Janicki & Kinter, 1971; Guarino
et ai., 1971). Contamination by, and the accumulation of radioactive
caesium have been studied by Reynier et al. (1970), and the uptake and
excretion of chromium have been studied by von Fouquier et al. ( 1973).
While chromium levels in the intestine and blood decreased after
45 days, it remained constant in the liver and was seen to accumulate
in the kidneys. DDD and DDT were found in A. dieffenbachi and
A. austraHs, but not in most of the other species of fishes examined
by Solly and Harrison (1972). Poisonous methyl mercury was detected
in A. rostrata (Freeman & Horn, 1973). Mercuric chloride accumulated
in the gills of A. anguilla after short-term exposure, but long-term
exposure resulted in an accumulation in the kidneys, spleen, liver and
brain as well (Bouquegneau, 1973a). With high concentrations of
mercuric chloride ( I ppm) the mercury had lethal effects on ion regula-
tion; at lesser concentrations adaptation sometimes occurred (Bou-
quegneau, 1973b). Blood counts and electrocardiograms both showed
changes under the influence of waste waters from the production of
aluminium oxide; heartbeat frequency and the number of erythrocytes
showed considerable decreases (Halsband & Halsband, 1971 ). Reference
is made on p. 345 to the possibility of certain tumours developing as a
consequence of contact with chemicals.
8
Trade and Processing
8. I International trade
With the exception of salmon and trout, hardly any species of food
fish has such a world-wide market, both for the deep-frozen and the
live product, as does the eel. It cenainly leads the market for live edible
fishes (see p. 241). The earliest reports of live transport from continent
to continent were in 1923 (Fischerbore, 17, 64-65, 1925. and 21, 357,
1929) when live eels were sent from Canada to Germany and Holland.
The quantity of eels involved was only small- about 100 kg - in
comparison with the export of around 7000 kg from Canada to the
USA. After 1949, Canada's principal customers were, in addition to the
USA, West Germany and the Benelux countries. However, even
the largest exports to West Germany were very insignificant; in 1957,
for example, they amounted to 300 kg (Eales, 1968), and in the last few
years, even exports to the USA have not been much higher.
The first consignments of Canadian eels aroused a great deal of
opposition since at that time, unlike today, the German market was not
so dependent on imports and there was a fear of the consequences of
competition from abroad. West German eel fisheries only provide
25 % of the country's total consumption (Koops, 1 967a). Table 40
shows the extent of Germany's imports from other countries. Deep-
frozen eels make up the greater bulk of these imports, though a small
proportion of eels is still transported live. I t is worth noting that even
eastern Asiatic countries send eels to Germany. By far the greatest pro-
.62
TRADE AND PROCESSING 363
Table 40 West Germany's eel imports (in tonnes)
(after Koops, 1967a)
,"'" ,",
Counfr), of origin Year
,..
1955
Denmark ISlI 1588 10 40
Holland '79 , 60
Sweden
Poland ''''" " 7
'"
' 43
,' ,
"
Ireland
Turkey
Great Britain
'7
" 80
70
F~tt
G,= , , "",
Austria
Spain , ,,
Switzerland
,
,,' ",,
Yugoslavia
..
Canada ' 97
, ,,'",
USA
Japan
Peoples Republic of China
New Zealand
Total U 71 3615 3119
portion of imported eels comes from European countries, Denmark in

particular. Unfortunately, Table 40 does not give one a completely
accurate picture because many imports come via a third country, e.g.
Holland or England. Imports from Poland, France and the Mediterra-
nean coasts appear to have increased quite considerably in the last few
years. In 1965 eel importS to Germany were of the order of 30 million
DM. Total imports from Poland rose from 20 tonnes of eels in 1952 to
129 tonnes in 1955, 2II tonnes in 1960 and 362 tonnes in 1964 (Koops,
1967 a).
Although Gennany is the principal single customer for eel imports,
a corresponding survey for the rest of Europe would show an even
larger trade. For example, isolated reports claim that 4' 5 tonnes of
A. australis were exported to Europe in 1965; New Zealand sent
100 tonnes (Desch. Fisch. Zrg., 13, 280, 19(7). Further details on
exports from New Zealand can be found in Teklenberg (1972). In the
three Egyptian lakes Edku, Menzaleh and Birollos near the Mediter-
ranean coast, a Dutch firm caught 25 tonnes of eels in 1962/3,58 tonnes
in 1963/4 and 90 tonnes in 1c)65/6, and shipped them alive to Europe
(Inform. Fischw. Ausland., 16, H. 6, 28, 1966).
THE EEL
8.2 Prices
It is clear that the upward trend in the cost of living does not exclude
the price of eels. Of course, average prices fluctuate from year to year
in relation to the number of eels that are caught, and clearly reflect
supply and demand. Between 1960 and 1966 the price per kilogranune
of eels at the marine fish market in Kie! showed the following trend:
6·03-6·60-HI-7·13-9·o-8·84-8·50 DM. If one compares the prices
(in cents per pound) of eels in Canada for every fifth year from 1920
to 1965, the result is as follows (Eales, 1968) : 8'9-8'9-8'7-6'4-4'9-
9'6-8'4-14'8-18'7-31'7. There is a sharp increase in prices after
1945 and a slight decrease in the 1930S.
Prices differ for eels of different sizes or 'categories', just as they do
for eels of different quality j variations in price are also seen between
broad- and narrow-headed eels, and between yellow and silver eels.
In Germany there is a greater demand for narrow-headed and silver eels
than for broad-headed eels, and the former thus realize higher prices.
From now onwards, EEe countries will be using the following size
groups for sorting eels: eels of category I are more than ISo g in weight;
category II eels weigh between 100 and 180 gj category III eels are less
than 100 g in weight. Other gradings, however, are still in use, e.g. in
Germany: 'Reisenaale' ('huge' eels), 'Gro/k Aale' ('large' eels), grades I
to III, or by weight (in 'Pfund', i.e, 500 g) per score of individuals, The
latter system originated in Denmark and dealers who import eels from
there are particularly familiar with it, Large eels, for example, would
probably weigh 30 Pfund per score, i.e. each eel would weigh 750 g;
small eels weigh 4 Pfund per score, i,e, about 100 g each, and so on. In
Germany the measures used for smoked eels are different from those
used for fresh or living eels, viz. I ('Riesenaale'), over 500 g per eel; II,
375 to 499 gj III, 250 to 374; IV, 175 to 249 gj V, 125 to 174 g per eel;
'Bund3ale' are smoked eels which are bound together with bast into
bundles weighing 100 g or even 125 g (Struck, 1965).
In 1967, 'GroBe' eels cost, on average, 9'24 OM per kg, class I eels
cost 10'56 DM, class II 7,80 OM and class III 4'72 DM per kg (see
Table 41 ). 'Riesen' eels were not available; eels weighing more than
100 g are rare in north Germany. Eels between 250 and 500 g appear to
be the most expensive; below and above that weight range they are
cheaper. The practice of selling smaller individuals at lower prices is
also found in the marketing of other species of fish. Extremely large
eels likewise fail to reach the top prices because they are awkward to
handle and prepare, or because they exceed the norm for curing.
TRADE AND PRO C ESS ING 365
Table 41 Average monthly price (in DM per kg) of eels sold in 1962 and 1963 (after
Herrmann, 1964), and in 1967 (prices arranged according to t he category
of eel, see p. 364) at the marine fish market in Kiel, as well as the average
monthly price (in U.S. dollars per kg) of eels sold at [he wholesale market
in T okyo (after Brown, 1969, from Fritzsche, [970)
KIEL TOKYO
OM/kg OM/kg 1967 Dollan lkg

