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Full text of "The British parasitic Copepoda"

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THE LIBRARY 

OF 

THE UNIVERSITY 

OF CALIFORNIA 

DAVIS 



THE 



EAY SOCIETY, 



INSTITUTED MBCCCXLIV. 




This volume is issued to the Subscribers to the Ray Society /or 
the Year 1912. 



LONDON 
MCMXIII. 



Digitized by the Internet Archive 

in 2007 with funding from 

IVIicrosoft Corporation 



http://www.archive.org/details/britishparasitic01scotrich 



THE 

BEITISH 
PAEASITIC COPEPODA 



THOMAS SCOTT, LL.D., F.L.S. 

AND 

ANDEEW SCOTT, A.L.S. 



VOLUME I 

COPEPODA PARASITIC ON FISHES 

TEXT 



LONDON 
PRINTED FOR THE RAY SOCIETY 

AND SOLD BY 

DULAU & CO., LTD., 37, SOHO SQUARE, W. 



1913 

mm A Mk ■ 1 



PBINTKD BY ADLARD AND SON 
LONDON AND DORKING 



THE AUTHORS 

DESIRE, WITH THE MOST SINCERE RESPECT, 

TO DEDICiVTB THIS WORK 

TO THE 

REV. CANON A. M. NORMAN, M.A., 

D.C.L., LL.D., F.R.S., F.L.S., ETC., 

IN GRATITUDE FOR HIS UNFAILING 

KINDNESS AND ASSISTANCE 

EXTENDED TO THEM FOR MANY TEARS. 



PREFACE 



Dr. Baird's work on 'The Natural History of the 
British Entomostraca,' published by the Bay Society 
in 1850, marked an important stage in British Natural 
History research, and forms the basis of the present 
volume. That work contains a complete account of 
the Copepoda found living as parasites on British 
fishes, so far as then known. Much has been done 
since then to extend our knowledge of these organisms, 
and many species have been added to the number 
recorded by Dr. Baird. His work naturally tended to 
produce such results, for it placed before British 
students, in a concise manner, a large amount of 
interesting information concerning these parasitic 
forms, and, by showing the successful results which 
had already been accomplished in other countries 
among such curious animals, led to an increased 
interest in their study. 

' A Monograph of the Free and Semi-Parasitic 
Copepoda,' by Dr. Gr. S. Brady, was published by the 
Ray Society in 1878-1880. That work, which con- 
sists of three volumes, marked a second important 
stage in the study of these minute crustaceans, and 
was an evidence of tlie increasing attention their study 
was receiving. But though several groups of the non- 
freeliving species were described in that Monograph, 
those found parasitic on fishes were expressly omitted. 
The author in his Introduction states : " The truly 
parasitic forms — fish-lice, etc. — are excluded as not 
coming within the scope of my work. They are of 
themselves sufficient to occupy an independent volume. 



Vlll PREFACE. 

and to afford abundant work to any naturalist wliose 
time and opportunities allow of his taking them in 
hand" (Vol. I, p. 2). The present Monograph is the 
result of an endeavour to enumerate and describe the 
species which Dr. Brady has excluded, and may there- 
fore be regarded as supplementary to that author's 
work. 

The Copepoda recorded by Dr. Baird as parasites 
on British fishes number thirty-four species ; this 
number is now increased to one hundred and thirteen. 
Many of the new records are scattered through various 
publications, and it has been considered by the Council 
of the Ray Society that, for the convenience of 
students, these should be brought together and 
combined with those described by Dr. Baird, and thus 
made more accessible. This work, which has been 
intrusted to us, we have endeavoured to carry out, 
and we hope, while conscious of its imperfections, that 
it may be found useful and tend to create a still 
greater interest in the study of these remarkable 
organisms. 

In the preparation of this Monograph we have been 
indebted for valuable assistance to numerous friends, 
and to none more so than the Eev. Canon A. M. 
Norman. Indeed but for his encouragement and 
assistance it is doubtful if we could have ventured to 
undertake it. We therefore embrace this opportunity 
of expressing our gratitude to him, not only for his 
helpfulness in this special work, but also for kindness 
shewn to us in many other ways. We are also 
indebted to Dr. Williamson and Dr. Bowman of tlie 
Scientific Staff of the Fishery Board for Scotland for a 
number of rare and interesting specimens; and to 
our friend Mr. John Lindsay (of Messrs. Blackwood & 
Sons, Edinburgh) for assistance in revising the proof- 
sheets and other useful help. Our grateful acknow- 
ledgments are also due to Mr. John Hopkinson, 
Secretary of the Eay Society, for the great labour and 
interest he has taken in seeing the work through the 



PREFACE. IX 

press. Our obligations to other friends who have 
assisted iis with information or specimens are grate- 
fully acknowledged in the body of the work. 

The drawings have, with some few exceptions, all 
been prepared from recent specimens, examined by our- 
selves, and wherever possible careful dissections have 
been made.* There are a few species, however, of 
which we have been unable to obtain specimens, such 
for example as Demoleus ^aradoxa, Levnxocera cypri- 
nacea, Brachiella pastiiiaca, and perhaps one or two 
others, figures of which have, with due acknowledg- 
ment, been reproduced from published works men- 
tioned in the sequel. In a considerable proportion of 
the species dealt with in this work the ovisacs are 
frequently greatly elongated, and we have followed 
Dr. C. B. Wilson in adopting the term "egg-strings" 
to describe them. The coloured figures of the para- 
sites represent them as observed by us in the particular 
specimen illustrated. The colours vary considerably 
according to the condition of the parasite, the place 
where it was attached, and the freshness of the host. 
External parasites lose their colours much more 
rapidly than those that are found under the oper- 
culum or attached to the gills, and it is not always 
possible to obtain perfectly fresh fish. 



THOMAS AND ANDREW SCOTT. 



Aberdeen, 

31st March, 1913. 



* Note by Thomas Scott.— The drawings and dissections refeiTed 
to above are entirely the work of my son Andrew Scott, who has 
otherwise assisted in the preparation of the Monograph. 



CONTENTS 



PAGE 
Inteoduction ....... 1 

Systematic List of the Species . . . . .30 

Descriptions op the Species . . . . .33 

Tribe I. CYCLOPOIDA 33 

Fam. i. Ergasilid^ . . . . .33 

Tribe II. CALIGOIDA 43 

Fam. II. Caligid^ . . . . . .44 

Fam. III. DicHELESTiiD^ ..... 105 

Fam. IV. Philichthyid^ ..... 137 

Tribe III. LERN^OIDA . . .141 

Fam. v. Lern^id^ ..... 141 

Fam. VI. Chondracanthid^ .... 166 
Fam. VII. Lernjeopodid^ .... 186 

Argulus foUacens ....... 227 

Tri^paphylus musteli (amended description of male) . . 229 

List of the Fishes on which Parasitic Copepoda were 

obtained ...... 230 

Bibliography of Memoirs referred to in the Text . 233 
Index , ... 248 

Errata . . . . . .252 



BRITISH PARASITIC COPEPODA. 



INTRODUCTION. 

Pauasitic habits are not peculiar to anj class of 
organisms, but are met with in all departments of life, 
vegetable as well as animal. Nevertheless there are 
certain groups both of plants and animals whose en- 
vironment seems to be specially favourable for the 
adoption of habits of a parasitic or semi-parasitic 
kind. Such habits seem to prevail to a considerable 
extent among the Crustacea, and particularly among 
the so-called lower forms belonging to that class. 
Many kinds of animals are required to act as hosts for 
these crustaceans, and curiously enough not a few of 
them belong to the same class as that to which the 
parasi-fces themselves belong. Many fishes are also 
infested by these crustacean parasites. Fishes live in 
a medium where on every side they are exposed to the 
attacks of all kinds of parasitic forms, so that very 
few of the fishes examined by us are found to be 
entirely free from these organisms. 

At a fish-market where quantities of fishes of various 
kinds are being landed, crustacean parasites may be 
frequently observed on the skin of the fishes, or found 
adhering to their fins, their gills, or on the inside of 

VOL. I. 1 



Z BRITISH PAKASITIC COPEFODA. 

the gill-covers; sometimes they may be found clinging 
to the roof and sides of the mouth, as well as on the 
tongue, especially the under side of it, and also in the 
nasal fossae. At other times they make excavations in 
the flesh of the fish, causing sores of a more or less 
serious nature, and they may even penetrate into 
the abdominal cavity, as occasionally happens with 
Lernaean parasites. 

The degree of parasitism varies greatly even among 
closely allied species. In somes cases the relationship 
of the crustacean to the fish is decidedly that of a 
parasite, while in others it would scarcely be accurate 
to describe the position of the crustacean to the fish 
as truly parasitic. Sometimes, however, it is more 
convenient to use the terms '^ parasite" or " parasitic " 
in the wider sense to avoid any confusion which might 
arise by trying to define in each particular case the 
degree of relationship of the one to the other, and it is 
in the wider sense that these terms are used here. 

The study of the Parasitic Crustacea is in some 
respects more difficult, if also more interesting, than 
that of the species which live under normal conditions. 
Their structure has become more or less altered by 
reason of their parasitic habits, and the forms which 
some of them assume are greatly at variance with 
those of free-living species. The forms of some of 
these parasites are so abnormal and grotesque that 
even experienced investigators have been deceived, and 
have failed to recognize their relationship to the 
Crustacea, and only by the study of their life-histories 
have their true affinities been determined. 

Among the crustacean parasites of fishes the Cope- 
poda are probably more numerously represented than 
any other of the crustacean Orders, and what we pro- 



INTKODUCTION. 6 

pose in the first volume of the present work is to give 
descriptions of the various parasitic Copepoda known 
to occur on British fishes. The second volume will 
contain the figures. 

We have not considered it necessary to enter into 
a lengthened discussion on the structure and develop- 
ment of the parasitic Copepoda ; this has already been 
done by Dr. Baird and some other writers. Mr. C. B. 
Wilson, in his various contributions dealing with the 
North American parasitic Copepods,has described them 
very fully both in their histological and physiological 
aspects. Moreover one of the authors of the present 
volume (A. Scott) has published a Memoir con- 
taining detailed accounts of a Lepeophtheirus and a 
Lerndea, representing two important and diverse types 
of the parasitic species.* 

G-ENERAL EeMARKS ON DEVELOPMENT, EtC. 

The following general remarks concerning the 
development and structure of these organisms are 
derived chiefly from the works referred to above. 

The Copepod parasites of fishes are roughly divided 
into two groups — viz., those which in the adult stage 
are free, that is, they can move about, at least to some 
extent, from one part of the fish to another; while 
the others are permanently fixed. Lepeophtheirus 
pectoraUs may be taken as representing the first, and 
Lerndea hranchialis the second. 

The two groups taken together form a series which 
exhibits remarkable variation in habits and structure. 
While there is little to distinguish some of the 
species at the one end of the series from free-living 
non-parasitic Copepods, those at the other end exhibit 

* Liverpool Marine Biology Committee, Memoir No. VI : ' Lepeophtheirus 
and Lernsea' (1901). 



4 BRITISH PARASITIC COPEPODA. 

forms so abnormal and grotesque tliat, as already 
observed, it is difficult without a knowledge of tlieir 
life-histories to regard them as belonging to the 
Crustacea. 

The young Copepod, no matter how abnormal it 
may be when adult, begins life as a free-swimming 
nauplius ; it is of an oval form, with a single median 
eye and three pairs of limbs arranged round the 
mouth; there are no frontal appendages and no 
proper mouth-organs. 

The young parasite passes through numerous 
ecdyses and increases in size. The various append- 
ages make their appearance in regular order, and 
the animal gradually takes on the adult condition. 
In the case of those parasites which retain their active 
habits throughout life, the appendages become com- 
pletely developed when the adult form is assumed. 
On the other hand, the appendages of the parasites 
that are more or less permanently attached to their 
host are comparatively well developed early in life, 
and then continue in a stationary condition or are 
gradually suppressed. One finds on carefully dissect- 
ing the head of an adult Leiiisea from the gills of its 
host that the four pairs of swimming-feet which were 
developed at an early period of its life are present, 
and retain the characters they had during the 
Cyclops stage, although they are not of any use to 
the animal. The four pairs of feet are buried with 
the head in the tissues, and if by any means the animal 
could detach itself from its host, they obviously would 
not enable it to swim through the water. In some 
parasites certain appendages used as anchors, such 
as the antennae and second maxillipeds, become 
greatly developed and are conspicuous even to the 



INTRODUCTION. 



naked eye. In others the cephalic segment becomes 
highly chitinised, and is frequently expanded into 
horn-like processes which are buried in the tissues of 
the host. Certain members of the Caligoida are able 
to live apart from their hosts and lead a more or less 
pelagic life in the water for a time, but whether adult 
females voluntarily detach themselves from healthy 
and living hosts is doubtful. They appear, however, 
to leave the host when it dies in an aquarium tank. 
One rarely finds Lepeojphtheirus pectoralis on dead 
flounders which have been allowed to remain in a tank 
for a day or two after death, although it was quite 
certain they were present when the host was alive. 
We have kept Lepeophtheirus pectoralis alive in vessels 
of sea- water for periods of six weeks after removal from 
flounders. Adult males of Galigus rapax are often 
met with in collections of plankton, but the females 
are less frequently noticed. The members of the 
genera GJiondracanthus and Clavella sometimes live for 
a couple of days after being removed from their hosts, 
but the Leriidea quickly perishes. It is almost certain 
that, with the exception of a few members of the 
Caligoida, all the copepod parasites of fishes die soon 
after the death of their host, even although the body 
remains in the water. On one occasion we dissected 
the spiracles of a large grey skate {Baia hatis) which 
had evidently died in the sea and was afterwards 
washed up partly macerated on the beach. A full- 
grown specimen of Charopinus dahnanni was found in 
one of the spiracles and was in a perfect condition, 
but there was no sign of life in it. 

The sexes are separate, the males as a rule being 
much smaller than the females. In many cases the 
males are simply parasites on the females, especially 



b BRITISH PARASITIC COPEPODA. 

those of Ghondracanthus, Lerneopoda, and Glavella, 
The fact that males are found upon egg-bearing 
females of these genera is due to their power of loco- 
motion having been lost when they reached the adult 
condition. When once they have settled down on a 
female and matured they are unable to change their 
position to any extent. Fertilization of the female is 
effected early in its life, before the metamorphoses are 
completed. The resulting embryos pass out from the 
two oviducts and remain attached to the external 
apertures either in single or multiserial columns, each 
being enclosed in a sac, until they hatch. In Ghondra- 
canthus zei the columns are very much compressed and 
resemble miniature oval biscuits. The period of incu- 
bation extends over several weeks. The young para- 
sites hatch out as free-swimming nauplii. These 
undergo metamorphoses, which in some forms, after a 
particular stage is attained, are retrogressive, finally 
leading to the adult condition. 

The following is a short account of the anatomy of 
two members of very different families, the Caligoida 
and the Lernaeoida. For fuller details the Memoir 
published by the Liverpool Marine Biology Committee 
and the reports by C. B. Wilson on North American 
Parasitic Copepods should be consulted. 

Tribe Caligoida. 
The species chosen to represent this tribe is Lepeo- 
phtheirus pectoralls (Miiller). This parasite is some- 
times very common on the fins and body of the 
flounder or ''white fluke" {Pleuronectes flesns). It 
also occurs on other members of the Pleuronectidge. 
Although Lepeophtheirus pectoralis does not confine 
itself to any particular part of the exterior of its host, 



INTRODUCTION. 7 

and though adult and immature specimens of both 
sexes may be found scattered all over the body, mature 
egg-bearing females are most frequently met with 
imder the pectoral fins. (See Plate A.) It is possible 
to collect a fairly complete series of stages of develop- 
ment from the one host. The average length of a 
mature egg-bearing female is one-fifth of an inch, and 
of a male one-ninth of an inch. The parasites attach 
themselves to the fish by means of their powerful 
second maxillipeds assisted by the antennae, and a 
decided pull has to be applied before they can be 
removed. By depressing the edge of the carapace 
and applying it closely to the skin, the parasite is able 
to increase its holding power to such an extent that 
the posterior portion can be torn off from the anterior 
without detaching it. The anterior portion when thus 
separated will sometimes remain alive for twenty-four 
hours. It is even capable of swimming about 
vigorously, but eventually its movements become 
erratic and finally it dies. Leipeophtheiriis ]jectoralis 
can be kept alive in sea water for about six weeks after 
removal from the host. The parasites live for a con- 
siderable time after the fish has been caught and re- 
moved from the water, if they are not allowed to dry up. 
Increase of temperature to 16° C. is fatal to them. 
Decrease of temperature to even below freezing-point 
does not appear to incommode them. If the aquaria 
are kept cool, the sea- water in which the parasites 
are placed after removal from the host need only be 
changed at long intervals. 

External Appearance and Structure. 
The animal is depressed dorso-ventrally and is 
divided into four parts. The first and usually the 



8 BRITISH PARASITIC COPEFODA. 

largest is almost circular in outline. It bears all the 
appendages with the exception of the fourth and fifth 
pairs of feet. This part is known as the cephalo-thorax. 
The second part is very small and represents the fourth 
thoracic segment of the pelagic Copepoda. It bears 
the fourth thoracic feet. The third part of the body 
is of variable size and shape, and is known as the 
genital segment. Its size, and shape depend largely 
upon the degree of maturity of the reproductive organs. 
The fourth part is short and narrow. It is only about 
one-fourth the width of the genital segment. It repre- 
sents the abdomen of the pelagic Copepoda, and bears 
at its apex two short papillae known as the f ureal joints 
or caudal stylets. 

Viewed from above, the cephalo-thorax is seen to be 
slightly convex and divided into four portions by im- 
perfect sutures. The frontal margin is indented. The 
greatest depth is in the middle line. In the centre of 
the hollow, situated on the ventral surface, is an oval- 
shaped opening with a chitinous fringe. This opening 
is apparently a rudimentary sucker, and is the remains 
of a median sucker which is greatly developed in the 
early life of the parasite. This rudimentary sucker can 
be traced in all the members of the Caligoida. No 
other suckers are present in the members of the genus 
Lepeophtlieirus. The frontal and lateral margins of 
the segment are surrounded by a transparent mem- 
brane with faint transverse lines. This membrane is 
simply an extension of the chitinous exoskeleton 
which covers the whole animal. The edges have fre- 
quently a serrated appearance, but this is due to the 
membrane being damaged. The eyes appear as a 
reddish spot in the living animal. They are placed on 
the dorsal surface midway between the anterior and 



INTRODUCTION. 9 

posterior margins. When the spot is examined micro- 
scopically it is found to consist of two distinct eyes 
closely approximated and enibedded in a mass of 
reddish-black pigment. The eyes are wholly under 
the carapace. Each eye has a simple, spherical, crys- 
talline lens beneath a thin cornea. Behind the lens 
lies a row of retinal cells of fairly large size and lined 
internally with a layer of pigment. A chitin division 
deeply pigmented with red separates the two eyes. 

The Appendages. 

There are twelve pairs of appendages, as follows : — 
One pair of antennules, one pair of antennae, one pair 
of mandibles, two pairs of maxillae, two pairs of maxil- 
lipeds, and five pairs of feet. The first three pairs 
of feet only are adapted for swimming. The fifth pair 
of feet is very small and is attached to the postero- 
lateral margins of the genital segment. A sixth pair 
of feet is usually present in the male, appearing as 
small papillae immediately behind the fifth pair. 

The antennules are placed on the frontal margin of 
the cephalo-thorax and are two-jointed. The basal 
joint is usually moderately broad and is furnished w^ith 
a number of plumose setse. The setae are well supplied 
with nerves and evidently act as sensory organs. The 
other appendages are all on the ventral surface. The 
first in order are the antennae. These are two-jointed. 
The apical joint is a strongly-prehensile claw which is 
used for holding on to the host. The mandibles are 
stylet-shaped and are enclosed in the suctorial mouth. 
The apical joint is flattened and curved. The inner 
margin is distinctly serrate at the distal end, the 
serrations being sometimes more or less irregular. 

The mandibles of all the copepod parasites of fishes 



10 BMTISH PAEASITIC COPEPODA. 

with the exception of Ghondracanthus are of a some- 
what similar type to that of Lepeophtheirus. The first 
maxillae are one-jointed and are situated near the 
lateral margins, slightly posterior to the base of the 
antennae. The second maxillae are also one- jointed and 
are placed at the sides of the suctorial tube which 
carries the mouth at its apex. The first maxillipeds 
consist of two-jointed appendages and are situated just 
behind the second pair of maxillae. The apical joint is 
claw-like and prehensile. The second pair arise mid- 
way between the first and the first pair of feet. They 
are composed of two joints. The basal joint is much 
inflated, and the apical is in the form of a powerful 
claw which closes upon the basal joint and provides 
the animal with a strong grasping appendage. The 
first three pairs of feet consist of an endopodite and an 
exopodite attached to a two-jointed protopodite. The 
endopodite of the first pair is rudimentary and is 
represented by a minute joint bearing a few setae. 
The exopodite is two-jointed. In the second pair both 
the endopodite and the exopodite are three- jointed. 
The protopodite of the third pair is greatly developed 
and forms a prominent lamella. The exopodite and 
endopodite are very small, the former being com- 
posed of two and the latter of three joints. Each of 
the first three pairs of feet is attached to a sternal 
plate. The protopodite of the fourth pair of feet is 
two-jointed and the exopodite is also two-jointed, but 
there is no trace of an endopodite. A strong chitinous 
plate with a bifid apex arises from the middle line 
between the second pair of maxillipeds. This is 
known as the sternal fork. Its function is not clearly 
established. It has been suggested that it acts as a 
crutch to raise the body of the animal from that of the 



INTRODUCTION. 1 1 

host to enable either the mouth-organs or swimming 
feet to have free play. The sternal fork, however, is 
not articulated, neither is it flexible.* 

The external openings are the mouth, the vulvae and 
apertures of the oviducts in the female, the vasa 
deferentia in the male, and the anus. The mouth is 
situated on the ventral surface of the cephalo-thorax, 
and is placed at the apex of a short, conical, movable 
tube. The vulvae are situated on each side of the 
middle line at the posterior end of the genital seg- 
ment and communicate with the receptacula seminis. 
They are difficult to see in the adult female, but each 
has frequently a spermatophore attached which indi- 
cates the position. The openings of the oviducts are 
in the same segment, but nearer the lateral margins 
and just under the fifth feet. The openings of the 
vasa deferentia are situated on the postero-lateral 
margins of the genital segment of the male. The anus 
is in the middle line at the apex of the abdomen. In 
addition to these more important openings, there are 
also apertures of pore-canals and glands on the 
anterior surface of the basal joint of the protopodites 
of the second and third pairs of feet, and also on the 
dorsal surface of the cephalo-thorax and abdomen. 
The opening in some cases is at the apex of a small 
papilla, and communicates with a sac in the interior. 
The function of these glands is uncertain, but it is 
possible that it is connected with the aeration of the 
blood. If a little methylene blue solution be added 
to the sea- water in which the parasites are living, a 
general staining takes place, but the glands are more 
deeply stained than the rest of the body. 

* Fig. 1, PL XIII, shows the general arrangement of the appendages in 
Lepeophtheirus. 



12 british parasitic copel'oda. 

Colour. 

The colour of the living animals varies with the 
position in which they live. On the dark side of the 
fish they are of a deep brown, almost black, colour. On 
the "white " side and under the fins they are nearly 
colourless, due to the contraction of the pigment-cells, 
which appear as brown spots under the microscope. 
The dark-coloured specimens soon become colourless 
when exposed to light. 

Alimentary System, Etc. 

The body- wall consists of (1) chitinous cuticle or 
exoskeleton, (2) cellular hypodermis, and (3) connective 
tissue laminae which line the integument, traverse the 
body-cavity, and support the alimentary canal and 
other organs. The only cavity left inside the body- 
wall is the system of lacunge in which the colourless 
blood flows. 

The mouth, already described, leads into a short 
narrow oesophagus, lined with a thin chitinous coat 
which is continuous with the exoskeleton. The oeso- 
phagus passes through the anterior part of the nervous 
system, and in a transverse section of that region 
appears as a minute pinhole. The stomach lies along 
the ventral surface and is lageniform in shape. At 
the anterior end it is produced into a short cascum, 
which extends over the posterior end of the oesophagus 
and terminates by opening into the intestine. The 
intestine is simply a direct continuation of the stomach 
and terminates in a short rectum leading into the anus 
at the apex of the abdomen. There are no convolu- 
tions in the alimentary canal. The wall of the whole 
alimentary canal is lined with a thin layer of chitin 



INTEOD POTION. 13 

continuous with the exterior. The wall of the stomach 
and intestines is marked by a series of transverse cod- 
strictions, giving it a crenate appearance which is 
easily seen in the living animal. When the animal is 
alive an intermittent movement of the alimentary canal 
is maintained. The action is wave-like. It begins at 
one end and passes slowly to the other. After con- 
tinuing in one direction for a time, it reverses and 
passes along the opposite way. The fluids in the 
alimentary canal are usually colourless. Sometimes, 
however, when taken direct from the fish and placed 
under the microscope, a reddish tint may be observed 
at the posterior end of the oesophagus. Two pairs of 
digestive glands are connected with the alimentary 
canal. The first and most important pair is situated 
in the anterior end of the cephalo-thorax. Each gland 
consists of three portions, two moderately large masses 
on the lateral margins just behind the antennules, and 
a median, smaller one, in front of the base of the 
mouth. The lateral masses are connected with the 
median one by a duct. A duct is given off by the 
median portion and passes posteriorly along the out- 
side of the oesophagus. It enters the caecum at the 
anterior end of the stomach. When the parasite is 
first removed from the host the glands are usually of 
a dark brown colour, but they soon become bleached. 
The product of the glands is a pale yellow fluid. The 
second pair of glands is much smaller than the first, 
and is situated close to the median line between the 
first and second pairs of feet. The glands are of 
a brown colour and shaped like a comma. A minute 
duct from each gland passes downward and then 
forward along the stomach, which it enters near the 
posterior end. 



14 british parasitic copkpoda. 

Circulatory System. 

There is no heart in LepeojMheirus, nor are there 
any proper blood-vessels. The circulation is wholly 
lacunar. The blood-streams simply pass through the 
spaces left among the internal organs and between 
the connective-tissue bands of the body- wall. The 
streams appear to have certain definite courses, but 
they are not uniform, continuous currents. The fluid 
progresses by successive jerks, due to the peristaltic 
movements of the alimentary canal. The blood is a 
clear fluid containing numerous colourless corpuscles 
which vary in size and shape. The corpuscles are 
able to accommodate themselves to the diameter of the 
spaces through which they pass. There are no inde- 
pendent organs of respiration, unless the pore-canals 
and glands in the basal joint of the protopodite of the 
second and third pairs of feet act as such. Hartog* 
and others have suggested that the blood is probably 
aerated from the sea- water contained in the alimentary 
canal by the method of "anal respiration." Further 
precise observations are, however, required to sub- 
stantiate this hypothesis, as the method is so entirely 
different from that in the higher Crustacea. 

Muscular System. 
The muscular system is a very complex one, as 
shown by the illustrations of the dorsal and ventral 
surfaces on Plate XIII. The members of the Cali- 
goida as a general rule are so very transparent, 
however, that the investigation of the musculature 
can be carried out with very little effort. An excel- 
lent description of the muscular system is given by 

* " The morphology of Cyclops and the relations of the Copepoda/' ' Trans. 
Linn. Soc. Lond/ (2), vol. v, pt. 1, pp. 1-46, 1888. 



INTRODUCTION. 15 

C. B. Wilson in his work on "North American Parasitic 
Copepods belonging to the Family Caligidae," Part 1.* 

Nervous System. 

The nervous system of G aligns and LepeopMheirus, 
and probably also of the other members of the Cali- 
goida, appears to be very similar. It is composed of 
two central ganglia and the paired nerves which arise 
from them. One ganglion, the snpra-oesophageal, lies 
above the oesophagus, and the other, the sub-oesopha- 
geal, immediately below it. The ganglia are practi- 
cally fused together by the wide commissures which 
join them. A very narrow opening is left in the 
centre for the passage of the oesophagus. These are 
the only ganglia, and they supply the various parts 
of the body with nerves. The supra-oesophageal 
ganglion is about half the size of the sub-oesophageal. 
It is produced on its dorsal surface into an optic lobe 
from which arises a distinct pair of optic nerves. 
Horizontal sections of the optic lobe show that the 
roots of these nerves cross each other, and each optic 
nerve, therefore, is supplied by fibres from both sides 
of the brain. The only other nerves that have their 
origin in the supra-oesophageal ganglion are those 
which supply the antennules and the antennae. The 
nerves of the antennules divide into a number of 
branches after they enter these appendages, and every 
seta on the surface of the basal joint and apex of the 
second branch is supplied with fibres. It is evident 
that they are important sensory organs. The sub- 
cesophageal ganglion is heart-shaped, and fully twice 
the size of the supra-oesophageal. It supplies the 
nerves to the remainder of the appendages. The 

* ' Proc. U. S. National Museum/ vol. xxviii, 1905. 



16 BRITISH PARASITIC COPEPODA. 

frontal and lateral margins give off seven pairs of 
nerves. Three other pairs arise from the posterior 
end and form a sort of spinal cord. There are thir- 
teen pairs of nerves altogether, as shown in Plate XV, 
fig. 9. Each nerve, after leaving the main trunk, 
sends out numerous branches which pass to the various 
muscles controlling the appendages enervated bj that 
nerve. There is considerable difficulty in tracing the 
endings of the branches when they pass amongst the 
muscles. 

Reproduction. 

The reproductive organs are paired, and as already 
stated the sexes are distinct. In the mature female 
the ovaries appear as comparatively large kidney- 
shaped organs on each side of the anterior end of the 
stomach. An oviduct arises near the anterior end of 
the ventral surface and passes posteriorly as a narrow 
tube until it enters the genital segment. It then 
expands rapidly and becomes convoluted, finally pass- 
ing out to the postero-lateral margins of the segment, 
where it opens to the exterior just under the fifth feet. 
Communicating with the oviduct near its extremity is 
a short semi-transparent cement-gland which secretes 
the enclosing membrane of the ovisac. The vulvae 
are situated near the middle line close to the junction 
of the genital segment with the abdomen. The vulva 
appears to be a simple opening leading into the vagina 
which expands into a receptaculum seminis. This is 
an elongated sac passing from the median line to the 
oviduct, which it enters alongside the duct of the 
cement-gland. 

The sexual organs of the male are very similar in 
position and arrangement to those of the female. 



INTRODUCTION. 17 

They consist of a pair of testes, a long vas deferens 
leading posteriorly from tliem, and a pair of spermato- 
phore sacs in the genital segment. The testes are 
oval bodies situated in a corresponding position to the 
ovaries in the female. The spermatophore is an oval 
body of a pale yellow colour filled with sperms. It is 
expelled from an opening near the posterior end of the 
genital segment. A short cement-gland is situated in 
the segment and is furnished with a duct which is 
connected with the anterior end of the sac. 

The female is fertilized early in life by the male 
attaching sperm atophores to the vulvae. The contents 
of these bodies appear to suffice for the fertilization of 
all the eggs which the female produces during its life. 
The genital segment of an unfertilized female is very 
small. It is only about one-fifth the size it attains in 
the mature condition. The eggs are fertilized from 
the store of sperms as they leave the oviduct, and are 
then enclosed in a thin chitinous tube which gradually 
lengthens as the eggs are expelled. This tube is the 
ovisac, and the eggs are arranged in a single column 
sometimes exceeding the length of the animal. The 
whole of the embryos in the tube practically hatch at 
once when the development is completed. The embryos 
hatch as nauplii about 0-46 mm. in length. The 
nauplii are free-swimmers for a time and then settle 
down as Cyclops forms upon their host. As soon as 
the Cyclops form settles it develops a thin chitinous 
filament from a median gland in the anterior end of 
the cephalo-thorax. The filament is inserted in the 
tissues of the host and the young parasite becomes 
firmly anchored. A median sucker which assists in the 
attachment is also developed at the same time. The 
genus ChalimuSj described by Burmeister in 1831, was 

VOL. I. 2 



18 BRITISH PAKASITIC COPEPODA. 

shown by Hesse and others to be only the cyclops stage 
of the Caligoida. The cyclops form grows, and the 
various appendages make their appearance in regular 
order. When the appendages are developed the 
filament separates at its junction with the frontal 
margin, and the parasite is free to move about over its 
host or to seek a new one. A notch is left in the 
frontal margin when the filament separates, and con- 
tinues all through the adult life. The male, at the 
conclusion of its cyclops stage, is practically fully de- 
veloped. The female remains in an immature con- 
dition until it is fertilized and the ova begin to pass 
down the oviducts. The genital segment then in- 
creases in size to its normal adult condition. 

Tribe Lekn^oida. 

The typical member of this tribe is Lerndea hran- 
chialis Linn. It presents one of the most marked 
examples of retrograde development which is to be 
found in the whole group of parasitic Copepoda. It 
is difficult to believe at a first glance that the parasite 
is a crustacean and that it is related to Lepeophtheirus. 
There is great excuse for the difficulty experienced by 
the earlier zoologists in deciding the true position of 
the members of this family in the animal kingdom. 
Nothing was then known about their life-history, and 
it is not surprising that Linnaeus included Lerndea 
amongst his Vermes. 

The adult female is found on the gills of various 
Gadoids, such as cod, haddock, and whiting. (See 
Plate A.) Immature (cyclops stage) males, and 
females with adult males attached, are to be found on 
the apex of the gill-filaments of flounders and plaice, 
sometimes in lars^e numbers. The adult female is 



INTRODUCTION. 19 

securely attached to its host by strong branched horns 
which are buried in the tissues of the gill-arches. In 
many cases the head will be found to have actually 
penetrated the ventral aorta. The specimens can 
only be removed from the host by careful dissection. 
Any attempt to remove them by force results in the 
head being left in the tissues. The parasite, when once 
fixed, remains in the same position throughout life. 
When it dies the softer parts decay, but the head 
remains embedded in the tissues of the host. The 
heads of dead parasites are frequently met with when 
dissecting-out living specimens. 

The adult female is cylindrical and is a little over an 
inch in length. It is unsegmented, but can be roughly 
divided into three parts — a globular head with anchor- 
like processes, a narrow neck, and a much swollen 
posterior part. The globular head coiTesponds to the 
anterior region of the cephalo-thorax in Le/peopJitheirus. 
It is furnished with three more or less branched horns 
which are outgrowths from the cephalo-thorax. The 
head is slightly curved downwards and terminates in 
a conical apex. The anterior portion of the neck 
represents the remainder of the cephalo-thorax and 
the fourth thoracic segment. The remainder of the 
neck and the greater part of the swollen mass behind 
correspond to the genital segment of Lejpeojplitheirus. 
The abdomen is represented by the terminal portion 
of the swollen part, which gradually tapers to a blunt 
end. The whole of the swollen part is bent into the 
form of the letter S. The neck is marked by fine 
transverse lines. 

The appendages are rudimentary, the greater number 
being entirely absent. The appendages present are a 
pair of maxillipeds immediately under the mouth and 



20 BRITISH PARASITIC COPEKODA. 

four pairs of feet at the anterior end of the narrow 
neck. The feet are exactly as they exist in the fully 
developed cyclops stage, both in size and structure. 
The protopodite is two- jointed. The exopodite of the 
whole four pairs is two-jointed. The endopodite of 
the first two pairs is also two-jointed. The third and 
fourth pairs of feet have no endopodite. 

The external openings are the mouth placed at the 
apex of the head, the openings of the oviducts on the 
ventral aspect of the S-shaped region, and the anus at 
the blunt apex of the abdomen. 

The colour of the living animal is a dark red, due 
to the contained blood. When removed from the fish 
and placed in sea-water the colour disappears. Speci- 
mens of Leriidea removed from their host and placed 
in sea-water do not appear to live longer than twelve 
hours. The parasites are simply inert sacs quite in- 
capable of movement. Occasionally they are covered 
with colonies of hydroids which may entirely obscure 
them. The exoskeleton consists of chitinous cuticle 
moderately thin and soft in the swollen part, but thick 
and hard on the neck and head. 

The mouth of the adult opens directly into the 
alimentary canal. The oesophagus and true stomach 
have entirely disappeared during the metamorphoses 
of the cephalo-thorax. A peristaltic movement of the 
intestine similar to that of Lepeophtheirus can be seen 
in the living animal. There are no indications of 
digestive glands, and it is probable that the digestion 
is intracellular. 

There is no heart in adult Lernaea and no move- 
ment of fluids that would indicate a blood-circulation. 
The animal is probably dependent on the blood sucked 
from the host for the oxygen necessary to maintain 



INTRODUCTION. 21 

life. It is possible that the early death after removal 
from the host is due to the inability to take up oxygen 
from the water. 

The muscular system is represented by a network of 
fibres betw^een the integument and the alimentary canal. 

There does not appear to be any nervous system in 
the adult. 

The reproductive organs of Lernxa, like those of 
Lepeophtheirus, are bilaterally symmetrical, but during 
the metamorphoses of the female the ovaries undergo 
a great change of position. They are displaced from 
the cephalo-thorax and pass into the genital segment, 
where they occupy a narrow region at the apex of the 
deep indentation. The two ovaries are practically 
fused together. The oviducts arise from their anterior 
ends, pass across the segment to its ventral surface, 
and then course along each side of the median line to 
the external openings. The cement-glands are nearly 
of the same length and breadth as the oviducts and lie 
underneath them. The posterior end communicates 
with the oviduct just inside the opening to the 
exterior. The ovisacs consist of long slender con- 
voluted tubes which may reach the length of eight 
inches when straightened out. The eggs are arranged 
in a single column. The female is fertilized shortly 
before the completion of the cyclops stage. It is 
probable that the sperms pass up the oviduct and 
fertilize the eggs very soon afterwards, as no trace of 
a receptaculum seminis can be made out in the adult. 

The young Lernsea hatches as a nauplius with three 
pairs of appendages similar to those of the Lepeo- 
phtheirus nauplii. It leads a short pelagic life and 
then settles on the gill-filaments of the flounder and 
plaice and occasionally other fishes. It attaches itself 



22 BRITISH PARASITIC COPEPODA. 

to the filament by a thin broad cliitinous plate and by 
the claw-like antennae. The young parasite assumes 
the Cyclops form and continues its progressive develop- 
ment. The various appendages and internal organs 
make their appearance. The male reaches maturity 
without altering from the cyclops form, and undergoes 
no further change. A considerable lengthening of 
the female genital segment accompanies the develop- 
ment of the various appendages. Fertilization takes 
place, and the young female severs its connection with 
the gill-filament. It leads a pelagic life for a time, 
and is occasionally found in surface-plankton. The 
males, unless accidentally separated, remain on the 
gill- filaments after the females have gone. 

The Cyclops form of Lernsea appears to cause con- 
siderable damage to the gills of its temporary host. 
The whole of the apex of the gill-ray assumes a tumid 
appearance, and a considerable number of the gill- 
filaments disappear from both sides. (See Plate A.*) 
When the fertilized female leaves the gills of the 
flounder or plaice the appendages and internal organs 
have practically completed their development. A pair 
of eyes similar in structure to that of adult Lepeo- 
phtheirus is present on the anterior region of the 
cephalo-thorax. The appendages are represented by 
one pair of four- jointed antennules, one pair of claw- 
like antennas, one pair of mandibles which are not 
enclosed in the mouth-tube, one pair of maxillae, two 
pairs of maxillipeds, and five pairs of feet. The body 
is divided into five distinct segments—the cephalo- 
thorax, three thoracic segments, and one terminal 
segment which represents the genital segment and 
abdomen. The whole of the terminal segment is 

* See also Plate XLII, fig. 8. 



INTRODUCTION. 23 

marked by fine transverse lines. The colour of the 
animal varies from dark violet to light red. The 
alimentary canal is similar in structure to that of 
the adult LepeojMheirus. A large digestive gland 
occupies each side of the cephalo -thorax. The 
nervous system of the Lernsean cyclops is practically 
similar to that of the adult Lepeojphtheirus. The 
ovaries occupy a normal position. 

At the conclusion of its pelagic life the Lernsean 
cyclops fixes itself to the gills of a fish, and the retro- 
gressive metamorphoses commence. The parasite 
buries its head in the tissues of the host and then 
develops three horns. The horns are simple at first, 
but by gradual division they acquire the adult type. 
The genital segment elongates very considerably. 
The eyes, antennules, antennae, mandibles, maxillge, 
and one pair of maxillipeds completely disappear. 
One pair of maxillipeds persists all through life, and 
is represented in the adult by a pair of small hooks 
placed under the mouth. 

The next phase, repres(3nted on Plate XLII, shows 
that the development of the horns, the disappearance 
of the various appendages, and the great lengthening 
of the genital segment are followed by a looping of the 
posterior region at that segment. This loop gradually 
expands, and finally takes on the adult condition. 

The Food of the Parasites. 

There appears to be considerable doubt regarding 
the precise food of the Copepod Parasites of Fishes. 
Some authors conclude that, because no red colour can 
be seen in the alimentary canal, such families as the 
Caligoida feed entirely upon the mucus of the host's 
body. It has been suggested, therefore, that Lepeo- 



24 BRITISH PARASITIC COPEPODA. 

phtheirus and allied forms are not parasites in the strict 
sense of the term, and may not be hurtful to their 
hosts. There is little doubt about the food of Lenidea. 
It is generally found in a position that is plentifully 
supplied with blood. Its mouth is buried out of the 
reach of mucus, and is often in direct communication 
with the main blood- stream entering the gills to be 
purified. Caligoids from the gill-chamber and the 
mouth seldom show any trace of red colour. Degene- 
rate forms such as Glavella and Ghondr acanthus, 
although actually living amongst the gill-filaments, 
are usually of a pale yellow colour, with occasionally a 
grey central streak indicating the alimentary canal. 
Lernanthropus Jcroyeri, which lives amongst the gill- 
filaments of Lahrax lupus, is nearly always of a deep 
brownish-red colour. It is said that mucus at the best 
is a poor food, but Lepeophtheiriis can live for upwards 
of six weeks in filtered sea- water without visible food 
of any kind. Mucus may not be a very sustaining form 
of nourishment, but as the Caligoids apparently do not 
require much food they may be able to obtain sufficient 
from the mucus. C. B. Wilson says : '' There is more 
mucus on the scales than anywhere else ; why should 
they choose the fins or inside of the operculum? " The 
fins and gill-chamber afford a certain amount of pro- 
tection to the parasites, and it is just as likely that 
they choose these places for shelter as for food. It 
has to be remembered, however, that the difference 
between the Caligoida and the obviously blood-sucking 
Lernsea is very great. The Caligoids are provided 
with large digestive glands which are entirely absent 
in the adult female Lernsea. When one finds blood in 
the alimentary canal of a parasite which has no digestive 
glands, and none in those which are provided with 



INTRODUCTION. 25 

sucli organs, it may reasonably be concluded that the 
character of the food will be rapidly altered. The 
fluid from the glands meets the food immediately it 
enters the stomach of the Caligoids and acts upon it at 
once. If blood be the food, the red colour may be 
destroyed, and give rise to the impression that Cali- 
goids are not blood-suckers. 