,,., ,,., GrQJse Aale Cal . I /I 11I 19I>4- 1<}66
(' Large' ulJ )
January 804 1'58

February 8·" 10·80 1'57
March 7·82 8'63 900 11 '90 9"56 5"46 1'73
April 8'08 6'89 9'26 11 ' 10 ,·60 4'46 2'02
M.y
June
July
6'98
7'26
8"26
6'54
7'26
10,06
10' 16
11 '00
11'4°
11·86
,.,.
860
8·66
,·66
,·90
1'92
1'7°
7'65 10'3° 5"40 1'7 1
August 7'74 7·61 9'76 10'5° 8·06 5' 56 1'55
,."
September 6'56 ,·06 9'26 6·00 ,60 1"48
,90
,."
October 6·82 6'59 7'40 8·", 6'46
November 7'.01 8"16 9'30 7' SO 4'70 1'46
December 7'7 2 7'97 10'00 1'78 4' SO "48
The price of Japanese eels varies according to their origin, i.e. whether
they come from lakes and rivers or from fish farm ponds. The price
per kg from 1955 to 1958 was 3'0 DM, with pond eels costing about
0'40 DM more than eels from natural waters (Koops, 1966b). 'The
higher price of pond eels is clearly due to the fact that the prices of eels
from river and lake fisheries are subject to greater seasonal fluctuations.'
In addition, the pond eel has more fat than do eels from lakes and rivers
(see below), though in Japan this may not necessarily be a sign of better
quality. In West Germany too, the eel 's origin has a certain influence on
the price; eels from the Baltic are considered to be of the highest
quality. Osmotic effects resulting from lower salinity of the Baltic may
be a decisive factor in this matter of quality (see p. 22). A similar situation
exists with regard to flat fishes (Tesch, 1956).
There are other factors besides quality which underlie fluctuations
over the year (Table 41 ). For example, in Germany a kilogramme of eels
costs very little in April and May, and even less in September and
October. These are the months when the greatest number of eels are
caught (see p. 249), the time when, in fact, the eels are changing from
winter to swruner quarters and vice versa (see p. 216) and the spawning
migration is just beginning (see p. 220) . Here, then, one can talk of an
'ecology of prices' as contrasted to the economy, which is controlled
by non-biological, purely economical requirements, e.g. monetary
THE EEL
value, competition from other foods, imports etc. Of course, the market
for eels is less affected by these economically detennined fluctuations
in price than is the market for other species of fishes. Live storage of
eels allows the 'season' to be extended for a relatively long time (see
p. 340). In addition, the high price of eels means that it is economically
worthwhile to store them deep frozen.
In japan there is a peak price period which lasts from March to june
and reaches a maximum between April and May; it is brought about
by the inadequate supply of eels during the spring months. In the
succeeding sununer months the quantity of eels available for sale
reaches its annual maximum and prices show a corresponding decline.
It is well known that the quantity of eels produced in japan is deter-
mined by pond culture (see p. 243). With this system one is continually
in a position to supply eels at any time during the sununer. The river
and lake fisheries, with their maximum catches in spring and autumn,
therefore fall sharply behind.
In comparison with German eel prices, those in japan were not
much lower in the years covered by Table 42. Indeed, if one takes into
account the fact that, on the German grading system, the majority
of eels produced in japan are in category III, and if one compares
German prices for category III eels with average japanese prices,
eels in japan are actually more expensive. Between 1955 and 1958
eel prices in Japan were half as high again as they were in Germany
(Koops, 1966b; see above). In 1965, Canadian eel prices were only a
third of those in Germany; before then, prices were even lower. In
western European countries other than Germany, prices were also
considerably lower.
In Germany and Japan the eel is considered a luxury food; it is a
fairly CODUnon food item in Denmark, Holland, England and Italy,
and in Spain as well, if one includes the Spanish people's predilection
for glass eels. In 1918 the English consumed almost 7000 tonnes, of
which only 1000 tonnes came from British waters (Koops, 1967a).
8.3 Quality
Although market prices are one index of quality, that parameter IS
better judged on the results of chemical analyses and food tests. For
the eel, as with most species of fish, the final criterion of quality is fat
content. Since more than 80% of eels in Germany are smoked (Struck,
1965), a high fat content, which is desirable for smoke curing, naturally
adds to the value of the fish.
TRADE AND PROCESSING
Table 42 shows fat content as based on the results of several studies.

Eels less than 30 em long contain relatively small amounts of fat,
usually little more than 5% ; water content is proportionately high.
Narrow-headed eels contain larger amounts of fat than do broad-
headed eels. For this reason, narrow-headed eels are used in smoking
if silver eels, which have an even higher fat content, are not available.
Unfortunately, the greater fat content of silver eels does not emerge
clearly from the figures given in Table 42. Eels from the Baltic are
particularly fatty, a fact which emerged from a study wherein river and
pond eels were compared (Table 42; Meyer-Waarden & Koops, 1968),
and is also illustrated in a comparison of eels from Wismar Bay in the
Baltic, and the inland lake of Dassow. Despite this difference in fat
content, eels from the inland lake were relatively heavier than those
from the Baltic, a finding that indicates a higher water content in the
freshwater individuals. Table 19 also shows that, on the whole, coastal
eels weigh less than inland eels. Pond raised eels are the heaviest of all
(Meyer-Waarden & Koops, 1968j Koops, 1971 ); their fat content,
however, is not above average in either Japanese or European eels
The tail region and the areas directly beneath the skin have the
highest fat content of all the body (Table 42), the viscera and the area
around the backbone the least amount of fat. Smoked eels, from which
the viscera have, of course, been removed, contain very little water and
are rich in protein, though they are not above average in fat content.
However, compared with other species of smoked fish, the eel contains
the most fat, an important reason for its high culinary value.
The fatty acid content of the fat is as follows: total fatty acids, 86'5% j
unsaturated fatty acids, 66'2% ; saturated fatty acids, 20'3%; glycerine,
I 1"4 %; unsaponified substances (cholesterol derivatives), 0'7 % , and
lecithin, 2·~ % . The iodine count of the fat is 97'4 in narrow-headed
eels and 105·8 in broad-headed eels (in other species of fish it is usually
well above 100, reaching a maximum of 161 in salmon) ; from this
one can conclude that eel fat is not composed of the glycerides of
unsaturated fatty acids to the same extent as it is in other species of
fish. The difference between narrow- and broad-headed eels is not
particularly marked and can clearly be attributed to the different
feeding habits (see p. 177) of the two forms (Wiehr, 1932). Data on the
effects of different foods on fatty acid composition are available for the
Japanese eel (Ando, 1968). Dabrowski et al. (1970) have compared the
cholesterol content of muscle and fat in the European eel with that
in 10 other species of freshwater fishes. Reichwald and Menzies (1973)
and Menzies and Reichwald (1973) have described the fatty acid compo-
368 THE EEL
sition of muscle lipids in the eel and compared it with that of freshwater
species. These authors have also described the same features in fresh
and smoked eels as compared with sturgeon, herring and spur-dog
( Squalus acancln'as) ; they found higher than average levels in the eel
and, in comparison with the seven other species studied, by far the
highest level of eicosenoic acids.
There is an improvement both in quality and taste as a result of
smoke-curing, and of CQurse such treatment also serves to preserve
the fish (Gehring & Wiinsche, 1965). The processes involved in smoking
have been described on many occasions (e.g. Breitenstein, 1954;
Visurgis, 1954; Struck, 1965 j Meike, 1970). Smoked eels are also tinned
for export (Lantz, 1966). Eels may be prepared in a wide variety of
ways, many of which include preservatives th~t prevent spoiling for a
limited period of time (Sparrenberg, 1934; Struck, 1965). Methods of
preservation used in New Zealand have been described by Teklenberg
([97 2 ) .
Table 42 Chemical composition of the body tissues. Data arranged according (0 the region of the body analysed, localities from which
the eels came, sizes of the eels, and their different stages of development
a" Different regions of the body, various sizes of eel (a ll the animals were from the Slelliner Hatf), and of smoked eels
Part of body Weight of Fo< Water Protei./ A,h Carbohydrate Remarks AUlhor
each indi- (%) ('Yo) ('Yo) (%) (%)
vidual (g)
H ead section 7" 18"3 6, ", 16"1 ,., after Meyer-Waarden, '943
Anal section 7" zS"9 S6"1 IS"1 '3 after Meyer-Waarden, '943
Tail section 7" 31"2 S'"1 88 '·9 after Meyer-Waarden, '943
Viscera 7" '9"3 63"7 14"1 ,., after Meyer-Waarden, '943
...
Head section 3' 12"4 67"4 I7 "S after Meyer-Waarden, '943 '">
Anal section 3' 13"4 67"6 10"4 "'7 after Meyer-Waarden, '943
Tail section ,6", ,., after M e yer-Waarden, '943 m
3' 14"6 6S"1 ">
Viscera 3' 9·' 73"3 '3"' after M eyer-Waarden , '943
Region around the " Z
backbone 29"1 S,"8 after M eyer-Waarden, '943
Adjoining region '"
7" 30 "S 62"9 after M eyer-Waarden, '943
Region beneath the skin 7" 40"6 4S"6 after Meyer-Waarden, '943 o
."'"
Sk.i;t 7" 9·7 S6"4 after M eyer-Waarden, 1943 n
26"4 S2"7 ,87 ' ·4 ,·8 smoked aftCf Soud, Fachmann & Kraut from ~
Meyer-Waarden, I<)6s ~
Z
-'"
b" Different d~vclopmental stages and forms
Developmental stage Weight Fal Dry malter Author

'"
or form (%) ('Yo)
Eel fry (elvers) 168mg 3·' 22"2 after Reuss & Weiland, 19'3, from Wiehr, '932
Eel fry (elvers) 4·' 21 "1 after Konig & Spliugerber, from Wiehr, 1932
Aalmont~e (elve rs) ,., 20"1 after Konig & Splitlgcrbcr, from Wichr, 1932
Young eel '48 '·4 19"6 after Wiehr, '932
Narrow-headed eel 27"3 40 "9 after Wiehr, 1932
Brood-headed eel 11 "9 547 after Wiehr, 1932 w
River eel 28"4 after Konig & Spliugerber, from Wiehr, 1932
-8'
Table 42 ConhnuuJ w
c, Differenl localities Cl
I...oaAlity '-"<,' Wright F" Remark, Al<thor
(=l (., (%)
Wismar Bay ,., narrow-headed male after Meyer-Waarden, 196~