Nothing appears to be known regarding the internal 
organs of the Copepod Parasites of Fishes with the 
exception of the Caligoida and Lernasa. When the 
anatomy of all the families has been investigated 
the food question may be settled. One or two para- 
sites on a fish may not be hurtful, but when the 
numbers increase they probably have an irritating 
effect, and finally, when they remain in one position 
for some time, the skin and tissues become badly 
lacerated, giving rise to tumours. The Medesicaste, 
which lives on the gills of various species of gurnards, 
occasionally has its head buried in a tumour-like 
swelling, but we have been unable to determine 
whether the tumour existed previous to the attach- 
ment of the parasite or was entirely due to the 
congestion of the blood-vessels caused by its attack. 
Gr. H. Drew, in a paper entitled " Some cases of new 
growths in Fish " (Journ. Mar. Biol. Assoc, N. s., 
vol. ix, No. 3, June 1912), describes this tumour-like 
swelling in the gurnard as a case of hsemangiomata, and 
regards it as probable that the parasite attacked the 
tumour after it had developed, but we have never come 
across the tumour without the parasite, or the remains 
of it, being present. It is quite as likely, therefore, 
that the haemangiomata of the gurnard described by 
Drew is caused by the parasite. (See also our remarks 
on Lernsea, p. 22.) 



SYSTEMATIC ARRANaBMENT. 

In this account of the Copepoda found parasitic on 
British fishes, the arrangement outlined by Professor 
G. 0. Sars in the introduction to voL iv of his great 
work on the ' Crustacea of Norway ' has, so far as it 
concerns these parasitic species, been followed by us. 

Professor Sars arranges the Copepoda into seven 
divisions or tribes, three of which are represented 
below — viz., the Cyclopoida, the Caligoida, and the 
Lernseoida. The Cyclopoida show considerable varia- 
tion in regard to their habits and manner of life ; some 
are entirely free-living (as Gijclops), others are asso- 
ciated with different organisms either as commensals 
or messmates, their commensal! sm tending in some 
cases in the adult stage towards a more or less true 
parasitic life. Several species belonging to this tribe 
are associated with fishes. 

The Caligoida, which are almost all parasitic on 
fishes, also exhibit considerable differences in their 
habits, but although none of them can be accurately 
described as free-living in the adult stage, some species, 
such as Galigus rapax, possess considerable freedom of 
movement, and have, in consequence, a tolerably wide 
distribution. In the majority of cases, however, the 
Caligoida, though not usually permanently fixed to 
their host, are limited in their movements, and do not 
apparently, to any extent, migrate from one fish to 
another ; and not only are they more or less confined to 
particular fishes but sometimes also to particular parts 
of the fish, hence such names as Galigus zei and 
Lepeoplithelriis pectoralis. In the first case the parasite 
is usually found only on the dory {Zeus faher), while 
in the second it occurs chiefly on the pectoral fins 
of flat fishes, such as plaice, flounders, and dabs. 

The Lernseoida differ from the other two tribes or 
divisions by the females in the adult stage being all 



SYSTEMATIC ARRANGEMENT. 27 

more or less permanently fixed to their host. This 
group of parasites was arranged by M. Edwards into 
three families — viz., the Chondracanthiens, the Lerneo- 
podiens, and the Lerneoceriens. The Chondracan- 
thiens " are fixed by the aid of stout foot- jaws armed 
with strong hooks." The Lerneopodiens have the 
second maxillipeds modified, and furnished, at their 
extremity, with a chitinous horn-coloured process, 
which, penetrating the gills, gill-covers, or some other 
part of the fish, forms a permanent attachment to it, 
while the head of the parasite is free and apparently 
possesses to a small extent a certain freedom of move- 
ment. In this family the male is very small and is not 
attached to the fish, but is usually found adhering to 
the body of the female, two or three males being some- 
times found on the same female. The third family of 
M. Edwards, the Lerneoceriens, are fixed to their host 
by having the head, sometimes with its appendages, 
buried in its tissues. This arrangement of Milne 
Edwards, slightly modified, was adopted by Dr. Baird 
in his ' British Entomostraca,' and is to some extent 
still adhered to. 

Various attempts have been made to frame a classifi- 
cation of the Copepoda which would include the para- 
sitic and semi-parasitic forms, but, with the exception 
of that of Professor Gr. 0. Sars, they have proved 
more or less unsatisfactory, and therefore, as already 
stated, we have, as regards these parasitic species, 
adopted the division proposed by Sars. Referring to 
the various attempts at classification, C. B. AYilson, 
in a recent article on the subject, remarks : "It 
requires about as thorough a knowledge of the Cope- 
poda to select intelligently from these various schemes, 
backed as they are by competent authority, as it 
would to construct an original scheme. Indeed most 
authors have apparently found the latter the easier, 
since each has propounded a scheme of his own."* 

* "The Classification of the Copepods/' by C. B. Wilson, in ' Zool. 
Anzeiger/ vol. xxxv, No. 20, 26th April, 1910, p. 611. 



28 BRITISH PARASITIC COPEPODA. 

Dr. Bassett-Smith in his excellent paper on " Para- 
sitic Copepoda found on Fishes," published in 1899,* 
follows, with some slight modifications, the classifica- 
tion proposed by Grerstaecker in Bronn's ' Tierreich,' 
1881, and as Sars' main divisions fit in fairly well with 
this arrangement it has been retained here. 

The Gydopoida. 

The Cyclopoida are represented among the Cope- 
poda parasitic on British fishes by a single family, 
the Ergasilidae, which comprises the three genera 
Ergasilus, Bomolochus, and Thersitina, 

The Galigoida. 

The Galigoida are represented by many genera and 
species ; they are arranged under three families — viz., 
the Caligidae, the Dichelestiidae, and the Philichthyida3. 
The Caligidae have been divided into the following sub- 
families : the Caliginae, Trebinge, Euryphorinse, Pan- 
darinae, and Cecropinae ; but with the exception of 
the Caliginae, which comprise Caligus, Lepeophtheirus, 
and one or two other genera, these subdivisions are 
each represented by only a small number of species, 
and it has, therefore, not been considered necessary to 
adopt these minor divisions here. 

The LerndBoida. 

The Lernaeoida also comprise a considerable number 
of genera and species, and these, like the Caligoida, 
are arranged under three families — viz., the Lernaeidae, 
the Chondracanthidae, and the Lernaeopodidae. These 
represent three tolerably distinct types, and exhibit in 
the adult female a more or less retrograde develop- 
ment, producing in many cases results, in regard to 

* " A Systematic Description of Parasitic Copepoda found on Fishes, with 
an Enumeration of the Known Species," in ' Proc. Zool. Soc. London,' 18th 
April, 1899. 



SYSTEMATIC ARRANGEMENT. 29 

their form and structure, of a remarkable kind, as 
indicated in the life-history of Lernsea already referred 
to. In the family Lernseidse the body of the adult 
female is greatly distorted, and the head is buried 
deeply in the tissues of the host; on the other hand, 
the mature male, which, as shown in Plate XLII, 
fig. 5, closely resembles the young female before it has 
begun to assume that abnormal development charac- 
teristic of the adult stage, does not, so far as known, 
undergo any further change, but continues during its 
short life to retain its normal structure and form. 
The Chondracanthidas differ from the other two 
families by the manner of their fixation, and also by 
the peculiar form of the mandibles, which are unlike 
those of any of the other parasitic groups described 
here. The Lernseopodidse are also distinguished by 
their mode of fixation, as shown elsewhere. 

The Linnsean species Lernsea asellina, which was 
ascribed by Blainville and Baird to Lernejitoma, and 
by Nordmann and others to GhondracantJms, was re- 
moved to a new genus, Oralien, by Bassett-Smith, on 
account of some peculiar structural differences. But 
while appreciating this, and after carefully comparing 
the characters of Oralien with those of the older genus 
Medesicaste Kroyer, we are inclined to think that the 
difference between them is scarcely of sufficient im- 
portance for the establishment of a new genus — a 
conclusion previously come to by Brian.* We have 
therefore transferred this Linnaean species to Kroyer' s 
genus Medesicaste. Moreover, as Sphyrion appears to 
be a true Lernasan rather than a Chondracanthian 
genus, with which family it is sometimes associated, 
it has been transferred by us to the family Lernseidae. 

* * Copepodi parassiti dei Pesci d'ltalia/ p. 94. 



SYSTEMATIC LIST 
OF THE SPECIES DESCRIBED IN THIS VOLUME. 



I 



Order COPEPODA. 
Tribe I. Cyclopoida G. O. Sars. 
Family i. ERGASILID^. 
Ergasilus Nordmann. 2. onosi T. Scott. 

L nanus Clans. 3. zeugopteri T. Scott. 

BoMOLOCHUS iSTordmann. 3. Thersitina Norman. 

1. so/ea^ Clans. 1. gras^eros <ei Pagensteclier. 

Tribe II. Caligoida G. O. Sars. 

Family ii. CALIGID^. 

Caligus O. F. Miiller. 
1. curtus O. F. Miiller. 



[NoGAUs Leach. 

1. ambiguus T. Scott.] 



2. mi7nmus Otto. 

3. rapax M. Edwards. 

4. ceidrodonti Baird. 

5. gumardi Kroyer. 

6. labracis T. Scott. 

7. zei Norman. 

8. hrevicaudatus A. Scott. 

9. ipelamydis Kroyer. 
10. diaphanus Nordmann. 

PSEUDOCALIGUS A. Scott. 

1. hrevipedes Bassett-Smith. 

SciiENOPHiLUS p. J. van 

Beneden. 
1. tenuis P. J. van Beneden. 
Lepeophtheirus Nordmann. 

1. pectoralis O. F. Miiller. 

2. novdmanni M. Edwards. 

3. hippoglossi Kroyer. 

4. thomsoni Baird. 

[ohscurus Bassett-Smith.] 

5. salmonis Kroyer. 

6. pollachii Bassett-Smith. 

7. sturionis Kroyer. 

LiJTKENiA Clans. 
1. asterodermi Clans. 



9. 



Demoleus Heller. 
1. paradoxus Otto. 

10. Trebius Kroyer. 

1. caudatus Kroyer. 

11. Elytrophora Gerstaecker. 

1. brachyptera Gerstaecker. 

12. DiNEMOURA Latreille. 

1. producta O. F. Miiller. 

13. EcHTHROGALEUS Stecnstrnp 

& Liitken. 

1. coleoptratus Guerin-Men- 

neville. 

2. liltTieni Norman. 

14. Phyllothreus Norman. 

1. cornutus M. Edwards. 

15. Pandarus Leach. 

1. bicolor Leach. 

16. Cecrops Leach. 

1. latreillii Leach. 

17. Orthagoriscicola Poche. 
1. muricata Kroyer. 

18. Philorthragoriscus Horst. 
1. serratus Kroyer. 



LIST OP SPECIES. 



31 



Family in 

19. DiCHELESTiTJM Hermann. 

1. ohlongum Abildgaard. 

20. Anthosoma Leach. 

1. crassum Abildgaard. 

21. Lernanthropus Blainville. 

1. hroyeri P. J. van Beneden. 

22. Hatschekia Poclie. 

1. Mppoglossi Kroyer. 

2. mulli P. J. van Beneden. 

3. labracis P. J. van Beneden. 

4. cluthse T. Scott. 

5. cornigera T. Scott. 

6. pygmasa T. Scott. 



DICHELESTIID^. 

23. Kroyeria p. J. van Beneden. 
1. lineata P. J. van Beneden. 

24. CoNGERicoLA P. J. van 
Beneden. 

1. pallida P. J. van Beneden. 

25. EuDACTYLiNA P. J. van 
Beneden, 

1. acuta P. J. van Beneden. 

2. acanthii A. Scott. 

3. similis T. Scott. 

4. ininuta T. Scott. 

5. insolens T. & A. Scott. 



27. 



29. 



80. 



36. 



Family iv. PHILICHTHTID^. 

26. Philichthys Steenstrup. 
1. xiphiae Steenstrup. 

Tribe III. Lernaeoida G. O. Sars. 
Family v. LERN^ID^. 



Lern^a Linnaeus. 

1. hranchialis Linnaeus. 

2. lusci Bassett-Smith. 

3. ininuta T. Scott. 

4. lumpi T. Scott. 

H^MOBAPHES steenstrup & 

Liitken. 
1. cyclopterinus O. Fabricius. 

H^MOBAPHOIDES T. & A. 

Scott. 
1. amhiguus T. Scott, 

Pennella Oken. 
1. orthagorisci E. P. Wright. 



31. Lern^ocera Blainville. 

1. cyprinacea Linngeus. 

32. Lern^enictjs Lesueur. 

1. sprattse Sowerby. 

2. encrasicola Turton. 

33. Trypaphylus Richiardi. 

1. musteli P. J. van Beneden. 

34. Rebeltjla Poche. 

1. edwardsi Kolliker. 

35. Sphyrion Cuvier. 

1. lumpi Kroyer. 



Family vi. CHONDRACANTHID^. 



Chondracanthus De la 

Roche. 

1. cornutus O. F. Miiller. 

2. aniiulatus Olsson. 

3. soleie Kroyer. 

4. flurse Kroyer. 

5. depressus T. Scott. 

6. limandie Kroyer. 



37 



7. clavatus Bassett-Smith. 

8. nodosus O. F. Miiller. 

9. zei De la Roche. 

10. lophii Johnston. 

11. merhtccii Holten. 

12. 07-natus T. Scott. 
Medesicaste Kroyer. 

1. asellinum Linnaeus. 



32 



ElUTISH PAEASITIC COPEPODA. 



Family vii. 

38. Thysanote Kroyer. 

1. impudica Nordmann. 

39. Chaeopintjs Kroyer. 

1. dalmanni Retzius. 

2. dubius T. Scott. 

3. ramosus Kroyer. 

40. AcHTHEEES Nordmann. 

1. percarum Nordmann. 

41. Lern^opoda Blainville. 

1. elongata Grant. 

2. galei Kroyer. 

3. cluthse T. Scott. 

4. salmonea Linnaeus. 

5. bidiscalis de Visme Kane. 

6. similis T. & A. Scott. 

7. lamjpri T. & A. Scott. 

42. Brachiella Cuvier. 

1. thynni Cuvier. 

2. rostrata Kroyer. 



LERN^OPODID^. 



3. insidiosa Heller. 

4. merluccii Bassett-Smitli. 

5. triglm Claus. 

6. ovalis Kroyer. 

7. bispinosa Nordmann. 

8. ^as^zViacaP. J. van Beneden. 

9. parTieri G. M. Thomson. 
43. Clavella Oken. 

1. uncinata O. F. Muller. 

2. rugosa Kroyer. 

3. dubia T. & A. Scott. 

4. emarginata Kroyer. 

5. scombri Kurz. 

6. brevicolis M. Edwards. 

7. stellata Kroyer. 

S. paradoxa P. J.vanBeneden. 
9. quadrata Bassett-Smith. 

10. canthari Heller. 

11. alata Brian. 

12. lophii M. Edwards. 



CLASS CRUSTACEA. 

SUB-CLASS ENTOMOSTRACA. 

Order COPE POD A. 
Tribe I. CYCLOPOIDA G. 0. Sars. 

Cephalothorax ovate and usually more robust than 
the abdomen. Antennules slender and elongated or 
tolerably short and stout; those of the male alike on 
both sides, sometimes modified for grasping. 

Antennae unbranched, or the outer branch rudimen- 
tary ; usually feebly armed, but sometimes stout or 
elongated and provided with strong terminal claws. 
Maxillipeds usually less developed than in the Cala- 
noida, but the second pair sometimes strongly uncinate. 

First four pairs of swimming legs as in the Cala- 
noida; fifth pair rudimentary, alike in both sexes, 
usually one- but sometimes two-jointed, rarely with 
more than two joints. Egg-strings two. 

The Cyclopoida found on British fishes all belong to tbe 
one family Ergasilidge, and to the three genera Ergasilus, 
Bomolochus, and Thersitina. The species comprised within 
these three genera are associated with fishes of various kinds. 
They cannot all be accurMtely described as parasites, some of 
them, such as Bomolochus solese, should rather be regarded as 
commensals or messmates; they appear to be able, within 
certain limits, to move about with considerable freedom. 
Their food appears for tlie most part to consist of the mucus 
exuded by the fish, and it is doubtful if they would long sur- 
vive if removed from their host. 

Family i. Ergasilid.e. 

Body more or less cyclopoid in form. First segment 
usually large, sometimes subglobose in the female. 
Antennules composed of five to seven joints. Antennae 

VOL. I. 3 



34 BRITISH PARASITIC COPEPODA. 

two- or three-jointed, more or less prehensile, and 
usually armed with terminal claws. Mandibles small. 
Maxillae rudimentary. First maxillipeds small and of 
a simple structure. Second maxillipeds well developed 
and jDrovided with strong terminal claws. First foui* 
pairs of thoracic legs biramose ; both rami of the first 
three pairs composed of three articulations, but in the 
fourth pair, while the inner ramus is three- jointed, as 
in the other three pairs, the outer may consist eithei* 
of three joints or only of two. Fifth pair small, one- 
branched, uniarticulate or rudimentary. Egg-strings 
two. 



Genus 1. ERGASILUS Isovdrnmin, 1832. 

Somewhat like Cyclops in general appearance. 
Cephalothorax elongated, segments five, the first 
seo'ment laro-e. 

Antennules composed of five or six joints. Antennge 
elongated and composed of three joints, terminal claws 
large, and forming powerful grasping organs. First 
four pairs of thoracic legs biramose ; both rami of the 
first three pairs three-jointed ; fourth pair with the 
inner ramus three- and the outer two- jointed. Fifth 
pair small or rudimentary. Egg-strings two. 



1. Ergasilus nanus P. J. van Beneden. 

(Plate I, fig. 1; Plate 11, figs. 1-5; Plate XL VIII, 

fig. 17.) 

1870. Ergasiltts nanus P. J. van Beneden. (16) Les Poissons des cotes 
de Belj^ique, leurs parasites et leurs commensanx, p. 27, pi. i, fig. 6 
(recorded and figured but not described in this work). 

1901. Ergasilus nanus T. Scott. (113) p. 122, pi. vii, figs. 1-8. 

Female, — Body somewhat expanded anteriorly, pos- 
terior extremity attenuated. Cephalic segment large, 
and if viewed from above, somewhat pear-shaped in out- 
line ; the widest part, which is near the anterior end of 
the segment, equal to fully half its length, and a shallow 



ERGASILUS NANUS. 6d 

but obvious constriction occurs near its middle. The 
remaining thoracic and the abdominal segments small. 

Antennules short, apparently only five-jointed, and 
sparingly setiferous ; the first joint about twice the 
length of the second, the fourth rather smaller than any 
of the others. Antennse tolerably slender and greatly 
elongated, composed of three joints, and furnished with 
moderately strong and curved terminal claws. Mouth- 
appendages apparently more or less rudimentary or 
obsolete. 

The first four pairs of thoracic legs all biramose and 
well developed ; the first three pairs with both the 
inner and outer rami three-jointed, but in the fourth 
pair while the inner ramus is three-, the outer is only 
two-jointed, the first joint being about twace as long- 
as the end one. The fifth pair small, and consisting 
each of a single one- join ted branch which is about 
three times longer than broad, with the lateral margins 
subparallel, and the distal end truncated and bearing- 
two or three moderately long apical setse. Caudal 
rami short. Egg-strings two, large, more than halt 
the lengfth of the animal and containine- numerous 
ova. Length, exclusive of egg-strings, 1-2 mm. 

Habitat. — Parasitic on the gills of the grey mullet, 
Mugil chelo (Giwier). Bay of Nigg, Aberdeen, in July, 
1900 {T. Scott). Swan Pool, Falmouth {A. M. Norman. 
1884). 

This appears to be a rare species ; though we have ex- 
amined several specimens of the grey mullet, we have only 
once observed it. Another species, E. sieholdii, found parasitic 
on carp, pike, and some other fresh-water fishes, appears to 
have a wide distribution on the Continent and may vet be 
obtained in Britisli waters. 

We have not seen the male of Ergasihis nanus, nor have we 
seen any published record of its occurrence ; the male of E. 
sieholdii, on the other hand, appears to be the form which 
has been most frequently met with. Though several other 
species of Er<jasilics are recorded, E. nanus is apparently the 
only British representative of the genus. 



36 BRLTISH PARASITIC COPEPoDA. 



Genus 2. BOMOLOCHUS Novclmann, 1832. 

Body elongated, subpjriform, and not unlike a 
Cyclops in general appearance. Antennules tolerably 
short and composed of about seven joints. Antennas 
small, three-jointed. Mandibles small, simple, and 
furnished with unequal tooth-like processes. Maxillae 
somewhat rudimentary. First maxillipeds small and. 
two-jointed, the end joint attenuated, and provided- 
wath a few marginal denticles and also with a mode- 
rately stout marginal plumose seta which reaches to 
about the apex of the joint. Second maxillipeds large, 
composed of two joints, and armed with a terminal 
claw% which is recurved in tlie female but not in the 
male. Swimming-feet all biramose and with both 
rami three-jointed ; the joints of the first pair lamelli- 
form, and furnished with spathulate and densely- 
plumose setae. Fifth pair of feet small and consisting 
of a single biarticulated l)ranch. 

The male does not differ greatly from the female, 
except in the structure and armature of the second 
maxillipeds. 

1. Bomolochus soleae Glaus. 
(Plate T, fig. 3 ; Plate II, figs. 6-9 ; Plate III, figs. 1-4.) 

1864. Bomolochus solex Claus. (33; Zeitscbr. f. wiss. ZooL, vol. xiv. 
p. 374, pi. XXXV. 

1893. Bomolochus solex T. Scott. (Ilia) Elevontli Annual Report 
Fishery Board for Scotland, pt. iii, p. 21'J. pi. v. 

1902. Bomolochus solex idem. (114) p. 288, pi. xiii, figs. 13-18. 

1906. Bomolochus soleai A. Brian. (21) Copepodi Parassiti dei Pesci 
d'ltalia, p. 31. 

1909. Bomolochus solese May E. Bainbridge. (3) Trans. Linn. Soc, 
ser, 2 (zoology), vol. xi, pt. 3, p. 45, pi. viii. 

Female. — Cephalic segment short, considerably ex- 
panded, widest in the middle, foi ehead flattened, sides 
rounded, length equal to rather more than half the 
width ; each of the two thoracic segments which follow 
are about equal to the length of the cephalic segment, 



BOMOLOCHUS SOLE^. o7 

but each segment is narrower than that which precedes 
it ; the other thoracic segments small, and partly con- 
cealed in dorsal view. Abdominal segments also 
small. 

Antennules moderately short and stout, composed of 
seven joints, and furnished with long plumose setae ; 
the joints subequal in length but the fourth rather 
longer than either the third or the fifth. Antennae 
three-jointed, middle joint short, end joint provided 
with three pectinated, setiferous appendages and a few 
apical setge. Mandibles small, but moderately stout, 
and armed with two small, oval, terminal plates serrated 
on the edges. First maxillipeds small, each with a 
stout, setiferous, terminal spine, from the side of which 
springs a minute accessory spinule. Second maxil- 
lipeds having the basal part considerably dilated, but 
the distal portion slender, curved, and claw-like. The 
first four pairs of thoracic legs Avith both the inner 
and outer ramus tiiree- jointed ; in the first pair both 
rami broadly lamelliform, somewhat distorted, and 
furnished with densely plumose setae, the middle joint 
rather shorter than the first or third. The second and 
third pairs with both rami longer and much narrower 
than those of the first pair, and furnished with densely 
plumose setee on the inner margins and short stout 
spines on the outer ; under the microscope these 
spines are seen to end in minute hooks. The inner 
ramus in the fourth pair narrower and rather longer 
than the outer, and bearing a single plumose seta on 
the inner distal angles of the first and second joints, 
and also three apical setae, the middle one being 
longer than the one on either side. Fifth pair con- 
sisting each of a single tw^o-jointed ramus, the first 
joint being very short, the other of moderate length 
and bearino^ one marsfinal and three terminal setae. 
Caudal rami short. Length about 1*3 mm. 

Habitat. — Found parasitic on the back of the common 
sole, Solea vulgaris Quen., and in the nasal fossae 
of various fishes, especially of cod-fishes : it has been 



38 BRITISH PARASITIC COPEPODA. 

found much more frequently in the nostrils of the cod 
than in those of any other fish we have examined. 
The following is a list of the fishes on which we have 
obtained this Bomolochus : — 

1. Lumpsucker, Cijclojiterus Inmpus L., in the 
nostrils, rather rare. 

2. Cod fish, Gadus callarins L., in the nostrils, fre- 
quent. 

3. Haddock, Gadus seglefimis L., in the nostrils, not 
common. 

4. Whiting, Gadus m£rJan(jvs L., in the nostrils, 
not common. 

5. Pollack or lythe, Gadus poUachius L., in the 
nostrils, moderately frequent. 

6. Ling, Mohta, m.olva L., in the nostrils, not com- 
mon. 

7. Plaice, Pleuronectes pJafessa L., in the nostrils, 
not very rare. 

8. Flounder, Pleuronectes flesus L., in the nostrils, 
rare. 

9. Common or black sole, Solea mJgaris Quen., on 
the back of the fish, not common. 

Bomolochus solese has been obtained at various places round 
the British Islands on one or other of the fishes mentioned. 
The specimens obtained on the ling-fish are ratlier larger 
than the typical form from the black sole or the cod-fish, but 
they do not appear to differ much otherwise. As many as 
twenty-nine specimens of Bomolochus have been obtained in 
the nostrils of a single large cod-fish captured in the Moray 
Firth. Frequently, not only adult males and females (the 
latter commonly with ovisacs), but also young ones in all 
stages of development, may be observed in the nostrils of the 
cod-fish, thus indicating clearly enout>-h that their presence in 
this curious domicile is not accidental, and that they find the 
conditions of life fairly satisfactory amid such surroundings, 
even though the accommodation is somewhat limited. This 
habit on the part of the Bomolochus is the more interesting 
when there is apparently notliing to hinder them from leaving 
the nostrils of the fish. They can move about freely amongst 
the mucus with which the nostrils are usually well supplied, 
and if they be removed and placed in a vesse) of clear sea- 



BOMOLOCHUS SOLE^. 39 

water they will be seen swimming or running about with 
nearly as much agility as a " free-swimming '^ species. It is 
fairly evident therefore that the presence of these Copepods 
in the nostrils of fishes is a matter of choice, and there is 
nothing to show that their presence is the cause of much 
inconvenience to the fish. Moreover, in this situation there 
are two points in favour of the Copepod — it has plenty of food 
of a kind, and it is well sheltered from its enemies. 

The male does not differ much from the female except that 
the second maxillipeds are armed with more powerful 
terminal claws. 

2. Bomolochns onosi T. Scott. 
(Plate I, fig. 4 ; Plate III, figs. 5-7.) 

1902. Bomolochns onosi T. Scott. (114) p. 289, pi. xiii, figs. 19-22. 

Female. — Body elongated, subpyriform ; the an- 
terior segment proportionally larger than the same 
segment in BomolocJius solese, being eqnal to nearly 
half the entire length of the cephalothorax ; the next 
three segments subequal in length, but each narrowei* 
than the one that precedes it. Abdomen narrow and 
moderately elongated, genital segment twice the 
length of the next one, other segments smaller, but 
the anal rather longer than either of the two preced- 
ing segments. Caudal rami about as long as the anal 
segment. 

Antennules moderately stout, but becoming some- 
what attenuated towards the distal extremity ; the 
first two joints tolerably large, the second being about 
one and a half times the length of the next, the third 
and fourth joints subequal and larger than those 
which follow, the penultimate joint smaller than any 
of the others ; the plumose setse with which the anten- 
nules are furnished, though somewhat similar to those 
on the antennules of Bomolochus solese, are apparently 
not quite so stout. Antennae, mandibles, and maxillas 
somewhat similar to the same appendages in Bumo- 
locJius solex. Second maxillipeds robust, and each armed 
with a tolerably stout and strongly curved terminal 
claw ; a slender seta also springs from the distal end 



40 BKITISH PARASITIC COPEPODA. 

of the inner margin of the first joint, and two smaller 
setae from near the middle of the second joint, while 
from the proximal half of the terminal claw there 
spring two slender bristles of unequal length, the 
longer one extending considerably beyond the end of 
the claw. 

Thoracic legs somewhat similar in structure and 
armature to those of Bomoloclms sohse. Length about 
1*3 mm. 

The species is colourless, with the exception of a 
slight chalky whiteness along the alimentary tract. 

Male. — The male of Bomoloclms onosi differs little 
from the female, except that the terminal claw of the 
second maxillipeds is more powerful, more evenly 
curved, and has its inner edge minutely dentated. 

Habitat. — Found parasitic on the inner surface of 
the gill-covers of four-bearded and five-bearded rock- 
lings, Onos (Motella) cimhrius and Onos {Motclla) 
mustelns. Firth of Forth, Moray Firth, and Bressay 
Shoal, east of Shetland {T. Scott). 

The species described above has a close resemblance to 
Bomolochus soleas, and might easily be mistaken for it. The 
most obvious difference is perhaps in the form and armature 
of the posterior maxillipeds; this difference is so marked that 
it may be detected even without dissection, when the creature 
is viewed from the ventral aspect witli the aid of a hand-lens. 

3. Bomolochus zeugopteri T. Scott. 
(Plate I, fig. 2 ; Plate III, figs. 8, 9.) 

1902. Bomolochus zeugopteri T. Scott. (114) p. 290, pi. xiii, figs. 23-25. 

Female. — Cephalic segment equal to the combined 
length of the next three segments, broadly rounded in 
front, rostrum not very prominent ; the next segment 
nearly as wide as the preceding one, and about half as 
long, the remaining segments narrower and shorter. 
Abdomen narrow, the genital segment about as long- 
as the combined length of the next three segments, 
the last two segments subequal and rather shorter 
than the second one. Caudal rami short. 



BOMOLOCHUS ZEUGOPTEEI. 41 

Antennules tolerably stout, shorter tlian the cephalo- 
tliorax and bearnig numerous and moderately stout 
plumose set^, first and second joints large, and, to- 
gether, about as long as the combined length of the 
next four, the third and fourth joints subequal, and 
the penultimate joint shorter than any of the others. 
Second maxillipeds tolerably stout, the terminal claw 
only slightly curved at the end, but at the base bent 
backward upon the preceding joint. Thoracic legs 
somewhat similar to those of Bomolochus onosi. Length 
about 0"84 mm. 

Xo males of this species have been observed. 

Habitat. — Found adhering to the back of a small 
Miiller's top-knot, Zeucfojjterus punctatus (BL), cap- 
tured near the mouth of the Clyde estuary in Septem- 
ber, 1897. About a dozen specimens, all females, and 
most of til em carrying ovisacs, were obtained ; they 
were adhering firmly on the rough dark-coloured side 
of the fish. 

This species is at once distinguished by the marked differ- 
ence in the structure and armature of the second maxillipeds ; 
it is also a smaller species than the other two described 
above. 

Genus 3. THERSITINA Nonnaii, 1905.* 

Syn. Thersitcs Pagenstecher 1861, a name preoccupied by Pfeiffer 
in 1855 for a genns of Mollusca, and by Spence Bate in 1857 for a 
genus of Amphipoda. 

Ergasilides having the cephalic segment in the female 
greatly enlarged and subglobular ; other thoracic and 
abdominal segments small. 

Antennules short, stout, and composed of five joints. 
Antennas also short, rather stout, and armed with a 
tolerably strong, claw-like, terminal spine, from the 
side of which springs a second and smaller one. 
Mandibles, maxillas, and first maxillipeds somewhat 
similar in structure to the same appendages in the 
Lichomolgidae. Second maxillipeds small, two- or 

* ' Museum Normanianum,' III, Crustacea, second edition, p. 41. 



42 BRITISH PARASITIC COPEPODA. 

three- joint eel, and bearing a few stout apical spines. 
Thoracic legs nearly as in Ergasilus. 

In the male the cephalic segment is not so much 
enlarged, and the body is more elongated. 

1. Thersitina gasterostei (Pagenstecher). 
(Plate XXV, figs. 1-6 ; Plate LI, fig. 6.) 

1861. Thersifes gasterostt i Pagenst. (94) vol. xvii, p. 118, pi. vi, figs. 1-9. 

1863. Ergasilvs gasterostei Kroyer. (71) p. 238, pi. xii. figs. 2, a — li. 

1892. Thersttes gasterostei Canu. (29) p. 245, pi. xxiii, figs. 13-18. 

1899. Ergasilus gasterostei Bassett-Smith. (8) p. 444. 

1900. Thersites gasterostei T. Scott. (112) p. 146, pi. v, figs. 1-7. 
1905. Thersitina gasterostei A. M. Norman. Museum Normanianum, 

III, Crustacea, second edition, p. 41. 

Female. — Cephalic segment considerably dilated, 
and, when viewed from above, concealing the remain- 
ing thoracic segments as well as part of the abdomen; 
genital segment of the abdomen longer than the 
combined leno^th of all the other abdominal seofments. 
Caudal rami short. 

Antennules very short, tolerably stout, and com- 
posed of five joints, which are sparingly setiferous. 
The formula shows their proportional lengths. 

Xumbers of the joints . . .12 3 4 5 

Proportional lengths of the joints 16 10 11 7 8 

Antennge short and stout, and each furnished with 
a tolerably strong terminal claw. Mandibles with a 
bilobed biting part. First maxillipeds short, simple in 
structure, and provided with a few small spines. 
Second maxillipeds small, basal part enlarged, ter- 
minal part more slender, forming a curved arm which 
is furnished with a few apical spines. 

The first three pairs of swimming-legs with both 
rami three-jointed, but the outer ramus rather shorter 
than the inner. In the fourth pair the outer ramus is 
only two-jointed, the elongated first joint consisting 
probably of two coalescent joints ; the inner ramus 
three-jointed as in the preceding three pairs. Length 
about 0*8 mm. (g-Q- of an inch). 

Habitat. — Parasitic on certain fresh- water fishes, 



'JHERSITIXA GASTEKOSTEI. 43 

usually on the inner surface of the gill-covers. On 
three-spined sticklebacks, Gasterodeus aculeatus, cap- 
tured in Sinclair Loch, Barra, Outer Hebrides, in 
May ] 894. On the same species of fish captured in 
the river Forth near Alloa in February 1896 ; and 
in brackish pools on Aberdeen links in 1 900. Taken 
also on a fifteen-spined stickleback, Gastrsea spinacJiia, 
captured in Loch Etive, West Coast of Scotland, in 
May 1896. 

This Copepod, which has been ascribed by some authors to 
the genus Ergasihts, does not agree very well with some of 
the characters of that genus. The typical female Ergasilus 
has the body tolerably elongated, but in Thersitina it is 
nearly globose; the structure and size of the antennae are 
also altogether different. On account of these and one or 
two other differences, as, for example, in the structure of the 
mandibles and second maxillipeds, we prefer to retain the 
species in the genus to which it was ascribed by Pagenstecher. 

Tribe II. CALIGOIDA G. 0. Sarn. 

Head usually in the form of a buckler or shield, 
with or without frontal plates : in some cases the 
frontal plates are provided with sucking disks. Thorax 
consisting of several, usually more or less distinct, 
rings or segments, with their margins sometimes pro- 
duced into flattened scales or plates. Abdomen or 
urosome composed of two or three segments, the 
genital segment being, in the female, sometimes con- 
siderably expanded (ex. Galifju^), or elongated (ex. 
Hatschekia). 

Mouth-apparatus consisting usually of a syphon 
provided with styliform mandibles and other accessory 
appendages. Usually there are also four pairs of 
thoracic limbs, but in some genera one or two pairs 
may be w^anting. A fifth pair, more or less rudimen- 
tary, is also sometimes present. 

The species comprised in this tribe are all, in the adult 
stage, parasitic on fishes. They are not permanently fixed 
to their host, but both the female and male possess a certain 



44 ■ BRITISH PARASITIC COPE POD A. 

freedom of locomotion, closely approaching in some cases 
that of the normal free-swimmers ; usually however their 
movements appear to be more or less restricted. They retain 
their position on the fish, at least to a considerable extent, by 
means of the strong terminal hooks with which certain of 
their appendages are furnished, such as the second pair of 
antennae, the maxillipeds, and one or two others. 

The young leave the eggs as free-swimming nauplii. 

The Caligoida comprise the three families Caligidae, 
Dichelestiidte, and Philichthyidae, the first two of which are 
represented by a considerable number of genera and species. 

Family ii. Caligidae. 

Carapace tolerably broad, more or less flattened, 
provided with anterior frontal plates. Free thoracic 
segments uncovered or sometimes more or less con- 
cealed by paired dorsal plates. Antennules small, tAvo- 
or tliree-jointed. Antennae forming short, movable, 
hooked claws. Mouth a suctorial beak formed by a 
modification of the upper and lower lips, and inclosing 
styliform mandibles. Second maxillipeds stout, with 
strong terminal claws. First four pairs of thoracic 
legs all biramose, or the first and fourth pairs may be 
uniramose. Fifth pair rudimentary. Eyes median, 
contiguous, sometimes obsolete. Egg-strings two, 
usually elongated. 

The male is generally smaller than the female. 

In some genera the mature stage is preceded by that of the 
chalimiis, or young stage, which is temporarily fixed to the 
host bv a more or less elongated slender frontal filament (r/. 
PI. IV; figs. 3-5). 

Genus 4. CALIGUS 0. F. MUller, 1785. 

Carapace large, flattened, shield-like Frontal plates 
provided with small but conspicuous sucking-disks or 
lunulse. Fourth pair of thoracic legs uniramose. Fifth 
pair rudimentary, and very small. Grenital segment 
in the female usually expanded and moderately large. 
Abdomen short or moderately elongated and composed 



CALIGUS. 45 

of one or two segments ; caudal rami short. Dorsal 
plates wanting. 

1. Caligus curtus 0. F. Miiller. 
(Plate XXIV, figs. 1, 2 ; Plate XXV, figs. 7-9.) 

1785. Caligus curtus O. F. Miiller. (86) p. 130, pi. xxi, fig. 1. 

1816. Caligus mulleri Leach. (74) p. 405, pi. xx. 

1832. Caligus bicuspidatus Nordmann, (89) p. 28. 

1850. Caligus dia'phanus Baird. (4) p. 269, pi. xxxiii, fig. 1. 

1863. Caligus yeglefi^u Kroyer. (71) p. 89, pi. vii, fig. 3, a-f. 

Female. — Carapace oval and fully half the entire 
length of the animal, rather longer than broad, widest 
behind and becoming gradually narrower towards the 
front, lateral margins slightly rounded, frontal plates 
tolerably large and furnished with conspicuous lunulge. 
Grenital segment subquadrangular, rather longer than 
broad, and equal to about one-fourth of the entire length. 
Abdomen narrow and scarcely equal to one-third the 
length of the genital segment. Caudal rami very 
short and furnished with about four short setae. 

Antennules short, the basal joint slightly expanded 
and its margin densely setiferous ; end joint narrow. 
The second maxillipeds consisting of a stout basal joint, 
and an end joint which is narrow and armed with 
two small terminal claws, one being larger than the 
other. Branches of the sternal fork short, moderately 
stout, and slightly divergent, also bluntly rounded 
at the end. Basal joint in the first pair of thoracic 
legs tolerably stout, and bearing a single, short, two- 
jointed branch ; the joints narrow and subequal, 
the first armed with a short spiniform seta on its 
outer distal angle ; the last with two short apical 
spines, and a moderately long one which is curved and 
claw-like. The fifth pair of legs very small and situated 
near the postero-lateral angles of the genital segment. 

Length about 10 mm., width equal to about half the 
length, but the size varies to some extent. 

The species is of a brownish or horn colour, which 
is apparently uniform, but when viewed with a hand- 



46 BRITISH PARASITIC COPEPODA. 

lens, irregular stellate markings are seen scattered 
over the surface of the carapace. 

Male. — In the male, the average dimensions of 
which, contrary to what is usual among the Caligidae, 
distinctly exceed those of the opposite sex, the cara- 
pace is proportionally larger and the genital segment 
smaller than in the female. The antenna and the 
second maxillipeds are more robust and form more 
powerful grasping organs. The form of the carapace 
also differs somewhat in being proportionally wider 
behind, and in tapering more towards the front. The 
total length of the male is about 13 mm. and the 
width 7*5 mm., but, as in the female, the size varies 
somewhat, ranging, according to C. B. Wilson, from 
13 to 20 mm. in length, and from 7*4 to 11*5 mm. in 
width of carapace. 

Habitat. — Parasitic on various kinds of fishes, but 
especially on Gadoids, and it has been ol)tained in 
abundance along with some other species on diseased 
cod-fishes. 

This species, first recorded by O. F. Miiller in 1785, is 
regarded as the type of the genus. It is, unlike Caligiis 
ra'pax, rarely captured as a free-swimmer, neither is it usual 
to find it on the gills or within the gill-covers of fishes, being 
more frequently observed adhering to, or moving about over 
the external surface of the fish. 

Caligus curtus is a common form around our shores, and 
appears also to be widely distributed in the North Sea and 
the North Atlantic. It has been recorded as abundant during- 
the autumn along the eastern sea-board of the New England 
States of North America.^ 

2. Caligus minimus Otto. 
(Plate IV, figs. 1, 2; Plate XXV, figs. 11-16.) 

1828. Caligus minimus Otto. (93) vol. xiv, p. 354, pi. xxii, figs. 7, 8. 

1826. Caligus minimus Risso. ( 1 05) p. 135. 

1840. Caligus minutus M. Edwards. (43) vol. iii, p. 450. 

1865. Caligus minutus Heller. (58) p. 163, pi. xiv, fig. 1. 

1899. Caligus minimus Bassett- Smith. (8) p. 447. 

* * North American parasitic Copepods/ by C. B. Wilson (Caliginas). 
' Proc. U. S. National Museum/ vol. xxviii, p. 581 (1905). 