" ,0" 4·' narrow-headed male afler Meyer-Waarden, 196~
"'9 narrow-headed male afler Meyer-Waarden, 196~
45 11 '1 narrow-headed male afler Meyer-Waarden, 196~
" ,0 28' 1 narrow-headed male after Meyer-Waarden, 196~
29'0 narrow-headed male after Meyer-Waarden, 1<)6~
37 90 27"8 narrow-headed male after Meyer-Waarden, 1965
"" "
,9 no 28·8 narrow-hC1ldc:d male after Meyer-Waarden, 1<)65
Lake Dassow (inland) ,6 6·' narrow-headed male after Meyer-Waarden, 1965
" 4·' narrow -headed male after Meyer-Waarden, 1965
"'9 narrow-headed male after Meyer-Waarden, 1965
9·' narrow-headed male afl er Meyer-Waarden, 1<)65
",0 ...0:
" 60 26·8 narrow-headed male afler Meyer-Waarden, 1965
'0 10'7 narrow-headed male aft er M eyer-Waarden, 1<)65 '"
37 <0, 2)'1 narrow-headed male after 'M eyer- Waarden, 1965
""
,9 n, 14'1 narrow-headed male afl(~r Meyer-Waarden, 1<)65
...'"'"
Greifswalder Bodden '9 4·' narrow-headed male after M eyer-Waarden, 1965
" narrow-headed male after Meyer-Waarden, 1<)65
narrow-headed male afler Meyer-Waardcn, 1965
"'9 ",0
,.,,.,
TO'7 narrow-headed male after Meyer-Waarden, 1965
" 37
" ,., narrow-headed male after Meyer-Waarden, 1<)65
,., narrow-headed male after M eyer-Waardcn, 1<)65
37 narrow-hC1lded male after M eyer-Waarden, 1<)65
"" "
39 narrow-hC1ldcd male after Meyer-Waarden, 1965
Baltic ) 1'0 from freshwcighl of silver eel fillets after Meyer-Waarden & Koops, 1968
River eels, Mosel 21'9 from freshweight of silver eel fillets after Mcycr-Waardcn & Koops, t968
Pond eels, Elb!: 27'0 from freshweight of silver eel fillets afte r Meyer-Waarden & Koops, 1<)68
Pond eels, Mosel 24'1
and 26·6 from fresh weight of silver eel fillets after Meyer-Waarden & Koops, t968
Pond eels, Japan 2)'8
and 24"3 from freshweight of silver eel fillets after M eyer-Waarden & Koops, 1968
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Index
Aalbrut, 18 diet, 182- 4

Acetylcholine synthesis, 37 diseases, 343-56
n-Acetylhistidine,37 egg size, 51
ACTH environmental influence on sex,
See Cortisol 53-4,57
Adenohypophysis, 42, 59 head width, 5
Adenosine triphosphatase, 25, 32 homing behaviour, 2 15-20
Adrenocortical homologue, 42, 64 migration, ' 33-240 passim
Adrenocortical steroids, 32 muscles, 21-2
Adriatic pigmentation, 14-19
eel migration, 86-7 pollution damage, 36 1
Aeromonas 'iqui/adem, 348 rhodopsin, 72
Aeromonas ( Pseudomonas ) puncwla, 346 , scales, 12-3, 186-7
347 sexual matu rit y, 50-1
Africa sound perception, 79
eel migration, 104, 111 - 14 spawning and larval forms, 83-102
Albinism, 359 sperm morphology, 50
Albumins, 39-40, 88 teeth, 20
Alimentary tract vertebral column, 8
length, 28-9 yield, 256-9
water and ion transport, 31 A. auslra/is, t 18-2 1
Alkaline phosphatase, 32 diet, 180
Ambisexuality, 49 growth rate, 189-90
American eel pollution damage, 361
See A. ros/rata sex and distance C(lvcred in migration,
Anemia, 38, 359 "3
Angling, 304-15 weight, 208
Anguilla 82-3 yields, 253
Anguilla ancestralis, 122-9 A. bicolor, 111-4, 122-9
A. anguilla A. borneensis, 122-9
blood albumins, 40 A . celebesensis, 122-9
chromosomes, 52 A. dieffenbachi, 118-21
continen t a l distribution, 102 - 10 diet, 176
423
INDEX
growth rate, 188-90, 198 growth rate, 190, 197

migration, sex and distance, 223 hide-outs, 2 I I
pollution damage, 361 homing behaviour, 215, 217
weight, 208 muscles, 22
yield, 253, 300 pectoral girdle, 6
A. interioris, 122-9 skull,4
A. japonica spawning grounds and larval forms,
chromosomes, 52 83-r02
diseases, 348, 351 subterranean migration, 234-5
egg size, 51 teeth,20
growth rate, :90 vertebral column, 7
head width, 5 Anguille jaune, 18
pancreas, 33 Anguille verte, 18
pee[orallins, 9 AnguilJidae, 82-3
pigment, 14 See Eel for general references
scales, 12 Anguilliformes, 83
sexual maturity, 51 Arginine vasotocin, 62
spawning grounds, t 14-8 Ascent
sperm morphology, 50 glass eels, 139-45
temperature effects on ascent, 142 young eels, 154-69
tidal effects on ascent, 143 Atlantic spawning grounds, 86-91
vertebral column, 6 Atrium, 35
A . marmorata, 111-4, 122-9 Auckland Island
distribution, 116-7 eel migration. 120
habitat and size, 213 Azores,
weight, 208 eel migration, 101
A. megastoma, 122-9
A. mossambica, III-4 Baits, fishing, 311-3
growth rate, 190, 205 Balaton, Lake
weight, 208 eel growth rate, 196
A. nebu/osa, 122-9 eel migration, 105
migration distance, 155 eel stocking, 270
growth rate, 188 Baltic Sea
scales, 12-13 eel migration, 135-6, 145, 232, 235-8
weight, 204 eel yield fluctuations, 244-6
A. Ilebulosa labiata, IIJ-4 Barriers, light, 322
weight, 208 Belorussian Lakes
A. nehulosa nebulosa eel yields, 269-70
growth rate, 198 Belts
weight, 208 fisheries in. 241
A. obscura, 122-9 Bermuda
A. reinhardti, 120, 12Z-9 eel migration, 95, 105-6
A. rostrata Bile salts, 35
blood albumins, 40 Biomass, 252-3
chromosomes, 40, 52 Black Sea
continental distribution, 102-10 eel migration, 86, 104
d iurnal effects on migration, 160 Blankaal, 18
egg size, 51 Blood, 38-41
food and body size, 176 albumins, 39-40
INDEX
flow eta, 59
effect of vasoactive agents, 37 juxtaglomerular, 42, 66
pressure Keys-Willmer,25
effects of vasoactive agents, 37 Cerebellum, 66
toxicity, 41 Cestodes, 355
Bones, intermuscular, 8 Chatham Island
Bonin Island eel migration, no
eel migration, 116 China Sea
Borneo, 122 eel migration, 114
Brain morphology, 66-8 Cholesterol,4 t
Branchionephritis, 351 Cholesterol derivatives, 367
Branchiostegal apparatus, 3 Chondrococcus coJumnaris, 35 I
Broadheaded eel Chromosomes, 40, 52, 100
classification, 4-5 Cilia, 74
food, 177-9 Civelles, 18
growth, 193 Colour vision, 71
heart weight, 36 Comacchio
price, 364 eel fisheries, 242, 295
weight, 205 eel growth rate, 196
Bronze eel Cones, 71
species differentiation, 18, 101 Conger conger
Bulbus arteriosus, 36 eyes, 72
Bulgarian rivers muscles, 22
eel migration, 104 vertebral column, 8
Bundgarn, 284 Conger marginatus, 22
Butterfly gates, 293 Conn, Lough
eel growth rate, 194-5
Calcium metabolism, 65 Continental shelf
Canada eel migration, 133
eel yields, 248-9 Continental slope,
Canary Islands eel migration, 114, 133
eel migration, 103 Contracaecum aduncum, 352
Canton River Coregonus nasus
eel migration, 117 competition with eels, 262-3
Capture, methods of, 19, 151-2, 162-3, Corpuscles of Stannius, 65, 359
27 2 -3 2 7 Cortisol, 61, 64
Cardiac Crustaceans
activity, 36 competition with eels, 263-5
rate, 36-7 Cryptocarion irritans, 351
Cardiovascular system, 35-8 CyStS, ovarian, 357
Caribbean Czechoslovakia
eel migration, 91 eel growth rate, 197
Caudal heart, 38
Celebes Danube
eel migration, 122 eel migration, 104, 105
Cells 000,3 61
aJpha, 59 DDT,361
chloride, 25 Deoxyoonicosterone, 51
club, 10 Dermis
42 6 INDEX
thickness, 10 pot, 273

Deropristis infiala, 356 pound nets, 281-5
Di~[hylstiJbesterol, 5 I quality, 366-8
Diparaoxyphenylhexan, 50 rake, 315
Diseases secondary sexual characteristics, 5-6,
bacterial,34 2 -51 ,88
cauliflower, 343-6 short-finned, 1:10
economic dfeCIS of, 343 smoked, 364, 367
monalitY, 344 spears, 315-7
treatment, 345 yield from, 317
freshwater eel, 346-7 stocking, 265-71
gill, 351 transport, 26-7
red fin, 347-9 traps, 273-85
saltwater ed, 347-51 permanent, 292-300
contcol of, 350-1 trawls, 300-4
damage from, 350 weirs, 292-7
virus, 342-5 I yield fluctations, 244-9
white-spot, 351 See also different eel species under
Dopamine, 37 Anguilla and under different growth
Drifting ed jug line, 307-8 stadia e.g. Glass eel, Silver eel
Drinking, 31 Egg'
artificial fertilization, 52
East Indies depth of laying, 95
eel migration, 122 hatching, 93
Erler Dam number, 52
eel stocking, 267 si2e, 50, 51
Ed Elbe, River
ancestry, 129-32 eel homing behaviour, 216-7
baskets, 273-7 Electrocardiogram, 37
capture methods. 272-327 Electrofishing, 304, 3t8-22
characteristics for identification, 81- Elvers
'"
compared with euryhaline and sreno-
haline teleostS, 37
classification, 18, 133
fishing for, 151 - 2
food intake, 148
competition with crustaceans, 26)-5 geographical origin and effect on sex
competition with other fishes, 256-63 determination, 58
diet, 169-184 length, 146-7
diseases, 342-51 pigmentation, 146-7
farming costs, 335 pollution effects, ' 53-4
fisheries development. 241-4 stocking, 267-9
fishing yields, 241-71 temperature control of pigmentation
fry, 18, 162 and length, 147
ladders, 164-8 weight and pigmentation, 147
live storage, 340-1 Eminentiae granulares, 67
long-finned, 1:10 Endocrine system, 58-66
ponds, 328, 330-2 Epidermis
population density and catch size, changes related to salinity, II
252-5 thickness, 10- 12
INDEX 427
Epineurals, 6 eel yields, 245-6
Epiphysis Gill, Lough
See Pineal organ eel growth rate, 194-5
Epipleurals, 6 Gills
Epithelium adenosine triphosphatase content, 25
electrolyte movement across, 25 apparatus, 3
olfactory, 74 arches, 23
Esterases, 118 blood clearance through, 38
European continental slope, 95 euryhalinity, 25-6
European eel filaments, 23
See A. anguilla lamellae, 23
Exophthalmia, 357 microcirculation, 24
Eye, 68-73 nitrogenous waste excretion, 26
changes in and sexual maturity, 68-70 osmoregulation, 24-5
Glasaal, 18
Fat content, 366-8 Glass eel, 18
Fatty acids, 367 age determination, 185
Feeding boxes, 335 alimentary tract, 28-9
Feeding quotient, 333, 340 arrival in Europe, 134-7
Fibres classification, 18
neurohypophyseal, 59 development, 93
neurosecretory, 61 export, 150-1
Fibromyoma, 357 fishing, 151-2
Fibrosarcoma, 357 food intake, 148
Fiji Islands Invasion, 140, 145
eel migration, 118 commercial use of, 149-53
Fins effect of west wind on, 246-8
anal,8 locomotion, 133-4
caudal, 9 losses, 268
malformation, 359 migration to fresh water, 139-49
dorsal,8 migration to inland wate.r, 140
pectoral, 6, 9 migration in sea, 133-9
rays, 14 migratory speed, 157
supports. 14 negative phototaxis, 140-2
Fish farming, 328-40 olfaction, 73, 77, 78
foods, dried, 333-4 pigmentation, 71 , 143
foods, fresh, 333 restocking in Japan, 118, 153
Fish passes, 164-5 skull,4
Fishing eyes, 293 spawning grounds, 86, 116, 119
Flavones. 16 stocking, 267-8, 332
Floats, fishing, 306-7 surface migration, 144
Florida Stream, 92 swimming speeds, 137
eel migration. 91, 239 temperature effects on ascent, 142
tidal effects on ascent, 142
Gall bladder, 35 transport, 152-3
Gas gland, 26 upstream migration, 143-4
Gastrointestinal tract, 28-32 yields, 150-1
German Democratic Republic Glomerular filtration rates, 41-2
INDEX
Golden eel Homing behaviour. 215-20