CALTGIJS MIXIMUS. 4/ 

1901. Cnligus minimus A. Scott. (107) vol. xv, p. 349, pi. i, figs. 1-8. 

1905. Caligus minimus T. Scott. (116) p. 109. 

1906. Caligus minimus Norman & T. Scott. (88) p. 205. 
1906. Caligus minimus A. Brian. (21) p. 36. 

Female. — Carapace suborbicular and scarcely equal 
to half the entire length ; frontal plates and lunulse 
large. Eyes conspicuous. Free thoracic segment 
narrow. Grenital segment rather less than a third of 
the total length, and equal in width to about two- 
thirds that of the carapace ; the postero-lateral angles 
somewhat produced into narrow rounded lobes. Abdo- 
men uniarticulate and narrow, becoming slightly 
expanded anteriorly, equal in length to about two- 
thirds that of the genital segment. Caudal rami 
about half as long as the abdomen. 

Antennules short ; end joint tolerably elongated and 
narrow. Sternal fork small, the stem moderately 
long with short and somewhat divergent branches. 
Fourth pair of thoracic legs tolerably elongated, basal- 
joint stout, ramus two-jointed, joints subequal, the 
first with a small spine on the outer distal augle, the 
second furnished with one moderately long and two 
short apical spines as shown in the figure (PI. XXV, 
tig. 15). The fifth pair consisting each of a minute 
setiferous plate near the postero-lateral corners of the 
genital segment. 

Male. — The male is larger than the female, and 
differs from it chiefly in the form of the genital 
segment; this segment is not much wider than either 
the free thoracic segment or the abdomen. The 
appendages are somewhat similar to those of the 
female, except that the second maxillipeds are con- 
siderably expanded distally, and are each armed with 
a strong claw, so that they form powerful grasping- 
organs. The branches of the sternal fork in the male 
are scarcely if at all divergent. 

Total length of the female about 5 mm., and of the 
male about 7 mm. 

Habitat. — Parasitic on the bass, Lalrax Inpus. 
Usually found inside the gill-covers, or in the mouth of 



48 BEITISH PARASITIC COPEPODA. 

the fish. Belfast {Thompson, 1856). Plymouth 
(Bassett- Smith). BarroAv Channel, Lancashire {A. 
Scott), Queensferry, Firth of Forth (T, Scott). 

The species does not appear to be very rare. 

Distribution. — European Seas. Mediterranean 
(Brian). 

Caligus 'minimus, judging from the occurrences of this 
parasite known to us, appears to be restricted almost entirely 
to the bass. We do not know of its occurrence on any other 
kind of fish. 

3. Caligus rapax M. Edwards.* 
(Plate IV, figs. 3-8; Plate VI, figs. 1, 2.) 

1840. Callgur^ rapax M. Edwards. (43) vol. iii, p. 453, pi. xxxviii, fig. 9. 

1840. Caligus elongatiis idem. (43) vol iii, p. 454. 

1850. Coligus rapax Baird. (4) p. 270, pi. xxxii, figs. 2 and 3. 

1861. Caligus raptax Stp. and Lkt. (127) p. 359. pi. ii. fig. 4. 

1900. Caligus rapax T. Scott. (112) p. 148, pi. v, figs. 13-19. 

1905. Caligus rapax C. B. Wilson. (145) p. 594, pi. vii, figs. 79-89. 

Female. — Carapace ovate, rather longer than broad; 
frontal plates slightly ronnded ; lunnlse large and con- 
spicuous ; thoracic area fully half the entire width of 
the carapace ; its posterior margin broadly rounded 
and produced somewhat beyond the narrowly-rounded 
lateral lobes. Free thoracic segment narrow, veiw 
short, and scarcely more than half the width of the 
next one. Genital segment quadri-lateral, about as 
long as broad and equal to al)outhalf the length of the 
carapace. Abdomen short, unsegmented, and about 
half the width and scarcely two-thirds the length of 
the genital segment. Caudal rami short, and furnished 
with moderately long plumose setse. 

Antennules tolerably large. Antennae small, simple, 
and armed with a small marginal tooth. Mandibles 
elongated and narrow, terminal portion incurved and 
toothed on the inner edofe. Maxilla small. First 
maxillipeds slender and furnished with two slightly- 
curved terminal claws of unequal length ; second 

* For a more detailed synonymy of this common species see 'North 
American parasitic Copepods,' by C. B. Wilson (Caliginse). 'Proc. U. S. 
National Mnsenm,' vol. xxviii, p. 568 (1905). 



CALIGUS RAPAX. 49 

maxillipeds moderately stout. Sternal fork con- 
spicuous, its rami narrow, tolerably elongated, slightly 
divergent, and with the ends blunt-pointed. 

The first three pairs of thoracic legs short, biramose, 
with both rami three-jointed; fourth pair one- 
branched, basal joint elongated, tolerably stout, and 
furnished with a small seta on the upper distal angle ; 
end portion narrow and consisting of two subequal 
joints, the first with the outer distal angle slightly pro- 
duced and bearing a short spiniform seta, the other 
with a spiniform seta near the middle of the outer 
margin and four at the apex, the second from the 
inside being elongated, slightly curved and claw-like. 
The fifth pair rudimentary, each consisting of a 
minute plate bearing a few small setae, and situated 
near the postero-lateral angles of the genital segment. 

Length about 5 to 6 mm. Egg-strings not greatly 
elongated. 

Male. — The male is smaller than the female ; the 
form of the carapace is somewhat similar in the two 
sexes, but the genital segment is narrow and scarcely 
wider than the preceding one. Abdomen as long as 
the genital segment and composed of two joints, the 
end joint being about twice as long as the other. 
Caudal rami as in the female. 

The antennae and second maxillipeds are stouter 
than those of the female, and are provided with 
stronger terminal claws ; the other mouth-appendages 
and the thoracic legs are similar to those of the 
female. Length about 4 to 5 mm. 

Habitat. — Found parasitic on various fishes, but 
perhaps most frequently on Gadoids. It is also not 
uncommon in plankton collections, among Calanoids 
and other free-swimming organisms; both males and 
females are frequently captured in this way, and the 
latter are generally without egg-strings. Owing to the 
wandering habits of this C aligns there is scarcely 
a fish in our seas on which it may not at one time or 
other be found. 

VOL. I. 4 



50 BfllTiSH PARASITIC COPEPODA. 

Caligus rapax is apparently widely distributed in the North 
Sea and North Atlantic, and is, according to C. B. Wilson, 
" the most common species of the genus on the north-eastern 
coast of the United States, having been taken from more than 
twenty-five different kinds of fish by many collectors working 
in the interests of the United States Fish Commission '' 
(' Proc. U. S. National Museum/ vol, xxviii, p. 571). 

A certain amount of variation occurs in the size of different 
specimens of this species, and also in the proportional dimen- 
sions of the different parts. The colour is also somewhat 
variable; it seems to change to some extent with the colour 
of the fish. 

4. Caligus centrodonti Baird. 
(Plate V, figs. 1-3 ; Plate XVIII, figs. 1-3.) 

1850. Caligus centrodonti Baird. (4) p. 272, pi. xxxii, figs. 6, 7. 
1863. Caligus abhreviatus Kroyer. (71) p. 61, pi. iii, figs. 3 a-h. 
1905. Caligus abhreviatus T. Scott. (116) p. 109, pi. v, figs. 1-6. 
1905. Caligus centrodonti C. B. Wilson. (145) p. 652, pi. xxvii, figs. 
333, 344. 

Female. — Length of carapace about equal to the width, 
and to two-thirds of the entire length of the animal, 
widest posteriorly, and becoming considerably narrower 
towards the front. Frontal plates large, lunulas pro- 
minent. Free thoracic segment very short. Genital 
segment short and subquadrangular in outline, the 
width equal to about one and a half times the length, 
and fully half as wide as the carapace ; greatest width 
near the anterior end ; lateral margins obliquely 
rounded, posterior margin slightly concave. Abdomen 
very short, small, uniarticulate, and scarcely reaching 
beyond the lateral lobes of the genital segment. 
Caudal rami also very short, wdth four or five short 
setse round their distal ends. Antennules moderately 
short. Sternal fork stout, with tolerably stout and 
slightly divergent rami. Maxillae and maxillipeds and 
also the first three pairs of thoracic legs resembling 
those of the species already described; the fourth 
pair elongated, the basal joint moderately stout and 
furnished with a lobulate process on its upper aspect ; 
the single two-jointed ramus somewhat slender; the 



CALIGUS CENTRODONTI. 51 

proximal joint of the ramus short and scarcely half the 
length of the other, and bearing a tolerably long straight 
spine on its outer distal angle ; the end joint narrow 
and provided with three terminal spines, the outer- 
most similar to that on the preceding joint, the middle 
one slender and nearly as long as the joint, and ter- 
minating in a curved claw- like extremity, the inner- 
most spine, a small one, appearing to be merely a 
prolongation of the inner distal angle of the joint. 
Fifth pair nearly obsolete. Egg-strings tolerably stout 
but not very elongated. Length about 4 mm. 

Male. — The male is somewhat similar to the female 
but larger. A specimen examined by us, which was 
found to be a male — not a female as stated in the 
Fishery Board's Report — measured five millimetres in 
total length. The carapace was suborbicular and 
distinctly larger iji proportion to the total length, 
while the genital segment was much smaller than in the 
female, and the posterior margin of this segment was 
also more deeply concave, so much so that the postero- 
lateral angles reached beyond, and enclosed the very 
short abdomen. The caudal rami were also very short. 

Antennules and frontal plates tolerably large. 
Second maxillipeds large, with the end-joint con- 
siderably expanded and provided with a strong, hinged, 
terminal claw, so as to form powerful grasping organs 
as shown in the drawing (PI. XVIII, fig. 2). 

Habitat. — Parasitic on the tail and fins of the sea- 
bream, Pagellus centrodontus, and of the ballan wrasse, 
Labrus bergylta. Dublin, 1837, on Pagellus centro- 
dontus (Baird). Plymouth, on Pagellus centrodontus 
(Bassett- Smith). Moray Firth, on Labrus bergt/lta 
{T. Scott). 

There can be no doubt that the species described by 
Kroyer in 1863 under the name of Galigus ahhreviatus is 
identical with Baird's C. centrodonti. The peculiar form of 
the genital segment, the very short abdomen and caudal 
rami, together with the structure and armature of the fourth 
pair of thoracic legs, establish the identity of the two forms. 



52 BRITISH PARASITIC COPE POD A. 

In Charles Branch Wilson's figure of the fourth pair of legs 
all the three spines with which the end joint is furnished 
spring from the apex, whereas in our specimen the outer 
spine springs from a notch on the outer margin a short 
distance from the apex. 

A young specimen representing the chalimus stage of this 
Galigus (PI. V, fig. 3) was obtained along with the adult. 
The siphon is still present showing a dilated, two-jointed 
base ; the carapace is elongate-ovate in outline, the free 
thoracic segment is concealed, and the abdomen is very short. 
The frontal plates also slope backwards at a considerable 
angle. 

The distribution of Galigus centrodonti, so far as known, 
appears to be somewhat limited. The ouly record, other than 
those for the British Islands, is that of Kroyer. The speci- 
men recorded by him as Caligus ahbreviatus was obtained on 
a Lahrus hergylta at Bergen in 1839. The species is de- 
scribed in C. B. Wilson's work on 'North American parasitic 
Copepods,' but the specimens, which were taken in the gill- 
cavity of Pagellus centrodontus, were sent from England by 
the Kev. A. M. Norman. 

5. Caligus gurnardi Kroyer. 
(Plate VIII, figs. 1, 2.) 

1863. Caligus gurnardi Kroyer. (71) ser. 3, vol. ii, p. 150, pi. ii, figs. 
3 a-g. 

1896. Caligus gurnardi Bassett-Smith. (7) p. 157. 

Female. — Carapace equal to about three-fifths of the 
total length, and rather longer than broad; frontal 
plates only slightly arcuate, lunulse rather small. 
Free thoracic segment distinct, small; genital segment 
obcordate, about half as long as the carapace, its 
greatest width about equal to the length, posterior 
margin sub truncated. Abdomen short, uniarticulate, 
length scarcely equal to twice the width. Caudal 
rami short. 

Antennules of moderate size, the end joint narrow 
and about as long as the proximal one. Sternal fork 
tolerably large with moderately long and somewhat 
divergent branches. Fourth pair of thoracic legs 
elongated with the ramus biarticulate. Length 6-7 mm. 



CALIGUS GURNAEDI. 53 

Male. — The male is somewhat similar to the female 
in general appearance but the genital segment is much 
smaller, its width being scarcely a third of the width 
of the carapace ; the segment is also not much longer 
than broad. The abdomen is composed of two joints, 
and the end joint is much shorter than the proximal 
one. The cephalothoracic appendages appear to be 
similar to those of the female except that the second 
maxillipeds are rather stronger, and the fourth pair of 
thoracic legs of proportionally greater length. 

Habitat. — Parasitic on Trigla spp., usually lodged in 
the gill-cavity. Found at Falmouth in 1884 {A. M. 
Norman). Plymouth on Trigla cuculu^ {B ass ett- Smith). 
Kroyer obtained this species on Trigla gurnardus. 

We have not seen this species, but it appears to have an 
extensive distribution ; besides being recorded from the 
Norwegian and British seas, C. B. Wilson mentions its occur- 
rence on fishes captured on the coast of Cahfornia, viz. on a 
king salmon taken at Monterey and on an elephant fish at 
La Tolla.^ 

6. Caligus labracis T. Scott. 
(Plate V, figs. 4, 5 ; Plate XVIII, figs. 4, 5.) 

1902. Caligus labracis T. Scott. (114) p. 291, pi. xiii, figs. 26-29. 
1904. Caligus labracis A. Scott. (109) p. 38. 

Female. — Carapace suborbicular, length and width 
about equal, lateral margins evenly rounded, frontal 
plates large, lunulas prominent. Free thoracic segment 
subquadrangular, considerably shorter than the cara- 
pace, and a little more than one fifth of the total 
length; the width of the segment exceeding the length 
by about one fourth. Abdomen short, uniarticulate, 
and about one half as long as the genital segment. 
Caudal rami very short. 

Antennules short. Mouth-appendages somewhat 
similar to those of aligns dia/plianus Nordmann. 
Sternal fork stout, with tolerably broad and slightly 

* * North American parasitic Copepods : A list of those found upon the 
fishes of the Pacific coast,' etc. ' Proc. U. S. National Museum/ vol. xxxv, 
p. 439 (1908). 



54 BRITISH PARASITIC COPEPODA. 

divergent branches, which have their ends obliquely 
truncated. Fourth pair of thoracic legs small, basal 
joint tolerably stout ; ramus short, two- join ted, and 
armed with sabre-like spines, the first joint with one 
on the outer distal angle, and the second with two at 
the apex, the spines moderately elongated, those at 
the apex being longer than the joint from which they 
spring, also with a minute tooth on the inner distal 
angle. The fifth pair consisting each of a minute 
setiferous plate on the postero-lateral angle of the 
genital segment. Egg-strings short, containing a 
comparatively small number of tolerably large ova. 
Length about 3*5 mm. 

Male. — The male is smaller than the female, being 
only about 2*6 mm. in total length. It does not differ 
greatly from the female except that, as usual, the 
second maxillipeds are larger and the genital segment 
is much smaller. Colour reddish-brown, broken up 
into minute patches of a lighter and darker coloration. 

Habitat. — Parasitic on the gills of a striped wrasse, 
Lahrus mixtus L., captured in the Firth of Clyde in 
January 1900, and sent to the Laboratory of the 
Fishery Board for Scotland at Aberdeen by Mr. 
Robert Duthie, the Fishery Ofiicer at Girvan. Also 
found on the gills of Lahrus mixtus and Lahrus rtiacu- 
latiis captured in the Irish Sea {A. Scott). 

7. Caligus zei Norman and T. Scott. 
(Plate VII, fig. 1 ; Plate VIII, figs. 3-9.) 

1906. Caligus zei Norman & T. Scott. (88) p. 206, pi. xxii, figs. 1-8. 

1907. Caligus zei A. Scott. (Ill) p. 93, pi. i. 

Female. — Carapace suborbicular, length and breadth 
nearly equal, margins only slightly arcuate. Frontal 
plates and lunulae of moderate size. Free thoracic 
segment about as long as broad, rather small. Genital 
segment subquadrate and equal to about a third of 
the entire length of the animal; length and width 
nearly equal, and the postero-lateral angles, which are 



CALIGUS ZEI. 00 

rounded, slightly produced so that the posterior margin 
between the rounded angles is more or less incurved. 
Abdomen short and uniarticulate. Caudal rami small. 

Antennules rather longer than usual, the second 
joint being about five times longer than broad and 
much longer than the first joint. Antennae sharply 
hook-formed. First maxillipeds elongated and slender, 
the second joint long and narrow, and furnished with 
two tolerably long spiniform and claw -like setge, the 
outer one being rather longer than the inner. The 
second maxillipeds moderately large ; the basal joint 
stout but the end one short and narrow, and armed 
with a tolerably stout terminal claw. Sternal fork 
small, with moderately long and slightly tapering 
branches which are also somewhat divergent. The 
fourth pair of thoracic legs moderately elongated ; 
basal joint narrow and about as long as the two- 
jointed ramus ; the joints of the ramus of nearly equal 
length, the proximal joint provided with a stout spine 
on the outer distal angle, but the end joint with four 
spines, one near the middle of the outer margin, and 
three at the apex, the middle one rather longer than 
the one on either side. The fifth pair, which are very 
small, situated near the postero-lateral corners of the 
genital segment. Length about 5*5 mm. 

Male. — The genital segment in the male is as usual 
considerably smaller than in the female ; it is narrow 
and scarcely twice the width of the abdomen. The 
abdomen, which is longer than in the female, is com- 
posed of two segments, the first being rather shorter 
than the second. The thoracic and other appendages 
are somewhat similar to those of the female except 
that the second maxillipeds are stronger. 

Habitat. — Taken forty years ago on the dory, 
Zens faher, at Polperro by Laughrin, and sent by 
him to A. M. Norman.* Several specimens were 
found attached to the skin of a Zeus fab er captured off 

* ' Crustacea of Devon and Cornwall/ by the Rev. A. M. Norman and T. 
Scott, p. 207 (1906). 



56 BRITISH PAEASITIC COPEPODA. 

New Quay Head on 16tli June 1906 ; also on speci- 
mens of the same fish captnrecl off Puffin Island in 
1908 ; on Zeus faher taken in Luce Bay, September 
1911 (A. Scott). 

This species m;iy be distinguished from those closely allied 
to it by the unusual length of the second joint of the anten- 
nules, by the form of the sternal fork, and by the structure 
and armature of the fourth pair of thoracic legs. 

8. Caligus brevicaudatus A. Scott. 

(Plate VI, fig. 3 ; Plate VIII, figs. 10, 11 ; Plate 
XLVIII, fig. 6.) 

1901. Caligus brevicaudatus A. Scott. (107) p. 349, pi. ii, figs. 7-10. 

Female. — Carapace suborbicular, rather wider pos- 
teriorly than in front, and equal to fully half of the 
entire length of the animal. Frontal plates large and 
slightly arcuate ; lunulge prominent. Free thoracic 
segment short and narrow. Grenital segment oblong in 
outline, rather longer than broad, and equal to about 
half the width of the carapace. Abdomen very short 
and uniarticulate. Caudal rami also very short, and 
furnished with a few plumose setae. 

Antennules moderately stout, end joint rather shorter 
than the basal one. Sternal fork moderately stout, 
and with the rami also stout and scarcely divergent, 
and rounded at the tips. Second maxillipeds tolerably 
elongated, and slender, with somewhat feeble terminal 
claws. Fourth pair of thoracic legs small, and com- 
posed of three joints ; basal joint tolerably stout, ramus 
slender and rather longer than the basal joint; the 
terminal joint also rather longer than the preceding 
one ; a small spiniform seta springing from the 
extremity of the basal joint on its upper aspect, and 
the first joint of the ramus provided with a similar 
seta on its outer distal angle ; the end joint with three 
aj)ical setse which differ considerably in length, the 
inner one being about as long as the joint from which 
it springs, the outer scarcely half as long, the middle 



OALIGUS BREVICAUDATUS. 57 

one intermediate in length to the others. Length 
about 5*3 mm. 

Habitat. — Parasitic on Trigla gurnardus and Trigla 
lucerna. Found adhering to the inside of the mouth 
of a grey gurnard captured in the vicinity of Piel, 
Barrow-in-Furness, in August lyOl. Found also on 
the inside of the gill-covers of sapphirine gurnards 
taken in Luce Bay in October 1910 (A. Scott). 

No males have been observed. 

This species resembles Caligus curtiis in general appear- 
ance, but differs in the form of the genital segment, in the 
very short abdomen and caudal rami, and in the structure 
and armature of the fourth pair of thoracic legs. 

9. Caligus pelamydis Kroyer. 

(Plate VII, figs. 2, 8 ; Plate IX, figs. 1-5 ; Plate 
LXXI, fig. 14.) 

1863. Caligus pelamydis Kroyer. (71) p. 50, pi. iv, fig. 4 a-g. 
1896. Caligus scomheri Bassett-Smith. (6) p. 11, pi. iii, fig. 2. 
1901. Caligus scomheri T. Scott. (113) p. 148, pi. v, figs. 9, 10. 

1905. Caligus pelamydis C. B. Wilson. (145) p. 594, pi. xiii, figs. 
154-161 ; pi. xiv, fig. 116 a. 

1906. Caligus scomheri A. Scott. (110) p. 52, pi. vi. 
1906. Caligus pelamydis Norman & T. Scott. (88) p. 206. 
1910. Caligus pelamydis T. R. R. Stebbing. (125) p. 558. 

Female. — Carapace suborbicular, length and width 
about equal and rather more than one-third the total 
length of the animal, lateral margins slightly arcuate ; 
frontal plates tolerably large, lunulse also of moderate 
size but not very prominent. Free thoracic segment 
short, narrow, somewhat dilated. Grenital segment 
ovate, equal to fully two- thirds the length of the 
carapace, lateral margins nearly straight distally, but 
rounded and converging towards the anterior end, 
posterior margin truncated, postero-lateral angles 
rounded. Abdomen narrow, about as long as the 
genital segment, indistinctly biarticulated, end joint 
short and rather narrower than the elongated proximal 
one. Caudal rami short and provided with several 
short plumose setas. 



58 BRITISH PARASITIC COPEPODA. 

Antennules moderately stout ; sternal fork rather 
small, narrow, and elongated, rami not divergent or 
but very slightly so. Fourth pair of thoracic legs 
stout, comparatively short, basal joint about as long as 
the ramus, which is composed of three short and 
moderately stout joints, the first being the largest, 
and having the outer distal angle produced so as to 
reach to near the end of the second joint, the first and 
second joints each provided with a moderately stout 
spiniform seta on the outer distal angle, the third joint 
of a triangular form and carrying three spiniform 
setae, the end one being rather stouter than the others ; 
the apex of the joint is produced slightly beyond 
the base of the end spine, forming a blunt-pointed 
knob fringed with minute setas. From the peculiar 
form of the first and third joints the marginal and 
terminal setae are crowded together, and as each seta 
is slightly longer than the preceding one, they impart 
a character to this species somewhat different from 
others, such as Caligus rapax. Fifth pair very minute. 
Egg-strings tolerably elongated. 

This species appears to vary in length : the. speci- 
men figured here measured about 5*5 mm., and 
C. B. Wilson gives the length of his specimens as 
3-3 mm. 

Male. — The male of this Caligus does not appear to 
have been previously met with, and we are inclined to 
consider that it is much rarer than the female. The 
specimen of the male figured on PL LXXI, fig. 14, is 
the first and only one we have yet secured. It repre- 
sents the result of the examination of nearly fifteen 
hundred mackerel. The specimen figured was found 
on the inside of the operculum of a mackerel caught 
in the northern part of the Irish Sea in July 1912. 
On that particular occasion two hundred fish were 
examined, and although several females were found 
only a single male could be detected. Caligus 
'pelamydis resembles the male of Caligus rapax in 
general appearance, and without careful examination 



CALIGUS PELAMYDIS. 59 

it might be readily passed over for that species. The 
abdomen is two-jointed and slightly shorter than the 
genital segment. The first joint is comparatively short 
and about equal to half the length of the second joint. 
The furcal joints are short and broad and nearly as 
long as the first abdominal segment. The specimen 
was not dissected, but so far as could be made out 
from the examination of the entire animal the various 
appendages are almost identical with those of the 
female. The fourth pair of feet of the female, which 
are quite distinct from those of any of the other 
members of the genus, form a quite reliable distin- 
guishing character. The fourth pair of feet of the 
male now illustrated are identical with those of the 
female. The living male was dark red in colour, and 
it was easily detected running over the inner surface 
of the operculum of its host. Length 2*9 mm. 

Habitat. — Parasitic on the belted bonito, Pelamys 
sarcla, and the mackerel, Scomber scombrus. Plymouth, 
on the inside of the gill-covers of mackerel {Bassett- 
Smith). Aberdeen {T. Scott). Irish Sea [A. Scott). 
Kroyer's specimens were obtained on Pelamys sarda. 

Distribution. — Galigns pelamydis has apparently a 
wide distribution in the seas of Europe. C. B. 
Wilson describes this species in his work on ' North 
American parasitic Oopepods,' and mentions the name 
of its host, but does not state if the fish was captured 
in American waters.* 

We think there can be no reasonable doubt that the form 
obtained by Kroyer on Pelamys sarda, and described by him 
in the work referred to under the name of Galigus 
'pelamydis, is identical with that from the mackerel; the 
general structure of the animal, the shape of the sternal fork 
and of the genital segment, and the structure and armature 
of the fourth pair of thoracic legs, are similar in both forms. 

* See Part I (The Caliginse), p. o9-4. 



60 BRITISH PARASITIC COPEPODA. 

10. Caligus diaphanus Nordmann. 
(Plate XVII, fig. 1 ; Plate XVIII, figs. 6, 7 ; Plate 
XL VIII, fig. 7.) 

1832. Caligus diaphanus Nordmann. (89) pt. 2, p. 26 (non C. dia- 
phanus, Baird). 

1863. Caligus diaphanus Kroyer. (71) p. 79, pi. vii, fig. 5 a-c. 

1894. Caligus isonyx T. Scott. (Ilia) 12tli Rept. Fishery Board for 
Scotland, pt. iii, p. 194. 

1896. Caligus diaphanus Bassett- Smith. (6) p. 156. 

1900. Caligus diaphanus T. Scott. (112) p. 149, pi. v, figs. 20-25. 

1904. Caligus diaphanus A. Scott. (109) p. 38. 

1906. Caligus diaphanus Norman & T. Scott. (88) p. 206. 

Female. — Carapace suborbicular and equal to about 
two-fifths of the entire length of the animal, and the 
width rather greater than the length. Frontal plates 
large, anterior margin slightly emarginate, lunula? 
rather small, partly concealed. The free thoracic 
segment short and narrow; genital segment sub- 
cordate and equal to about two-thirds of the length of 
the carapace, truncated posteriorly, and the postero- 
lateral corners bluntly rounded. Abdomen composed 
of two joints, and nearly as long as the genital 
segment, the end joint small. Caudal rami small. 

Antennules of moderate size. Mandibles slender, 
elongated, with the end joint incurved and distinctly 
serrated on the inner margin. The fourth pair of 
thoracic legs short and stout, the ramus, which is 
composed of three short joints, has both the first and 
second joints provided with a stout seta of moderate 
size on the outer distal angle, while the end joint has 
three similar apical setae ; all the setae plumose and of 
nearly equal size, except that the inner one is rather 
longer than the others. Fifth pair minute, and, as in 
the case of other species, occurring as minute setiferous 
plates near the postero-lateral corners of the genital 
segment. Length about 4*5 mm. 

Male. — The carapace of the male is rather wider in 
proportion to its length ; the genital segment is small 
and scarcely twice the width of the abdomen, it 
becomes narrower towards the proximal end, and the 
postero-lateral corners are somewhat angular. The 



CALIGUS DIAPHANUS. 61 

abdomen is composed of two subequal, angular 
segments, and is only equal to about one-sixth of the 
entire length. 

Habitat. — Parasitic on various species of gurnards 
{Trigla). Belfast (W. Thompson). Found on the inner 
surface of the operculum of Trigla hiruiuio and Trigla 
(luculus at Plymouth (Bassetf- Smith). On Trigla spip., 
Firths of Forth and Clyde {T. Scott). Irish Sea 
{A. Scott). 

The species is apparently not uncommon, but we have 
only met with it on gurnards. 

Genus 5. PSEUDOCALIGUS A. Scott, 1901. 

Carapace large and scutiform ; frontal plates 
moderately prominent, and provided with lunulse. 
Genital segment aiid abdomen as in Galigus. Cephalo- 
thoracic appendages also similar to those of Galigus, 
except that the fourth pair of thoracic legs are some- 
Avhat rudimentary, and consist each of a single small 
joint, furnished with a few minute terminal bristles. 

This genus has, in its general appearance, a close 
resemblance to Galigus. 

1. Pseudocaligus brevipedes (Bassett- Smith). 
(Plate IX, figs. 6-9 ; Plate X, figs. 1, 2.) 

1896. Galigus brevipedes Bassett-Smitli. (6) p. 11, pi. iii, fig. 1. 

1901. Pseudocaligus brevipedes A. Scott. (107) p. 350, pi. ii, figs. 1-6. 

1902. Pseudocaligus brevipedes T. Scott. (114) p. 291. 

1906. Pseudocaligus brevipedes Norman & T. Scott. (88) p. 207. 

Female. — Carapace suborbicular, about as long as 
broad, and equal to rather more than half the entire 
length of the animal. Frontal plates well developed, 
lunulas of moderate size. Free thoracic segment small. 
Genital segment subquadrangular, and about half as 
long as the carapace, the length rather less than the 
width, and the lateral margins nearly straight, the pos- 
terior end also subtruncated, and the postero-lateral 
corners bluntly rounded. Abdomen consisting of a 



62 BIUTISH PARASITIC COPEPODA. 

single joint, somewliat longer than broad, and scarcely 
equal to one-third of the length of the genital segment. 
Caudal rami of moderate size. 

Antennules tolerably stout, and composed of two 
subequal joints. Antennae with the end joint in the 
form of a strongly-hooked claw. Sternal fork small 
with tolerably short and slender rami which are 
moderately curved and somewhat divergent. The 
fourth pair of thoracic legs consisting each of a single 
joint, small and rudimentary and furnished with three 
spiniform setae, two of them short and subequal, and 
the other tolerably elongated and plumose. The fifth 
pair in the form of minute setiferous plates near the 
postero-lateral corners of the genital segment. Length 
about 3*6 mm. 

Male. — The male is rather smaller than the female, 
and differs from it chiefly by the small size of the 
genital segment. The carapace is also rather wider in 
proportion to the length, while the second maxillipeds 
are larger and form more powerful grasping organs. 

Habitat. — Parasitic on the three-bearded rockling, 
Onos (Motella) tricirrata Brun., and usually found 
adhering to the inner surface of the gill-covers. 
Plymouth {Bas sett- Smith). Irish Sea, Barrow Channel, 
and at Port Erin, Isle of Man (A. Scott). Dunbar, at 
the mouth of the Forth estuary, and near Aberdeen 
{T. Scott). 

This species is easily distinguished by the rudimentary 
character of the fourth pair of thoracic legs in both sexes. 
From the records of the species published hitherto it seems 
probable that the distribution of the Pseudocaligus will be 
coextensive with that of the fish mentioned. 

Genus 6. SCIiENOPHILUS P. /. va.n Beneden, 1852. 

Carapace somewhat similar to that of Galigus. 
Frontal plates distinct and provided with lunulge. 
Free thoracic segment small. G-enital segment narrow 
and considerably elongated. Abdomen slender and 
of great length. Caudal rami short. Mouth append- 



SOI^NOPHILUS. 63 

ages somewhat as in Caligus, but the second maxilli- 
peds very large. Sternal fork absent. Thoracic legs 
nearly as in C aligns, 

1. Scisenophilus tenuis P. J. van Beneden. 
(Plate XI, figs. 1-6.) 

1852. Scienophilus tennis, P. J. van Beneden. (13) p. 464, and plate. 

1861. Scie7wphilus tenuis idem, (15) p. 148, fig. 21. 

1896. Scimwphilus tenuis Bassett-Smith. (6) p. 156. 

1906. Sciaenojphilus tenuis Norman & T. Scott. (88) p. 207. 

Female. — Carapace small, suborbicular, length and 
width about equal. Frontal plates distinct, lunulae 
small. Free thoracic segment small ; genital segment 
elongated and narrow, but increasing in width gradu- 
ally though slightly towards the posterior extremity, 
the greatest width being equal to about one fourth of 
the length, and the length equal to about three times 
the length of the carapace, the distal end slightly 
emarginate and the postero-lateral corners bluntly 
rounded. Abdomen extremely long and slender, fully 
twice the length of the genital segment and about as 
long as, and not much thicker than, the egg-strings. 
Caudal rami short, and provided with four to six 
apical setge. 

Antennules small, two-jointed, and similar to those 
of aligns ; mouth -appendages also similar. The 
second maxillipeds stout and elongated, and armed 
with strong curved terminal claws. The first pair of 
thoracic legs slender, and bearing at their distal end 
three strong and curved setiferous spines ; only the 
second pair of legs distinctly biramose. The fourth 
pair elongated and uniramose, and consisting each of 
two tolerably long joints, the terminal one being fur- 
nished with a few marginal and apical setse. Total 
length about 14 mm. (fuU}^ half an inch). 

We have not seen the male of this species. 

Habitat. — Parasitic in the branchial chamber of the 
maigre, Scisena aquila (or Scidena vmhra), Plymouth 
{Bassett'Sniith). 



64 BRITISH PARASITIC COPEPODA. 

Dr. Bassett-Smitli obtained four specimens of this curious 
parasite on the inside of the operculum of a maigre captured 
at Plymouth. Though this is the only British record of 
Scissnophihis we know of, Professor P. J. van Beneden states 
that the species is very common on the maigre, and he has 
seen such a great number of specimens crowded together that 
they resembled tufts of moss (" formant des touifes semblables 
a une mousse'^). This fish is not common in the British 
seas, and, according to Aflalo, is taken only casually in the 
mackerel nets, and this probably is the reason why such a 
common parasite as this appears to be has not been more 
frequently recorded. 

Sci8B7iophilus tenuis, so named from its narrow and elon- 
gated form, has at first sight a close resemblance to a Hat- 
schekia {Clavella), but the form and structure of the carapace 
reveal at once its near relationship with Caligus. 

Genus 7. LEPEOPHTHEIRUS Nordmann, 1832. 

Free thoracic segment without dorsal plates. Genital 
segment simple. Abdomen one- or two-segmented. 

Mandibles serrated only on the inner edge. Second 
maxillae small, furcate, the branches acuminate, not 
simple and spine-like as in Caligus, Fourth pair of 
thoracic legs and other thoracic appendages as in 
Caligus, 

Both the females and the males have a general 
resemblance to those of Caligus, but the frontal plates 
are without lunulae. 

1. Lepeophtheirus pect oralis (0. F. Mliller). 

(Plate X, figs. 3-4 ; Plate XII, figs. 2-3; Plates XIII, 
XIV, XV; and Plate XVI, figs. 1-3.) 

1777. Lernsea pectoi'aUs O. F. Mliller. (85) Zool. Danica, p. 41, pi. 
xxxiii, fig. 7. 

1838. Caligus pectoralis Kroyer. (70) (ii), p. 8, pi. vi, fig. 4. 

1832. Lepeophtheirus pectoralis Nordmann. (89) p. 30. 

1847. Caligus pectoralis Thompson (129), p. 247. 

1850. Lepeophtheirus pectoralis Baird. (4) p. 275, pi. xxxii, fig. 10. 

1900. Lepeophtheirus pectoralis T. Scott. (112) p. 150, pi. v, figs. 
26-31. 

Female. — Carapace suborbicular, length and width 
nearly the same, and scarcely equal to half the entire 



LEPEOPHTHEIRUS PECTORALIS. 65 

length of the animal. Frontal plates of moderate size, 
lunuljB wanting. Free thoracic segment small. Genital 
segment subqnadriform and equal to about one-third 
of the entire length of the animal, with its postero- 
lateral angles rounded. Abdomen short, with a slight 
constriction near the middle which in certain posi- 
tions gives it the appearance of being obscurely two- 
segmented. Caudal rami very short. 

Antennules of moderate size. Antennae armed with 
strong, terminal, hooked claws. Mandibles small, 
slender, and with the inner edge of the distal joint 
distinctly serrated. Posterior maxillipeds strong and 
provided with stout terminal claws. Sternal fork 
moderately stout, with short and scarcely divergent 
rami, each ramus somewhat expanded in the middle 
and thence tapering to the pointed extremity. Fourth 
pair of thoracic legs short ; the basal joint tolerably 
stout, and the ramus consisting of two subequal joints 
— the first with a small spine on the outer distal angle, 
and the second with three short terminal spines, each 
of the two inner spines being rather longer than the 
one in front of it. The fifth pair extremely small and 
situated near the postero-lateral angles of the genital 
seofment. Lenolh about 5 mm. 

Male. — The male is little more than half the size of the 
female, but the carapace is proportionally larger, being 
equal to nearly two-thirds of the entire length of the 
animal. Genital segment small ; abdomen also small 
and uniarticulate. 

Habitat. — Found usually adhering to the underside 
of the pectoral fins of certain flat fishes, as plaice, Pleu- 
ronectes ]jlatessa, dabs, Pleuronectes limanda, and one 
or two others. Belfast {W. Thompson). Plymouth 
(Bassett- Smith). Irish Sea {A. Scott). Firths of 
Forth and Clyde {T. Scott). 

This is one of the more common and easily recognized of 
the species of Lepeophtheirus, but though widely dispersed in 
the seas of Europe it does not appear to have yet been 
recorded from American waters. 

VOL. I. 5 



66 BUiTISH PAKASITIC COPEPODA. 

2. Lepeophtheirus nordmanni (M. Edwards). 
(Plate XII, fig. 1 ; Plate XVI, figs. 4-11.) 

1840. Caligus nordmanni M. Edwards. (43) p. 455. 

1847. Caligus nordmanni W. Thompson. (129) p. 248. 

1850. LeiJeophtheirus nordmanni Baird. (4) p. 275, pi. xxxiii, fig. 1. 

1865. Lejoeophtheirus nordmanni Heller. (58) p. 180, pi. xvi, figs. 1, 2. 

1884. Lepeophtheirus nordmanni Rathbun. (97a) Proc. U. S. National 
Mns., vol. vii, p. 487. 

1900. Lepeophtheirus nordmanni T. Scott. (112) p. 151, pi. v, figs. 
32-37. 

1905. Lepeophtheirus nordmanni 0. B. Wilson. (145) p. 623, pi. xix. 
figs. 223-233. 

1909. Lepeophtheirus nordmanni E. Y. Elwes. (45) p. 19. 

Female. — Carapace suborbicular, about as long as 
broad and equal to about half the entire length of the 
animal. Frontal plates narrow, without lunulse. Free 
thoracic segment small, somewhat rhomboid in shape. 
Grenital segment obovate, considerably narrower than 
the carapace, and equal to fully one-fourth of the entire 
length of the animal ; length and width about equal, 
and with the postero-lateral corners forming rounded 
lobes so that the space between is deeply incurvated. 
Abdomen short, consisting of a single segment with a 
slight constriction near the middle. Caudal rami short. 
Antennules of moderate size. Antennae armed 
with tolerably long and strongly-hooked terminal 
claws. Mandibles slender and elongated, end joint 
comparatively short and finely serrated on the inner 
margin. Second maxillipeds moderately stout and 
elongated, and provided with strongly-hooked terminal 
claws. Sternal fork with the basal portion narrow, 
the rami also comparatively narrow and elongated, 
and somewhat divergent. Fourth pair of thoracic 
legs tolerably elongated, basal joint not very stout and 
rather shorter than the ramus, which consists of three 
subequal joints ; the first and second joints of the 
ramus each provided with a short spine on the outer 
distal angle, the end joint with three moderately long 
apical spines, the outermost being scarcely so long as 
the other two. Length about 12 mm. Egg-strings 
considerably elongated. 



LEPEOPHTHEIllQS NORDMANNI. 67 

Male. — The male, which is only about half the size 
of the female, has the carapace proportionally rather 
longer, and the frontal plates are more distinct. The 
free thoracic segment is also rather longer and the 
genital segment narrower than in the female, and the 
postero-lateral corners of the genital segment have each 
two small angular processes, the one slightly anterior 
to the other. Abdomen short and about one third the 
length of the genital segment, and composed of two un- 
equal joints — the proximal one being small. Caudal 
rami short and bearing several short setae. 

Habitat. — Parasitic on the short sun-iish {Orthagoris- 
cus mola). Coast of County Antrim, Ireland, 1848 
{W. Thomioson). Polperro, Cornwall {A. M. Norman). 
Plymouth {B ass ett- Smith). Aberdeen {T. Scott). 
Found also on a short sun-fish captured off Berry 
Head {E. V. Mives). 

L.nordmanni does not appear to be very commonly met with ; 
the sunfish when captured is usually subjected to some rough 
handling ere it reaches the market and the parasites get 
rubbed off. 

C. B. Wilson records this species from both the xltlantic 
and Pacific Coasts of North America. 

3. Lepeophtheirus hippoglossi (Kroyer). 

(Plate VI, figs. 4, 5; Plate XVII, fig. 2 ; Plate 
XVIII, figs. 8-10.) 

1838. Caligiis hippoglossi Kroyer. (70) R. l,vol. i, p. 625, pi. vi, fig. 3. 

1850. Lepeophtlieirus liippoglossi Baird. (4) p. 276, pi. xxxii, fig. 
12 ( ? ). 

1850. Lepeophtheirus ohscurus Baird. (4) p. 277, pi. xxxii, fi.g. 11 iS). 

1863. Lepeophtheirus hippoglossi Kroyer. (71) p. 131, pi. vi, fig. 5 a-d. 

1900. Lepeophtheirus hippoglossi T. Scott. (112) p. 151, pi. v, figs. 
38-42; pi. vi, figs. 1,2. 

1905. LepeopJitheirus hippoglossi C. B. Wilson. (145) p. 625, pi. xx, 
figs. 234-243. 

Female. — Carapace somewhat longer than broad and 
equal to rather more than half the entire length of the 
animal, lateral margins gently and evenly arcuate. 
Frontal plates narrow, without lunulge. Free thoracic 
segment small. Grenital segment fully half the length 



68 BRITISH PARASITIC COPEPODA. 

of the carapace, width about three fourths of the 
length, lateral margins only slightly arcuate. Abdomen 
short, composed of a single segment ; caudal rami 
short and bearing a few apical setae. 

Antennules small. Antennse short and moderately 
robust, and armed with a stout sinuately-curved 
terminal hook. Mandibles slender but scarcely so 
much so as in Lepeojjhthelrus nordmanni. Second 
maxillipeds stout and provided with short but strong 
terminal claws. The sternal fork with the basal part 
somewhat expanded, the rami short, tolerably broad 
and divergent, each ramus truncated at the end and 
split into two portions by a longitudinal sinus, the 
inner portion being much narrower than the outer. 
Fourth pair of thoracic legs tolerably elongated, basal 
joint stout, ramus composed of three joints, the 
middle one being rather longer than the first or third ; 
the outer distal angle of the first joint ending in a 
bluntly-rounded knob, the second joint bearing a short 
spine on its outer distal angle, and the end joint three 
terminal spines, the innermost of the three spines being 
longer than the others and finely serrate along the 
outer margin. The fifth pair very small and rudimeu- 
tary. Length about half an inch (12*5 mm.), but 
varying somewhat in different specimens. Egg-strings 
slender and elongated. 

Male. — Carapace orbicular, as long as broad, and 
equal to about two-thirds of the entire length of the 
animal. Free thoracic segment very small, and 
slightly produced on each side. Grenital segment very 
small, with a pair of minute setiferous processes on 
each side at the postero-lateral corners. Length about 
6*5 mm. 

Habitat. — Parasitic on the backs of large halibut, 
Hijypoglossus vulgaris Fleming. Berwick Bay (Dr. 
Johnston). Polperro, Cornwall {A. M. Norman). 
Aberdeen {T. Scott). Shetland (0. B. Wilson). 

Distrilmtion. — JN'orth Sea ;' Greenland, Iceland, 
Massachusetts (Cape Ann), U.S.A. 



LEPEOPHTHEIRUS THOMPSONI. 69 

4. Lepeophtheirus thompsoni Baird. 

(Plate XVII, fig. 3 ; Plate XXV, fig. 10 ; Plate 
XLVIII, figs. 1-3.) 

1850. Lepeophtheirus thompsoni Baird. (4) p. 278, pi. xxx, fig. 2. 

1851. Caliyus gracilis P. J. van Beneden. (11) p. 90, pi. ii, figs. 1-7. 
1861. Caligus branchialis Malm, mscr. $ Steenstrup & Liitken. 

(127) j3. 362, pi. ii, fig. 3. 

1863. Lepeophtheirus rhomhi Kroyer. (71) p. 118, pi. v, fig. 5. 

1885. Lepeophtheirus gracilis Cai'us. (29a) Prodr. Faunae Mediter- 
ranese, p. 359. 

1900. Lepeophtheirus thompsoni T. Scott. (112) p. 152, pi. v, figs. 
43-45. 

1900. Lepeophtheirus ohscurus idem. (112) p. 153, pi. vi, figs. 16-19. 

1905. Lepeophtheirus thompsoni C. B. Wilson. (145) p. 619. pi. xviii, 
figs. 212-219. 

Female. — Carapace suborbicular and equal in length 
to about two-fifths of the entire length of the animal. 
Frontal plates arcuate, and tolerably prominent. Free 
thoracic segment small. Genital segment large and 
nearly as long as the carapace, widest at the distal end, 
which is about three-fourths the width of the cara- 
pace, but the width gradually decreases anteriorly to 
where the proximal end rapidly contracts to form a 
narrow neck ; lateral margins slightly arcuate, or 
nearly straight, postero-lateral corners somewhat pro- 
duced into broadly-rounded lobes. Abdomen toler- 
ably elongated and equal to about two-thirds the 
length of the genital segment, a slight constriction 
near the posterior end giving it the appearance of an 
indistinct joint. Caudal rami short. 

Antennules about two-thirds the length of the frontal 
plates. Antennae rather slender. Second maxillipeds 
tolerably stout. Sternal fork robust with the rami 
somewhat expanded, widest in the middle and tapering 
slightly towards each end, also moderately far apart 
and scarcely divergent. Fourth pair of thoracic legs 
small, basal joint tolerably stout, and with the ramus 
composed of three joints, the outer distal angle of the 
first joint forming a blunt-pointed knob, that of the 
second joint bearing a small spine, the third armed 
with one short and tw^o moderately long and spiniform 
apical setse. Fifth pair minute. Length about 8*5 mm. 



70 BRITISH PARASITIC COPEPODA. 

Male. — The male is only about half the size of the 
female. The carapace is somewhat elliptical in out- 
line, rather longer than broad, and equal to about two- 
thirds of the entire length of the animal; lateral 
margins evenly rounded. Genital segment suborbicular 
and equal to about one-third of the length of the 
carapace. Abdomen about half as long as the genital 
segment. Total length about 4 mm. 

Habitat. — Parasitic on the gills of turbot, Rhombus 
maximus (L.). North of Ireland (TF. Thompson). 
Polperro, Cornwall {A. M. Norman). Plymouth 
(Bassett- Smith). Firths of Forth and Clyde, Aber- 
deen, &c. {T. Scott). Irish Sea {A. Scott). 

C. B. Wilson records Lepeophtheir^cs thompsoni from La 
Jolla;, Cahfornia, U.S.A.^ 

[Lepfeop)htheirits obscuvus Bassett-Smith. 

As there is some ambiguity concerning this form, 
we quote the following remarks by A. M. Norman and 
T. Scott, in ' Crustacea of Devon and Cornwall,' 
p. 208 : " Mr. Bassett-Smith (' Journ. Marine Biol, 
xlssoc.,' vol. iv, 1896, p. 157) records a species from 
the brill {BJiombus Idevis) taken at Plymouth, and 
refers it with doubt to Le/peophtheirus obsciirvs Baird, 
which was obtained from the same species of fish. 
Subsequently Bassett-Smith figures what he then 
styles Galigus obscurus Baird (* Ann. and Mag. Nat. 
Hist.,' ser. 6, vol. xviii, pi. iv, fig. 2), and not onl}^ is 
his parasite there referred to the genus Galigus^ but 
the figures show the presence of sucking -disks. 
Whether the form he found was a Lepeophtheirus or a 
OaJigvs, it was certainly not Lepeophtheirus obscurus of 
Baird. In Bassett-Smith's third paper {' Proc. Zool. 
Soc.,' 1899, p. 456) he referred it back again to the 
genus Lepeophtheirus. It is just possible, if the species 
he found was a Lepeophtheirus, that it may have been 
the female of Ij. appendiculatiis Kroyer (' Naturhist. 
Tidssk.,' ser. 3, 1863, p. 207, pi. vi, fig. 4, a-i), which 

* ' Proc. U. S. National Museum/ vol. xxxv, p. 441 (1908). 



LEPEOPHTHEIKUS OBSCUEUS. 71 

lias a branched furcula, but the former was taken on 
the brill, while the latter was found on Baia clavata.^' 
A LepeophtheiruH found on the brill by T. Scott, and 
doubtfully ascribed by him to L. ohscnrus Baird, seems 
to be merely a form of L. thompsoni, not deserving- 
even varietal rank. Bassett- Smith's L. ohscurus is 
probably similar, for in a copy of the ' Ann. and Mag.' 
paper referred to above kindly presented by the author, 
the lunulge shown in figure 2, Plate 4, are deleted and 
the words " no lunula " are added at the side ; so also 
the bifurcate appearance of the rami of the sternal 
fork shown in figure f on the same plate may like- 
wise be due to an inadvertence.] 

5. Lepeophtheirus salmonis (Kroyer). 
(Plate XVII, fig. 4; Plate XVIII, figs. 11, 12.) 

1837-38. Caligus salmonis Kroyer. (70) vol. i. pi. vi, fig. 7 a-c ; vol. ii, 
pp. 13-18. 

1850. Lepeophtheirus stromii Baird. (4) p. 274. pi. xxxii, figs. 8, 9. 

1863. Lepeophtheirus salmonis Kroyer. (71) p. 211, pi. xvii, figs. 1 a, h. 

1900. Lepeophtheirus stromi T. Scott. (112) p. 152. pi. vi, fig«. 3-8. 

1905. Lepeophtheirus salmonis C. B. Wilson. (145) p-. 640, pi. xxiv. 
figs. 294-300. 

Female. — Carapace rather longer than broad, and 
equal to about three-sevenths of the entire length of 
the animal, lateral margins slightly and evenly arcuate. 
Frontal plates not very clearly defined, the margin 
convex, and without lunulse. Free thoracic segment 
very small. Genital segment tolerably large and of 
an oblong form, rather longer than broad, its length 
being about a third less than that of the carapace, the 
lateral margins only slightly arcuate but the postero- 
lateral corners produced into rounded lobes, watli the 
space between them deeply incurved. Abdomen 
narrow, elongated and un segmented, and equal to 
about one-fourth of the entire length of the animal ; 
distal end slightly constricted so as to have the appear- 
ance of an indistinct joint. Caudal rami very short. 

Antennules rather small. Antennge tolerably robust, 
and furnished with a moderatelv strono- terminal hook. 



72 BEITISH PARASITIC COPEPODA. 

Maxillae short and stout, and with the extremity dis- 
tinctly bifurcated. Second maxillipeds robust and 
armed with short and stout terminal claws. Basal 
portion of the sternal fork short, the rami also short, 
somewhat expanded and bluntly rounded at the tip ; 
slightly divergent and separated from each other by a 
comparatively wide semicircular space. The fourth 
pair of thoracic legs somewhat like those of Lepeoph- 
theirus hippoglossi, but rather more robust, the first 
joint of the ramus, which is rather longer than the 
second or third, with the outer distal angle in the form 
of a blunt knob covered with microscopic bristles, the 
second joint provided with a short spine on the outer 
distal angle, and the third with three spiniform apical 
setae, the inner one being the longest. Egg-strings 
very long and slender. Length 14-16 mm. 

Male. — The carapace of the male is rather longer 
than broad and more than half the entire length of 
the animal. Frontal plates narrow. Free thoracic 
segment of moderate size, its length rather less than the 
width. Genital segment ovate, longer than broad, and 
equal to about one-third of the length of the carapace. 
Abdomen rather narrower than the genital segment, 
and about a third shorter. Caudal rami about half as 
long as the abdomen. 

The thoracic appendages are somewhat similar to 
those of the female, except that the second maxillipeds 
are proportionally stronger. Length 6-7 mm. 

Habitat. — Parasitic chiefly on salmon, Salmo salar, 
L., but it also occurs on one or two other species of 
the Salmonidse. Berwick (Dr. Johnston). Ireland 
{W. Thompson). Polperro, Cornwall {A. M. Norman). 
Plymouth (Bassett- Smith). Scottish coasts {T. Scott). 
Irish Sea (A. Scott). 

This is one of the more widely distributed of the Cahgid^e. 
It is recorded from the coasts of Alaska, Labrador, and 
from other parts of the North American sea-board, as well as 
from the coasts of Europe. The parasite does not appear to 
survive long after the fish enters the fresh water. The dorsal 
surface in this species has a curious metallic lustre different 



LEPEOPHTHEIRUS SALMONIS. 73 

from most of the others of the same genus. Males appear to 
be comparatively scarce. 

6. Lepeophtheirus poUachii Bassett- Smith. 

(Plate XVII, figs. 5, 6; Plate XVIII, figs. 13-15; 
Plate XL VIII, figs. 4, 5.) 

1896. Lepeophtheirus pollachius Bassett-Smitli. (6) p. 12, pi. iv, fig. 1, 

1899. Lepeojjhtheinis pollachii idem.. (8) p. 455. 

1900. Lepeophtheirus pollachii T. Scott. (112) p. 153, pi. vi, figs. 9-18. 
1905. Lepeophtheirus inominatus C. B. Wilson. (145) p. 656, pi. 

xxviii, figs. 345-352. 

Female. — Carapace oval, rather longer than broad, 
and equal to rather more than a third of the entire 
length of the animal, lateral margins slightly arcuate. 
Frontal plates not very prominent. Free thoracic seg- 
ment small. Grenital segment subquadrangular, length 
about a fourth less than the width at the proximal 
end, and a third less than the length and width of the 
carapace ; widest posteriorly ; the lateral margins 
nearly straight and the postero-lateral corner sub- 
angular. Abdomen fully as long as the genital seg- 
ment, and indistinctly biarticulated, the end joint being 
rather narrower than the other. Caudal rami very small. 

Antennules of moderate size. Antennae with toler- 
ably elongated, strong and abruptly hooked terminal 
claws. Second maxillipeds moderately short and robust, 
terminal claw stout and strongly curved. Sternal 
fork small, rami divergent, and bluntly rounded at the 
tips. Fourth pair of thoracic legs not reaching to the 
end of the genital segment ; ramus somewhat slender 
and consisting of three subequal joints ; the first and 
second joints each provided with a small spine on the 
outer distal angle, but the spine on the first joint 
sometimes wanting ; the end joint furnished with three 
spiniform apical setae of unequal length, the innermost 
being the longest. The entire lengtli of the specimen 
represented by the drawing on PL XVII, fig. 5, is 
8 mm. Eofof-strino's lono- and slender. 

If ale. — The cai-apace of the male is equal to half the 
entire length of the animal. The genital segment is of 



74 BRITISH PAEASITIQ COPEPODA. 

an oval form, not mucli longer tlian, and abont twice the 
width of, the following segments ; its distal margin is 
rounded, with a minute setiferous plate on each side 
representing the fifth pair of legs. Abdomen com- 
posed of two subequal but indistinctly segmented 
joints. Length about 4*5 mm. 

Habitat. — Found for the most part on lythe, Gadus 
pollachius, adhering to the inside of the mouth, and 
rarely on other fishes. Plymouth {B ass ett- Smith). 
Falmouth (^4. M. Norman). On salmon captured at 
Polperro, Cornwall (Laughrin in 'Museum Normania- 
num'). Girvan, Firth of Clyde, and at Aberdeen Fish- 
Market (T. Scott). Irish Sea {A. Scott). 

This species does not appear to he so common as some of 
the others. 

7. Lepeophtheirus sturionis (Kroyer). 
(Plate XVIII, figs. 16-19 ; Plate XXI, fig. 1.) 

1837. Caligus sturionis Kroyer. (70) pi. vi, fig. 6. 

1840. Lepeophtheirus sturionis M. Edwards. (43) vol. iii, p. 457. 

1863. Lepeophtheirus sturionis Kroyer. (71) p. 139, p], xvii, fig. 4. 

1905. Lepeophtheirus sturionis T. Scott. (116) p. 110, pi. v, figs. 7-14. 

Female. — Carapace suborbicular and equal to about 
two-fifths of the entire length of the animal. Frontal 
plates not very prominent. Free thoracic segment 
not very clearly defined. Genital segment somewhat 
pyriform, widest posteriorly, becoming gradually 
narrower towards the front, then suddenly contract- 
ing to about the width of the free thoracic segment, 
length about equal to the width at the posterior end, 
and to about two-thirds of the length of the carapace ; 
bluntly rounded at the postero-lateral corners. Ab- 
domen tolerably elongated and narrow, being nearly 
three-fourths as long as the genital segment, slightly 
constricted near the distal end, the constriction forming 
a kind of false joint. Caudal rami very short and 
somewhat rudimentary. 

•Basal joint of the antennules tolerably expanded. 
Antennae robust, and armed with large and strong 



LEPEOPHTHEIRUS STURIONES. 75 

terminal claws, which have their ends bent at nearly 
a right angle to the basal part. Mandibles small, 
and resembling those of Leyeophtheirii^ pectoralu. 
Second maxillae stout, and dividing at the apex into 
two slightly bent teeth. Second maxillipeds mode- 
rately stout and elongated, and armed with short but 
strong terminal claws. Sternal fork very stout, with 
short rami which are stout at the base and taper to a 
tolerably sharp point, the rami distinctly divergent. 
Fourth pair of thoracic legs robust, ramus moderately 
elongated and composed of three joints, the middle 
joint being rather longer than the first or third ; the 
outer distal angle of the first joint terminating in a 
minute tooth, a tolerably stout spine springing from 
the outer distal angle of the second joint, the end joint 
armed with apical spiniform seta3 of unequal length, the 
innermost being considerably longer than the other two, 
and the outermost the shortest. Length about 14 mm. 

We have not seen the male of this species. 

Habitat. — Found as a parasite on the sturgeon, 
Acipensev sturio Linn. Obtained on a sturgeon 
captured about sixteen miles south-east of Aberdeen 
and landed at the Fish-Market there, 29th December 
1904. We are indebted to Dr. Alexander Bowman 
for this specimen. 

This species does not appear to be a very common one. 

Genus 8. LUTKENIA Clam, 1861. 

Syn. Cecropsina Heller, I860. 

Carapace obcordate, or nearly round, without frontal 
plates. Antennules two-jointed. Fourth ring of the 
thorax covered by small dorsal plates. Genital seg- 
ment prolonged backwards in the form of lobes. 
Abdomen short, not jointed, and terminating in two 
small caudal plates. 

The first pair of thoracic legs consisting of one or 
two branches, but the inner branch, when present, very 
smalL The next three pairs two-branched. Both 



76 BRITISH PARASITIC COPEPODA. 

branches of the second and third pairs two-jointed, and 
of the fourth pair one-jointed. 

1. Liitkenia asterodermi Claus. 
(Plate XIX, figs, i-9 ; Plate XXIII, fig. 4.) 

1864. Liitkenia asterodermi Clans. (33) p. 365-383, pi. xxxiv. 

1865. Cecropsina glabra Heller. (58) p. 209, pi. xix, figs. 1, 2. 

1906. Liitkenia asterodermi Norman & T. Scott. (88) p. 210, pi. xx, 
fi^s. 9, 10; pi. xxiv, figs. 1-8. 

1906. Liitkenia asterodermi A. Brian. (21) p. 48. 

Female. — Carapace suborbicular or obcordate, with 
the posterior margin deeply hollowed out. Last 
thoracic segment produced into a leaf -like bifurcated 
dorsal expansion, the distal end forming two rounded 
lobes which are separated by a tolerably deep and 
narrow sinus ; the width of the plate at the proximal 
end equal to about two-thirds the width of the cara- 
pace ; the anterior portion of the outer margin sloping 
in a slightly arcuate line from the middle towards the 
front corners which are distinctly angular. Genital 
segment produced backwards into two contiguous lobes 
which completely overhang and conceal the abdomen 
and caudal rami, so that they can only be seen from 
the ventral aspect. 

Abdomen somewhat lozenge- shaped, broader than 
long, the length being three-fifths the width. Caudal 
rami nearly contiguous, broadly lamelliform and 
scarcely half as long as the abdomen. 

Antennules very small, second joint slender and 
nearly as long as the first. Antennas armed with 
strong hook-like terminal claws. First maxillipeds 
slender and furnished with terminal claws, two of 
which are small and subequal and the others rather 
longer and stouter. Second maxillipeds large, basal 
joint stout with one or two nodulous processes on the 
inner aspect, the terminal claw large, strong, and 
sickle-shaped. First pair of thoracic legs small, bi- 
ramose, inner ramus minute, simple, and not half the 
length of the first joint of the outer ramus. The outer 
ramus two-jointed, the end joint very ^mall, its distal 



LUTKENIA ASTERODERMI. 11 

end truncated and provided with three or four minute 
apical spines. In the second and third pairs both rami 
two- jointed ; the first joint of the outer ramus consider- 
ably larger than the second; in the inner ramus the 
first joint is the smaller one. In the fourth pair, also 
biramose, both rami only one- jointed, the inner ramus 
being very minute. 

Habitat. — Parasitic on Luvarus imperialis Haf. 
Three specimens of this TAlthenia were sent to the 
Eev. A. M. Norman by Laughrin about 1863, which 
had been obtained by him from a specimen of Luvarus 
captured off Polperro, Cornwall. ('Crustacea of Devon 
and Cornwall,' by Norman and T. Scott, 1906, p. 210.) 

Distribution. — Mediterranean ; apparently very rare 
in the British Seas. 

Genus NOGAUS Leach, 1819.^ 
Syn. Nogagus M. Edwards, 1840. 

The genus Nogaus is now generally regarded as unsatis- 
factory, it comprises uiales only, some of which have already 
been recognized as belonging to more than one genus, of which 
the females had previously only been known ; and it is con- 
sidered probable that as our knowledge of the Copepod para- 
sites of fishes increases and their relationships and life-history 
are better understood, all the males ascribed to Nogaus will ere 
long be removed from it and the genus itself become obsolete. 
There are however, a few of the Nogaus males whose relation- 
ship is still doubtful, and it will be better to leave these 
where they are till they can be disposed of satisfactorily. 
Among these doubtful forms is the one described below. 

In Nogaus the frontal plates are without lunulaB, but other- 
wise the forms ascribed to this genus have a general resem- 
blance to Caligus. Steenstrup and Liitken divided Nogaus (or 
Nogagus) into two groups, the principal differences between 
them being as follow. In species belonging to the first 
group the four pairs of swimming legs are biramose, and the 
rami are all two-jointed ; the abdomen also is two-jointed. In 
those belonging to the second group, while the rami of the 

* The name " Nogaus " was used by Dr. Leach in 1819 ; it was afterwards 
changed to Nogagus by M. Edwards in 1840, but we find Dr. Baird still using 
the original word "Nogaus " in his 'British Entomostraca ' at p. 282 ; and 
C. B, Wilson in his recent work on ' North American parasitic Copepods of 
Fishes ' also adopts this form of the name. 



78 BRITISH PARASITIC COPEPODA. 

first three pairs are two-jointed, the fourth has only one- 
jointed rami, and the abdomen consists of a single segment. 
The form described below differs from both these groups, but 
partakes to some extent of the characters of both. 

Nogaus ambiguus T. Scott. (Provisional name.) 
(Plate XX, figs. 1-8.) 

1907. Nogagus amhiguus T. Scott. (117) Twenty-fifth Annual Report of 
the Fishery Board for* Scotland, Part III, p. 217, pi. xv, figs. 10-17 ( c? )• 

The cephalic shield in this form is of an oval outline, the 
front is somewhat narrowly rounded but the frontal plates 
are moderately large. The last two thoracic segments are 
subequal, their width is about one-third of that of the 
cephalic shield at its widest part, and the last segment is 
truncated behind. The abdomen consists of a single small 
subtriangular segment blunt-pointed at the apex ; and the 
caudal rami, wliich are short but moderately wide, are fur- 
nished with tolerably long plumose setae. 

The antennules, which are of average size, are provided 
with long plumose hairs. The antennae, mandibles, and 
maxillae are of the usual Caligus type. The first maxillipeds 
are elongated, and are each armed with a long and powerful 
terminal claw which has a moderately stout seta at its base, 
nearly as in Nogaus lunatics (Stp. & Liitk.), a species which 
the present form resembles in some other particulars. The 
second maxillipeds are short and very stout, and are each 
fitted with a stout claw which forms, with the tuberculated 
palm, a strong grasping organ. All the four pairs of swim- 
ming legs are short and biramose ; in the first three pairs 
both the outer and inner ramus are two-jointed and of nearly 
equal length, and the end joints of both rami bear tolerably 
long, densely plumose setae round the inner margin and end ; 
there are also a few short spines on the outer margin. The 
fourth pair are rather small, the inner ramus is biarticulate 
as in the other three pairs, but the outer ramus is composed 
of a single, somewhat club-shaped joint with three long 
plumose setae round the distal end of the inner margin, and 
with four spines — three small ones and a moderately large 
terminal spine — on the exterior edg'e. Length about 5'5mm. 

Habitat. — Parasitic on a piked dog-fish, Squahts acanthias, 
captured in the North Sea in 1902. 

This form has a somewhat close resemblance to N. lunatuH 
described by Steenstrup and Liitken [' Bidrag til Kundskab,^ 
p. 389, pi. ix, fig. 17, 1861). 



DEMOLEUS. 79 

Genus 9. DEMOLEUS Heller, 1865. 

Syn. Caligus Otto (not O. F. Miiller). 

Carapace suborbicular, divided into three parts by 
two longitudinal sutures, middle portion subquadrate, 
lateral portions narrow, produced backwards into pro- 
minent rounded lobes. Frontal plates distinct, with- 
out lunulse. Eyes conspicuous, close together. First 
and second free thoracic segments small; the next larger 
and prolonged backwards so as to form small dorsal 
plates in the female but which are wanting in the 
male. Grenital segment elongated in the female, sub- 
quadrate in the male. Abdomen short, not jointed, 
covered dorsally with a foliaceous lamina. Caudal 
appendages large. 

1. Demoleus paradoxus (Otto). 
(Plate XII, figs. 4, 5.) 

1828. Caligus paradoxus Otto. (93) p. 352, pi. xxii, fig. 5. 
1861. Nogagus grandis Steenstrup & Liitken. (127) p. 386, pi. x, 
fig. 19. 

1865. Demoleus paradoxus Heller. (58) p. 199, pi. 19, fig. 3. 

Female. — Carapace suborbicular and equal to about 
a third of the entire length of the animal ; frontal 
plates distinct, lunulse wanting, postero-lateral lobes 
considerably produced behind. First and second free 
thoracic segments small, and just filling the space 
between the produced lateral lobes of the carapace; 
the first segment with, and the second segment with- 
out, lateral processes. The next segment less than 
lialf the width of the other two and furnished with 
two small dorsal plates. Grenital segment elongated, 
length equal to fully twice the width, and divided 
posteriorly by a deep median sinus into two lobes 
which are rounded at the end. Abdomen very small, 
triangular, not seen from the dorsal aspect. Caudal 
rami large, extending beyond the end of the genital 
segment and bearing a few minute spiuiform setas. 

Antennules two-jointed, partly concealed by the 



80 BRITISH PARASITIC COPEPODA. 

frontal plates ; antenna small with weak terminal 
claws. The terminal claws of the second maxillipeds 
of moderate size. The first four pairs of thoracic legs 
all biramose, and the rami biarticulated and furnished 
with plumose setse, the first and fourth pairs with very 
small basal joints, those of the second and third con- 
siderably expanded. Egg-strings very long and slender, 
and looped so as to appear as if they each consisted of 
three strands. 

Male. — The male, which is of the usual Noqaus form, 
is somewhat similar to the female, but the suborbicular 
carapace is proportionally larger, being equal to at 
least three-sevenths of the total length. The first 
free thoracic segment fills the space between the lateral 
prolongations of the carapace, and is slightly produced 
into small rounded lobes which are contiguous with 
the lobes of the carapace. The next segment is small, 
while the third (the fourth counting the carapace) is 
without dorsal plates. Genital segment oblong, rather 
longer than broad, width equal to fully one-third the 
width of the carapace, and having the postero-lateral 
corners slightly produced and rounded. Abdomen 
small, two-jointed ; caudal rami large and lamelliform, 
longer than broad and furnished with small spiniform 
setse on their truncated distal extremities. Cephalic 
appendages somewhat like those of the female, but the 
second maxillipeds are short and tolerably stout, bear- 
ing stout terminal claws ; the thoracic legs are also 
similar to those of the female. Length variable, 
about 13 to 16"5 mm. 

Habitat. — Parasitic on dog-fishes. On a dog-fish 
captured off the North of Ireland. Belfast Bay (IF. 
TJiompson, ' Nat. Hist, of Ireland,' vol. iv, 1856)". 

We have not seen this species. Our figure of it is repro- 
duced from that of Otto referred to above. 

C. B. Wilson, after a critical examination of the characters 
of Nogagus grandis Steenstrup and Liitken, remarks it is 
" fairly certain ^' that this Nogagus is the male of Demoleus 
paradoxus (Otto). 



TEEBIUS. 81 

Genus 10. TREBIUS Kroyer, 1838. 

Eesembling Lepeophtheirus in its general form and 
in the absence of lunulae, and also generally in the 
structure of the body and its appendages, except that 
the fourth pair of thoracic legs are biramose in both 
the male and female. The second maxilla tolerably 
long with the extremity pointed or slightly bifurcated. 
The first maxillipeds rather stronger than in Lepeo- 
'phtlierni>i or Galigus, but the second pair less powerful 
than those of the two genera mentioned. 

Several species of Trehius have been described, but the one 
recorded below is the only species represented in the British 
fauna. 

1. Trebius caudatus Kroyer. 
(Plate XXII, figs. 1, 2: Plate LIV, figs. 1-11.) 

1838. Trehius caudatus Kroyer. (70) p. 30, pi. i, fig. 4. 
1850. Trehius caudatus Baivd. (4) p. 280, pi. xxxiii, fig. 3. 
1900. Trehius caudatus T. Scott. (112) p. loo, pi. vi, figs. 20-26. 
1907. Trebius caudatus C. B. Wilson. (147) p. 681. pi. xv, figs. 11-13 ; 
pi. xvi, figs. 14-22. 

Female. - — Carapace suborbicular, rather longer 
than broad, and equal to about a third of the entire 
length of the animal. Frontal plates narrow, without 
lunulge. Eyes conspicuous and close together. Free 
thoracic segment small. Grenital segment oblong, 
rather longer than broad, its width equal to about 
three-fourths of the length, and to about two-thirds 
the width of the carapace, the lateral margins nearly 
straight, its posterior end truncated, and the postero- 
lateral corners rounded and provided with three small 
but stout marginal spines. Abdomen elongated, 
narrow, and composed of three segments, its length 
about equal to that of the carapace, the proximal joint 
longest, being rather more elongated than the next 
two combined ; the second about twice the length of 
the third joint, the articulation between the second and 
third joints not very clearly defined. Caudal rami 
short and furnished with a few plumose apical set^. 
VOL. I. 6 



82 BRITISH PARASITIC COPEPODA. 

Antennules small. Antennae armed with strong 
terminal hooks. Mandibles similar to those of 
Lepeojphtheirus but the end joint somewhat stouter and 
more distinctly toothed. Second maxillge with the 
endopodite slightly bifurcate. Second maxillipeds 
tolerably elongated, not very robust and provided 
with rather weak terminal claws. Sternal fork 
small, with the rami simple, short, and slightly 
divergent. The four pairs of thoracic legs all 
biramose ; the first pair with both rami two-jointed, 
but the rami of the other three pairs composed of 
three joints ; the basal joint of the fourth pair short 
and tolerably expanded, the rami also short and of 
nearly equal length, the inner being rather the shorter, 
both rami with the inner margins fringed with 
tolerably long plumose setae. Fifth pair nearly 
obsolete. Length about 10 mm. 

Male. — Carapace orbicular and equal to about half 
the entire length. Genital segment small, ovate, 
somewhat longer but not much wider than the free 
thoracic segment. Abdomen biarticulate, narrow, and 
about as long as the genital segment ; proximal joint 
rather shorter than the end one. Length about 4*5 mm. 

Habitat. — Parasitic on skates, rays, dogfishes, &c. 
Belfast {W. Thompson). Polperro, Cornwall {A. M. 
Norman), Plymouth (B ass ett- Smith). Irish Sea 
(A. Scott). Firths of Forth and Clyde and at Aber- 
deen (T. Scott). 

Not very rare. The distribution of Trebius caudatvs 
appears to be somewhat limited; it is mentioned by C. B. 
Wilson in his ^ North American parasitic Copepods,' but the 
specimens, he states, were collected off the coast of Shetland. 



Genus IL ELYTROPHORA Gerstaeclcer, 1853. 

Female. — Carapace rounded, frontal plates distinct 
but without lunulse. First three thoracic segments 
fused with the head ; fourth segment with two dorsal 
plates. Genital segment lobed posteriorly. Abdomen 



ELYTROPHORA. 83 

two-jointed, joints siibequal, without wings. Caudal 
rami tolerably large. Antennules two - jointed. 
Antennas uncinate. Mouth-organs somewhat similar 
to those of Galigus. Thoracic legs, four pairs, all 
biramose. In the first pair both rami two- jointed ; 
both rami of the second and third pairs three-jointed, 
but in the fourth pair, while the outer ramus is three, 
the inner is only two-jointed. 

Ifa/e.— Somewhat similar to the female, but rather 
smaller. 

1. Elytrophora brachyptera Gerstaecker. 

(Plate XIX, fig. 10; Plate XXIII, figs. 1, 2; Plate 
XXXI, figs. 1-6.) 

1853. Elytrophora brachyptera Gerst. (48) p. 60, pi. iii, figs. 1-14. 
1863. Ar7i£eus thynyii Kroyer. (71) p. 157, pi. viii, fig. 5 a-g. 
1865. Elytrophora brachyptera Heller. (58) p. 189, pi. xvii. 
1896. Elytrophora brachyptera Bassett-Smith. (6) p. 12, pi. iv, fig. 3. 

Female. — Carapace orbicular, scarcely equal to half 
the entire length of the animal, or as 5 to 11. Frontal 
plates distinct, without lunulse. The fourth thoracic 
segment carrying two small dorsal plates which are 
somewhat widely apart in front, but gradually 
approach each other behind; their outer margins 
incurved, and the posterior margins, which are 
rounded, slightly overlapping the anterior edge of 
the genital segment. Genital segment ovate and 
moderately tumid, nearly half as wide as the carapace 
and about one and a half times longer than broad, 
the lateral margins slightly arcuate, and the postero- 
lateral corners produced into short rounded lobes. 
Abdomen biarticulate, narrower than the genital 
segment, first joint with the postero-lateral corners 
somewhat produced and rounded, anal segment sub- 
orbicular with a minute posterior lobe in the median 
line; caudal rami obovate, expanded towards the distal 
end, and bearing four or five apical plumose setas. 

Antennules small. AntennaB strongly uncinate. 
Mandible and maxillae somewhat like those in Trebius. 



84 BRITISH PARASITIC COPEPODA. 

Second maxillipeds armed with strong terminal claws. 
Thoracic legs four pairs, all biramose ; both rami of 
the first pair two -jointed, the outer ramus tolerably 
elongated, but the inner very small ; both rami of the 
second and third pairs composed of three joints, but in 
the second the rami are nearly of equal length, with the 
end joints very small, while in the third the outer ramus 
is distinctly shorter than the inner. These three pairs 
of thoracic legs all liberally supplied with densely 
plumose setae. In the fourth pair the outer ramus 
stout, and composed of sub-equal joints, both the first 
and second joints having a strong and slightly-curved 
spine on their outer distal angle, while the end joint 
has three similar spines on its outer margin, and three 
or four small ones on the inner margin; the inner 
ramus, which is situated close behind the outer, small, 
and composed of two subequal joints ; the basal joint 
of the fourth pair also tolerably large, and gibbous 
below. Length about 11*5 mm. 

Male. — The male, though somewhat similar to the 
female in its general appearance, is smaller, and the 
second maxillipeds are more powerfully clawed ; and 
while the genital segment is only about half as large, 
the abdomen is rather longer than in the female. The 
postero-lateral corners of the penultimate segment of 
the abdomen are angular, while the anal segment has 
a subquadriform outline. The caudal rami are furnished 
with four setae longer and more densely plumose than 
in the female. Length about 9 mm. 

Habitat. — Parasitic on the tunny, Orcynns thynnu^, 
Plymouth {B as sett 'Smith). Outer Hebrides {Dr. Alex. 
Boivman). We are indebted to Dr. Bowman for speci- 
mens of this interesting species. 

Distribution. — Euroj^ean seas. 

The species is not an uncommon one on the tunny. 



DINEMOTJRA. 85 

Genus 12. DINEMOURA Latreille, 1829. 

Syn. Dinematura Burmeister, 1833. 

Female. — Carapace suborbicular, deeply excavated 
posteriorly ; frontal plates narrow. The first free 
segment of the thorax with small lateral lobes; the 
next segment narrow with or Avithout rudimentary 
dorsal plates ; the third segment with dorsal plates of 
tolerable size, separated by a narrow median fissure, 
and overlapping the anterior portion of the genital 
segment. The genital segment of an oblong form 
and with the postero-lateral corners produced into 
short, broadly-rounded lobes. Abdomen small and 
uniarticulate ; caudal rami tolerably large and 
foliaceous. Between the genital segment and the 
abdomen there is a very small joint with two dorsal 
plates and furnished below with a pair of rudimentary 
legs.* 

Antennae short and moderately stout, and armed 
with strong hooked terminal claws. Mandibles long 
and very slender, and provided with a few minute 
teeth near the tip. Second maxillae slender, three- 
jointed ; first maxillipeds also slender, and furnished, 
each, with an apical claw and two small lateral pro- 
cesses. Second maxillipeds moderately stout. Swim- 
ming-legs all l)iramose ; first pair with both rami 
two-jointed ; those of the second and third pairs 
three-jointed, while in the fourth pair both rami con- 
sist of tolerably large one-jointed foliaceous plates. 

Male. — Carapace proportionally wider than in the 
female. Second free thoracic segment without dorsal 
plates. The dorsal plates of the third segment small 
and overlapping only a small portion of the genital 
segment. Genital segment cuneiform, wider towards 
the distal end. Abdomen very n arrowy biarticulate ; 
caudal rami large. Swimming-legs biramose, both 

* C. B.Wilson appears to be the first to give an accurate description of this 
part of the animal which he names the sixth segment. Cf. 'North American 
Parasitic Copepoda,' * Proc. U. S. National Museum/ vol. xxxiii, pp. 374, 376 

(1907). 



86 BRITISH PARASITIC COPKPODA. 

rami of the first and fourtli pairs composed of two 
joints, and those of the second and third pairs of 
three joints ; the rami of the first three pairs are 
furnished with tolerably long plumose setae while those 
of the fourth pair are armed with spines. 
The only British species is that described below. 

1. Dinemoura producta (0. F. Midler). 

(Plate XXII, fig. 3 ; Plate XXVI, figs. 1-3 ; Plate 
XXVII, figs. 1-8.) 

1785. Caligus productus O. F. Miiller. (86) p. 132, pi. 21, fig. 3. 

1829. Binemoura producta Latr. (38) Cuv. Regne Anim., vol. iv, p. 127. 

1835. Pandarus lainnse Johnston. (65) p. 203. 

1850. Dinemoura lamnse Baird. (4) p. 286, pi. xxxiii, fig. 7. 

1853. Nogagus productus Gerst. (48) Wiegmann's Archiv fiir Natnr- 
gesch., vol. xix, p. 63, pi. iv, figs. 1-10. 

1857. Dinematura elongata P. J. van Beneden. (14a) Bull. Acad. 
Roy. Belg., p. 231, pi. xxiv. 

1861. Dinematura ^'iroducta Stp. & Liitk. (127) p. 34, pi. vii. fig. 13. 

1900. Dinematura producta T. Scott. (112) p. 156, pi. vi, figs. 27-31. 

1907. Dinematura producta C. B. Wilson. (148) p. 380, pi. xxiii. 

Female. — Carapace suborbicular and equal to about 
a third of the entire length of the animal; width 
slightl}^ exceeding the length ; lateral margins boldlj 
rounded ; frontal plates narrow ; eyes small, but quite 
distinct, nob quite close together and distant from the 
front margin about a tenth part of the total length. 
The first free thoracic segment furnished with small 
lateral plates which inclose the next segment between 
them, these two segments together filling the entire 
space between the produced postero-lateral lobes of 
the carapace. The third segment of the thorax carry- 
ing a dorsal plate which is equal to about one-sixth of 
the entire length of the animal and is as wide as the 
genital segment the anterior part of which it overlaps ; 
the antero-lateral corners of the plate subangular, 
and also divided from behind forward into two lobes 
by a narrow median fissure which extends to near its 
base, each lobe being boldly rounded at the posterior 
end. G-enital segment oblong, considerably longer 



DINEMOURA PEODUCTA. 87 

than the segment last described and equal to nearh^ 
two-thirds of the width of the carapace ; the postero- 
lateral corners of the segment produced backwards into 
moderately narrow lobes with rounded ends and hav- 
ing a space between them equal to about the width of 
one of the lobes ; a shallow median groove also extend- 
ing along nearly the whole length of the segments as 
indicated in the drawing (PL XXII. fig. 3). The space 
between the lobes occupied by a small plate provided 
with two slightly divergent appendages which scarcely 
reach to the end of the abdomen. Abdominal segment 
small, subquadraugular, concealed in dorsal view. 
Caudal rami in the form of broad foliaceous plates 
rather longer than broad, and having one marginal, 
and three short, tolerably stout, spiniform setae. There 
is also a small segment, intermediate between the 
genital segment and abdomen, which is provided with 
lateral uniarticulate and somewhat rudimentary appen- 
dages, but this segment is seen only from the ventral 
aspect. 

Antennules small. Antenna short and stout and 
armed with strongly-hooked terminal claws. Mandi- 
bles elongated and extremely slender, with a few 
minute marginal setse near the tip. First maxillipeds 
each furnished with a terminal and tolerably elon- 
gated hook-like appendage and one or two short 
accessory processes. Second maxillipeds short and 
stout, but somewhat rudimentary in structure. Swim- 
ming-legs all biramose ; both rami in the first pair 
two-jointed, the first joint of the outer ramus consider- 
ably expanded, and its outer distal angle reaching 
forward to the middle of the short end joint and 
terminatiug in a short stout spine ; the inner ramus 
very small and the joints subequal. The fourth pair 
large and foliaceous, each ramus about twice as long as 
broad, and furnished with a few minute spines round 
the distal end. Length exclusive of egg-strings about 
20 mm.; egg-strings alone sometimes reaching to 80mm. 

We have not seen the male of this species. 



88 BEITISH PARASITIC COPEPODA. 

Habitat. — Parasitic usually on the porbeagle shark, 
Lamna cornuhica. Berwick Bay, 1834 {Dr. Johiisfon). 
Polperro, Cornwall {A. M. Norman). Moray Firth — 
off Dunrobin, and at Aberdeen Fish-Market (T. Scott). 
Near Shetland (C. B. Wilson). On a thrasher shark, 
Alopias vuljjes, at Buddon, Firth of Ta}^ in 1887 
{Prof. d'Arcy W. Thowpson). Recorded also from the 
Grreenland shark, Scymnus glacialis. 