See Yellow eel Hooks. fishing, 304-5
Gonadotroph cells, 59 damage from. 359-60
Gonadorrophin Hormones
chorionic, 50 artificially induced maturity, 50- I
mammalian, 5 I renal regulation, 41
Gonads, 43-58 uterotonic. 61
development, 43-50 Hyperphagia
fatty degeneration, 43-4 effects on pancreas, 33
identification, 43 Hypoglycemia, 34
weight, 50, 51 Hypophysis
Great Lakes anatomy, 61
eel growth, 197 morphology, 67
eel migration, 107 reltionships with hypothalamus, 61
Green eel weight. 59
classification, 18 Hypothalamus, 61
Greenland Hypural plates, 9
eel migration, 91, 106
Gro!3e eels, 364 Iceland
Growth, 184-208 eel migration, 91, too
Grunaal, 18 Ijsselmeer
Guayana eel migration, 140,221 . 231
eel migration, 91 Jchthyophthiriodts browni, 351 - 2
Gulf of Mexico Jchthyophthirius multifilii5, 35 I
eel migration, 107 Illinois River
Gulf Stream, 92 eel migration, 107
eel migration, 91, 98,135 Indo--Pacific eels. 11 0-129
G U[ ontogeny, 29-30 diseases, 352
Gymnorhorax peull;, 22 Injuries
fish hook, 359-60
Haematocrit, 38 pathological,356-9
Haematopoiesis, 39, 41 pollU[ion, 361
Haemoglobin sewage, 361
content, 38 turbine, 360
electrophoretic panerns, 40,100 Interrenal tissue
ontogeny of synthesis, 38 Set AdrenOCOrlical homologue
species differentiation, 130 Intersexuality, 49, 53
Hassal corpuscles, 65 Iodine content, 367
Head width Iris, 68
characteristics, 4-5 Islets of Langerhans, 32
diet, 177- 9 Isoaglutinin system, 40
growth, 193 Isoenzymes, 118
weight, 205 Isotocin, 62
Hearing, 79-80
Hearl,35-8 Japan
caudal,38 eel farming , 329-35
Heligoland Bight eel fisheries, 243
eel migration, t35 eel restocking, 11 8, 153
Hermaphrodites, 56-7 Japanese eel
INDEX 429
See A.japonica Macrophthalmus, 73, 357
Java Magnetic fie ld
eel migration, 122 influence on migration, 211}-20
Jaw, upper, 1-3 Malagasy
spawning ground, near, 1 11-2
Kaupichlhys diodorllus, 22
Malate dehydrogenase, 100, 118
Kenya
Malawi, Lake
eel migration, 113
eel migration, 113
Kidneys, 41-2, 66
Malnutrition
Kuroshio Current, 92
effects on gonad function, 57
eel migration, 114-5
Marketing categories, 364, 366
Labrador Medulla oblongata, 67
eel migration, 107 Mediterranean
Labrador Stream, 91-2 arguments for and against spawning
Lactate dehydrogenase, 118 area, 86-8
Larvae, 83-102 eel growth rate, 196
length frequency and age groups, eel migration, 136
94-95 Metamorphosis
metabolic rate, 95 length changes, 146-8
stage length, 94-5 skin changes, 1I, 146-7
Lateral line sense organs, 79-80 weight changes, 147-8
Length Migration
yellow eels, 173-7, 184-208 elvers, 145-9
young eels, 154-6 European and American eels, 83-110
Lens, 68 glass eels, 133-145
Leptocephalus Indo-Pacific eels, 110-129
lack of blood pigment, 38 olfaction, 78-9, 98
locomotion, 133-4 sex and distance from coast, 56
olfaction, 73 thyroidal activity, 63
pigmentation, 14 young eels, 154-62
skull, 4 Mississippi
LeplOcephalw brevirostris, 83 eel migration, 107
Light sensitivity, 68, 322 Moluccas
Line fishing, 304-15 eel migration, 124
Lines Montee, 18
angling, 306 Moringua edwardsi,
jug, 307-8 pectoral girdle, 6
long-, 308-13 skull,4
yields, 3rt Moringua javanica, 22
Liver, 34-5 M02ambique
ammonia production, 26, 34 eel migration, 112
double, 359 Muscles, 21-2
Locomotion, 133-4 chemical composition, 22
neural influences, 67 proteins, 118
Lunar periodicity, 161-2,224-6 Myomere counts, 88-9, 120
Lutschu Island
eel migration, II6 Naring, 313-5
Lymph hearts, 37 yields, 314
Lymphatic system, 37 Narrowheaded eel
430 INDEX
classification, 4-5 eel migration, 120