Distribution, — European waters. Atlantic coast of 
North America. 



Genus 13. ECHTHROGALEUS Steenstriip c^- Liltken, 

1861. 

Female. — Carapace, antennae, and mouth-appen- 
dages nearly as in Phyllortkragorisctis, but the dorsal 
plates of the last thoracic segment larger. The genital 
segment, which is also large, extending backwards so 
as to overlap and conceal wholly or partially the 
abdominal segment and caudal rami ; the postero- 
lateral lobes of the genital segments rounded at the 
end and separated by a tolerably deep sinus which 
may be narrow as in E. coleoptratns or moderately 
wide as in E. dentindatus. 

Thoracic limbs as in Phyllorthrag oris ens except that 
the inner ramus of the second and third pairs are 
composed of two instead of three articulations. 

'' Male. — Carapace like that of the female but pro- 
portionately larger, frontal plates more prominent. 
Lateral lobes of second thoracic segment corresponding 
to the first pair of dorsal plates in the female ; no 
lobes on the third segment; a rudimentary pair on 
the fourth segment wdiich are closely appressed to the 
anterior margin of the genital segment. The latter 
smaller than in the female, w^ith one pair of legs at or 
just in front of the posterior corners. Abdomen 
small and two-jointed; anal lamina large and armed 
with plumose setse. Appendages as in the female." 
{C. B. Wilso7i.) 



ECHTHEOGALEUS COLEOPTEATUS. 89 

1. Echthrogaleus coleoptratus (Guerin). 
(Plate XXII, fig. 4.) 

1829-1843. DmematuracoJeoptrata Gnerhi. (55) pi. xxxv, fig. 6. (1840). 

1835. Pandarus alatvs ("M. Edwards") Johnston. {QQ) p. 202, two 
text-figs. 

1850 Dinemoura alata Baird. (4) p. 285, pi. xxxiii, figs. 8, 9, 

1861. Echthrogaleus coleoptratus Stp. & Ltk. (127) p. 380, pi. viii, 
fig. 15. 

1900. Echthrogaleus coleoptratus T. Scott. (112) p. 156, pi. vi, fig. 52. 

1907. Echthrogaleus coleoptratus C. B. Wilson. (148) p. 367, pi. xix. 

1910. Echthrogaleus coleoptratus T. R. R. Stebbing. (125) p. 559. 

Female. — Body oblong, fully twice as long as broad. 
Carapace suborbicular and equal to about a third of 
the entire length of the animal ; frontal plates toler- 
ably large and distinct. First two thoracic segments 
short and narrower than the carapace ; dorsal expan- 
sion not greatly developed. Dorsal plates of the third 
segment much enlarged, wider behind than in front, 
and covering rather less than half the genital segment, 
their posterior margins obliquely and sinuately trun- 
cated so that the inner corners extend further back- 
ward than the outer, and though the inner margins of 
the plates come close together in the middle line they 
do not overlap, their margins being even, not serrated, 
and their surface smooth and ornamented with a 
number of small pellucid impressed circular markings 
arranged in a more or less regular pattern, which, 
along with the form of the plates, gives them a fairly 
close resemblance to the elytra of certain coleopterous 
insects. Genital segment tolerably large, extending 
backwards considerably beyond the dorsal plates 
described above, becoming somewhat narrower pos- 
teriorly and being divided into two lobes by a narrow 
and deep median cleft or sinus ; the inner edges of 
the lobes closely appressed and not overlapping, and 
their posterior margins rounded and reaching to about 
the end of the caudal rami ; the abdomen, which is 
concealed in dorsal view by the genital segment, com- 
paratively small and sub-quadriform in outline, the 
width being somewhat greater than the length ; caudal 



90 BUITISH PARASITIC COPEPODA. 

rami lamelliform, broadly ovate, the ends subtruncate, 
scarcely extending beyond the genital segment, and 
provided with a few apical spinules. Intermediate 
between the genital segment and abdomen is a small 
rounded plate concealed in dorsal view. 

Antennules rather slender, not prominent; antennas 
furnished with strong, hooked, terminal claws. Mouth- 
tube moderately short and slender. Second maxillipeds 
short, moderately stout, and furnished with powerful 
terminal claws. Other mouth-organs as in Binemoura. 
Swimming-legs short and biramous ; first and second 
pairs with both rami two-jointed ; third pair with the 
outer ramus composed of three and the inner of two 
joints ; while in the fourth pair both rami consist of a 
single foliaceous joint. The fifth pair of thoracic legs 
represented by a small spine-like process on the under 
side and near the posterior end of the genital lobes. 

Length 11 to 13 mm. Egg-strings long and slender. 

Habitat. — Found parasitic on the porbeagle shark, 
Lamna coimuhica. Berwick Bay (Dr. Johnston), 
Aberdeen Fish-Market and near Fair Island between 
Orkney and Shetland {T. Scott). From both the por- 
beagle and the blue shark, Carcharias glaucus, taken 
at Polperro, Cornwall {A. M. Norman). 

The distribution of this Echthmgaleus is extensive and 
includes the seas of Europe, the Atlantic and Pacific coasts 
of Nortli America, and the coast of South Africa. 

2. Echthrogaleus liitkeni (Norman). 
(Plate XXIX. fig. 1 ; Plate XXX, figs. 1-8.) 

1869. Nogagus liitkeni Norman. Last Rept. on Dredging among the 
Shetland Isles; Brit. Assoc. Rept. for 1868, p. 300. S 

1906. Echthrogaleus liltheni Norman & T. Scott. (88) p. 213, pi. xxii, 
figs. 1-9. 

In the " Last Report on Dredging among the 
Shetland Isles," the Rev. A. M. Xorman records and 
describes a fish parasite under the name of Nogagus 
liltheni. The specimen, wdiich was procured by Dr. 
Saxby, was found on a skate. Two other specimens 



ECHTHROGALEUS LUTKENl. 91 

were subsequently obtained at Polperro in Cornwall, 
but the name of the fish on which they occurred has 
not been recorded. All these specimens were males. 
In dealing with this group of fish parasites it is some- 
times difficult, if males only are available, to determine 
the species to which the specimens belong ; a careful 
study, however, of the examples referred to, leaves 
little doubt that they are the males of an Echtliro- 
(jaleits. 

The following is the description of E. liitkeni as 
given in ' Crustacea of Devon and Cornwall ' : — 
" The cephalosome is much rounded, its breadth 
much greater than the length ; the hinder corners of 
the lateral area incurved, w^ell rounded, and reaching 
backwards to the end of the first of the two exposed 
segments of the metasome. First segments of meta- 
some Avith lateral expansions broader and longer than 
in the following short segment. The urosome consists 
of three segments ; the first, or genital segment, is sub- 
quadrate with slightly arched sides, longer than broad; 
the second segment very short, the terminal rather 
broader than long; the uropodal lamina are as long as 
the two preceding joints, and of an ovate form. 

" The antennules have the first joint much longer 
than the second ; the second joint has one spine on 
the hinder margin. The antennse have the penulti- 
mate joint stout, the last long and gradually attenuated, 
only very slightly curved, bearing a single seta on the 
inner face. The first maxillipeds have the claws 
setose. The second maxillipeds have the terminal 
joint very broad and stout, obliquely truncate distally, 
with a nodule, and areolated disk at the commence- 
ment of the palm ; the finger short and stout. The 
first feet have both rami composed of two joints ; the 
inner branch terminates in three setse, the outer in 
four ; the first joint of this outer branch has one spine 
on the outer margin, and the second joint three. The 
second, third, and fourth feet are alike in general 
character, though differing slightly in the number of 



92 BRITISH PARASITIC COPEPODA. 

setse and external marginal spines. The fiftli feet, 
which are situated under the genital segment, are 
minute, one-jointed, bearing two or three short setae. 
" Polperro, two specimens, but the record of the fish 
on which they were found was not given. The type 
specimen was taken on a skate at Shetland." (See 
pp. 213-214.) 

Our figure of the species is reproduced from that in the 
work referred to. We have not, ourselves, obtained this 
parasite. 

Genus 14. PHYLLOTHREUS Norman, 1903. 

Syn. Phyllophora M. Edw. (name preoccupied). 

Body depressed. Carapace broadly cordate, without 
frontal plates. Thorax furnished with three pairs of 
broadly-rounded overlapping laminae, which extend 
considerably on either side of the carapace. Abdo- 
minal region two-segmented ; genital segment tolerably 
short and broad ; distal segment small, rounded, and 
provided with short lateral processes. 

Antennules small. Antennae assuming the form of 
large hooks which project in front of the carapace. 
All the thoracic legs biramose and lamelliform. Egg- 
strings slender and elongated. 

The name Phyllophora employed by M. Edwards for this 
genus Avas preoccupied by Thunberg in 1812 (for a genus of 
Orthoptera), and by Gray in 1838. 

1. Phyllothreus cornutus (M. Edwards). 
(Plate XIX, figs. 11-18; Plate XXIII, fig. 3.) 

1840. Phyllophora comuta M. Edw. (43) p. 471, pi. xxxviii, figs. 13, 14. 

1899. Phyllophorus cornutus Bassett-Smitli. (8) p. 465. 

1903. Phyllothreus cornutus Norman. (87) Ann. and Mag. Nat. Hist. 
(7), vol. xi, p. 368 (April). 

1906. Plujllothreus cornutus Norman & T. Scott. (88) p. 212, pi. xxiv. 
figs. 9-17. 

Female. — Carapace subcordate, wider posteriorly, 
greatest width equal to about one and one-tbird times 
the length, lateral margins slightly arcuate, converging 



PHYLLOTHREUS CORNUTUS. 93 

towards the front which is truncated and without 
frontal plates ; posterior edge deeply emarginate, and 
with the lateral corners rounded. Thorax provided 
with three pairs of leaf-like and suborbicular over- 
lapping plates, which extend on each side to consider- 
ably beyond the margin of the carapace and reach 
backwards to a distance equal to the length of it. 
Abdominal region distinctly narrower than the cara- 
pace and composed of two segments; the genital 
segment suborbicular, its length being rather less than 
the width, and bearing, at the corners of the truncated 
hinder margin, ovate processes which represent the 
fifth pair of feet ; the last segment consisting of two 
small transversely-obovate plates from between which 
issue the two slender and elongated egg-strings. 

Antennules small, two-jointed, end joint narrow^ 
much shorter than the proximal one, and bearing a 
few apical setse and a minute spine near the middle of 
the lower margin; the antennae are represented by 
large hooks which project considerably in front of the 
carapace and form powerful grasping organs. First 
maxillipeds with the end joint narrow and provided 
with three terminal spines, the middle one being the 
largest, tips of all three slightly hooked ; second 
maxillipeds cheliform. Thoracic legs short, biramous, 
and more or less lamelliform ; first pair v^ith outer 
ramus one-jointed and slightly geniculated, and the 
inner tw^o-jointed ; second pair with both rami two- 
jointed ; while in the third and fourth pairs both 
rami are uniarticulate. All the rami are devoid of 
setae and bear only a few small spines. Length about 
13 mm. Male unknown. 

Habitat. — Parasitic on the blue shark. Gar charms 
glaucus. Two specimens were collected by William 
Laughrin at Polperro, Cornwall, many years ago, and 
were sent by him to the Rev. Canon A. M. Norman. 
'' The only other previously known habitat of the 
species was, according to Milne Edwards, Tongatabu 
in the Friendly Islands, whence the type specimens 



94 BRITISH PARASITIC COPEPODA. 

came ; this is therefore a remarkable instance of wide 
distribution " (A. M, Norman). 

We are greatly indebted to Canon Norman fur the privilege 
of examining* the specimens referred to, and for permitting us 
to reproduce the figure of it published in ' Crustacea of Devon 
and Cornwall/ 

Genus 15. PANDARUS Leach, ]816. 

Female. — Body moderately broad and elongate. 
Carapace ovate, wider behind than in front ; frontal 
plates of moderate size ; posterior margin not very 
deeply excavated. Thorax provided with three pairs 
of dorsal plates, the first pair lateral and of an 
elongate-ovate form, the second pair between them in 
the form of broadly-rounded lobes separated from 
each other by a moderately deep median sinus; the 
third pair nearly as wide as the carapace and prolonged 
backwards so as to cover a considerable portion of the 
genital segment ; the dorsal plates of the genital 
segment with the inner lateral margins contiguous and 
apparently coalescent, and produced posterioidy into 
rounded or angular lobes having between them a 
flattened suborbicular median plate concealing the 
abdomen. 

Antennules biarticulate and provided with an 
adhesive disk at the base. Antennge armed with 
terminal claw-like hooks, and also furnished with 
adhesive disks or pads somewhat similar to those at 
the base of the antennule;s and maxillipeds. Thoracic 
legs biramose ; rami of the first three pairs biarticu- 
late, the outer ramus of the first pair more or less 
abnormal in form and the joints sometimes coalescent ; 
in the fourth pair the rami consisting of a single joint. 
Caudal rami subtriangular and divaricate, usually seen 
projecting — one on each side — beyond the plate at the 
end of the genital segment. 

Male. — The male has a close general resemblance to 
the male of Lejjeophtheirus, the antennules, which are 
provided with adhesive disks as in the female, differ in 



PANDARUS. 95 

being armed with terminal claws ; the maxillipeds are 
also furnished with small terminal but distinct claw- 
like hooks. The other mouth-appendages are some- 
what similar to those of the female. All the four 
pairs of thoracic legs are biramose, with bi articulate 
rami, and they differ from those of the female in being 
provided with moderately long and densely plumose 
setse. 

1. Pandarus bicolor Leach. 

(Plate XXI, fig. 2; Plate XXII, figs. 5-6; Plate 
XXVI, figs. 4-19 ; Plate LVIII, figs. 1-8.) 

1816. Pandarus hicolor and hoscii Leach. (74) pp. 405, 406, pi. xx, 
figs. 1 and 2 and figs. 1-10. 

1840. Pandarus fissifrons M. Edwards. (43) p. 470. 

1850. Pandarus hicolor and hoscii Baird. (4) pp. 288, 289, pi. xxxiii, 
fig. 6. 

(?) 1854. Nogagus augustulus Gerst. {^). (48) p. 193, pi. vii, figs. 17 
and 18 (cT). 

1900. Pandarus hicolor T. Scott {^ & $). (112) p. 157, pi. vi, figs. 
33-38 (cJ & ?). 

1907. Pandarus hicolor C. B. Wilson. (148) p. 400, pi. xxvii. 

1907. Nogagus latus T. Scott. (117) p. 216, pi. xv, figs. 1-9 (c^, im- 
mature). 

Female. — Body oblong, width equal to leather more 
than a third of the entire length. Carapace widest 
posteriorly, lateral margins slightly arcuate, converg- 
ing towards the front, anterior margin with a small 
median notch, frontal plates tolerably distinct, postero- 
lateral corners not greatly produced. Thorax provided 
with three pairs of dorsal plates ; first pair lateral and 
of an ovate form and inclosing the second pair between 
them ; the second pair terminating in broadly-rounded 
lobes which scarcely reach beyond the ends of the 
first, the combined width of the two pairs being 
nearly equal to that of the carapace ; the posterior 
margins of the two pairs of plates are nearly in a 
straight line and have the appearance of four subequal 
lobes ; the third pair about as wide as the carapace and 
prolonged backward to near the middle of the genital 
segment, separated posteriorly by a moderately deep 
median sinus and terminating in two broadly-rounded 



96 BRITISH PARASITIC COPEPODA. 

lobes. Dorsal plates of the genital segment consider- 
ably produced, their inner margins contiguous and 
coalescent ; posterior corners prolonged into rounded 
lobes, the space between the lobes being filled by a 
suborbicular lamina which entirely conceals the 
abdomen, while the caudal rami are seen projecting 
one on each side of the median plate in the form of a 
triangular process. 

Antennules small, partly concealed by the frontal 
plates ; antennas also small, terminal claws slender 
with an accessory spine on the inner margin. 
Adhesion-disks four pairs. First maxillipeds slender, 
joints subequal, terminal claws unequal ; second 
maxillipeds considerably dilated. Thoracic legs 
biramose ; the inner ramus in the first pair biarticulate, 
the end joint being longer than the proximal one, and 
with the apex broadly rounded and bearing a few 
setse on the lower half of the inner margin; outer 
branch longer than the inner and composed of two 
partly or wholly coalescent joints, the proximal part 
being distinctly wider than the distal portion, which 
is abruptly geniculated and tapers towards the apex ; 
second and third pairs with both rami two- jointed ; 
while in the fourth pair each ramus is composed of a 
single joint. Length about 10 mm. 

Male. — The male has a general resemblance to the 
male of a Lepeo])htlieiriis. The antennules are provided 
with adhesive disks as in the female, but differ in 
being armed with terminal claws ; the second maxilli- 
peds are also furnished with distinct though small 
claw-like hooks. All the four pairs of thoracic legs 
are biramose with biarticulate rami ; the rami also 
differ from those of the female in being provided with 
moderately long and densely plumose setse. The 
genital segment is subquadriform and equal to scarcely 
one-fourth of the total length, and the abdomen is 
composed of two segments. Length about 6 mm. 

Habitat. — Parasitic on various dog-fishes and sharks: 
Galens canis [Squalus galeushmii.) ; Carcharias glaucus ; 



PANDARDS BICOLOR. 97 

Scyllmin catulus ; and Squalus mustelus. Falmouth 
1849 {W, P. Cocks— see Baird). Torcross, Devon (Dr. 
Leach). Plymouth {A. M. Norman and Bassett- Smith). 
Fh'th of Clyde, Aberdeen, Moray Firth (T.' Scott). 
Shetland (G. B. Wilson). Irish Sea (A. Scott). 

• This appears to be the only British species of Pandarus 
recorded hitherto. Pandarus hoscii Leach is considered to 
be only a pale-coloured variety ; an apparently similar variety 
— pale-coloured — is recorded in the 26th ' Annual Report of 
the Fishery Board for Scotland/ Part III, p. 74, pl.iii (1909). 
It was obtained along with specimens of the typical form on 
a number of Squalus acanthias captured on the West Coast 
of Scotland in March 1908. Pandarus fissicornis M. Edw. 
is regarded by Kroyer as synonymous with the present species, 
and Canon A. M. Norman considers that the species described 
by Gerstaecker under the name of Nogagus angustulus is the 
male of this Pandarus. The iemale of the typical form is 
ornamented on the back with dark-brownish or almost black 
coloured patches. The carapace is usually coloured, and so 
are the second and third pairs of thoracic plates. 



Genus 16. CECROPS Leach, 1816. 

Female. — Carapace oval, robust, distinctly notched 
in front and deeply excavated posteriorly. Frontal 
plates coalescent with the carapace. Second thoracic 
segment with tolerably large rounded lateral lobes, 
and the last one with a pair of short dorsal plates. 
Genital segment small but provided with considerably 
expanded dorsal plates, larger than the carapace, and 
reaching backward so as to conceal the short caudal 
rami. Abdomen small, somewhat flattened. 

Antennules small, two- jointed. Antennas moderately 
stout, each terminating in a strong hook. Mandibles 
and other mouth-organs of the usual caligoid type. 
All four pairs of thoracic legs biramose. Both rami 
of the first three pairs composed of two joints, and 
those of the fourth pair of one joint. 

Male. — The male, which is smaller than the female, 
is similar to it, except that the genital segment wants 

VOL. I. 7 



98 BEITISH PAEASITIC COPEPODA. 

the large dorsal plates wliicli the female possesses. 
The frontal sinus is also not so pronounced. 

1. Cecrops latreillii Leach. 

(Plate XXI, figs. 3, 4; Plate XXVII, figs. 9, 10; 

Plate XXVIII, figs. 1-7.) 

1816, Cecrops latreillii Leach. (74) p. 20, and five figures. 

1850. Cecrops latreillii Baird. (4) p. 293, pi. xxxiv, fig. 1. 

1857. Cecrops latreillii Hoeven. (61a) Mem. d'Entomol. de la Soc. 
Entom. des Pays-bas, vol. i, p. 67, pis. iii & iv. 

1892. Cecrops lalreillii A. Scott. Trans. Nat. Hist. Soc. Glasgow, 
vol. iii, pt. 3, p. 266. 

1900. Cecrops latreillii T. Scott. (112) p. 157. 

1907. Cecrops latreillii C. B. Wilson. (148) p. 468, pis. xxxviii & xxxix. 

1909. Cecrops latreillii E. V. Elwes. (45) p. 20. 

1910. Cecrops latreillii T. R. R. Stebbing. (125) p. 558. 

Female. — Carapace oval, stout, length and width 
nearly equal, deeply emarginate in front, and with the 
posterior margin deeply excavated. Second and third 
thoracic segments apparently coalescent and forming- 
one segment which bears a dorsal plate with broadly- 
rounded lateral lobes. The last thoracic segment with 
a tolerably large dorsal plate, the front margin of 
which is nearly straight, but posteriorly of a semi- 
circular outline and reaching to near the middle of the 
genital segment; a narrow and moderately deep cleft 
or sinus dividing the plate in the median line at its 
posterior margin. The dorsal plates of the genital 
segment nearly twice as long as the carapace and 
extending backward so as to conceal the abdomen and 
caudal rami ; their inner margins contiguous and 
apparently coalescent, forming a single plate which is 
wider than the carapace and terminates posteriorly in 
two equal and broadly - rounded lobes ; the genital 
segment covered by these coalescent plates narrow and 
flattened, and the abdomen and caudal rami small. 

Antennules two-jointed, end joint small. Antennae 
armed with strong, hooked, terminal claws. Second 
maxillipeds stout, terminal claws strong and hook- 
like. All the four pairs of thoracic legs biramose and 



CECROPS LATRKILLII. 99 

both rami two-jointed, except in tlie fourth pair, the 
rami of which are both uniarticulate. The outer 
ramus in the first three pairs stouter than the inner, 
and the first joint larger than the end one, and 
carrying on its outer distal angle a short but stout 
spine. The outer ramus in the fourth pair small, 
but the inner considerably expanded and so also is 
the basal joint of this pair. Caudal rami short and fur- 
nished with a few apical setaB. Length about 25 mm. 

Male. — The male differs from the female in being 
smaller, and it also wants the large posterior dorsal 
plates of the genital segment. Some of the thoracic 
legs are also slightly modified. Length about 16 mm. 

Habitat. — Parasitic on the gills of the short sun-fish, 
Orthagoriscus mala. Recorded from many parts of 
our coasts. South of Ireland, Dublin, Coast of 
Antrim, &c. {G. J. Allman—Aug. 1848, B. Ball, W. 
Thomjpson, — see Baird, p. 293). Polperro {A. M. 
Norman). Plymouth {Bassett- Smith). Falmouth 
(Cocks). Solway, Jan. 1857 (/. Steiuart). Firth of 
Forth (A. Scott). Lerwick, Shetland; Mallaig, W. 
coast of Scotland (T. Scott). On a short sun-fish 
captured off Berry Head (Major E. V. Elives). 

In the female the egg-strings are very long, but instead of 
projecting externally they are twisted upon each other in 
numerous loops and lie concealed in the hollow space 
between the abdomen and the large buckler-shaped last 
segment of the thorax. The specimens we have seen have 
all been obtained from the gills of the short sun-fish, where 
large numbers may sometimes be found crowded together. 
The general colour of the parasites is yellowish, but the 
colour of the strong terminal hooks of the maxillipeds is 
dark brown, nearly black. 

Cecrops latreillii appears to be one of the most widely 
dispersed species of the CaligidaB; in addition to the 
European records of this parasite, it has also been reported 
from both the Atlantic and Pacific coasts of North America, 
and Gr. M. Thomson states that several specimens were 
obtained from the mouth of a sun-fish {Orthagoriscus mola) 
which was caught in Otago Harbour (Otago Museum)."^ 
* 'Trans. N. Z. Institute/ vol. xxii (1889), p. 362. 



100 BRITISH PAEAISITIC COPEPODA. 

Genus 17. ORTHAGORISCICOLA Poche, 1902. 

Syn. Lxmargus Kroyer. (Name preoccupied for a genus of fishes.) 

Female. — Carapace, outline trapezoidal, consider- 
ably narrower in front, posterior corners rounded, 
length equal to about three-fourtlis of the width at 
the widest part. The first and second thoracic seg- 
ments very short and fully half as wide as the carapace. 
Third segment provided with an expanded dorsal 
plate, which overlaps a considerable portion of the 
genital segment and is divided posteriorly by a median 
cleft into two broadly-roimded lobes. G-enital segment 
enlarged, dorsal plates greatly expanded, contiguous, 
slightly overlapping each other in the median line, 
and, viewed from the dorsal aspect, entirely concealing 
the abdomen and caudal rami. 

Antennules tolerably elongated and composed of 
three joints. Antennae, mandibles, and maxillipeds 
somewhat similar to those of FJnlorthragoriscits. All 
the thoracic legs biramous ; the first and second pairs 
both somewhat similar to tlie normal type observed in 
this group of parasites, but in the third and fourth 
pairs the rami consist of broad one-jointed laminae 
almost devoid of spines or setee. 

Male. — The male resembles the female but is 
rather smaller. The structure of the appendages 
does not greatly differ except that the antennge and 
second maxillipeds are armed with stronger terminal 
claws ; the rami of the third pair of thoracic legs are 
also biarticulate. 

Kroyer estabhshed this genus in 1837 under the name of 
Lsemargiis, but it happened that only a short time before the 
same name had been sfiven to a genus of fishes by Henle 
(1837). 

1. Orthagoriscicola muricata (Kroj^er). 
(Plate XXI, %. 5; Plate XXVIII, figs. 8-18.) 

1S37. Lxmargus muricatus Kroyer. (70) p. 487, pi. v. figs. a-e. 
1850. Lxmargus muricatus Baird. (4) p. 295, pi. xxxiv, figs. 3 & 4. 
1857. Livmargus muricatus Hoeven. (61a) Op. cit. p. 11, pi. iv, figs. 
1-10, 12, 14, 15. 



ORTHAGORISCICOLA MURICATA. 101 

1861. Lpemargus muricatus P. J. van Beneden. (15) p. 129, pi. xix, 
figs. 1-4. 

1892. LEemargus muricatus A. Scott. Trans. Nat. Hist. Soc. Glasgow, 
vol. iii, pt. 3, p. 266. 

1900. Lsemargus muricatus T. Scott. (112) p. 158, pi. vi, figs. 39-42. 

1902. Orthagoriscicola muricata Poche. (97) p. 14. 

1907. Orthagoriscicola muricata 0. B. Wilson. (148) p. 473, pis. xl & xli. 

1909. Orthagoriscicola muricata E. Y. Elwes. (45) p. 20. 

1910. Orthagoriscicola muricata T. R. R. Stebbing. (125) p. 559. 

Female. — Carapace trapezoidal in outline, much 
wider behind than in front, postero-lateral corners 
broadly rounded, lateral margins minutely serrated, 
posterior margin slightly incurved, a number of 
minute spines scattered over the dorsal surface of the 
carapace, and the frontal plates apparently fused with 
it. First and second thoracic segments short and about 
half as wide as the carapace, attenuated at the sides 
and without accessory dorsal plates. The next seg- 
ment provided with a broad dorsal plate which overlaps 
a considerable portion of the genital segment and is 
divided by a median cleft into two broadly-rounded 
lobes the margins of which are minutely serrated. The 
genital segment furnished with two large broadly- 
expanded plates the inner margins of which somewhat 
overlap each other ; the plates suborbicular in outline 
and together fully one and a half times the width of 
the carapace, also entirely concealing the abdomen and 
caudal rami ; their posterior margins broadly rounded 
and conspicuously serrated. 

Antennules of moderate length and composed of 
three joints, the first as long as the other two com- 
bined. Antennge short, armed with stout and strongly- 
hooked terminal claws. Mandibles long, stylet-shaped, 
and minutely serrate at the distal end. First maxilli- 
peds small, provided with short but moderately broad 
terminal claws serrated on the margins. Second 
maxillipeds strong ; end joint with one or two marginal 
processes on its inner aspect and provided with a 
strong curved terminal claw considerably shorter than 
the joint to which it is articulated. The thoracic 
legs all biramose, the rami of the first and second pairs 



102 BEITISH PARASITIC COPEPODA. 

two-jointed, but those of the third and fourth pairs 
uniarticulate, tolerably large, and lamelliform. Length 
about 20 mm. 

Male, — The male does not differ greatly from the 
female, but it is only about half or three-quarters the 
size, and the dorsal plates of the thorax and genital 
segment are proportionately smaller ; the dorsal plates 
of the genital segment have also their inner margins 
apparently coalescent except at the posterior end, 
where they are separated by a moderately deep sinus; 
they scarcely extend so far back as in the female, so 
that the caudal rami are exposed. 

The antennae and the second maxillipeds are also 
stronger and form more powerful grasping organs than 
those of the female ; the rami of the third pair of 
thoracic legs are not foliaceous, but, like the first and 
second, are biarticulate ; they differ however in the 
inner ramus being small and somewhat rudimentary. 
The fourth pair are similar to the fourth pair in the 
female. Length from 10 to 15 mm. 

In both sexes the colour is dull yellow. 

Habitat. — Parasitic on the short sun-fish, Ortha- 
goriscus mola. " Specimens taken off Plymouth were 
given to A. M. Norman a great many years ago by the 
late Mr. C. Spence Bate."* Moray Firth {T. Edward), 
Firth of Forth, October 1890 {A. Scott), Dr. Baird 
records this parasite on the authority of W, Yarrell, 
but does not mention the locality. On a short sun- fish 
captured off Berry Head, 1898 {Major E. V. Elwes, 
Babbacombe). 

Distvihution. — European Seas. South Africa 
{T, B. B. Stehhing). New Zealand {G, M. Thoivson), 
The Atlantic and Pacific coasts of North America 
(C, B. WUso7i). 

* 'The Cmstacea of Devon and Cornwall,' 1906, p. 211. 



PHILOKTHEAGORISCUS. 103 

Genus 18. PHILORTHRAGORISCUS Horst, 1897. 

Syn. Binemahira Kroyer, in part. 

Female. — Carapace suborbicular, width somewhat 
exceeding the length. First thoracic segment fused 
with the head; second and third united together, 
forming one segment, provided with small lateral ex- 
pansions. Fourth segment furnished with a large 
dorsal plate, overlapping fully the half of the genital 
segment, about as wide as the carapace and divided 
into two suborbicular lobes by a median fissure which 
extends nearly to the base of the plate. The genital 
segment also furnished with a bilobed dorsal plate, 
nearly as wide as that of the fourth segment, and, 
viewed from above, almost concealing the com- 
paratively small abdomen. Caudal rami short and 
tolerably broad. 

Antennules two- jointed and tolerably elongated. 
Antennse stout and armed with large hook-like terminal 
claws. Mandibles long and slender. First maxilli- 
peds composed of two joints and provided with three 
terminal spines, two of them being of moderate length 
and claw-like. Second maxillipeds stout and bearing 
strong terminal claws. First four pairs of swimming- 
legs biramose ; both rami of the first three pairs bi- 
articulate. Caudal rami short. 

Male. — The carapace of the male is much larger 
than the rest of the body; width greater than the 
length, dorsal surface grooved as in Pandarus. Second 
and third thoracic segments coalescent, and furnished 
with a pair of small lateral plates. Fourth segment 
with very small dorsal plates, which are coalescent 
along the middle line, while the posterior margin with 
its intermediate sinus is similar to that of the genital 
segment in the female of Peris.^ojjus. Abdomen as in 
the female but more exposed behind the genital seg- 
ment. Caudal rami rather smaller than in the female. 

Antennules proportionately longer; the anteunae 
also enlarged, their terminal claws projecting well in 



104 BRITISH PARASITIC COPEPODA. 

front of the carapace. Other appendages somewhat 
similar to those of the female. 

We are indebted to Major E. V. Elwes, Babbacombe, for the 
privilege of examining and describing the male, lie having 
kindly sent it to us for that purpose along with specimens 
of the female. 

Philorthragoriscus serratus (Kroyer). 
(Plate XXIV, figs. 3, 4; Plate XXVII, figs. 11-24.) 

1863, Binematura serrata Kroyer, (71) p, 176, pi, viii, fig. 4 a-i. 
1897, Philorthragoriscus serratus 'H.ovB.t. (63a) Notes Ley den Museum, 
vol, 19 N, *, Note xiv, p, 137. pi, vii, 

1901, Dinematura serrata T, Scott, (113) p. 125. 

1906. Philorthragoriscus serratus A. Brian, (21) p, 53. 

1907. Philorthragoriscus serratus C, B. Wilson. (147) p. 479, pis. 
xlii & xliii, 

1909. Philorthragoriscus serratus E, V. Elwes. (45) p. 20. 

Female. — Carapace, which is generally described in 
the generic definition, with the lateral margin denticu- 
late. The antero -lateral corners of the dorsal plates of 
the fourth segment produced outward into sharp teeth, 
and both lobes with their posterior margins also denti- 
culate. The outer ramus in the first four pairs of 
thoracic legs considerably larger than the inner, the 
first joint being longer than the entire inner ramus, and 
having a small tooth on its outer distal angle ; the end 
joint small and bearing a few small spines and setae on 
its rounded apex ; the joints of the inner ramus sub- 
equal, and the end joint somewhat expanded and 
furnished with three apical setse. Basiopodites of 
the second and third pairs dilated, rami small. The 
basiopodite of the fourth pair also considerably en- 
larged, and both rami small, uniarticulate, and some- 
what rudimentary, but the inner much smaller than the 
outer ramus. Egg-strings long and slender. Colour 
yellowish. Length about 7 mm. 

Male. — See under generic definition. Length 5 mm. 

Habitant. — Taken on a short sun-fish, Orthagorisciis 
mola (L.), at Banff in 1862 by the late Thomas 
Edward. On a short sun-fish captured off Berry 
Head, 1898 {E. V. Elwes). 



PHILORTHRAGORISCUS SEKRATIJS. 105 

Bistrihiition. — European seas. Atlantic coast of 
North America (C B. Wilson). 

The species will he found recorded under the Rev. A. M. 
Norman's MS. name Monima fimhriata in the appendix to 
Smiles' ' Life of 'Jliomas Edward ' (p. 437, 1876), as one of 
the many creatures that keen-sighted naturalist added to the 
fauna of Scotland. The same species was also found on a 
short sun-fish captured by Mr. C. Beadle, off Berry Head, in 
August 1908, and presented by him to the Torquay Natural 
History Society. On this sun-fish Major E. Y. Elwes obtained, 
among other interesting parasites, four female and one male 
Philorthragoriscus serratus which he very kindly permitted us 
to examine. 

Family iii. DiCHELESTiiDiE. 

Body generally elongated, head moderately small. 
Free thoracic segments usually simple, but sometimes 
provided with dorsal plates. Abdomen usually small 
or rudimentary. 

Antennules slender, moderately elongated, and com- 
posed of several joints, rarely short with two or three 
joints. Antennae armed with terminal claws and 
generally projecting more or less beyond the edge of 
the cephalothorax. Mouth parts somewhat similar to 
those of the Caligidge. Thoracic legs usually four 
pairs, frequently short, stump-like or suppressed, or 
with the posterior limbs transformed into lamelliform 
plates. Eye single, median, or absent. Genital organs 
as in the Caligidse. Sexual differences not very 
marked, but the male usually smaller than the female, 
and both for the most part capable of a certain amount 
of locomotion. 

Genus 19. DICHELESTIUM Hermann, 1804. 

Head obtuse, body elongated, consisting of four 
distinctly- articulated segments without dorsal plates. 
Genital segment tolerably elongated. Abdomen small. 
Caudal rami not very prominent. 

Antennules slender and composed of eight joints. 



106 BRITISH PARASITIC COPEPODA. 

Antennae large, as long as the antennnles, and cliele- 
form or uncinate at the end, usually projecting 
forward beyond the front of the head. The first and 
second pairs of thoracic legs small and biramose, but 
the fourth pair each composed of a single one-jointed 
lamelliform plate. 

The male is somewhat similar to the female, but 
smaller. 

1. Dichelestium oblongum ( Abildgaard) . 
(Plate XXXI, figs. 7-18; Plate XLV, figs. 4, 5.) 

1794, Caligus ohlongus Abildgaard. (1) vol. iii, p. 52, pi. v, figs. 4-11. 
1804. Dichelestium sturionis Hermann. (61) p. 125, pi. v, figs. 7 & 8. 
1838. Diclielestiwm sturionis Kroyer. (70) vol. i, p. 299, pi. ii, fig. 5, 5a. 
1836. Dichelestium stumonis Rathke. (99a) Act. Akad. Leopold 
Carol, vol. xix, p. 127, pi. xvii, figs. 1-17. 

1840. Dichelestium sturionis M. Edw. (43) p. 485, pi. xxxix, fig. 4. 

1905. Dichelestium sturionis T. Scott. (116) p. Ill, pi. v, figs. 17-24; 
pi. vi, figs. 1-6. 

1906. Dichelestium oblongum Nonnan & T. Scott. (88) p. 215. 

Female. — Body elongated, narrow ; carapace nearly 
as broad as long and somewhat rhomboid in outline, 
widest behind the middle, and with the sides bluntly 
angulated ; it is also obscurely lobed in front and 
narrowed and truncated behind. Thoracic segments 
four, the first and second short and subequal, but 
the first produced laterally into short rounded lobes 
directed backwards, the second having the sides also 
lobate but produced slightly outwards. The third 
segment, which is rather longer than the preceding 
one, widest behind, and with a slight constriction in 
the middle. The fourth segment similarly constricted, 
and somewhat longer than the third. The genital 
segment about one and a half times the length of the 
preceding segment and tapering slightly towards the 
posterior end. Abdomen short. Caudal rami small ; 
egg-strings long and narrow\ 

Antennules moderately short and slender and com- 
posed of eight subequal joints. Antennae stout, and 
extending prominentl}^ in front of the carapace ; indis- 
tinctly three- or four-jointed, the end joint slightly 



DICHELESTItJM OBLONGUM. 107 

reflexed and furnished with a small apical claw which 
can be folded inwards so as to impinge against a 
rounded knob and thus form a tolerably powerful 
grasping-organ. Mandibles somewhat like those 
of C aligns except that the basal part is rather 
stouter. Maxilla small, biramose, primary branch 
stout, tapering towards the distal end, and bearing 
two slender apical setge ; secondary branch very small. 
The first maxillipeds with the first and second joints 
of nearly equal length, but the first is more robust ; end 
joint very small and bearing a few short spines and 
setse. Second maxillipeds short, robust, and furnished 
with stout terminal claws. Thoracic legs short and 
stout, and the first and second pairs biramose. Both 
rami of the first pair indistinctly two-jointed, the 
proximal joint of the outer ramus having a small spine 
on its outer distal angle wdiile the end joint has five 
spines : all moderately stout, on its rounded apex ; 
the inner ramus, which is shorter than the outer, 
carrying two terminal spines. The second pair 
similar to the first but the outer branch rather stouter 
and the spines shorter ; the inner branch also propor- 
tionally rather shorter. The fourth pair one-branched 
and each branch consisting of a single uniarticulate, 
lamelliform plate, rather longer than broad, with a few 
minute prickles round the distal end. Length 17 to 
18 mm., but varying somewhat. 

Male. — The male bears a close resemblance to the 
female, but is considerably smaller, and the genital 
segment is proportionally shorter. In the second 
pair of thoracic legs the inner branch is very short 
and broad, and bears on its outer aspect a small flat- 
tened plate. The fourth pair are also proportionally 
shorter and broader. Length about 13 mm. 

Habitat, — Parasitic on the gills of the sturgeon, 
AcijpenHer stiirio. On a sturgeon captured at Polperro, 
Cornwall, in 1867 {A. M. Norman). On a sturgeon 
captured about sixteen miles S.E. by E. of Aberdeen 
in December 1904 {Dr. AlexBoivman). On a sturgeon 



108 BRITISH PARASITIC COPEPODA. 

captured near Barrow-in-Furness, Lancashire {A. 

Scott). 

The structure of the mouth-organs of Dichelestiiim, and 
especially of the mandibles, shows a somewhat close relation- 
ship with the Caligid86. The parasite seems to be peculiar 
to the sturgeon, and to have a distribution coextensive with 
that of the fish. 



Genus 20. ANTHOSOMA Leach, 1816. 

Cephalothorax of considerable size, in the form of a 
narrow ovoid buckler having attached to it near the 
mouth a pair of large foot-jaws armed with strong 
hooks. Genital segment furnished with elytraform 
appendages. Thoracic feet, three pairs, all folia- 
ceous. 

The two sexes are somewhat similar except that, 
in the female, the genital segment and abdomen are 
almost entirely concealed by the elytraform append- 
ages, but are more or less exposed in the male. 

Anthosoma crassum (Abildgaard) . 
(Plate XXIII, figs. 5, 6.) 

1794. Caligus crassus Abildg. (1) vol. iii, p. 54, pi. v, figs. 1-3. 
1816. Caligus imhricatus Risso. Hist. nat. Crust, des Environs de 
Nice, p. 162, 'pi- iii. fig- 13. 

1816. Anthosoma smithii Leach. (74) p. 406, pi. xx, fig. 1. 

1838. Anthosoma smithii Kroyer. (70) vol. i, p. 295, pi. ii, figs. 2 &2a. 

1850. Arithosoma smithii Baird. (4) p. 296, pi. xxxiii. fig. 9. 

1861. Anthosoma crassum Stp. & Ltk. {(^). (127) p. 397, pi, xxii, fig. 24. 

1905. Anthosoma crassum T. Scott. (116) p. 112, pi. v, figs. 15 & 16. 

1906. Anthosoma crassum Norman & T, Scott. (88) p. 214. 

Fem^ale. — Tolerably elongated, and, when seen from 
above, ovate in general outline but narrowed in front. 
The head and a considerable portion of the thorax 
covered by a brownish-coloured horny shield Avhich 
gradually expands towards the posterior end, and the 
junction of the thorax with the head marked by an 
obscure constriction. The remaining portion of the 
thorax, and also the abdomen and caudal rami, covered 



ANTHOSOMA CRASSUM. 109 

entirely by two large foliaceous elytraform circular 
plates, the inner margins of which partly overlap each 
other on the dorsal aspect, these plates being orna- 
mented with numerous minute scattered punctures or 
depressions. 

Antennules moderately short, and composed of 
six sparingly-setiferous joints; the antennae stout, 
three-jointed, longer than the antennules, and armed 
with strong terminal hook-like claws. First maxilli- 
peds slender and feebly armed, apparently consisting 
of three joints, the end one being small and of a 
peculiar shape ; second maxillipeds short, very stout, 
and furnished with strong terminal claws. Thoracic 
legs in the form of thin and broadly-foliaceous plates 
each with a distinct notch on the inner margin, con- 
cealing the genital segment as well as part of the 
abdomen. Caudal rami narrow and moderately elon- 
gated. Length about 15 mm. Egg-strings long and 
slender, reaching in length from 40 to about 50 mm. 

The shield is of a chitinous texture, the colour on 
the sides is yellowish but merging into blackish-brown 
along the middle and towards the front ; the elytra- 
form plates and thoracic feet, which also appear to be 
slightly chitinous, are whitish with a slight tinge of 
yelloAV. 

Male. — The male has a close resemblance to the 
female, except that the large dorsal plates which cover 
the posterior portion of the female are wanting. 

This interesting species was found on what was supposed 
to be a porbeagle shark {Lamna comubica) captured off the 
coast of Scotland in October 1904, by one of the trawling 
steamers which make only short runs from Aberdeen and 
which are locally known as " short trippers,^^ Two specimens 
of the Anthosoma were obtained — the one described here and 
a smaller one, probably a male. We are indebted to the 
kindness of Mr. Irvine, Aberdeen, for the specimen here de- 
scribed. Dr. Baird states {op. cit., p. 299) that a specimen 
was " discovered sticking to a shark (the Lamna comubica) 
thrown ashore at Exmouth, Devonshire, by T. Smith, Esq., 
of the Temple, who sent it to Dr. Leach, British Museum." 



110 BRITISH PARASITIC COPEPODA. 

Anthosoma crassum lias apparently a wide distribution in 
both the north and south hemisphei^es. Mr. G. M. Thomson 
of New Zealand, after describing the species, states that 
^^ numerous specimens were taken from the upper jaw of a 
porbeagle shark, Lamnn- coroiuhica (Otago Museum). Also a 
number from the same kind of shark taken at Napier by A. 
Hamilton '' ; and he adds that " according to Gould this 
species has been taken on the mackerel-shark, Lamna imnc- 
tata, on the coast of Massachusetts, U.S.A."* 

Genus 21. LERNANTHROPUS Blainville, 1823. 

Body broadly ovate, depressed. Cephalon oblong 
or pyriform, with the sides incurved. Neck distinct; 
the posterior part in the female covered by a large 
dorsal shield sometimes divided into two portions by 
a transverse constriction, and prolonged backwards so 
as to conceal more or less completely both the genital 
segment and the abdomen; in the male the dorsal 
shield not being so large as to conceal the abdomen. 

Antennules slender, composed of five or more — 
usuallv seven — articulations, and sometimes furnished 
with supplementary appendages. Antennae robust 
and armed with strong terminal hooks. Other mouth- 
organs somewhat similar to those of the Caligidse. 
Thoracic legs biramose ; the first and second pairs 
very small and rudimentary; the third and fourth 
tolerably large ; rami lamelliform, those of the last 
pair usually, and sometimes those also of the others, 
forming elongated appendages with their distal ends 
more or less attenuated. 

Lernanthropus kroyeri P. J. van Beneden. 
(Plate XXIX, figs. 2-5; Plate XXX, figs. 9-17.) 

1851. Lernanthropus hroyeri P. J. van Ben. (11) p. 102, pi. iii, figs. 7-9. 
1858. Lernanthropus hroyeri Glaus. (30) p. 18, pi. ii, figs. 15-19. 
1864. Lernanthropus hroyeri Nordmann. (90) p. 508, pi. vii, figs. 5-8. 
1879. Lernanthropus hroyeri Heider. (56) vol. ii, p. 90, pi. v, figs. 
72-73. 

1896. Lernanthropus hroyeri Bassett-Smitli. (7) p. 159. 
1904. Lernanthropus hroyeri A. Scott. (109) p. 43. 

* 'Trans. N. Z. Institute/ vol xxii (1889), p. 366. 



LERNANTHROPUS KROYERI. Ill 

Female. — The body, which is oblong in shape, some- 
what flattened and rather wider towards the posterior 
end, and divided into three unequal, but moderately 
distinct, portions; the head, which is the shortest, 
equal to about one-fourth of the entire length of the 
animal, exclusive of the posterior appendages, widest 
behind and tapering slightly on each side forwards to 
the flatly-rounded forehead ; the next two segments 
larger and subquadriform, but the last segment about 
a third longer than the middle one. The genital 
segment and abdomen, which are small, entirely con- 
cealed in dorsal view ; caudal rami moderately stout 
and elongated. 

Antennules moderately short and composed of seven 
joints, basal joint robust, the others small ; a slender 
two- or three-jointed appendage springing from near, 
but a little behind, the base of each of the antennules, 
and reaching to fully beyond their apex. Antennae 
short, robust, and furnished with short but strong 
terminal claws; mandibles and maxillge slender; 
maxillipeds short, stout, and strongly uncinate. 
Thoracic legs biramose ; first and second pairs very 
small and rudimentary, the inner ramus uniarticulate 
and rather more robust than the outer, Avhich consists 
of one or two small joints. 

Other appendages occur on the ventral aspect which 
are referred to by Steenstrup and Llitken and others 
as the third and fourth pairs of legs. The third pair 
consist each of a one-jointed linguliform appendage 
which reaches to near the base of the next pair ; the 
fourth pair have each two branches; they form elon- 
gated one-jointed appendages which reach backwards 
considerably beyond the posterior end of the body; 
they are nearly of equal length, and taper towards the 
blunt-pointed extremity. 

Colour dark red. Length about 21 mm. 

Male. — The male, which is only about half the size 
of the female, has a general resemblance to it, but the 
head is proportionally larger, the dorsal shield is 



112 BRITISH PARASITIC COPEPODA. 

smaller, and the urosome and caudal rami are not 
covered. The third and fourth pairs of ventral 
appendages are also more prominent. 

Habitat. — Parasitic on the gills of the bass or 
sea-perch, Labrax lujms. Plymouth {Bassett- Smith). 
Barrow Channel, Lancashire {A. Scott), 

Genus 22. HATSCHEKIA Poche, 1902. 

Syn. Clavella Kroyer (1838) and others but not Oken, 1815. 

Body elongated and narrow. Cephalon distinct, 
small, usually rounded. Thorax short, obscurely 
biarticulated, without dorsal plates. Grenital segment 
much elongated, narrow, and subcylindrical, five or 
six times the length of the cephalothorax. Abdo- 
men and caudal rami very short or obsolete. 

Antennules small and composed usually of not more 
than six articulations. Antennae short and stout, 
with strong terminal hooks, sometimes and perhaps 
generally with a slender spiniform or other appendage 
at the base of each antenna. Mandibles and maxillae 
small. First maxillipeds apparently obsolete. Second 
maxillipeds slender and uncinate. Thoracic legs two 
pairs, very short, and two-branched. 

Professor van Beneden remarks that this jj-enus is perfectly 
characterized and easy to distinguish by the length of the 
body ; by the three pairs of appendages that follow the 
antennas ; and by the two pairs of short biramose feet. 

1. Hatschekia hippoglossi (Kroyer). 
(Plate XXXIII, figs. 3, 4; Plate XXXIV, figs. 8-11.) 

1838. Clavella hippoglossi Kroyer. (70) p. 196, pi. ii, fig. 3. 

1829-1843. Clavella hippoglossi Guerin, (55) Icon, du Regne Anim., 
Crnstaces, pi. x, fig. 7. 

1840. Clavella hippoglossi M. Edwards. (43) vol. iii, p. 494. 

1851. Clavella hippoglossi P. J. van Beneden. (11) p. 100, pi. iii, 
figs. 5 & 6. 

1900. Clavella hippoglossi T. Scott. (112) p. 159, pi. vii, figs. 1-6. 

Female. — Body slender, elongated, and subcylin- 
drical. Cephalon small, of an oval form, width greater 



HATSCHEKIA HIPPOGLOSSI. 113 

tlian the length, sides rounded. Between the cephalon 
and the genital segment is a short neck, narrow and 
indistinctly segmented, and provided with rounded 
lobes on each side. Genital segment considerably 
elongated and rather narrow, its width scarcely equal 
to one-fourth of the length, and with the postero- 
lateral corners produced into rounded lobes about 
equal in size to the small abdomen that is intermediate 
between them and which is also rounded ; the pos- 
terior end has thus a trilobed appearance. Caudal 
rami extremely small. 

Antennules short, rather stout, and composed of 
five small joints which are sparingly setiferous. 
Antennge tolerably large and armed with strong 
terminal hooks and with a small appendage at the 
base. Mandibles small, elongated, tapering distally, 
and provided with a few small teeth at the distal 
end of the inner margin. Maxillae very small but 
comparatively stout, and furnished with two or three 
tooth-like processes. Maxillipeds elongated and 
slender. Thoracic legs two pairs, short and biramose, 
both rami appearing to be composed of two somewhat 
indistinct joints. Length about 9 mm. ; length of head 
and neck combined 1*5 mm. Egg-strings long and 
slender. Colour resembling that of the gills to which 
the parasite was adhering. 

Male. — The male is much smaller than the female 
but has a general resemblance to it. The cephalon is 
proportionally rather larger and the thoracic ring is 
distinctly segmented. The genital segment is also 
proportionally much shorter than in the female, being- 
only about twice as long as broad. 

Antennules short, composed of five joints. Antennae 
provided with large and strong hooks, projecting well 
in front of the cephalon. Caudal rami narrow and 
more prominent than in the female. Length 1*5 mm. 

Habitat — Parasitic on the gills of halibut, Hippo- 
glossus vulgaris ; not uncommon. The reddish 
coloured, thread-like egg-strings may sometimes be 

VOL. J. 8 



114 BRITISH PARASITIC COPEFODA. 

observed projecting beyond the edges of the gill- 
filaments. Frequently observed on halibut brought 
to the Fish-Market at Aberdeen. 

The males are apparently very rare. Though many speci- 
mens of liahbut have been examined, only one male specimen 
of this species has been met with. 

2. Hatschekia muUi (P. J. van Beneden). 
(Plate XXXII, figs. 1-11.) 

1851. Clavella mulli P. J. van Beneden. (11) p. 99, pi. iii, figs. 3 & 4. 
1896. Clavella mulli Bassett-Smitli. (7) p. 159. 
1906. Clavella mulli Norman & T. Scott. (88) p. 215. 

Female, — Body narrow, elongated, and sub-cylin- 
drical ; head small, subglobose, but the width rather 
greater than the length ; front somewhat lobate and 
without any horn-like processes. Neck very short, 
the thorax, which is indistinctly segmented, merging 
almost imperceptibly into the genital segment ; on the 
dorsal aspect of the thorax, and on each side of the 
median line, a small rounded protuberance, seen best 
in lateral view (PL XXXII, fig. 2). Genital segment 
elongated and narrow, slightly expanded in the middle 
and tapering gradually towards both ends, greatest 
width equal to about one-fourth of the entire length 
of the animal; posterior end trilobed, lateral lobes 
small and distinct, and produced slightly outw^ards, 
the middle one broad and not very prominent. 

The cephalothoracic appendages comprise the 
antennules, which are moderately small and composed 
of three spinulose joints, the last one wath also a few 
apical spines ; the antennge, composed each of a single 
large and moderately long basal part, and furnished 
wdth a stout terminal hook-like claw ; there is also at 
the base of each antenna a small secondary appen- 
dage — probably a sense-organ (PI. XXXIII, fig. 5). 
Mandibles small w^itli the extremity slightly curved 
and hook-like. Maxilla? small, with both the internal 
and external rami considerably attenuated. Second 
maxillipeds moderately slender and somewhat similar 



HATSCHEKIA MULLI. 115 

to those of HatscheJcia hippoglossi. Thoracic legs two 
pairs, each composed of a stout two-jointed basal part, 
which is furnished with two short two-jointed rami. 
Egg-strings long and slender. Length of adult female 
exclusive of egg-strings 3*4 mm., and including egg- 
strings about 10 mm. 

No males have been observed. 

Habitat. — Parasitic on the gills of red mullet, Mullus 
harbatus. Recorded from Plymouth by Dr. Bassett- 
Smith. Observed also on the gills of four specimens 
of red mullet sent to us from Looe, Cornwall, by 
Mr. 'N. M. Richards of that town, and on one from 
Plymouth sent by Dr. E. J. Allen, Director of the 
Plymouth Laboratory. It was also on Mullus barbatus 
that Professor van Beneden obtained his specimens of 
HatscheJcia mulli. 

A feature which appears to be peculiar to this species, and 
which was also noticed by van Beneden, is the secondary 
process at the base of each antenna : this process differs from 
that of any of the other species known to us, 

H. mulli is readily distinguished from other members of 
the genus recorded here by the dorsal humps on the thoracic 
region, and the small lateral lobes at the posterior end. A 
young* female is represented in dorsal view at fig. 3, and it 
also shows traces ot the characters by which the species is 
distinguished. 

3. Hatschekia labracis (P. J. van Beneden). 
(Plate XXXIII, fig. 2 ; Plate XXXIV, figs. 6, 7.) 

1870. Clavella labracis P. J. van Beneden. (16) pp. 45 & 46, pi. i, fig. 4. 

1901. Clavella labracis T. Scott. (113) p. 127. 

1902. Clavella labracis T. Scott. (114) p. 292, pi. xiii, figs. 10-12. 
1904. Clavella labracis A. Scott. (109) p. 40. 

Female. — Cephalothoracic segment, seen from the 
dorsal aspect, rhomboid or diamond-shaped, the lateral 
margins produced so as to form distinct angular pro- 
jections ; the width of the segment across the lateral 
angles rather greater than the length and equal to 
about a fifth of the entire length of the animal. Genital 
segment elongate-ovate, about three times as long as 
the cephalothorax, widest in the middle and with the 



116 BRITISH PAEASITIC COPEPODA. 

lateral margins slightly convex. Abdomen and caudal 
rami very small. 

Antennules short, moderately stout, and consisting 
of about five joints, the first large and equal to nearly 
all the other joints combined, and its lower distal angle 
produced downwards in the form of a strong hook ; 
the three end joints subequal and shorter than the 
preceding one ; several short, dagger-like spines 
springing from the upper margin of the various 
joints. Antennae about as long as the antennules, 
two-jointed and armed with stout terminal claws; a 
small process in the form of a knob occurs at the base 
of each antenna. Mandibles and maxillae small and 
simple in structure. Maxillipeds moderately slender, 
composed of two (or three) joints, and with the 
extremities uncinate. Thoracic legs two pairs, both 
biramose and somewhat similar in structure ; the basio- 
podite consisting of two moderately stout joints and 
both rami also two-jointed, the inner ramus in both 
pairs being rather more robust than the outer one; the 
proximal joint of the outer ramus the largest and 
bearing a small spine on its outer* distal angle, the 
joints of the inner ramus also unequal but the proximal 
one the smaller. Caudal rami short. Length about 
1 mm. Egg-strings fully as long as the animal, and 
with proportionately few but rather large ova. Colour 
similar to that of the gills of the host. 

The male of this species has not yet been observed. 

Habitat. — Parasitic on the gills of the striped wrasse, 
Lahrus mixtits Linn., and the ballan wrasse, Labrn.s 
bergylta Ascan. (Lahrits maculatus Bloch). Firth of 
Clyde {T. Scott). Irish Sea {A. Scott). Professor P. J. 
van Beneden states that HatscheJcia labracis is abundant 
on both of the species of fish mentioned above. 

This Hatschekia, though distinct enough, is very small and 
easily overlooked, but as the egg-strings are of a somewhat 
hghter colour than the gills of the fish, and tolerably 
elongated, they help to reveal the presence of the parasite. 



HATSCHEKIA CLUTH^. 117 

4. Hatschekia cluthae (T. Scott). 
(Plate XXXIII, fig. 1 ; Plate XXXIV, figs. 1-5.) 

1902. Clavella cluthse T. Scott. (114) p. 292, pi. xii, figs. 26-31. 

Female. — With a general resemblance to the female 
of H. labracis (van Beneden) recorded above, but 
much larger, and the lateral margins of the cephalic 
segment evenly rounded instead of being angular as 
in that species. The genital segment, the form of 
which is also somewhat different from that of H. 
labracis, becoming gradually wider posteriorly, so that 
the widest part near the hinder end is about twice as 
wide as it is immediately posterior to the cephalon ; the 
segment narrowing convexly and somewhat rapidly 
behind the widest part to the obscurely-angulated ex- 
tremity. Abdomen and caudal rami very small. 

Antennules apparently only four-jointed, moderately 
stout and furnished with a few minute spines, the basal 
joint about as long as the next three taken together, 
but the end joint very small. Antennae fully as long 
as the antennules, and composed of two joints, each 
provided with a short but strong terminal hook-like 
claw. Maxillae very small and simple in structure, 
consisting each of a minute papilliform basal joint, 
bearing threesmall spines. Maxillipeds small, two- or 
three-jointed, and furnished with small terminal claws. 
Both pairs of thoracic legs biramose and somewhat 
similar in structure ; the basiopodite tolerably stout 
and composed of two joints; the rami also two- 
jointed, and armed with small terminal spines; the 
first joint of the outer ramus larger than the second and 
carrying a small spine at its outer distal angle, while 
on the other hand the end joint of the inner ramus is 
tlie larger one ; the second basal joint also with a 
small spine on its inner distal angle. 

Caudal rami very short ; the egg-strings also toler- 
ably short and slender. Colour somewhat similar to 
that of the gills of the fish. Length about 1*5 mm. 
exclusive of the egg-strings. 



118 BRITISH PARASITIC COPEPODA. 

Habitat. — Parasitic on the gills of Jago's goldsinny, 
Ctenolabrns rupestris Linn., East Locli Tarbert (a 
branch of Loch Fyne) {T. Scott). 

The male has not yet been oloserved. 

The more obvious characters by which this species may be 
distinguished from H. lahracis appear to be its larger size, the 
different form of the cephalon and genital segment, and the 
structure and armature of the antennules. 

5. Hatschekia cornigera T. Scott. 
(Plate XXXV, figs. 1, 2; Plate XXXVI, figs. 1-5.) 

1909. Hatschekia cornigera T. Scott. (118) p. 74, pi. iii, figs. 1-7. 

Female. — Body narrow and greatly elongated. Head 
proportionally very small and expanding laterally into 
broadly -rounded lobes, also produced backward, on 
the median dorsal aspect, into a blunt-pointed spur- 
like process; this segment being distinctly limited, 
by a constriction, from the thorax, which is narrow 
where it joins the head. 

No distinct separation between the thorax and the 
genital segment, the one merging gradually into the 
other. Genital segment very long and narrow, and of 
about the same width throughout — the width being 
only equal to about one - sixth of the length ; the 
segment near its posterior end tapering to the narrow 
and obscurely bilobed extremity ; abdomen very small 
and not clearly defined. 

Antennules short, moderately stout, and composed 
of five articulations which are sparingly setiferous. 
Antennae large and armed with strong and hook -like 
terminal claws. Mandibles and maxillse somewhat 
similar to those of H. cluthse. Maxillipeds tolerably 
elongated and slender, with a short spine arising from 
the inner aspect and near the proximal end of the 
second joint, the end joint terminating in a pair of not 
very strong and slightly-curved spines. Both pairs 
of thoracic legs biramose ; the basiopodite consider- 
ably enlarged and composed of two articulations ; the 



HATSCHEKTA CORNIOERA. 119 

outer and inner rami also two-jointed and provided 
with a few apical spines. Caudal rami inconspicuous 
or obsolete. Length about 2*5 mm. Egg-strings 
scarcely as long as the genital segment. Colour 
similar to that of the gills of the fish. 

No males have been observed. 

Habitat. — Parasitic on the gills of sea bream, 
Pagellns centrodontns De la Roche, captured in the 
I^orth Sea and landed at the Fish-Market at Aberdeen 
during the earl}^ months of 1908 {T. Scott). 

This species is very slender in proportion to its length, and 
in this respect it has a resemblance to Kroyeria ; the head is 
also proportionally very small. When in situ on the gills of 
the fish, the parasite is not very conspicuous and may be 
easily overlooked. The horn-like process on the dorsal aspect 
of the cephalon, best seen in lateral view, is perhaps one of 
the more distinctive features of the species. 

6. Hatschekia pygmaea T. Scott. 
(Plate XXXV, figs. 3-7 ; Plate XXXVI, figs. 6-9.) 

1907. Clavella labracis (?) P. J. van Beneden, A. Scott. (Ill) p. 95,pl. iv. 

Female. — Antennules comparatively short, stout, 
and composed of five articulations, the first being 
robust and equal to about two-fifths of the entire 
length of the antennule, the next about half the size 
of the first, but the remaining three considerably 
smaller and subequal ; the first and second joints 
furnished with a number of short spatulate hairs on 
their upper aspect ; the first joint also with a short 
and stout spiniform seta on the lower distal angle, a 
minute seta springing from the lower margin of the 
third joint, while the end one has several similar setas 
on its lower margin and apex and two small hooks on 
the upper distal angle as shown in the drawing 
(PL XXXV, fig. 4). Antennae tolerably elongated 
and slender, and terminating in strongly-curved apical 
hooks. Mandibles and maxillae rudimentary, the 
former having apparently no serrated apex. Both 
pairs of thoracic legs biramose, small and more or less 



120 BRITISH PARASITIC COPEPODA. 

rudimentary ; in the first pair both rami two- jointed, 
the outer ramus of the second pair also consisting of 
two joints but the inner one three-jointed. Caudal 
rami very small and inconspicuous. Length about 
1 mm. Colour similar to that of the gills of the fish. 
Egg-strings tolerably short and containing compara- 
tively few but moderately large ova. 

No males have been observed. 

Habitat. — Found adhering to the gills of Crenilahrm 
melops (Linn.), the connor or goldsinny, captured in 
Luce Bay, Solway Firth {A. Scott). 

This species, which was described in 1907, was at first 
doubtfully assigned to Clavella (or Hatschekia) lahracis van 
Beneden; the examination, Iiowever, of additional specimens, 
obtained subsequently, showed that certain differences which 
had been recognized in the first specimens were fairly constant, 
and were moreover sufficiently important to make it necessary 
to regard it as distinct from others that have been described. 

In its general appearance this species has a resemblance to 
the Hatschehia lahracis of van Beneden. It differs from that 
species in the head being rounded at the sides — not angular 
as in that species — in the armature of the antennules, and in 
the terminal claws of the antennae being much less strongly 
hooked, as well as in one or two other anatomical details. At 
the base of each antenna there is a small knob-like process 
similar to that on the antenna of Hatschehia lahracis. This 
fish is not the same as Jago's goldsinny, but belongs to a 
different species of the Wrasse family. 

Genus 23. KROYERIA P. J. ran Beneden, 1853. 

Syn. Lonchidiam Gerstaecker, 1854. 

Cephalic segment moderately broad and plate-like, 
and furnished at the posterior end with movable styli- 
form processes, projecting backwards. Three distinct, 
free thoracic segments, without lobes or dorsal plates. 
Genital segment long and narrowly cylindrical. Ab- 
domen short, unsegmented in the female, and termi- 
nating in two lanceolate setose furca. Antennules 
composed of seven distinct joints. Antennae very 
stout and cheliform. Mandibles and maxillae rudimen- 



KROYERIA. 121 

tary and resembling those of Eudactylina. Second 
pair of maxillipeds large and furnished with powerful 
claws. Thoracic legs, four pairs, well developed and 
biramose; each ramus three-jointed, and furnished 
on the inner margin with long plumose setae. 

The male has a general resemblance to the female 
but is considerably smaller. The thoracic and ab- 
dominal segments are also narrower than in the female, 
and the abdomen is distinctly three- join ted. 

1. Kroyeria lineata P. J. van Beneden. 
(Plate LXX, j&gs. 1-13.) 

1853. Kroyeria lineata P. J. van Beneden. (13c) Bull. Acad. Roy. 
Belg.. vol. XX, p. 94. 

1861. Kroyeria lineata idem. (15) Rec. sur les Crust. Belg., p. 149, 
pi. xxii. 

1858. Kroyeria lineata Glaus. (30) Beitrag Parasit. Crust., p. 24, pi. ii. 

1880. Kroyeria lineata Yalle. (141a) Boll. Soc. Adriat. Sc. Nat. vol. vi, 
p. 65. 

1885. Kroyeria lineata Carus. (29a) Podrom. Faunae Meditm-ranese, 
p. 364. 

1899. Loncliidium lineatum Bassett-Smitb. (8) Proc. Zool. Soc. 
Lond. 1899, p. 473. 

1906. Kroyeria lineata A. Brian. (21) Copepodi Parassiti dei Pesci 
d'ltalia, p. 67, pi. v, fig. 3. 

Female.— lje\igt\\, exclusive of the f ureal setge, 
4-5 mm. Cephalic segment moderately broad and 
flattened and having the antero-lateral margins con- 
cave, giving rise to a broadly-rounded forehead. 
Posterior margin furnished on each side with a strong 
movable spine, the apex of which is somewhat clieli- 
form, and reaches to the end of the first free segment 
of the thorax. The eyes, two in number, situated in 
the front portion of the segment, a V-shaped band of 
chitin with a pointed end traversing the segment from 
front to rear (PI. LXX, fig. 1). The three thoracic seg- 
ments short and quadrangular in outline, the second 
being distinctly shorter than the other two. The third 
segment considerably longer than the first and nearly 
twice the length of the second, with its lateral margins 
slightly inflated posteriorly. All three segments pro- 
vided with dorsal and lateral bars of chitin. Genital 



122 BRITISH PARASITIC COPEPODA. 

segment very long and narrowly cylindrical. Its 
distal end, in dorsal view, distinctly truncated, and in 
lateral view broadly rounded and somewhat gibbous 
(PL LXX, fig. 10). Abdomen short and narrow, joining 
the ventral surface of the distal end of the genital 
segment ; uniarticulated, but in some specimens two 
slight constrictions appear which may correspond to 
the distinct segments seen in the male. Furcal joints 
moderately long and narrow, and about four times 
longer than broad. Egg-strings short and cylindrical, 
containing only a single row of eggs. 

Antennules short and seven-jointed; moderately 
setiferous. Basal joint nearly as long as the combined 
lengths of the next five, and its distal end crossed by 
three indistinct lines as shown in the drawing (fig. 2). 
Distal end of the last joint provided with a short, but 
stout, sensory filament. Antennge short and stout, 
and armed with powerful chelse which are used for 
securing the parasite to its host, and are easily visible 
in the entire animal, even witliout dissection and by 
using an ordinary hand magnifier. 

Mandibles stylet-shaped, with the distal end of the 
inner margin serrate as in Eudactylina. Maxillse 
bilobed, very small and somewhat rudimentary. The 
distal end of each lobe furnished with two setge. 

First maxillipeds small and cheliform, the claw -like 
end joint moderately broad and spoon-shaped; the 
distal end of the upper margin of the joint over which 
the claw folds slightly serrate and ciliated (fig. 6). 
Second maxillipeds large, and forming powerful pre- 
hensile organs, their terminal claw strongly curved 
and nearly as long as the joint from which it springs. 

The four pairs of thoracic legs all somewhat similar 
in structure. They are biramose, and the rami are 
three-jointed as shown by figs. 8 and 9 on PL LXX. 
Basal plate of the first pair produced at its distal 
angles into minute tubercles, but in that of the second, 
third, and fourth pairs the angles are produced into 
moderately long and stout spines which are quite pro- 



KROYERTA LTNEATA. 123 

minent in the entire animal. No trace of a fifth pair 
of feet. 

Male. — Length exclusive of the f ureal setae 2*7 mm. 
The cephalic segment is similar to that of the 
female in every respect except in the shape and length 
of the movable spines at its distal end. The spines 
are evenly curved to the apex, and do not reach the 
end of the first thoracic segment. The first two 
segments of the thorax are nearly of equal size. The 
third segment is almost twice as long as the second, 
and its proximal lateral margins are slightly expanded. 
The genital segment is narrow and cylindrical, and 
distinctly longer than the combined lengths of the 
three thoracic segments. The abdomen is composed of 
three clearly-defined segments. The first segment is 
about half the length of the genital segment. The 
second and third are of nearly equal length, and are 
each about equal to two-thirds of the length of the 
genital segment. The furcal joints are almost similar 
to those of the female. 

The various thoracic and other appendages are 
similar to those of the female, and therefore need not 
be described. A rudimentary fifth pair of feet repre- 
sented by three minute setse can be detected by care- 
fully examining the lateral margins of the genital 
segment. 

Habitat. — In the hollows between the gill-rays of 
male specimens of Galeorhinus galeus or tope, captured 
in the Irish Sea in the vicinity of King William Bank, 
off the north end of the Isle of Man, April 1912. We 
have not yet met with this parasite on the gills of 
female specimens of the tope, although a considerable 
number of them have been examined. Much care is 
required to detect the parasite, as it is usually entirely 
hidden in the hollows between the gill-rays. The 
presence of the parasites is occasionally revealed by 
the egg-strings projecting beyond the ends of the 
gills. Several specimens of both sexes may sometimes 
be found on the gills of a single fish. 



121^ BRITISH PAEASITIO COPEPODA. 

The genus tind species were both described by P. J. van 
Beneden in 1853. In the following year G-erstaecker 
described a closely allied parasite under the niMne Lonchiditom 
aculeatum. Bassett-Sniith in his work ^A Systematic De- 
scription of Parasitic Copepoda found on Fishes/ sets aside 
tlie generic name given by van Beneden and adopts Ger- 
staecker's Lonchidium. Brian restores van Beneden^s name 
in his work ' Copepodi Parassiti dei Pesci d^Italia/ and as we 
have been unable to find any satisfactory reason to support 
Bassett-Smith in changing the generic name, we have fol- 
lowed Brian in adhering to van Benedeu^s name Kroijeria. 
Spence Bate used the name Kroyera to distinguish a genus of 
Amphipoda in 1857, but Sars points out in his ' Crustacea of 
Norway/ vol. i, that the name had already been appropriated 
in zoology. Brian apparently regards Gerstaecker's Lon- 
chidium aculeatum to be nearly identical with Kroyeria 
lineafa. We kept a number of specimens of both sexes alive 
in sea-water for a time, but were unable to detect them making 
any attempt to swim. 

Gemis 24. CONGERICOLA P. J. van Beneden, 1854. 

Syn. Cycnus M. Edwards (1840), preoccupied by Hiibner for a genus 
of Lepidopteva. 

Head rounded, small ; free thoracic segments two, 
without lobes or dorsal plates ; genital segment elon- 
gated, narrow, and subcylindrical ; abdomen short ; 
caudal rami setiferous. Antennules composed of about 
six articulations ; antennse two-jointed and armed with 
terminal claws. Second maxillipeds slender. Thoracic 
legs four pairs, all biramose. 

The only British species known to us is that described 
below. 

1. Congericola pallida P. J. van Beneden. 
(Plate XLI, fig. 1.) 

1854. Congericola pallida P. J. van Beneden. (14) vol. xxi.pt. 2, p. 583. 
1861. Congericola 'pallida idem. (15) p. 148, pi. xxiii. 
1896. Ci/cwwspfl//icZ«s Bassett-Smith. (7) p. 159. 

1900. Cycnus pallidas T. Scott. (112) p. 160. 

1901. Cycnus palUdus A. Scott. (107) p. 350. 

1906. Congericola pallida Noi-man & T. Scott. (88) p. 215. 

Female. — Cephalic segment small and suborbicular. 



CONGERICOLA PALLIDA. 125 

Free thoracic segments two, very short, and without 
lobes or dorsal plates. G-enital segment elongated, 
narrow, and subcylindrical. Abdomen short ; caudal 
segments small and setiferous. 

Antennules short and composed of about six joints, 
but one or two of the articulations are not very 
distinct; antennae two-jointed, and provided with 
strong terminal hooked claws. Mouth-appendages 
somewhat similar to those in HatscheJcia. Thoracic 
legs four pairs, biramose, and somewhat rudimentary. 
Colour pale reddish, scarcely so deep-coloured as the 
gills of the fish. Length, exclusive of egg-strings, 
about 4 mm. ; egg-strings slender and elongated, about 
twice as long as the animal. 

This species, in its general appearance, has a some- 
what close resemblance to HaUclielda, 

The male has not been observed. 

Habitat. — Parasitic on the gills of conger eels. 
Conger vulgaris Cuv. {Conger niger, Risso). Plymouth 
(Bassett- Smith). Firth of Clyde and at Aberdeen {T. 
Scott). Lancashire coast {A. Scott). 

This parasite is not uncommon on the gills of large speci- 
mens of the conger, and sometimes occurs in considerable 
numbers. Thirty individuals have been taken from the gills 
of a single fish. The species has been recorded from the 
Adriatic (Dr. Graeffe) as well as from the coast of Belgium 
(P. J. van Beneden) . 



Genus 25. EUDACTYLINA P. /. van Beneden, 1853. 

Female. — Cephalothorax composed of five segments ; 
cephalic segment obtuse, wider behind than in front ; 
thoracic segments without dorsal plates or lateral 
processes. Abdomen, including the genital segment, 
considerably shorter than the cephalothorax and com- 
posed of three segments. Antennules short, stout, 
and usually five-jointed, basal joint usually provided 
with strong hooked spines ; antennee three-jointed and 
armed with stout terminal claws. Mandibles and 



126 BEITISH PARASITIC COPE POD A. 

maxillae small, somewhat similar in structure to those 
of HatscheJda. First maxillipeds small, composed of 
three joints, and provided with a small terminal claw ; 
second maxillipeds of moderate size, armed with 
terminal claws, strong and cheliform. First four 
pairs of thoracic legs short, biramose ; rami of the 
first pair two- or three-jointed ; those of the third and 
fourth pairs are usually both composed of three joints; 
but though in the second pair the inner ramus is some- 
what similar in structure to that of the next pair, the 
outer one is more or less modified. Fifth pair con- 
sisting each of a small uniarticulate lamina ; caudal 
rami short. 

Male. — The male someAvhat resembles the female, 
but differs in being smaller and in the antennules being- 
armed with one or two moderately strong terminal 
hooks. The second maxillipeds are not cheliform but 
resemble in structure those in HatscheMa ; the outer 
ramus of the second pair of thoracic legs is not modi- 
fied, and there is also a difference in the form of the 
caudal rami. Moreover the abdomen, including the 
genital segment, is in the male nearly as long as the 
cephalothorax. 

1. Eudactylina acuta P. J. van Beneden. 
(Plate XXXVII, fig. 1 ; Plate XXXVIII, figs. 1-5.) 

1853. Eudactylina acuta P. J. van Beneden. (13b) Bull. Acad. Roy. 
Belgique, vol. xx, pt. i, p. 235. 

1861. Eudactylina acuta idem. (15) p. 150, pi. xxv. 

1892. Eudactylina acuta Canu. (29) p. 52, pis. 2 & 3. 

1902. Eudactylina acuta T. Scott. (114) p. 293, pi. xii, figs. 20-25. 

1904. Eudactylina acuta A. Scott (109) p. 40. 

Female. — Body slender and elongated ; the cephalo- 
thorax consisting of five distinct segments, and the 
abdomen of three, including the genital segment. 
Cephalic segment about equal in length to the next 
two segments combined ; the first of these two seg- 
ments smaller than the second, the fourth and fifth 
subequal. Abdomen small and only a little more than 



EUDACTYLINA ACUTA. 127 

one-fourth the length of the cephalothorax ; caudal 
rami short and broadly ovate. Antennules short, 
stoutj and composed of about five joints, and armed 
with several strong spines; a large strongly-curved 
spine springing from the upper distal angle of the 
second joint and reaching to near the end of the next 
joint ; a stout but shorter spine on the lateral aspect 
of the same joint and also one or two spiniform setge ; 
a tolerably stout, elongated, and nearly straight spine 
springing from the upper distal end of the following 
joint, and immediately below, another, also moderately 
stout, but only half the length of the first ; the pen- 
ultimate joint also provided with a short spine, and a 
few spiniform setse at its distal end ; the terminal joint 
very small. Antennae small, but armed with stout 
terminal claw-like spines ; also one or two stout spines 
on the inner aspect of the first and second joints. 
Mandibles slender, elongated; first maxillipeds small, 
three-jointed, and furnished with a minute terminal 
claw. Second maxillipeds of moderate size, each 
armed with a tolerably powerful, terminal chela which 
forms an effective grasping-organ, having a somewhat 
close resemblance to similar appendages in Pseudotanais 
-^a genus belonging to Isopoda-Chelifera. 

The first four pairs of thoracic legs all biramose ; in 
the first, third, and fourth pairs both the inner and 
outer ramus distinctly three-jointed ; in the second 
pair the inner ramus composed of three subequal 
joints, but the outer somewhat abnormal in structure, 
the first joint comparatively large and fully as long as 
the entire inner ramus, while the distal extremity 
appears to consist of two minute coalescent joints; 
the basiopodite in all the four pairs composed of two 
unequal joints, the second being the smaller; the 
inner margin of the second basal joint in the first pair 
fringed with about five very short but stout spinules ; 
the first and second joints of the outer ramus each 
with a small dagger- shaped spine on the outer distal 
angle, and the third joint with two similar spines and 



128 BRITISH PARASITIC COPEPODA. 

a moderately elongated seta at tlie apex ; the inner 
ramus also provided with two spiniform apical setae, 
and both rami furnished with marginal spinules as 
shown in the drawings ; the armature of the rami of 
the third and fourth pairs somewhat similar to that 
described above, but in the second pair, while the 
armature of the inner ramus does not differ materially 
from that of the others, the outer ramus is devoid of 
either spines or setae. Fifth pair small, uniarticulate, 
lamelliform, and broadly ovate, situated on the lateral 
aspect and near the distal end of the last thoracic 
segment, each furnished with about three apical 
setae and their outer margins fringed with minute 
spines. Caudal rami short, nearly twice as long as 
broad, with two short stout spines at the apex, a 
small seta on the lower half of the outer margin, 
and a fringe of minute prickles on the lower inner 
margin. Length about 2*5 mm. Egg-strings toler- 
ably slender and elongated. Colour similar to that of 
the gills, but the egg-strings are not so dark red. 

No males have been observed. 

Habitat. — Parasitic on the gills of the angel-fish, 
Bhina squatina (Linn.). Moray Firth and Firth of 
Clyde {T, Scott). Irish Sea {A. Scott). Not rare. . 

2. Eudactylina acanthii A. Scott. 

(Plate XXXVII, figs. 2, 3 ; Plate XXXVIII, 
figs. 6-12.) 

1901. Eudactylina acanthii A. Scott. (108) p. 14. 

1902. Eudactylina acanthii T. Scott. (114) p. 296, pi. xiii, figs. 1-9. 
1904. Eudactylina acanthii A. Scott. (109) p. 41. 

Female. — Body moderately stout, cephalothorax 
equal to five times the length of the abdomen, in- 
cluding the genital segment ; the first cephalothoracic 
segment about one and a half times the length of the 
next, the second and third subequal, the fourth rather 
larger than the third or the following segment. Ab- 
domen very short, consisting of three segments, the 



EUDACTILINA ACANTHII. 129 

genital segment being fully as large as the other two 
combined. 

Antennules short, tolerably stout, and tapering 
towards the distal end ; first two joints large and 
together equal to more than half the entire length of the 
antennule ; the curved spine at the distal end of the 
second joint much smaller than the spine similar to it 
on the antennules of Eudactylina acuta, and the strong, 
elongate, nearly straight spine at the distal end of the 
third joint in that species wanting here. Antennae 
somewhat similar to those of Eudactylina acuta, but 
the stout spines on the first and second joints wanting. 
Second maxillipeds cheliform, other mouth-appendages 
similar to those of the species mentioned above. 

The inner ramus two-jointed in all the four pairs of 
thoracic legs ; the outer ramus of the first pair con- 
sisting apparently of only one joint, or of two subequal 
and coalescent joints, those of the next three pairs 
appearing to be composed of three joints, but the 
articulation between the second and third joints not 
so distinct as that between the first and second ; in the 
first pair the inner ramus is provided with a number 
of short but stout spines chiefly on the exterior aspect, 
and the outer fringed with minute setse ; a small spine 
also springing from a notch near the middle of the 
outer margin and two or three small setse from the 
distal end. The inner ramus in the second pair some- 
what similar to, but rather stouter than, the inner 
ramus of the first pair ; the outer ramus three-jointed, 
but the second and third joints appearing to be co- 
alescent ; both rami fringed on the exterior edge with 
small prickles, the outer margin of the first basal joint, 
which is somewhat expanded, forming a rounded 
gibbose projection, also fringed with similar prickles, 
which are scattered sparingly over portions of all the 
joints. The remaining two pairs somewhat similar in 
structure to the second but rather more robust and 
less spiniferous. The fifth pair resembling those of 
Eudactylina acuta, but differing slightly in their foi-m 

VOL. I. 9 



130 BRITISH PARASITIC COPEPODA. 

and armature. Caudal rami moderately stout, fully as 
long as the last abdominal segment ; two short spines 
springing from the outer margin of each ramus and 
three from the apex, the middle seta being the longest. 
Length about 2 mm. Colour similar to that of the 
gills of the fish. 

No males have been met with. 

Habitat. — Parasitic on the gills of the piked dog- 
fish or spur-dog, Squalus acanthias Linn.( = Acaiithias 
vulgaris Risso). Beaumaris Bay, Anglesey, Septem- 
ber 1901, and other parts of the Irish Sea {A, Scott), 

This Eudactylina is frequent on the gills of Squalus acan- 
thias captured in the Irish Sea, and it may probably also occur 
on Scottish specimens of the same fish. Eudactylina acuta 
has been recorded from both the angel-fish and the piked 
dog-fish by van Beneden and Dr. Cnnu, but though the 
Eudactylinse obtained by these authors from the two fishes 
mentioned may belong to the one species, those found para- 
sitic on the dog-fishes taken in Beaumaris Bay, &c., certainly 
differed from the specimens obtained on angel-fishes captured 
on the English and Scottish coasts and examined by us. 

3. Eudactylina similis T. Scott. 
(Plate XXXVII, figs. 4, 5; Plate XXXIX, figs. 1-17.) 

1902. Eudactylina similis T. Scott. (114) p. 295, pi. xii, figs. 1-19. 

Female. — First cephalothoracic segment about one 
and a half times as long as the next, but the length of 
that segment, and of the two that follow, nearly the 
same ; the last segment rather smaller than any of the 
others. Abdomen, including the genital segment, 
short, being only about one-third as long as the 
cephalothorax. 