food, 177- 9 North Atlantic Stream, 92
growth rate, 193 eel migration, 91, 95
heart weighl, 36 North Sea
price, 364 eel yield fluctuations, 244, 248
weight, 205 Nostrils, 72-3
Natal
eel migration, lIZ Oculomotor nerve, 68-9
Neagh, Lough Oesophagus
eel growth, 204 mucous membrane, 28
eel migration, 231 Oestradiol propionate, 50
Nematodes, 352 Oestradiol dipropionate, 51
N~oconger tJtrmi/ormis Oestradiol monobenz.Gate, 51
pectoral girdle, 6 Olfaction, 73-9, 98-9
skull,4 O lfactory attractantS, 77-8
Neoplasms, 357 Olfactory bulb, 74
Nephron histology, 41 O lfactory repellants, 76-7
Nets Olfactory rosette, 73
drag, 301 Olfactory sensitivity, 74-6
fyke, 277-81 Ontario, Lake
round, 279-80 eel growth, 204
winged,178 Oocytes
lift,3 I S number, 52
pound,181-5 Oogenesis, 52
Pomeranian, 284 Optic lobes, 66
yields, 181 Optic neves, 67
stow, 285-92 Orientation
anchored. 286-8 magnetic fields, 219-20
otterboard, 288-9 olfaction, 76, 78-9, 98-9
staked,186 temperature, 97
yields, 290-1 visual, 99
nammd Osmoregulation
yields, 301 gills, 24-6
trawl, 300--4 gut, 29-30
e1ectrofishing with, 304, 321-1 parasympathetic nervous system, 67
yields, 303-4 pituitary function, 59-61
Neurohypophysis, 42. S9 urohypophysis, 63
New Caledonia Otoliths, 185-7, 192
eel migration, 120 Ovaries, 43-50
New Guinea Oxygen
eel migration, 122, 126 dissociation, 39
New Zealand panial pressure, 39
eel migration, 113. II9-121 saturation, 39
Newfoundland uptake, 23-4, 26
eel migration, t 07 Oyashio current, 92
N iagara falls eel migration, 116
eel migration. 107
Noradrenaline, 37 Pacemaker system, 35-6
Norfolk Islands Pacific eels, 1I4-29
IND EX 43 1
Pancreas, 32-4 Puis fishing, 301
Papilloma, 345 Pytl wnichthy s sp.
Parasites, 351 -6 pectoral girdle, 6
Pareja (twin trawl), 301 skull, 4
Pectoral girdle, 3, 6
Rearing techniques, 33 2
Peptides
Red blood cells
neurohypophyseal, 62
Petersen young fish trawl, 86 dimensions, 38
effect of environmental salinitY, 39
Phenol flavour , 341
Pheromones, 78 Red Sea
Phototaxis, negative , 140--2, 161, 1 eel migration, 104
3 7, Reisenaaie, 365
3" Reis sner fibre complex, 68
Phylogeny, 130
Renin system, 42, 66
Phytoplankton, 337
Pigmentation Res piration
branchial, 11 , 24,2 6
anomalies, 359
environmental influence, 18- 19 cutaneous, II, 24
swim blad der, 26
external, 14-1 9
Respiratory exchange
internal, 14
surface area, 23
retinal, 71
Respiratory organs, 23-8
sexual development, 19
temperature correlation, 14, 143 Respiralory rates , 23
Resting pools, 330--1
Pineal organ, 16
Retina, 68-7 0
Pituicytes,61
RheotaXis, 99, 139, 154, 157, 233
Pituitary gland, 42, 58-6 2 -4
Rhodesia
function, 59
eel migralion, 113
morphology , 59
Rhodopsin, 68, 72
Plasma
Rhombencephalon, 67
electrolyte concentrations, 39
Rhone Delta
osmolarities, 39
Plat ten, Lake eel migration, 87
Rods (eye anatomy ), 70--1
eel stocking , 267
Pneumatic duct , 26 Rods, fishing, 305 -6
Poland Roggiin, Lake
eel stocking, 267
eel fisheries, 243
Pollution Russian dam s
effects on elver ascent, 153- 4 eel growth rate, 195 - 6
Russian rivers
types, 361 -2
eel migration, 10 5
Ponds
eel restocking, 110
eel feeding, 333 -5
fattening , 332 St . Lawrence river
systems, 330 -2 eel growth, 204
yields, 330--2 Sakrow, Lake,
Porphyropsin , 72 eel growth, 205
Prices Salinity
categories, 364- 6 effect on epidermis, I t
ecology of, 365 effect on migration, 233- 4
Prolactin, 32, 61 effect on red blood cells , 39
Proteins, 40-- I, 118 effect on sex determination, 57
432 INDEX
Salmonids, meteorological influen~ on migration,

competition with eels, 260 226-30
Saproiegnia parasirica, 352-3 rrUgration, 88. 96, 220-40
Sargasso Sea migration at sea, 235-9
eel migration, 95. 97. 102 migration direction, 238
Satzaale, 18 migration distance, 240
Scales ontogeny, 10
development, 12 oocyte weight, 51-2
use in age determination, 186-7. 192 pancreas, 34
Schleswig-Holstein rainfall influence on migration, 227
eel yields, 245- 6, 269 river current influen~ on rrUgration,
Schmidt, Johannes, 86-91, 99. 102-4 "7
Schokker, 286 salinity influence on rrUgration, 233-4
Seines. 300-1 seasonal variation in migration, 220-4
Serum sex and rrUgration, 222-3
cholesterol,41 speed, 239-40
proteins, 40- I subterranean migration, 234-5
transferins. 40, 87. 100 temperature influence on migration,
urea, 40 23 2 -3
Sewage. 361 tidal current influen~ on rrUgration,
Sexual development, 43-58 228-30
eye siu, 69 water level influence on rrUgration,
pigmentation, 19 227- 8
Sexual differentiation weight, 207
distribution and. 54-6 wind influence on rrUgration, 230-2
effects of environmental salinity, 57 yield, 220, 245, 248, 250, 295-6
effects of population density, 53 Sinkers, 306
effectS of temperature, 57 Skeleton, 1-9
environmental factors, 53-7 axial,4
gonads, 43-4. 49. 52-3 decalcification, 8
growth rate, 187-93 Skin
head shape, 5-6 gas exchange, 11 -2,2)
migratory paneros and, 222-3 mucus production, 12
Silver eel ontogeny, 10
barometrical influence on migration,
".
structure and function, 10-2
tactile sense, 80
classification, 18 Skull, 1-6
depth when migrating, 232-3 ontogeny, 4-5
development. 19 spot, 15-6
diurnal influen~ on migration. 224 Snigs
effect of light, 326-7 See Young eel
eyes, 72-3 Snoods, 305
growth rate. 190-3 Somatotroph, 59
gut, 29-30 South Equatorial Current, 92
hydrolOgical influence on migration, eel migration, 127
226-30 Spawning
length,223 salinity influence on, 97-8
lunar periodicity influence on migra- seasonal influence on, 128
tion, 224-6 temperature influence on, 97, 123
marking for recapture, 235 Spawning grounds
INDEX 433
depth, 95-7, 128 effects on glass eel ascent, 142
European and American eels, effects on pigmentation, 14, 143
86-102 effects on silver eel migration, 232-3
Indo-Pacific eels, 110-129 effect on young eel migration, 159
Spears, 315-7 Teratology, 357-8
Species, differences between, 81 - 132 Testes, 43
Sperm morphology, 50 development, 48-9
Starvation, 33 Tethys Sea
Sterrhurus musculus, 352 eel distribution, 131
Stocking, 265-71 fossil eels, 131
elvers, 267-9 Thigmotaxis, 317
glass eels, 267-8, 332 Thymus gland, 65
Japan, 118, 153 Thyroid gland, 62-3
Stomach,28 Thyrotroph, 59
mucous membranes, 28 Torus semicircularis, 67
Storage Trade, international, 362-3
weight loss in captivity, 341 Transdeamination,34
Straits of Gibralter, Transferins, 41 , 87, 100
eel migration, 86 Traps, 273-85
Straits of Korea cylindrical, 281
eel migration, 114 light, 322-7
Straits of Messina permanent, 292-300
eel migration, 86 refuge, 317-8
Subcommissural organ, 68 shelter, 318
Suez canal stream and river, 297-300
eel migration, 103 yields, 298-300, 323
Sumatra Trematodes, 355
eel migration, 122, 127 Trinidad
Swimbladder eel migration, 91
histo1ogy,27 Tropical eels, 122-9
inflammation, 357 Trout
structure, 26-7 competition with eel, 257-61
Swimming speeds, 102, 137 yields from farms, 330
Synahorin,51 Tucker's hypothesis
arguments for and against, 99-102
Tahiti Tumours
eel migration, 118 fibroepithelial, 345
Taiwan kidney, 357
eel migration, 114 Tunis, Lake
eel ponds, 331 eel migration, 147
Teeth Twin trawl (Pareja), 301
shape and ontogeny, 19-20
specific differences in, 20, 100 Ultimobranchial bodies, 64
vomerine, 2, 20 Urohypophysis, 63
Telemetry, 102, 219, 229-30, 233, Uropurygius knighli, 22
238-40
Temperature Valencia
effects on eel yields, 247-8 eel growth rate, t96
effectS on elver pigmentation and Vascular beds, 37
length, 147 Ventricle, 35-6
434 INDEX
Vertebrae diet and habitat, ISo