Antennules stout, and somewhat similar in structure 
and armature to those of Eudactylina acuta, but the two 
principal spines have each a fringe of minute prickles 
along the upper edge, and the penultimate joint is 
proportionally shorter ; antennae tolerably stout, elon- 
gated, and composed of four articulations, the first 
and second joints each provided with a stout but 



EUDACTYLINA SIMILIS. 131 

moderately short spine on the inner aspect, while the 
end joint bears two curved spines which are stout and 
claw-like ; the second joint rather shorter than the 
first or third, and the terminal one very small. 
Mandibles slender and moderately elongated, with the 
distal half of the end joint serrated on the inner edge. 
Maxillae small, bilobed, principal lobe stout, rather 
longer than broad, and provided with apical setse of 
unequal length, one being nearly twice as long as 
the others; secondary lobe elongated, narrow, and 
bearing a moderately long apical seta. First maxilli- 
peds small, three- jointed, and furnished with a short 
but rather stout terminal claw. Second maxillipeds 
tolerably large and strongly chelate; the end joint 
considerably expanded and its postero-lateral corners 
more or less produced, one of them extending into a 
spoon-like process, while to the other is articulated 
a strongly-curved claw, the apex of which impinges 
against the spoon-like process of the opposite angle, 
thus forming an effective grasping organ. 

First pair of thoracic legs somewhat similar to those of 
Eudactylina acuta except in the following particulars : 
the second basal joint is furnished with two small but 
stout spines on the inner distal angle instead of a 
fringe of stout marginal spinules ; the first joint of 
the outer ramus is proportionally considerably larger 
than either the second or third joints, the inner 
ramus is more slender, and the articulations are more 
unequal. The second pair are similar in structure to 
the same pair in Eudactylina acuta , but the outer ramus 
is proportionally and distinctly larger. ' The third and 
fourth pairs are somewhat similar in the two species ; 
the fifth, however, are larger and proportionally broader 
than those of the species named, the length does not 
greatly exceed the width, and there is also a corre- 
sponding difference in their outline. Caudal rami 
about twice as long as the anal segment of the 
abdomen, and moderately wide but becoming narrower 
towards the apex, each bearing three small spiniform 



182 BEITISH PARASITIC COPEPODA. 

setae — one at the apex and two on the lower half of 
the outer margin. Length about 3 mm. 

Male. — The male is smaller than the female, the 
body is more slender, and the abdomen is about as long 
as the cephalothorax. Antennules tolerably stout and 
composed of about seven articulations ; the first and 
second joints are short but considerably dilated, the 
second is provided with a stout and strongly curved 
spine which springs from the upper distal angle ; the 
remaining joints are all nearly of equal width and 
much narrower than the basal joints, the third joint is 
about as long as broad, the fourth and fifth are very 
short, the sixth is rather longer than the fourth and 
fifth combined, while the end joint is nearly three 
times the length of the sixth ; the third and fifth 
joints are each furnished with a stout and moderately 
long straight spine, and a shorter spine also occurs on 
the distal end of the sixth joint, while the last joint 
is armed with a strong terminal hook and a slender 
sensory filament. Antennas and mouth-appendages 
like those of the female except that the second maxilH- 
peds are not chelate but provided with a simple 
terminal claw. The first four pairs of thoracic legs 
have both rami three-jointed, but the articulation 
between the second and third joints of the outer ramus 
of the first pair is not clearly defined ; the inner ramus 
of the second pair is provided with a tolerably 
elongated and slightly curved spine which springs 
from the inner distal angle of the first joint ; the outer 
ramus of the second pair is normal in structure. Both 
rami of the third pair are moderately slender and 
setiferous. Fifth pair somewhat similar to those 
of the female. Caudal rami narrow, each with two 
apical and one marginal spiniform setae. Length about 
2 mm. Colour similar to that of the gills of the fish. 
Egg-strings moderately slender and elongated. 

Habitat. — Parasitic on the gills of the starry ray, 
Baia radiata Don. The fishes on which this parasite 
was obtained were captured east of the Shetland 



EUDACTYLINA SIMILIS. 133 

Islands in May and off Aberdeen in November 1901 
(T. Scott). 

This species lias a general likeness to Eudactylina 
acuta van Beneden, but differs from it in several 
respects. 

4. Eudactylina minuta T. Scott. 
(Plate XXXVII, figs. 6, 7 ; Plate XL, figs. 1-9.) 

1904. Eudactylina minuta T. Scott. (115) p. 275, pi. xvii, figs. 1-11. 

Female. — Body slender ; the ceplialothorax about 
four times as long as the abdomen ; the first cephalo- 
thoracic segment scarcely equal in length to the next 
two segments combined ; the last two segments of the 
thorax rather more dilated than the others ; all the 
segments rough with minute scattered hairs, especi- 
ally along the dorsal aspect. 

Antennules short, stout, and apparently consist- 
ing of five joints ; the first three joints large and 
subequal, the fourth narrow and only about a third of 
the length of the preceding joint, end joint very small ; 
the second joint furnished with a stout and strongly- 
curved spine on the upper distal angle, two smaller 
spines springing from the distal half of the upper 
margin of the third joint ; no terminal claw, but only 
a few apical setse. Antenna moderately elongate, 
three- jointed, and having a general resemblance to 
those of Eadacti/lina acuta. ; the second joint, which is 
about equal to the third one, having the lower distal 
angle produced downwards in the form of a short 
stout spine, while the end joint terminates in a strong 
curved claw. Mandibles and maxillae somewhat 
similar to those of Eudactylina acuta. First max- 
illipeds moderately stout and armed with a stout 
and nearly straight terminal claw, while the end 
joint has a row of minute coarse denticles along its 
inner edge. Second maxillipeds large and strong and 
somewhat similar in structure to those of Eadactijlina 
similis, but the extremity of the claw which impinges 



134 BRITISH PARASITIC COPEPODA. 

against the lower spoon-like process forms a rounded 
apical expansion. Both rami in the first pair of 
thoracic legs distinctly two-jointed and moderately 
stout, the inner ramus, which is larger than the outer, 
sparingly fringed with minute setae and provided with 
tw^o apical spines of unequal length ; the outer and 
shorter ramus with a fringe of minute setae on the 
outer margin of the first joint, the end joint bearing 
several spines round its outer margin and apex, the 
inner spine of moderate length but the others small. 
The second pair with a general resemblance to those 
of both Eudadylina acuta and Eu. similis ; the inner 
ramus, which is distinctly three-jointed, considerably 
smaller than the outer one, and the first joint having 
a longitudinal row of small spines extending obliquely 
upwards from near the distal end of the outer margin, 
while the end joint bears two small apical spines of 
unequal length; the outer ramus stout, tolerably 
elongated and indistinctly three-jointed, end joint 
small, about as broad as long, and bearing a spiniform 
seta on its margin and two minute prickles on the 
rounded apex ; two short spines, each with a thickened 
base, springing from near the distal end of the outer 
margin of the first joint, this joint being elongated 
and slightly arcuate ; a pseudo-articulation may be 
noticed between the two short spines and extending 
partly across the joint. The third and fourth pairs 
nearly alike and resembling those of Endacfylina 
similis except that the inner rami have a number of 
scattered spinules on their outer aspect ; the outer 
rami each rounded at the extremity and carrying a 
single elongated terminal seta ; they are also provided 
with a few small spines on the outer margin of the 
second and third joints, while a fringe of minute prickles 
extends along the margin of the first joint. The fifth 
pair, which are broadly foliaceous, ornamented with 
several transverse rows of minute spines and also 
furnished with three apical setae. Caudal rami short, 
about as long as the last abdominal segment, and bear- 



EUDACTYLINA MINUTA. 135 

ing short, moderately stout apical spines, with two 
setas on the outer margin. Length a little over 1 mm. 
Egg-strings short, bearing a small number of mode- 
rately large ova. Colour reddish, resembling the colour 
of the gills of the fish. 

IN'o males observed. 

Habitat. — Parasitic on the gills of a sting ray, 
Trygon pastinaca Linn., captured in the Dornoch 
Firth in October 1903 (T. Scott). The fish was sent to 
Dr. Fulton, Scientific Superintendent, Fishery Board 
for Scotland, who kindly placed it at our disposal. 

5. Eudactylina insolens T. and A. Scott. 
(Plate LXXI, figs. 1-12.) 

Female. — Length exclusive of the f ureal setae 1*8 
mm. Body moderately robust and cylindrical, and 
having no appearance of segmentation between the 
third and fourth free thoracic segments as is usual in 
the members of the genus. The cephalic segment, 
seen from above, comparatively large and somewhat 
quadrangular in outline. Frontal margin broadly 
rounded and terminating on each side in a distinct 
knob-like projection. Lateral margins for some dis- 
tance behind the knob nearly straight. First thoracic 
segment deeply constricted a little behind the middle. 
Second segment decidedly larger than the first, and its 
lateral margins somewhat inflated. Third segment 
large and cylindrical, tapering slightly towards the 
posterior end. Abdomen very short and three-jointed, 
rather less than one-seventh of the length of the entire 
animal. Genital segment comparatively large and 
swollen, and equal to the combined length of the 
second and third segments together with the furcal 
joints. Second and third segments short and of nearly 
equal length. Furcal joints short and about as long as 
the last abdominal segment, their length equal to twice 
the width. Each furcal joint furnished with one 
marginal and four apical setae, which are very small. 



136 BRITISH PARASITIC COPEPODA. 

The frontal end of the cephaHc segment and the second 
and third abdominal segments as well as the fnrcal joints 
have the dorsal surface covered with fine spinnles. 

Antennules short, stout, and six-jointed, resembhng 
those of Eudactylina aeanthii. The fourth joint very 
distinctly shorter than any of the others. The second 
joint furnished with a single stout curved spine. 

Antennae also similar to those of Eudactylina aeanthii, 
except that the inner margin of the second joint is 
produced into two short teeth, and the terminal claw 
is much stouter. 

Mandibles and maxillae somewhat rudimentary in 
structure and very similar to those of Eudadylin 
aacnta. 

The first pair of maxillipeds resembling those of the 
other members of the genus in general appearance, but 
the end joint is narrowly ovate in shape and its inner 
margin is furnished with a row of fine teeth. Terminal 
claw slender. The second pair of maxillipeds large and 
strong, and forming powerful chelse. They are nearly 
similar in structure to those of Eudactylina minuta. 

The first pair of feet biramose. Both branches 
short, stout, and two-jointed. The second joint of 
each branch much shorter than the first. The armature 
of the joints similar to that of the first pair of Evdacty- 
lina minuta. The second pair of feet decidedly different 
from the second pair of the other members of the 
genus known to us. Inner branch very short and three- 
jointed. Outer branch also three-jointed but having a 
distinctly uncinate appearance, and being about three 
times the length of the inner branch. The first joint of 
the outer branch large and swollen ; equal to four times 
the combined length of the second and third joints. 
The second joint short and wide. The outer margin 
produced into a bluntly-rounded point. The third 
joint very short and narrow, and appearing to be 
loosely articulated to the second joint. It requires 
some care to detect it, as it appears to be easily folded 
over the surface of the second joint. The outer margin 



EUDACTYLINA INSOLENS. 137 

of the first joint strongly chitinized. The third pair 
of feet with the branches of nearly equal length and 
both branches three- jointed. The fourth pair of feet 
with a three- jointed outer branch and a two- jointed 
inner branch. The first joint of the outer branch large 
and swollen. The second and third joints short and of 
nearly equal length. The inner branch about half the 
length of the outer one and its two joints of nearly 
equal length. The fifth pair large and foliaceous and 
similar to those of Eudactylma minida, except that 
there are no spinules on the surface. 

Habitat. — Four specimens, all females, were found 
on the gill filaments of a male tope, Galeorhinus 
galeus, captured in the vicinity of King William Bank, 
off the north of the Isle of Man, Irish Sea, April 
1912. The topes from which Eudacfylina insohns and 
Kroyeria cineata were obtained, were caught in the 
trawl of the Lancashire and Western Sea Fisheries 
steamer, and landed at Piel, Barrow-in-Furness, along 
with other material, by Captain Wignall. 

We were inclined at first to regard this Eudactylina as a 
form of Eudactylina acanthii, but a close examination showed 
that there were decided differences. The strong uncinate 
appearance of the long outer branch of the second pair of 
feet can be easily seen by examining the entire animal when 
lying on its side. That character, along with the differences in 
the structure of the first, third, and fourth pairs of feet, readily 
separate Eudactylina insoleris from Eudactylina acanthii or 
any of the other members of the genus. The incomplete 
segmentation of the thorax, which gives the species only three 
free segments instead of four, may be abnormal and due to 
uncompleted development, but as the whole of the appendages 
appear quite normal we prefer in the meantime, in the 
absence of further material, to regard the form as distinct. 



Family iv. Philichthyid^. 

Female. — Body elongated, more or less segmented 
but without articulated locomotive appendages. Fre- 
quently furnished with lateral processes, which may 



138 BRITISH PARASITIC COPEPODA. 

be slender and flexuose, or in the form of spines. 
Antenna3 and mouth-organs more or less rndimentary. 

Male. — Body slender, distinctly segmented, and 
furnished with several cephalothoracic appendages, 
including antennules and antennae; first and second 
maxillipeds and two pairs of thoracic legs all bira- 
mose ; the first abdominal segment also sometimes 
provided with a pair of limbs. The first maxillipeds 
with strong terminal hooks forming powerful grasping 
organs. Abdomen usually composed of about eight 
segments. 

Habitat. — The species belonging to this Family 
occur free in the mucus canals and sinuses of various 
fishes. 



Genus PHILICHTHYS Steenstrup, 1861. 

Female. — Head small, rounded. Body elongated 
and distinctly segmented ; no dorsal plates, but the 
body, including the head, furnished with a number of 
lateral and ventral processes, moderately slender, and 
more or less curved inwards upon the ventral aspect. 
The egg-strings, which are of moderate length and 
thickness, extending alongside the body, and are partly 
enclosed and supported by the curved lateral and 
ventral processes. 

There is in the female a single median eye- spot. 

Male. — The male is much smaller than the female. 
The body is slender and distinctly segmented. The 
anterior part of the body consists of three segments, 
the first moderately large and bluntly rounded in front, 
the other smaller. The posterior portion is also 
segmented, very narrow, and considerably longer than 
the front part. 

Antennules slender, composed of six articulations ; 
antennae two-jointed and provided with two hook-like 
setae. The first maxillipeds moderately large, the 
second feeble. 

Only one species is known. 



PHILICHTHYS XIVRIM. 189 

1. Philichthys xiphiae Steenstrup. 
(Plate XLV, figs. 2, .3.) 

1862. Philichthys xiphice Steenstrup. (126) p. 295, pi. ii. 

1864. Philichthys xiphix Bergsoe. (17) p. 87, pi. 13. 

1877. Philichthys xiphiai C. Yogt. (142) p. 29, pi. ii, figs. 13-15. 

Female. — Body elongated and somewhat tumid, and 
having a rugged or coarse appearance ; consisting of 
numerous segments and furnished with many append- 
ages which vary in shape and size. Anterior portion 
of the cephalothorax consisting of three small seg- 
ments, the one in front being extremely minute ; a 
minute median eye-spot present on the dorsum of the 
second segment and a mouth-aperture beneath ; the 
body immediately posterior to this front portion of the 
cephalothorax becoming more tumid and expanding 
equally on each side so as to form a nearly circular 
disc, the width of which is fully more than a third of 
the entire length ; the remaining portion of the body 
for the most part subcylindrical and composed of 
numerous annulations, the penultimate segment small 
and only about half the w4dth of the preceding one ; the 
anal segment broadly subtriangular and with tw^o 
small knobs on the dorsal aspect, one on each side 
of the median line, and a blunt-pointed, finger-like 
process extending backwards from the middle of the 
posterior margin. Genital segment the sixth from the 
posterior end. 

Body furnished with about fifteen appendages on each 
side, a few ventral but most of them lateral and all soft 
and uniarticulate, those on the posterior half of the 
body more or less incurved and hook-like. Egg-strings 
tolerably elongated and thick, attached to the genital 
orifices by the middle, extending forward as well as 
backward along the ventral aspect, and enclosed and 
supported by the curved lateral and ventral processes ; 
the egg-strings not extending beyond the posterior end 
of the body. Colour pale purple, consisting of numerous 
minute purple spots on a white ground. Length 
varving from 6 to 36 mm. — {Bergsoe.) 



140 BRITISH PAEASITIC COPEPODA. 

Male. — Slender, elongated, becoming gradually at- 
tenuated posteriorly. Body distinctly segmented, rings 
free and mobile. First segment of the cephalothorax 
scutiform, about as broad as long and equal in length 
to the next two segments taken together ; third seg- 
ment larger than the second and armed with a strong, 
slightly curyed spine on each postero-lateral angle. 
Abdomen slender ; anal segment longer and narrower 
than any of the preceding segments. Caudal rami 
yery slender, elongate, rather shorter than the anal 
segment, and proyided with two apical setae. 

Antennules yery slender and consisting of six indis- 
tinct joints. Antennae moderately stout, two- join ted, 
and furnished with two terrpinal, slightly hooked setae. 
In the first maxillipeds the basal part is greatly 
enlarged and bears strong claw-like terminal spines ; 
the second pair are small and feeble. Locomotary legs 
two pairs, short and biramose ; both pairs haye the 
outer ramus two- and the inner one-jointed ; the rami 
are subequal in length but the outer ramus is rather 
more robust, and the proximal joint is very short; 
both pairs are liberally supplied with spines and setae. 
Length about 4 mm. 

Habitat. — Living freely in the mucous canals in 
the head of the sword-fish (Xiphias r/ladins). On a 
sword-fish captured off Lowestoft in 1892 ('Cambridge 
Natural History,' vol. iy, p. 73, footnote). 

Distribution. — Mediterranean. jSTew Zealand (G. M. 
Thomson). 

Dr. S. F. Harmer, who examined the sword-fish and ob- 
tained the specimens of Philichthys described above, has sent 
ns the following note, which he kindly permits us to use : — 

" Some years ao^o (1892), when examining a specimen of 
sword-fish which had been taken off Lowestoft, it occurred to 
me to look for Philichthys xiphise, which 1 readily found in the 
place in which it has been described as occurring, namely, 
in the frontal bones. ... I speak from memory, but I 
think I am rightinsayinof that one of the parasites was found 
in each frontal bone.^' — {S. F. Harmer.) 



LERN^OIDA. 141 

Tribe III. LERN^OIDA. 

Mouth usually suctorial. Thorax not articulated, 
or only obscurely so. Thoracic legs, when present, of 
a more or less rudimentary structure. Body some- 
times, as stated by Baird, " very outre in appearance."* 
Egg-strings two, stout and of moderate length, or some- 
times slender and greatly elongated and straight or 
more or less twisted. Male usually very small. 

The species belonging* to this tribe are all more or less 
permanently fixed upon their hosts ; but, as stated by Baird, 
it is in general " only the adult female of the Lerna3ida3 that we 
are in the habit of observing, and in an animal whose organs 
of motion and perception for the most part are merely rudimen- 
tary, and whose existence is strictly stationary, the manner of 
life must be very simple.^t They are usually arranged under 
the three families Lern^idee, Chondracanthidae, and Lernseo- 
podge, which differ, and, for the most part, are characterized 
by the manner in which the parasites fasten themselves to the 
fish on which they live, which also presupposes corresponding 
differences in at least some of the other appendages, as well 
as in the relation of the sexes. (See remarks on Lernsea in 
the Introduction.) 

The young Lernsea leaves the egg as a free-swimming 
nauplius, which, in general appearance, is very similar to the 
larva of Cyclops. 

Family v. Lern^id^. 

Body of the mature and fixed ovigerous female more 
or less cylindrical, and unsegmented but roughly 
divided into three parts — a globular head with anchor- 
like processes either simple or branched; a narrow, 
cylindrical, and sometimes more or less flexuous 
neck connecting the head with the posterior part of 
the body or genital segment ; the genital segment 
usually more or less swollen, and straight or sig- 
moid. Egg-strings two, slender, elongated, sometimes 
twisted into two involved masses, or forming more or 
less regular spirals. 

* * British Entomostraca/ p. 307. f 0]^. cit., p. 318. 



142 BRITISH PARASITIC COPEPODA. 

In the young but sexually mature stage the Lernseidae do 
not differ greatly from the preceding families. The sexually 
mature young are furnished with two pairs of antennee ; the 
posterior pair usually end in strong hooks which project 
beyond the forehead ; the maxillipeds are small and feeble, 
and the four pairs of thoracic legs are tolerably developed. 
The abdomen is usually rudimentary, and a median eye- spot 
is present. In the larval stages the young approximate more 
closely to Cyclops and are more or less free-swimming. 

Genus 27. LERN^A Linnaeus, 1767. 
Head globular, slightly recurved, with the apex 
rounded or conical, and provided usually with three 
chitinous, more or less branched and liorn-like append- 
ages, two of them lateral and one median and dorsal. 
Thorax in the form of a moderately long, slender, and 
fiexuous neck, which gradually becomes merged into 
the genital segment. Genital segment moderately 
enlarged, elongated and subcylindrical, and usually- 
more or less abruptly bent upon itself, somewhat like 
the letter S. Egg-strings elongated and slender, and 
twisted into involved roundish masses under the pos- 
terior portion of the genital segment. Several of the 
cephalothoracic appendages becoming, in the fixed stage 
of the female, degenerate and rudimentary or obsolete ; 
the first maxillipeds however retain their form, and 
are furnished with terminal hooks ; the four pairs of 
thoracic legs are also persistent, and may be seen on 
the ventral aspect behind the subglobular head with the 
aid of a low-power objective. 

The various species of Lerntea are securely anchored to the 
host by the cephalic horns. 

1. Lernaea branchialis Linnseus. 
(Plates XLII & XLIII.) 

1767. Lernsea hranchialis Linn. (78) vol. i, pt. 2, p. 1092. 
1850. Lernsea branchialis Baivd. (4) p. 344, pi. xxxv, fig. 12. 

1900. Lernxa branchialis T. Scott. (112) p. 161, pi. vii, figs. 11, 12. 

1901. Lernsea branchialis A. Scott. (108) p. 33, pis. iv and v. 

Female. — Head apiculated or conical, slightly re- 
curved and provided with strong, branching, horn-like 



LERN^A BEANCHIALIS. 143 

appendages; neck narrow, of moderate length, and 
somewhat flexuose ; genital segment usually tolerably 
elongated, somewhat swollen and tapering slightly to 
the bluntly-pointed distal extremity ; also abruptly 
folded upon itself in the form of the letter S. Egg- 
strings long, slender, and twisted into roundish 
masses under the posterior portion of the genital 
segment. 

Cephalothoracic appendages rudimentary or obso- 
lete, except the first pair of maxillipeds, which are 
situated near the apex of the head immediately behind 
the mouth ; the four pairs of thoracic legs situated at 
the proximal end of the neck, and being exactly as they 
exist in the cyclop s stage, both in size and structure. 
The first and second pairs biramose and both rami 
biarticulate, but the inner ramus wanting in the third 
and fourth pairs while the outer is composed of two 
joints. The whole animal when stretched out, and 
exclusive of the cephalic horns and the egg-strings, 
measuring about 40 mm., but the size varies to some 
extent. Colour dark red, due to the contained 
blood. 

Cephalic horns usually fixed in the gill-arches of 
the fish, the tissue of which they simply penetrate 
when the animal settles on the host, and then become 
branched, thus securing a firm anchorage. The 
branches in this species assume a more or less dicho- 
tomous arrangement, and in this respect differ from 
the other species to be described. 

Male. — The male is very small, and resembles the 
young female in general appearance. Having reached 
the cyclopoid stage it undergoes no further change, and 
is then sexually mature. 

Habitat. — The female in the adult fixed stage is 
usually parasitic on the gills of various Gadoids, as cod- 
fishes, haddocks, and whitings, and is not uncommon ; 
while young females with males attached to them may 
be found on the gill-filaments of flat-fishes such as the 
flounder, Pleuronectes flesus. The following are some 



144 BRITISH PARASITIC COPEPODA. 

of the localities where this Lerndea has been obtained : 
Dublin and Belfast Bays {W. Thompson). Polperro 
and Falmouth {A. M. Norman). Plymouth {BassetU 
Smitli). Various parts of the Scottish and Lancashire 
coasts {nob.). Coasts of Northumberland and Durham 
(Brady ^ Norman). Four adult female Lernsea hranclii- 
alis were obtained from the gill-arches of twenty-four 
Gallionymus li/ra and one on the gill-arches of a Gentro- 
notus c/imnellus which were captured in the Irish Sea 
in 1910 (.4. Scott). 

2. Lernsea lusci Bassett- Smith. 
(Plate XLIV, figs. 1, 2.) 

1896, Lernsea lusci Bassett-Smith. (6) p. 13, pi. iv, fig. 6. 
1904. Lernsea lusci T. Scott. (115) p. 277, pi. xvii, figs. 12, 13. 

Female. — Horns of the cephalon unequally developed, 
stout, and not so branching as in Lernsea branchialis ; 
very short, except the one which springs from the dorsal 
aspect of the cephalon, this branch, which is tolerably 
elongated, extends outwards at nearly right angles 
to the neck, and is narrow and linguliform with the 
margins irregularly lobed and reflexed ; the other horns 
very short and terminating in one or two rudimentary 
branches. Neck moderately short and thick, merging 
posteriorly and directly into the genital segment ; this 
segment being proportionally considerably swollen and 
at first abruptly reflexed and bent upon itself, except 
at the distal end, where, in marked contrast to Lernsea 
hrancliialis^ it is only slightly curved. The twisted egg- 
strings are also proportionally less slender than those 
of that species. 

The antennae and other cephalothoracic organs 
appear to be somewhat similar to those of Lerrtaea 
hranchialis. Length of the female represented by the 
drawing (PL XLIV, fig. 1) only a little over half an 
inch (or about 15 mm.), measuring from the head to 
the end of the genital segment. Colour dark red. 

Habitat. — Parasitic usually on the gill-arches of 



LERNJiA LUSCr. 145 

brassies or whiting-pouts, Gadus luscus (Linn.). Found 
on the gills of this species of fish at Plymouth 
{Bassett- Smith). Found on the gills of a brassie cap- 
tured 10 miles off Aberdeen in January 1901 {T. Scott). 
Frequent on the gill- arches of brassies, Irish Sea (A. 
Scott). 

A Gadus luscus sent from the Fish-Market at Aberdeen 
had one of these parasites adhering* to it; in this example 
the head of the parasite was buried in the tissues in the 
abdominal region, behind and a little below the base of the 
pectoral fin — a somewhat unusual position for a Lemma. 

3. Lernsea minuta T. Scott. 
(Plate XLIV, fig. 3.) 

1900. Leryisea minuta T. Scott. (112) p. 161, pi. vii, fig. 13. 
1904. Lernsea minuta, A. Scott. (109) p. 42. 

Female. — Head moderately enlarged, not very clearly 
defined, and merging into the rather short and stout 
terminal appendage which is somewhat distorted and 
furnished with small marginal papillge ; two very short 
lateral appendages also present, with their ends ob- 
scurely bifurcate. Neck very short, narrow, and some- 
what abruptly joined to the genital segment ; this 
segment, which is sigmoid, having the proximal half 
considerably swollen, but tapering somewhat towards 
the bluntly -rounded and slightly-recurved extremity. 
Egg-strings as in Lerndea branchialis but rather stouter. 
Antennae and other cephalic organs rudimentary or 
obsolete ; thoracic legs four pairs, small, but quite dis- 
tinct and situated immediately behind the lateral 
cephalic horns. Length of the specimen represented 
by the drawing (PI. XLIV, fig. 3) a little over 7 mm. 
Colour dark red. 

Habitat. — Parasitic on the gill-arches of speckled 
gobies, Gohius minatus G-mel. On Gobius miiiiUiis 
from the Solway in November 1899 (T. Scott); and 
on specimens of the same kind of fish captured in the 
Irish Sea {A. Scott). 

VOL. I. 10 



146 BRITISH PAEASITIC COPEPODA. 

4. Lernaea lumpi T. Scott. 
(Plate XLIV, fig. 4.) 

1901. Lemxa lumpi T. Scott. (113) p. 128, pi. vii, fig. 12. 

Female. — Head globular, furnished with three very 
short, simple, spine-like horns, two of them lateral and 
one dorsal. Mouth seen as a small papilliform promi- 
nence on the ventral aspect. Head separated from 
the neck by a shallow constriction which is not an 
articulation. Neck long, moderately slender, slightly 
flexuose, and somewhat wrinkled. Grenital segment, 
though rather stouter than the neck, proportionally 
less swollen than in Lernaea branchialis, neither is it 
sigmoid as in that species but doubled round in the 
form of a hook; a distinct constriction also present 
between that part where the egg- strings are attached 
and the caudal portion of the body. Clusters of egg- 
strings small and more or less twisted, as is usual in 
Lernsea. Antennae and mouth-appendages appearing to 
be somewhat similar to those of other species of Lcrnasa. 
Length about two inches (50 mm.). Colour dark red. 

Habitat. — Parasitic on the gill-arches of lumpsuckers, 
Gi/clopterus lumpus. Found on a lumpsucker cap- 
tured in the salmon nets at the Bay of Nigg near 
Aberdeen, 29th March 1900. 

Only a single specimen of this somewhat curious parasite 
was obtained, though dozens of lumpsuckers have been exa- 
mined ; the head and a portion of the neck measuring about 
three-quarters of an inch penetrated the tissues of the fish. 

The structure of this species seems in some respects to 
approach more nearly to that of Pennella than is the case with 
the adult Lernsea hranchialis, the body is recurved to a much 
smaller extent, and the cephalic horns are greatly reduced in 
size ; its hold on the fish might therefore be correspondingly 
weakened, but its fixation is rendered secure by having a 
larger proportion of the neck enclosed in the tissues of the 
fish. 

It sometimes happens that the Lernsea dies while still 
attached to the living fish, and in that case, though the 
genito-abdominal part of the parasite's body disappears, the 
fish seems to be unable to get quit of the tougher and more 



LERN^A LlIMPr. 147 

chitinous neck and head which may remain fixed to the 
fish for a considerable time. It is interesting to note the 
difference in the form of the body of the two closely allied 
genera Fennella and Lernsea : in the one the body is elongated, 
cylindrical, and straiglit or nearly so; in the other it is bent 
back upon itself, forming one or two more or less abrupt loops. 

Genus 28. HJEMOBAPHES Steeristrup k Liltlcen, 

1861. 

Parasites having a general resemblance to Lenisea. 

Head rectangular, flattened, longer than broad and 
without lateral horn-like appendages. Two short, 
more or less distinct thoracic segments immediately 
posterior to the head, each bearing a pair of rudimen- 
tary bilobed appendages, followed by two pairs of legs, 
biramose and biarticulate. Neck elongated, the 
anterior end abruptly reflexed so that the head 
becomes pendulous, and near the flexure are two short, 
lateral, bluntly-pointed horns. Grenital segment sig- 
moid, proximal half swollen, distal half narrow^er and 
with the apex bluntly pointed, a pair of short lateral 
processes occurring on either side over the origin of 
the egg-strings. Egg-strings very long and slender, 
but twisted up into regular convoluted spires. 

1. Haemobaphes cyclopterina (0. Fabr.). 
(Plate XLIY, figs. 5-7.) 

1780. Lernxa cyclopterina O. Fabr. Fauna Groenlandica, p. 337. 
1822. Lernaeocera cyclopterina Blainv. (24) vol. 95, p. 376. 
1837. Lermea cyclopterina Kro,yer. (70) p. 502, pi. v, fig. 4. 
1840. Lernsea cyclopterina M. Edwards. (43) vol. iii, p. 529. 
1861. Heemobaphes cyclopterina Steenstrup & Liitken. (127) p. 405, 
pi. xiii, fig. 30. 

1900. Hsemobaphes cyclopterinus T. Scott. (112) p. 162, pi. vii. fig. 14. 

Female. — Head small, rectangular, without horn-like 
projections ; two short, more or less distinct thoracic 
segments crowded behind the head, each furnished 
with a pair of rudimentary bilobed appendages. Two 
pairs of biramose legs with two-jointed rami also 
present, the rami being more or less setiferous. Xeck 
long and slender, and bearing midway between the head 



148 BRITISH PARASITIC COPEPODA. 

and genital segment two short lateral horns or pro- 
cesses ; at about this point a more or less abrupt 
flexure occurring, so that the head becomes pendulous. 
The proximal half of the genital segment somewhat 
swollen, the terminal part narrow and ending in a 
bluntly-pointed apex. A pair of short processes on 
each side of the genital segment over the origin of the 
egg-strings. Egg-strings, which form regular con- 
voluted spires, situated at the termination of the 
swollen portion of the genital segment. Length about 
12 mm. Colour dark red, similar to that of Lernaea, 

Habitat. — Parasitic on the gill-arches of various 
fishes. On the gills of a pogge, Agonus cataphractus^ 
obtained by Mr. Peter Jamieson in the stomach of a 
large codfish landed at Dunbar, Haddingtonshire, in 
April 1891. Firth of Forth, on the gills of a pogge 
in February 1892, and on the gills of a butter-fish, 
Centronotus gunnellus, captured in 1901 (T. Scott), 
Irish Sea {A, Scott). 

The following other fishes are mentioned by Steen- 
strup and Liitken as hosts for this parasite : Cyclo})- 
terus spinosus, Cottus scorpms, G. bubalis, and C. groen- 
landlcus, Sebastes novvegicus, Centronotus fa sciatus^ and 
Gadus nfierlangiis . * 

Genus 29. HJEMOBAPHOIDES T. ^' A. Scott. 

Syn. Hiemohaphes T. Scott (not Stp. & Liitk.). 

Resembling Hddniobaphes Stp. & Liitk., but the head 
furnished with branched chitinic horns ; neck ver\^ 
short, the posterior portion of the genital segment 
more produced, compressed, and expanded at the end. 

1. Hsemobaphoides ambiguus (T. Scott). 
(Plate XLIV, fig. 8.) 

1900. Hwmohaphes ambiguus T. Scott. (112) p. 162, pi. vii, fig. 15. 

Female. — Head not very clearly defined, and pro- 
vided with branched cartilaginous horns, which are 

* ' Parasitiske Copepoder/ p. 65 (1861). 



HiEMOBAPHOIDES AMBIGUUS. 149 

short and stout. Neck very short or nearly obsolete, 
genital segment flexuous ; the anterior portion swollen 
as in Hasmobaphes, but the distal portion narrow, and 
prolonged so that it at least equals in length the 
proximal part; this narrow elongated part also flat- 
tened, and, at the extremity, abruptly expanded, the 
margins somewhat irregular in outline and inflexed, 
and the lateral lobes at its proximal end moderately 
prominent. A distinct though quite shallow median 
groove extending along the dorsum of the swollen half 
of the genital segment. Egg-strings forming regular 
convoluted spires as in Hdemobaphes. Colour reddish- 
brown. Length about 11*5 mm. 

Habitat. — Parasitic on the gills of spotted dragonets, 
Callionymus maculatus Bonap. On spotted dragonets 
captured in the Solway Firth and in the Firth of Clyde. 

In a sample of fifty-five Callionymus macidatus captured 
in the Firth of Clyde in October 1901, fifteen specimens of 
Hsemohaphoides amhigtius Siud eight of Chondr acanthus ornatus 
were obtained. In most instances the specimens of the two 
species occurred singly and on different fishes, but in several 
cases two specimens of the same species or a specimen of each 
occurred on the gills of a single fish. For example, a spotted 
dragonet 73 mm. in length had a Chondr acanthus on one side 
and a Hsemohaphoides on the other ; another dragonet had a 
Chondr acanthus and a Hsemohaphoides on the same side ; a 
third had two Chondr acanthus on the same side, while the 
other side was free of parasites; and a fourth had a Hxmo- 
haphoides on each side but no Chondr acanthus. It was also 
noticed that when only one parasite occurred on a fish it was 
frequently on the right side — the fish resting on its ventral 
surface and with its head toward the observer. It was further 
noticed that though seventeen specimens of the common 
dragonet [Gallionymiis lyra) captured at the same time and 
place were examined, no parasites were observed on them. 



Genus 30. PENNELLA Oken, 1815. 

Female. — Body greatly elongated and slender. 
Head tolerably large, globose or nearly so, studded 
with minute tubercles and usually provided with horn- 



150 BRITISH PARASITIC COPEPODA. 

like appendages which extend obliquely backwards 
or at nearly right angles to the median line of the 
body. Thoracic region forming a more or less elon- 
gated and slender neck. Anterior portion of the 
genito-abdominal region elongated and moderately 
stout, and bearing near its distal end two tolerably long 
egg-strings. The terminal Dr caudal portion of the 
region more or less distinctly annulated and carrying 
along its ventral surface numerous fascicles of bristle- 
like and more or less branching appendages. 

Antennules short, small, setiferous. Antennae small, 
uncinate. Mouth-aperture near the anterior end on 
the ventral aspect. Thoracic legs four pairs, situated 
close behind the head and with only a small interval 
between each pair. The first and second pairs bi- 
ramose, but the others one-branched ; the rami all 
two- jointed, and all the thoracic limbs small. 

In the immature Fennella the head is narrow and 
subcylindrical, and the horns are usually wanting. 
Both pairs of antennae though small are quite distinct, 
the second pair are provided with strong terminal 
claws and form effective grasping organs ; the second 
maxillipeds are also stout and strongly clawed; a 
considerable space intervenes between these append- 
ages and the thoracic legs. 

"Male minute and not elongated " {B as sett- Smith). 

M. Edwards remarks that the male is very small, nearly 
spherical, and possesses anteriorly a conical sucker, furnished 
with a few styliform appendages ; and that on the inferior 
aspect there are two pairs of very large subcheliform claws 
by means of which it is enabled to hang on to the female.* 
We have not seen the male. 

1. Pennella orthagorisci E. P. Wright. 
(Plate LI, fig. 2.) 

1829-1843. Pennella filosa Gnerin-Meneville. (55) p. 11, pi. ix. fig. 3. 
1861. Pennella filosa Steenstriip & Liitken. (127) pi. xiv. fig. 31. 
1870. Pennella orthagorisci E. P. Wright. (150) p. 42, pi. i, figs. 1-6. 

* ' Hist. Nat. Crust./ vol. iii, p. 522. See also G. M. Thomson, ' Trans. 
New Zealand Institute/ vol. xxii, 1889, p. 368. 



PENNELLA OBTHAGORISCI. 151 

1889, Pennella orthagoHsci Giard. (50) p. 82. 

1899. Pennela filosa Bassett-Smith (part). (8) p. 483. 

1905. Pennella filosa T. Scott. (116) p. 113. 

1905. Pennella ortliagm-isci T. R. R. Stebbing. (124) p. 119. 

1906. Pennella filosa Norman & T. Scott. (88) p. 216. 

1908. Pennella filosa Cuvier? A. Brian. (21a) p. 8, text-figs. 2 «& 3. 
1910. Pennella orihagorisci T. R. R. Stebbing. (125) p. 256. 

Female. — Body elongated and slender; head mode- 
rately large, globose or nearly so, and provided with 
three horn-like appendages on the ventral aspect; 
the middle horn very small and rudimentary but the 
others of moderate size and projecting obliquely back- 
wards. Neck elongate, slender, smooth, and equal to 
about one third of the entire length of the animal. 
Genital segment moderately stout and about as long 
as the neck ; obscurely annulated and bearing at the 
distal end two long and very slender egg-strings. The 
terminal segment, or post-abdomen, which is fully half 
as long as the genital segment, bearing along the 
ventral side fascicles of branching cartilaginous ap- 
pendages, each fascicle dividing into two or three 
principal branches, which are again subdivided irregu- 
larly and in a bifurcate manner into long slender 
filaments, but including also a few which are short or 
undeveloped. Colour of the animal blood-red. Length 
90 to 100 mm. {^ to 4 inches). 

Habitat. — Parasitic usually on the short sun-fish, 
Orthagoriscus mola. 

The Rev. A. M. Norman, whose kindness we have 
experienced on numerous occasions, has permitted us to 
examine and figure a Pennella sent to him many years ago 
(about 1862) by TJiomas Edward of Banff. The specimea 
was found on a short sun- fish captured in the Moray Firth, 
and is referred to in Smiles^ Life of Edward among the 
numerous other natural history records at the end of that 
work, under the name of Pennella fibrosa, that name being 
no doubt a misprint for '\fllosa " ; unfortunately this speci- 
men wanted the head. Some years previous to the publica- 
tion of Smiles' Life of Edward^ Dr. E. P. Wright described 
under the name of Pennella orihagorisci specimens found on 
short sun-fish captured in Cork Harbour in 1869"'^ ; he also in 

* 'Ann. & Mag. Nat. Hist./ ser. 4, vol. v, p. 42, pi. i, figs. 1-6 (1870). 



152 BRITISH PARASITIC COPEPODA. 

the same paper states that Dr. Baird informed him that he had 
^' examined a specimen of Pennella from a sun-fish captured 
at Megavissy, Cornwall^ which he (Dr. Baird) refers to 
P. filosa Linn.^' We have little doubt that the specimens 
obtained at the places mentioned were all referable to the 
same species. 