abdominal, 6 diurnal variation in food intake, 170--1
calcium phosphate content, 8 effects of light on, 326-7
caudal. 6 effects of salinity on migration, t 60--t
crystalline apatite content, B environment and growth, 187-92
use in age determination, 187 eyes, 69-70, 7Z
Vertebral colunm, 6-8 feeding discrimination, 181-4
calcium phosphate content, 8 food and body size, '74-7
crystalline apatite content, 8 food and head size, 177-9
curvature, 357 growth rate, 186, 194
shortening, 358 head width and growth, t93-4
Vertebral counts, 86, 99,120 homing ~haviour, Z15-20
Vibrio anguillarum, 346, 347 interspecific difference in growth,
Vilm, Lake 197-8
eel stocking, 266 length and weight, 199-208
Viremia, 345 negative phototaxis, 209
Vitamin A. 72 olfaction, 73, 98
Vitreous humour, 68 pancreas, 33
Volga River seasonal variation in diet in temperate
eel growth rate, 195 species, '7'-3
seasonal variation in diet in tropical
Weight species, 173
correlation with head width, 205 seasonal variation in food intake,
correlation with length, 206-7 169-70
seasonal variation, 204 sexual differences in growth, 187-92
sexual differentiation, 204-5 survival in air, 213-5
Weirs, 292-7 swimming activity, 171
self-fishing, 293-4 tube hide-outs, z09
stake, 284 yields, 248, 250, 299-300
yields, 295-7 Yellow Sea
West Germany eel migration, 114
eel yields, 248 Young eel
Wind characteristics, 154
effect on glass eel invasion, 146-8 diurnal effects on migration, 159-60
Windemere, Lake effects of lunar periodicity on migra-
eel migration, 231 tion, 161-2
Worth, Lake effects of salinity on migration, 160-1
eel growth rate, 195 invasion
Xanthochromatism, 18,359 commercial use of, 162-9
length, 154-6, 160
Yangtse-Kiang River migration, 154-62
eel migration, 117 migratory speed, 157
Yellow cd stocking, 266
age and growth, 184-208 temperature effects on migration, 159
burrows, 209 trapping, 162-3
cardiac rate, 36-7 yields, 168-9
classification, 18, 169
depth distribution, Zl 1-37 Zambesi River
diet and food availability, 179-81 eel migration, 11 3

F.-W. Tesch Dr. Agr. (Auth.), P. H. Greenwood D.sc. (Eds.) - The Eel - Biology and Management of Anguillid Eels-Springer Netherlands (1977)

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THE EEL

F.-W. TESCH, Dr. agr.

Translated by Jennifer Greenwood

ENGLISH EDITION EDITED BY

P.H. GREENWOOD, D.Se.

This first English-language edition of Der Aal by F.-W. T esch, has

ISBN-13: 978-94-009-5763-3 e-ISBN-13: 978-94-009-5761-9

© 1977 Chapman and Hall Ltd.

Distribution in the U.S.A. by Halsted Press,

Library of Congress Cataloging in Publication Data

Translation of Der Aal.

I. Body Structure and Functions [

I. 1.2 Pectoral girdle 6

I.7.2 The thyroid gland 62

2. Eel Species, their Developmental Stages and

3· Post-Larval Ecology and Behaviour 133

B The 'Yellow eel' stage to the 'Silver eel' stage 169

4· Fishing Yields 241

4.2.5 Catch variations in Canada 248

5· Fishing Methods 272

6. Eel Culture and Live Storage 328

7· Diseases, Parasites and Injuries 342

8. Trade and Processing 363

Dr Tesch's wide ranging account of anguillid eels impinges on the

From a physiological viewpoint the eel is a parcicuiarly popular

in classification within the family Anguillidae (see p. 100) . The basis-

A comparative study of the skull and axial skeleton in Anguilla

therefore be considered suitable as a ready means for distinguishing

I. 1.2 Pectoral girdle

1.1.3 Vertebral column

TabJe I Comparison of the vertebral columns of A . anguilla and C. conger (after

Autogenous neural arches : Vertebrae nos. 1St-5th lSI-16th or 17th

R elatively high neural procencs 2nd, and less 1st-16th or 17th

Double structure of the parapophysis

Undulating surfa<;e Structure on

Parapophyses and ventral posuygapophyses in the

The last abdominal vertebrae and a large number of

vertebrae. According to Smith & Castle (1972), the distribution of

1.2 Skin and musculature

1.2. I Structure and function of the skin

Jakubowski ( Ig60b) compared the skin of seven teleost species, among

contrasting with species such as the armoured catfish Corydoras palaea-

exchange or small exchanges of electrolytes. Jakubowski (I960b)

Fig. 7 Photographs of eel scales (after Rahn, 1955a).

changes that progress in a caudo-rostral direction can be seen in the

a certain degree of rostral pigmentation are already in evidence in

Table 2 Development of pigmentation in A . anguilla (abridged from Strubberg,

I Larva, fully grown lep tocephalus (Fig. 36.1)

appearance of which is certainly of great physiological significance; this

Fig. 10 Teeth of the premaxi1lo.-ethmovomerine bloc in A. anguilla (Photograph :

During ontogenetic development the teeth undergo considerable

bony fishes and members of the Salmoniformes. Considering the

A number of workers have examined the physiological mechanisms

typified by species such as Fundulus, a killifish, is termed 'osmoconfor-

counter-current exchange situation may readily explain the respiratory)

1.4 Feeding and organs of digestion

1.4. I The gastrointestinal tract

Fig. 14 Cross-section through A. anguilla. ab, Swimbladder; c, Visceral cavity;