The name Pennella filosa sometimes applied to the sun-fish 
parasite appears to have been given to it under the supposi- 
tion that it and the swordfish Pennella were identical ; thus 
far, however, there is no satisfactory evidence to show that 
they are so, and, till such evidence is forthcoming, the name 
Pennella filosa (Linn.), as pointed out by Steenstrup and 
Lutken, should be retained for the form found on the sword- 
fish. Percival Wright's Pennella orihagorisci should also for 
similar reasons be accepted for the Pennella of the sun-fish. 
The Eev. T. R. R. Stebbing, in Part III of his work on 
" South African Crustacea" (1905), in a short discussion on 
Pennella, remarks that '^the description given by Linnasus 
can scarcely be said to have any specific value apart from 
the name of the host, so that no injury is done him by leav- 
ing his specific name in abeyance until a Pennella infesting a 
Xiphias has been again observed. For the parasite of the 
sun-fish an appropriate name is available, which appears to 
have escaped recent attention.""^ 

In January 1908, M. Alexandre Brian published an in- 
teresting paper entitled *'Note preliminaire sur les Cope- 
podes parasites des poissons provennnt des campagnes 
scientifiques de S.A.S. le Prince Albert 1^^ de Monaco ou 
deposes dans les collections du Musee Oceanographique.^' 
This Note forms Bull. No. 110 of the Oceanographical Insti- 
tute. The author describes with illustrative figures an adult, 
and several immature, female Pennellm under the name of 
" Pennella filosa Cuvier '^ which were obtained on a sun-fish 
captured in lat. 39° 56' 10" N., long. 34° W., 19th July 
1887, and which seem to be identical Avith Wright's P. 
orihagorisci. He also describes a '^ Pe7inella sp.?" A single 
adult obtained " dans la peau du ventre d'un Xiiolilas gladius " 
captured at Saint-Jean-sur-Mer, 11th March 1905. May 
this Pennella not be referable to the species described by 
Linnaeus ? It is more than twice the size of the adult female 
recorded from the sun-fish ; the shape of the head and the 
form and size of its two horn-like appendages are also very 
different. The length of this specimen is described as 212 

* ' Marine Investigations in Sotith Africa/ " South African Crustacea/' 
pt. iii, p. 118. 



PENNELLA ORTHAGORISCI. 153 

millimetres (= about 8 J inches), whereas the adult female 
from the sun-fish is only 93 millimetres (about 3|- inches). The 
head of the young female of the sun-fish Pennella, as shown 
by Brian, is narrow and subcylindrical, fully three times 
longer than broad, and the horn-like processes are wanting ; 
the abdominal appendages are also more or less rudimentary, 
and the posterior annulations appear to be more distinct. 

The free-swimming larvae of Pennella have a close resem- 
blance to those of Lernsea hranchialis and exhibit the near 
relationship between these genera. Dr. Al. Mrazek, in a 
paper " Ueber Bacculus Lubb. und Hessella Br. Ein Beitrag 
zur Anatomie der Lerngeiden,^' ^ gives a detailed description 
of the anatomy of these larval forms. 

Pennella balsenopterse Koren & Danielssen. 

Specimens of a giant species of Pennella found attached to 
a finner whale [Balsenoptera muscuhts (Linn.)) were presented 
to Sir William Turner, F.R.S., Edinburgh, by Mr. Chr. 
Castberg, the manager of a Norwegian Whaling Company 
which has a fishing-station at Ronasvoe, in the north of 
Shetland. A full and interesting description of the species, 
illustrated with four plates, is published by Sir William 
Turner in the ' Transactions of the Royal Society of Edin- 
burgh,^ vol. xli, Part 2 (No. 18), 1905. 

This somewhat remarkable species does not come within 
the scope of a Monogiaph devoted to the parasites of fishes, 
but we nevertheless refer to it here on account of the author's 
interesting remarks on the history of the peculiar genus to 
which the species belongs, and because the description given 
of the species is also generally applicable to the one found on 
the sun-fish. 

The largest specimen obtained by Sir William Turner 
measured 294 mm. (nearly lU inches) in length, but, as 
the author remarks, even larger specimens have been recorded 
by Koren and Danielssen; one of these is said to have 
measured 320 mm. (12 J inches) in length. In this species the 
head is provided with three slender chitinous horns of unequal 
length which extend horizontally outwards at nearly right 
angles to the body, the dorsal horn being usually the shortest ; 
and in this and a few other respects Pennella halxnopterse 
differs from the sun -fish parasite. It seems however to be 
still a moot-point whether all these different Pennellpe are to 
be regarded as valid species. 

« ' Sitzungsberichte d. konigl. bohm. Gesellsch. d. Wiss./ Math.-nat. 
Classe (1895), xliv, pp. 1-17, 2 plates. 



154 BRITISH PARASITIC COPEPODA. 

Another Cetacean Pennella, P. crassicornis Steenstrup 
& Liitken, was fouud attached to a bottle-nose whale 
(Uyperoodon rostratus (Miill.)) captured south of the Faroes 
in 1855.^ In this form the horn-like appendages of the 
cephalon are short and rather stout, and the whole animal is 
much smaller. 



Genus 31. LERNiEOCERA Blainville, 1822. 

Body long and slender^ head not distinctly de- 
fined, provided ^'itli horn-shaped appendages wliicli 
are simple and more or less symmetrical in form. 
Mouth situated at the apex of a small conical lobe in 
the median line and projecting slightly forward of the 
base of the lateral horn-like appendages. Genital 
segment much elongated, becoming more or less 
thickened posteriorly, and somewhat curved at the 
distal extremity. Abdomen very small or obsolete. 
Egg-strings straight and of moderate length. 

1. Lernaeocera cyprinacea (Linn.). 
(Plate L, figs. 1-5.) 

1761. Le'i'mea cyprinacea Linn. (77) vol. ii, No. 2100, pi. xi, fig. 2. 
1783. Lemxa cyprinacea Barbut. (5) vol. i, p. 3, pi. vii, fig. 3. 
1783. Lerrnea (/) esocina Hermann. (61) vol. xix, p. 44, pi. ii, fig. 6. 
1822. Lernseocera cyprinacea Blainv. (24) vol. xcv. p. 377. 
1832. Lermwocera (.^) cyprinacea Xordm. (89) p. 123. jdI. vi, figs. 1-8. 
1840. Lernseocera {?)eoscina M. Edwards. (43) vol. iii, p. 527. pi. xl, 
figs. 13-15. 

1840. Lernseocera cyprinacea idem. (43) vol. iii, p. 527, pi. xl, fig. 16. 
1850. Lemeoce7'a cyprinacea Baird. (4) p. 343, pi. xxxv, fig. 13. 
1868. Lemfeocera (/) esocina Clans. (34) vol. xxxi, p. 530. 

Female. — " Head furnished with four horn-shaped 
appendages, which are somewhat long and slender. 
The two outer or posterior are bifurcated ; the anterior 
simple. 

" The thorax is very slender anteriorly, forming a 
long neck, but becomes much broader posteriorly, and 
when it terminates in the small abdomen, appears 
obliquely truncate. The ovigerous tubes are cylin- 

* ' Bidrag. til Kundskab cm detaabne Havs Synltekrelts og Lernseer,' 
p. 76 (separate copy), 1861. 



LERNiEOCERA OYPEINACEA. 155 

drical and rather long. The length of the whole 
animal is about eight lines." 

" Hab. — Found on the sides of the carp, bream, 
and roach in many of our ponds and rivers in great 
abundance " (Barhtd). " I have not seen any specimens 
of this species " {Baird, ' Entomostraca,' pp. 343, 344). 

As we have not been able to obtain specimens of this 
Lernxocera, Dr. Baird^s description and remarks are here 
reproduced. Fig. 5 on PL L is also reproduced from his 
work. 

Dr. Baird does not inchide Hermann's Lernsea esocina in 
his list of synonyms, but seems to regard it as a different 
species. He says : " Linnaeus was the first who noticed any 
of the animals belonging to this genus. In his ' Fauna 
Suecica/ 1746, he describes a species found in Sweden on a 
carp; a species which Barbut, in 1783, ascertained to be 
British. Hermann also, in 1 783, describes and figures another 
species, and several have since that time been added to the 
list. . . . The genus, as established by Blainville, con- 
tains two species, which have been separated from it by 
Kroyer and M. Edwards ; but still it has been retained in a 
restricted sense by all succeeding authors." M. Edwards 
apparently recognizes the two species, Lerneocera cyprinacea 
and L. esocina;^ Kroyer (1863) on the other hand does not 
mention either, but records Lernseocera jphoxinacea Kollar, 
and describes two n. spp. from American waters. 

Linnaeus, after giving a brief description of his Lernsea 
cyprinacea, adds, " Habitat in Plscinns freqnens supra corpus 
Cyprini carassii^t ; and Pennant, in speaking of CypHnus 
carassius, which he also calls the " Crucian Carp," says " it is 
common in many of the fish-ponds about London and other 
parts of the south of England but I believe is not a native fish." J 

From the remarks of Dr. Baird it is evident that the 
Lernsea recorded by Barbut is the species described by 
Linnteus, and that it also was found parasitic on fishes 
belonging to the Cyprinida3 ; probably on the '^Karauschen " 
or crucian carps, which were according to Pennant common 
at thnt time in many fish-ponds about London. 

Nordmann in 1832 records a Lernseocera cyprinacea, five 
specimens of which were found "in der unteren Kinnlade des 
Hechtes," or fresh-water pike, Esox lucius, a kind of fish very 

* ' Hist. nat. Crust.,' vol. hi (1840), p. 527. 

t / Systema Naturae/ ed. xii, vol. i, pars ii, p. 1093 (1767). 

J 'British Zoology/ vol. iii, p. 364 (1776). 



lo6 BRITISH PARASITIC COPEPODA. 

difPerent from the Carp family ; and his figures of the para- 
sites of this fish, while in general agreement with that of 
Dr. Baird, represent what appears to us to be a different 
species from the one figured by the English author. Nord- 
mann^s figures show a more robust animal, with shorter and 
thicker cephalic horns, the distal extremity is also more 
decidedly deflexed, and the ovisacs are short and saccate."^ 
Probably this is the Lernaeocera esocina of Hermann and not 
L. cypriiiacea (Linn.). We prefer therefore to retain mean- 
while Linnets species name, cyprinacea, for the Leriifeocera 
recorded by Dr. Baird. 

Genus 32. LERNiEENICUS Lesueur, 1824. 

Body elongated, slender, obscurely or non-seg- 
mented. Head somewhat expanded, sometimes pro- 
duced into a cone-like process in front, and usually 
provided with two or three stiff cartilaginous horns, 
projecting outwards or more or less obliquely back- 
wards. Thorax very narrow, forming a kind of neck, 
and merging posteriorly into the somewhat stouter 
abdomen. Abdomen without penniform appendages. 
Antennules small, obscurely biarticulate, and more or 
less setiferous. Antennae small but strongly chelate, 
and similar to those of Pf'nndla. Thoracic legs very 
small, and situated immediately behind the head, each 
pair slightly apart from the other; the first and 
second pairs biramous and the others uniramous, and 
all the rami two-jointed. Egg-strings long and slender. 

1. Lernaeenicus sprattae (Sowerby). 
(Plate XLVI, figs. 1-5.) 

1806. Lertuva spratta Sowerby. Brit. Miscellany, vol. ii, p. 17, 
pi. Ixviii. 

1840. Lermeonema monilaris M. Edwards. (43) p. 525, pi. xli, fig. 5. 

1850. Lerneonema spratta Baird. (4) p. 341, pi. xxxv, fig. 10. 

1850. Lernivonema bairdi Salter. (106) (ser. 2), vol. vi, p. 86, pi. vii, 
fig. 1. 

1865. Lern<eonema monilaris Heller. (58) p. 248, pi. xxv, fig. 4. 

1868. Lermeenicus spratta Olsson. (92) p. 46. 

* This figure of Nordmann's is reproduced on PI. L, fig. 1. If com- 
pared with Dr. Baird's figures of L. cyprinacea on the same plate (fig. 4), 
the difference referred to will be readily observed. See also Selago's figure, 
which resembles Baird's and is also reproduced on this plate (fig. 5). 



lernj!:enicus spratt^. 157 

1900. Lernxenicus sprattie T. Scott. (112) p. 161, pi. vii, figs. 7-10. 
1907. LerniEenicus sjprattye A. Scott. (Ill) p. 94, pi. ii, figs. 1-5. 

Female. — Body elongated, slender ; head somewhat 
enlarged, and provided with two stiff horn-like pro- 
cesses, one on each side and extending obliquely back- 
wards. Thorax, between the head and the genital 
segment, slender and neck -like, with a number of 
minute constrictions, giving to this part of the body a 
moniliform appearance ; genital segment moderately 
elongated and increasing somewhat in thickness pos- 
teriorly. Abdomen rudimentary and not very clearly 
defined, apparently destitute of appendages. 

Antennules short, distinct, setiferous, indistinctly 
two-jointed, the end joint being shorter than the 
other ; antennge very short, stout, strongly chelate 
as in PenneUa; other mouth-organs somewhat rudi- 
mentary. Thoracic legs small, situated immediately 
posterior to the head ; first and second pairs biramose, 
rami two-jointed; the next two pairs uniramose, the 
rami being also two-jointed. In fresh or living speci- 
mens the colour of the genito- abdominal segment is 
greenish. Egg-strings long and slender, usually 
exceeding the length of the body, which, exclusive of 
the egg-strings, measures about 18 mm. 

Habitat. — Usually found parasitic on the eyes of 
sprats, Clupea sixnitta, and sometimes also on other 
parts of the body. Occasionally more than one para- 
site will be found adhering to the same eye. London 
Market 1848 {W. Wing, see Baird). Youghal, Ireland 
(IF. Thomiison 1852). Observed on a sprat in Leith 
Docks 1890 (/. Scott). Xot uncommon on sprats 
captured along the Lancashire coast {A. Scott). One 
perfect specimen taken at Plymouth (Ba^f sett- Smith). 

A sample of six hundred sprats captured with the shrimp 
trawl off Blackpool in 1910 yielded fourteen Lernseeniciis 
Sfrattpe. The eye of one of the sprats examined had three 
specimens of Lernseenicus fixed to it. The head of the 
parasite is entirely buried in the tissues of the host, and can- 
not be removed except by dissection. The parasite when 
fixed to the eye appears to cause partial or total blindness. 



158 BK'ITISH PARASITIC COPEPODA. 

2. Lernseenicus encrasicola (Turton). 
(Plate XLVI, figs. 6-U.) 

1807. Lermea encrasicola Turton. (141) vol. i, p. 137, No. 108. 
1850. Lerneonema encrasicola Baird. (4) p. 341, pi. xxxv, fig. 11. 
1868. LerniEenicus encrasicola Olsson. (92) p. 46. 
1877. Lernseenicus encrasicola Richiardi. (102) vol. iii. fasc. i. 
1907. LerniEenicus encrasicola A. Scott. (Ill) p. 93, pi. ii, figs. 6-9. 

Female. — Closely resembling Ler7iasenicas Kprattse, 
and may be easily mistaken for it. Among the more 
obvious points of difference are the following : — (1) 
The horn-shaped appendages of the cephalon, instead 
of pointing obliquely backwards, stand out at nearly 
right angles with the median line of the body. (2) 
" The neck is long and slender, quite smooth and 
destitute of the constrictions which mark so decidedly 
the preceding species" {Baird). (3) It is usually 
found attached to the body of its host, the head some- 
times penetrating into the abdominal cavity. 

The appendages of the cephalon and thorax do not 
appear to differ greatly from those of Lernseenicus 
sprattse. Length about 27 mm. 

No males of either species have been observed. 

Habitat. — Parasitic on the anchovy, Enc/raulis 
encrasicholns, and sprat, Clupea spratta. On a sprat 
captured at Youghal (B. Ball, W. Thompson). '' Found 
attached to the bodies of Glnpea encrasicholns and 
sprattus frequently, in Swansea Bay" (TF. Turton). 
Attached to the body of a sprat (/. Donhledaj/, 
British Museum).* A broken specimen, probably 
belonging to this species, was taken from a Clupea 
alosa at Plymouth {Bassett- Smith). On the eye of a 
young pollack at Falmouth (Cocks; cf. 'Crust, of Devon 
and Cornwall' by Norman and Scott, p. 216, 1906). 

"A laroe catch of sprats was taken off Blackpool on 19th 
February 1906 by the Lancashire Fisheries Steamer, and 
a few hundreds of them were landed at Piel. A careful 
examination of these was made, and one sprat with two of 
the above mentioned parasites attached to it and another 
with one were found. The parasites were embedded in the 
* These records are from ' British Entomostraca/ p. 342 (1850). 



LERNiEENlCUS ENCEASICOLA. 159 

tissues at the anterior end of tlie dorsal fin. On dissecting 
one of the specimens out, it was found that the head of it had 
penetrated into the visceral cavity" [A. Scott). ^ 

M. Marcel Baudoin, in his article on the parasites of the 
sardine^t describes under the name of Lernseenicus sardinse 
a form which he has found adhering to this Clupeoid. There 
is, as stated by Aflalo, a British Pilchard-fishery on the south- 
west coast, J and the same fish has occasionally been captured 
in Scottish waters, § but no British specimen of this Lernxenicits 
has yet been observed. Though, however, many pilchards 
are captured each season, this Lernseenicus if present may be 
easily missed, or mistaken for the more common L. sprattse. 
One of the more obvious characters of the species being the 
shape of the head, as this is buried in the tissues of the 
host, sometimes at the side of the eye as in Lernseenicus 
s'prattse and sometimes near the dorsal fin, it is only by careful 
dissection that it can be obtained for examination. The neck 
next the head is extremely slender, and therefore the external 
portion of the parasite is easily broken oiT, while the head 
remains entirely concealed. 

Genus 33. TRIPAPHYLUS Bichiardi, 1878 

Syn. Lerneonema, P. J. van Beneden (in part). 

Female. — Body greatly elougated, slender, non- 
segmented. Head rounded and provided with stiff 
cartilaginous horns. Thorax very slender, forming 
an elongated neck. Genito-abdominal segment narrow 
at the proximal end but becoming enlarged posteriorly 
and furnished with two long and slender distal pro- 
cesses. Antennules and other cephalic appendages 
somewhat similar to those of Jjernaeenicus, 

Male. — Body divided into two nearly equal portions ; 
the anterior portion large and carrying three pairs of 
appendages ; the cephalon rather obscurely defined, 
and the abdomen not so robust as the cephalo-thoracic 
portion. The cephalothoracic appendages also some- 
what rudimentary. 

* * Report for 1906 on the Lancashire Sea- Fisheries Laboratory at the 
University of Liverpool, and the Sea-fish Hatchery at Piel,' No. xv, p. 94. 

t "Les parasites de la Sardine," 'Eevue scientifique,' 5^ ser., vol. iii, No. 
23, p. 715, with text-figures (1905). 

X 'Natural History (Vertebrates) of the British Isles,' p. 398 (1898). 

§ ' Fishes of the Firth of Forth/ by Dr. Parnell, pp. 320-322 (1838). 



160 BRITISH PARASITIC COPEPODA. 

1. Tripaphylus musteli (P. J. van Beneden). 

(Plate XLV, fig. 6 ; Plate LI, fig. 1 ; Plate XLIX, 

figs 1-7.) 

1851. Lerneonema Tnusteli P. J. van Ben. (12) vol. xviii, p. 287, pi. 
No. 8, figs. 1-7, c? & ? . 

1851. Ije7meo7iema musteli idem. (11) vol. xvi, p. 125, pi. 6, figs. 11-14. 

1877. Lerneonema musteli Yogt. (142) p. 69, pi. iii, fig. 11. 

1878. Tripaphylus musteli Ricliiardi. (103) p. xx. 

1885. Tripaphylus Tnusteli Cams. (29a) Prodr. faunse Mediterr., p. 372. 
1899. Lernssenicus musteli Bassett-Smith. (8) p. 485. 
1904. Lernaeenicus musteli A. Scott. (109) p. 41. 
1906. Ti'ipaphylus musteli A. Brian. (21) p. 87. 

Female. — Body greatly elongated and slender, non- 
segmented and with the surface quite smooth. Head 
rounded and furnished with cartilaginous horns. 
Thorax very slender, rather weak and somewhat 
flexuous. Grenito-abdominal portion narrow at the 
proximal end where it joins the neck, but becoming 
gradually enlarged from the front backwards, and 
assuming nearly the form of a spindle. Body pro- 
vided posteriorly with two long slender appendages 
nearly equal in length to the genital segment ; these 
appendages, though in communication with the body- 
cavity, have no connection with the egg-strings, which 
have their origin immediately behind the base of the 
prolongations referred to and are nearly twice their 
length. 

Head with a considerable portion of the neck 
buried in the tissues of the gill-arch of the fish. 
Colour of the parasite deep red. Length as given 
by van Beneden : Body exclusive of the abdominal 
appendages 45 mm. ; abdominal appendages 15 mm. 
in length and the egg-strings 23 mm. The body in 
its greatest width measures 8 mm. — this specimen 
wanted the head. The specimen figured here (PI. LI, 
fig. 1), which is also minus the head, measures 28 mm. 
exclusive of egg-strings. 

Male. — The male of this species has the body 
divided into two unequal portions : the anterior por- 
tion is large, and carries three pairs of appendages — 



TRIPAPHYLUS MUSTELI. 161 

viz. one pair of antennge and two pairs of feet ; the 
posterior is much more narrow, and rounded, and 
carries at the end two rounded tubercles which pro- 
bably represent the fifth pair of feet ; the body may 
also be divided into head, thorax, and abdomen. On 
the sides of the cephalic portion we distinguish a 
pair of setiferous antennae showing feeble indications 
of articulation; the mouth is terminal and fringed 
with minute setaB. The mandibles and maxillse are 
somewhat rudimentary. The two pairs of maxillipeds 
are comparatively Avell developed, and form prehen- 
sile appendages similar to what is found in the males 
of many of the sedentary female fish-parasites. 

The thorax is dilated (or bulging), smooth and 
even on the surface ; it carries two pairs of feet 
singularly formed, and nearly as dilated and elongate 
as the abdomen. The feet of both pairs are con- 
solidated in all their length as the organs of adhesion 
in the females of the Lerneopodians ; it is only at the 
end that they are divided. The anterior pair is 
biramose ; the outer ramus is two-jointed, and is ter- 
minated by a double hook for clinging. The other 
pair is longer; the outer ramus shows three articu- 
lations terminated by two hooks, as in the outer ramus 
of the preceding pair, but on the base of the hooks is 
a small additional tooth which renders these organs of 
adhesion more effective. 

It will be seen from the above description and the 
drawing PI. XLV, fig. 6) that there are some of 
the Lernsean males considerably removed from the 
ordinary type. 

Habitat. — Parasitic on the gills of the smooth hound, 
Musteliis vulgaris {Sqnalus mustelus). Irish Sea {A. 
Scott). 



VOL. I. 11 



162 BltlTISH PARASITIC COPEPODA. 

Genus 34 REBELULA Foche, 1902. 

Syn. Lophura KolKker (name preoccupied for a genus of birds). 

Head subcylindrical, narrow, in the same straight 
line as the neck and furnished with two bilobed or 
slightly branched processes at its base.* Neck usually 
slender and elongated. G-enital segment large, inflated, 
somewhat flattened dorsally, and having small, slightly 
obscure circular depressions on the dorsal and ventral 
aspects, and provided posteriorly with two bundles of 
filiform appendages, one on each side of the abdomen. 
Abdomen small and obscurely lobate. Egg-strings 
tolerably elongated, and containing numerous small 
ova. 

1. Rebelula edwardsi (Kolliker). 
(Plate XLV, fig. 1 ; Plate LI, fig. 5.) 

1853. Lophura edwardsi Kolliker. (68a) Zeitschr. f. wiss. Zool., vol. 
iv, p. 359. 

1860. Lophoura edwardsi Clans. (30a) Wiirzburger naturwiss. Zeitschr., 
vol. i, p. 20. 

1865. Lophura edwardsii Comalia. (36a) Atti della Soc. Ital. d, Sci. 
Nat., vol. ix, p. 1, pi. i. 

1902. Behelula edwardsii Poche. (97) p. 20. 

1906. Behelula eciwarcfsu Brian. (21) p. 90, pi. xix, fig. 1 ; pi. xxi, fig. 5. 

1908. Behelula edwardsi Brian. (21a) p. 15, text-fig. 6, a, b. 

Female, — Head subcylindrical, narrow, moderately 
short, and, with part of the neck, buried in the 
tissue of the host in the neighbourhood of the dorsal 
fin. Neck tolerably long and narrow. Genital seg- 
ment stout, suborbicular, or pyriform, somewhat flat- 
tened dorsally, and provided posteriorly with two 
bundles of slender digitiform filaments, one on each 
side of the abdomen. Abdomen small, obscurely tri- 
lobed. Egg-strings tolerably elongated, containing 
numerous small ova, and springing from the base 
of the abdomen, inside the digitiform appendages. 

The specimens examined by us were of a reddish 

* The description of the head of Rehelula given in the definition of the 
genus is taken from complete specimens found on macrurid fishes captured 
in the Bay of Bengal and in the Malay Archipelago, and representing two 
apparently distinct species of this curious genus of parasites. 



UEBELULA EDWAliDSI. 1 08 

colour, but they had been for some time in a preserva- 
tive fluid. The smaller of two specimens measured 
from the posterior end of the neck to the extremity of 
the digitiform processes about 12 mm., the other was 
about twice that size. 

Habitat. — Parasitic on species of Macruridae. One 
on a Macrurns {Lepidoleprus) coelorhipichus Risso, cap- 
captured off the south-west of Ireland by H.M.S. 
' Research ' at Station I, depth 200 fathoms, 10th July 
1899 {A. M. Norman).^ One in a gathering collected 
by means of a small trawl, at a depth of 1448 metres 
in lat. 58° 43' N., long. 9° 6' W., 23rd August 1910 
(Fishery Steamer' Goldseeker'). Host doubtful: this 
specimen appears, in some way, to have become 
detached from its host. In both specimens the head 
was Avanting. 

The following measurements are from the specimen 
from the west of Scotland : — 

Length of neck . . . .12 mm. 

Length of genital segment . . 15 ,, 

Width of genital segment at widest 

part 5 „ 

Length of posterior appendages . 9 ,, 

Length of egg- strings . . about 52 ,, 
In this specimen the genital segment is somewhat 
different in shape, and rather more elongated than the 
other, but otherwise they are similar, and appear to 
belong to the same species. 

Brian lias recorded a female Rehelula\ which was found 
adhering to a Macruru^ about 14 inches in lengtli captured 
lat. 45° J 3' N., long. 3° 06' W., at a depth of 358 metres, in 
July 1903. Tliis parasite, which he, with some doubt, 
ascribes to R. edwardt^ii., is not unlike the specimen from the 
south-west of Ireland described by us. He gives a figure of 
the specimen but was unable to show the head, as the anterior 
part was buried in the tissues of the fish; the visible portion 
measured about 12 mm. 

The author remarks: " J^ai cru devoir determiner cette 

* Canon Norman, with his usual kindness, permitted us to make a draw- 
ing of this specimen (see PI. LI, fig. 5). 

t 'Bull, de I'Institut Oceanographique,' No. 110, p. 15 (1908). 



104 BRITISH PARASITIC COPEPODA. 

espece avec quelque doute, n^ayantpn examiner tout le corps, 
qui dans sa portion anterieure est cache et enfonce dans les 
muscles du Macrurus. Les parties visibles du copepode 
comme Tabdomen et ses prolongements dioitiformes montrent, 
cependant, des caracteres specifiques qu^on pent considerer 
comme probablement identiques a ceux de Tespece bien 
connue : R. EdivardsiKoU., qui n'est indiquee jusqu^a present 
que pour la Mediterranee." As we have indicated, there is 
apparently more than one species of Rehehda, but identifica- 
tion is difficult if the specimen is incomplete. 

Genus 35. SPHYRION Cuvier, 1830. 

Syn. Lestes and Lesteira Kroyer. 

Head more or less expanded transversely, small and 
bulb-like but sometimes of considerable dimensions, 
and separated from the genital segment by a smooth 
and slender neck, cylindrical and inoderately elon- 
gated. Genital segment of moderate size, smooth, 
ovate, subglobose, or bulbiform, and furnished pos- 
teriorly with two tolerably large bunches of vesicles 
resembling clusters of grapes, one cluster being on 
each side of the abdominal lobe. Antennules rudi- 
nlentar3^ Thoracic limbs apparently suppressed. 
Egg-strings straight, tolerably elongated. Colour 
pale red. Male unknown. 

The original definition of the genus by Cuvier is sufficiently 
correct, with the exception of his description of the posterior 
appendages which he calls faiscean de poils. The Rev. 
T. E. R. Stebbing remarks that these grape-like clusters 
" are appended to the genital segment probably with a 
branchial function." "^ 

1. Sphyrion lumpi (Kroyer). 
(Plate LI, figs. 3, 4.) 

1845. Lestes lumpi Kroyer. Danmarks Fisk, vol. ii. p. 217. 

1863. Lesteira lumpi Kroyer. (71) p. 325, pi. xviii. fig. 5, a-g. 

1869. Lesteira lumpi Steenstrup. (126a) p. 182, pi. ii, fig. 5. 

1899. Sphyrion lumpi Bassett-Smitli. (8) p. 489. 

1901. Sphyrion lumpi T. Scott. (113) p. 128, pi. vii, fig. 13. 

Female, — Cephalothorax rather small and scarcely 

* ' Marine Investigations of South Africa/ Crustacea, pt. 1, p. 60. 



SPHYRION LUMPI. 165 

equal to a quarter of the entire length of the animal, 
somewhat cordiform or crescent-shaped, and consisting 
of two lateral projections directed slightly forward 
and expanded into rounded wing-like processes — one 
on each side of the small cephalon or head. A neck, 
tolerably elongated, slender and cylindrical, joining 
the genital segment to the anterior end of the body. 
Genital segment of moderate size, broadly ovate or 
obscurely heart-shaped and slightly flattened; the 
width, which is nearly the same as, or which slightly 
exceeds, the length, equal to fully one and a half times 
the Avidth of the cephalothorax ; its lateral margins 
boldly rounded and the posterior margin nearly 
straight; a small median lobe on the posterior margin 
representing the terminal part of the abdomen, and 
on either side of it a tolerably large cluster of vesicles 
resembling a bunch of grapes, each cluster fully half 
the size of the genital segment. Egg-strings straight 
and moderately elongated. Length of the animal ex- 
clusive of the egg-strings about 40 mm. 

The following measurements are taken from a 
fairly perfect specimen : — Length of head, 6 mm. ; 
of neck, 15"5 mm. ; of genital segment, about 12 mm. ; 
appendages of genital segment, 7' 5 mm. 

Antennules small and feeble. Thoracic limbs appa- 
rently obsolete, mouth-organs rudimentary. 

The male has not been observed. 

Habitat. — An imperfect specimen Avas taken from a 
lumpsucker {G y clop tern. s lumpus) captured in the nets 
of the salmon fishers at the Bay of Nigg near Aberdeen 
in April, 1900; and a fine, nearly perfect specimen 
was presented to one of the authors by Mr. Irvine of 
Aberdeen who obtained it on one of a number of cat- 
fislies {Ana;vrMclias lujms) landed at the Aberdeen Fish- 
Market but captured in the North Sea, probably out- 
side the Scottish area, by a Norwegian Trawler. It 
is from this specimen that the description given above 
was prepared. The Bay of Nigg specimen was minus the 
head, probably owing to the fish having been roughly 



166 BRITISH PARASITIC COPEPODA. 

handled; it also appeared to be somewhat immature, 
as the appendages of the genital segment were not 
fullj developed; they resembled those of a young 
female represented by figure 5c on plate xviii of 
Kroyer's work, *Bidrag til Kundskab om Snyltekreb- 
sene ' (1863). Kroyer's specimens of Sphyrion lumpi 
were found fixed on the tail of a Ci/cJopterus lumpus 
from Iceland. 

A species from the southern hemisphere, Sphyrion Isevi- 
gatum Guerin-Meneville, described by G. M. Thomson, New- 
Zealand, and also recorded from South African waters 
by the Rev. T. R. R. Stebbing, exhibits a remarkable 
difference in the form and dimensions of the cephalothorax, 
which, unlike that of Kroyer's species, is distinctly larger 
than the genital segment, while its shape, instead of being 
regular, is more or less distorted. 

The cephalothorax of Sphyrion, like that of Lernsea, 
Hsemohaphe>i, and other members of the family LernaBidaB, 
penetrates, and is wholly enveloped by, the tissues of the 
host. 

The nauplius of Sphyrion, ns shown by Kroyer, is a free- 
swimming organism somewhat similar to that of Lemma 
hranchialis. 

Dr. Bassett- Smith records, but somewhat doubtfully, the 
occurrence of Sphyrion lumpi at Dun gen ess. (Cf . ' A syste- 
matic Description of Parasitic Copepods found on Fishes,' 
p. 488.) 



Family vr. CHONDRACANTHiDiK. 

Female. — Body usually more or less incompletely 
segmented, sometimes furnished with outgrowths in 
the form of lobes or prolongations, and frequently 
with the head Avholly or partially immersed in the 
tissues of the host. Antennules short, two- or three- 
jointed, or rudimentary. Antennae small and armed 
with simple but sometimes powerful terminal hooks. 
Mandibles falciform, expanded at the base, and taper- 
ing to the distal end, which is usually more or less 
attenuated and incurved ; furnished with two rows of 
minute prickles, usually marginal, which extend from 



CHONDEACANTHID^. 167 

the base to near the apex. Thoracic legs rudimentary 
and usually in the form of unsegmented lobes. Egg- 
strings two, usually short, and containing numerous 
ova, but sometimes elongated and more or less 
twisted. 

Male. — Very small and adherent on the female by 
means of hooked appendages. Cephalothorax dis- 
tinct; limbs more or less articulated. Abdomen 
segmented. 

In the Chondracanthidae the type of mandible is distinctly 
different from that of the Cjiligida9 or the Lern^opodidse ; the 
biting part is composed of a single moderately elongated 
piece, and is falciform in shape, broad at the base and 
tapering to the pointed and often attenuated distal extremity ; 
these biting parts are articulated to a moderately stout base, 
and curved towards each other at an abrupt or almost a 
right angle, and both margins are fringed from the base 
almost to the apex with minute serratures. This type of 
mandible is rarely met with in species belonging to any of 
the other families described here. 



Genus 36. CHONDRACANTHUS De la Roche, 1811. 

Syn. Chondracanthus and Lementoma Baird. 

Female. — Head usually small, not articulated to the 
thorax, separated by a constriction which may be 
distinct or not clearly defined. Thorax short, mode- 
rately narrow, and indistinctly bisegmented. Genital 
segment proportionally large, slightly flattened, and 
divided usually into two portions by a transverse con- 
striction, more or less distinct, but sometimes indis- 
tinct, and with the postero-lateral corners prolonged 
into narrow processes which may be short or mode- 
rately elongated. Abdomen very small, situated 
between the lateral processes and composed of one 
or two segments. 

Antennules moderately large, more or less con- 
spicuous in front and somewhat rudimentary in 
structure. Antennae very short and armed with 
strong terminal hooks. Mandibles falciform, broad 



168 BRITISH PARASITIC COPEPODA. 

at the proximal end, but tapering to the more or less 
attenuated distal extremity; both margins fringed 
with minute prickles or teeth. First maxillipeds 
small and provided with a straight terminal spine 
which is sometimes serrated. Thoracic limbs two pairs, 
biramose; rami rudimentary. Caudal rami obsolete. 

Male. — Very small. Cephalothorax considerably 
dilated. Abdomen small, more or less segmented. 
Maxillipeds fairly well developed, and fitted for 
grasping; other appendages somewhat rudimentary. 

1. Chondracanthus cornutus (0. F. Miiller). 

(Plate XLVIT, figs. 1,2; Plate LII, fig. 5 ; Plate LIIl, 

figs. 1-9.) 

1777. Lermea comuta O. F. Miiller. (85) p. 124, pi. xxxiii, fig. 1. 

1815. Anops comuta Oken. (91) t. iii. 

1816. Entomoda conmita Lamarck. (72) vol. iii, p. 233. 
1822. Lernentoma comuta Blainville. (24) j). 441. 

1832. Chondracanthus cwnutus Nordmann. (89) p. 111. pi. ix, figs. 
5-10. 

1850. Let-nentoma comuta Baird. (4) p. 328, pi. xxxv, fig. 2. 

1851. Chondracanthus cornutus P. J. van Beneden. (11) p. 108, pi. iv, 
figs. 1-4. 

1863. Chondracanthus cornuius Kroyer. (71) p. 249, pi. xiii, figs. 7 a-8. 

1877. Chondracanthus coi'nutus C. Vogt. (142) p. 78, pi. vi, figs. 4-8. 

1900. Chondracanthus cornutus T. Scott. (112) p. 164, pi. vii, figs. 19- 
31. 

1906. Chondracanthus cornuius Norman & T. Scott. (88) p. 217. 

Female. — Head ovate, somewhat longer than broad ; 
thorax narrower ; genital segment rather wider than 
the head, moderately elongated, flattened, and usually 
with a constriction — sometimes indistinct — dividing it 
into two subequal portions ; the postero-lateral corners 
of the segment produced backwards into straight and 
narrow prolongations, their length being rather less 
than the width of the segment. Abdomen very small, 
bisegmented and rather shorter than the lateral pro- 
longations of the genital segment. 

Antennules robust and tolerably large but showing 
scarcely any structure. Antennae short and armed 
with strong terminal hooks. Mandibles stout, falci- 
form, attenuated towards the distal extremity, and 



CHONDRACANTHUS CORNUTUS. 169 

fringed along both sides with minute bristles. First 
maxillipeds with the basal part moderately stout, but 
the end joint rather slender, and its inner edge 
serrated. Thoracic legs short and very rudimentary. 
Length about 6 mm. 

Male. — The cephalothorax in the male is consider- 
ably enlarged, and apparently unsegmented ; dorsum 
boldly rounded ; abdomen composed of three or four 
segments. Caudal rami small. Antennules conspicuous 
but showing little structure. Second maxillipeds 
armed with small but effective and claw-like terminal 
hooks. Length scaj:'cely 0*5 mm. {^o of an inch). 

Habitat. — Parasitic, usually inside the gill-covers of 
plaice {Pleuronectes jplatessa). The species is tolerably 
frequent, and has been recorded from various parts of 
the British coasts. In the drawing (PI. XLVII, 
fig. 2), a male " m " is seen clinging to the female. 

Some variation has been observed in specimens belonging 
to this species as shown by figs. 1 and 2 in PI. XLVII, some 
being longer and narrower than others and with the thoracic 
legs less developed, but these seemed to be the only differ- 
ences between them. 

2. Chondracanthus annulatus Olsson. 

(Plate XX, fig. 9 ; Plate XLVII, fig. :3 ; Plate LVI, 

figs. 8-10.) 

1868. Chondracanthus annulatus Olsson. (92) p. 30, pi. ii, figs. 13-15. 

1880. Chondracanthus laevirajse Delia Yalle. (141a) Boll. Soc. Adriat. 
Sc. Nat., vol. vi, p. 73. 

1900. Chondracanthus annidatus T. Scott. (112) p. 164, pi. vii, figs. 
46-51. 

1909. Chondracanthus inflatuH Bainbridge. (3) p. 47, pi. ix, figs. 9-15. 

1910. Cho7idr acanthus annulatus Norman & Brady. (87a) p. 156. 

Female. — Body elongated, moderately narrow and 
cylindrical, about four times longer than broad, and 
only slightly flattened. Head small but well defined ; 
the thorax also fairly well marked and the head and 
thorax combined equal to about one-third of the entire 
length of the animal. Genital segment, which con- 
stitutes the other two-thirds, showing a slight constric- 



170 BRITISH PARASITIC COPEPODA. 

tion near the middle, with the postero-lateral corners 
produced into short bluntly-rounded lobes ; a small 
median projection on the dorsal aspect which conceals 
the proximal part of the abdomen. Abdomen, which 
is bisegmented, reaching to about the end of the 
lateral lobes, and imparting a trilobed appearance to 
the posterior end of the genital segment. 

Antennules considerably enlarged but showing very 
little structure, and bearing a few small apical spines. 
Antennae short, stout, and armed with bluntly -pointed 
claws. Mandibles similar to those of Chondracanthus 
cornntus; maxillipeds also somewhat similar to those 
of the same species. Both pairs of thoracic legs short 
and rudimentary. Length about 14 mm. Colour pale 
red. Egg-strings tolerably elongated. 

Male, — The male of this species is proportionately 
larger than that of the one previously described ; the 
cephalothorax also greatly enlarged. Antennules 
short, not very robust, uniarticulate and furnished 
with a few short marginal spines and a fascicle of 
tolerably long apical setas. Antennae short, stout, and 
armed with short but strong terminal hooks. Second 
maxillipeds strong, biarticulated, and provided with 
terminal claws. Other thoracic appendages rudi- 
mentary. Abdomen composed of three or four seg- 
ments, but one or two are not very clearly defined. 
Length about 3 mm. 

Habitat. — Parasitic on the gills of large blue (or 
grey) skates {Baia haMs), captured in the North Sea 
and landed at the Aberdeen Fish-Market {T. Scott). 
North Shields {Miss M. Lehonr)* 

3. Chondracanthus soleae Kroyer. 
(Plate XLVII, fig. 4; Plate LIII, figs. 12-15.) 

1838. Chondracanthus solese Kroyer. (70) vol. ii, p. 139, pi. iii, fig. 4. 
1863. Choyidracanthus solese idem. (71) p. 256. 

1900. Chondracanthus solese. T. Scott. (112) p. 165, pi. vii, figs. 41-45. 
1906. Chondracanthus solese Norman & T. Scott. (88) p. 217. 

* See Norman & Brady's 'Crustacea of Northumberland and Durham,' 
p. 156 (1909). 



CHONDRACANTHUS SOLE^. 171 

Female. — Body tolerably robust ; head subtrian- 
gular, narrow in front, widest behind, its length 
scarcely equal to the width. Thorax short, composed 
of two fairly-distinct segments ; the head and thorax 
combined, equal to rather more than half the length of 
the genital segment ; this segment robust, somewhat 
flattened and divided by a distinct transverse constric- 
tion into two nearly equal portions ; the postero- 
lateral corners of the distal portion produced back- 
wards into straight and narrow prolongations the 
length of which is more than half the width of the 
segment. Abdomen very small, biarticulated, and 
considerably shorter than the lateral prolongations of 
the genital segment. 

Antennules uniarticulate, conspicuous, but not very 
robust, and furnished with a few minute subapical 
setge. Antennge short, stout, and ending in short 
claw-like spines. Mandibles and other mouth-appen- 
dages somewhat similar to those of Glwndra canthus 
cornutus. Both pairs of thoracic legs tolerably elon- 
gated, but rudimentary in structure. Length about 
8 mm. Egg-strings about the same length. 

Mrde. — The male of this species is very small and 
similar to that of OhondracaiitJuis cornutn.'^ ; the appen- 
dages are also similar except that the antennules have 
the basal joint considerably enlarged and subtrian- 
gular, while the end joint is very small and bears a 
few minute setae. 

Habitat. — Parasitic on the gills of the black sole 
(Solea vvlgaris), and only occasionally on other flat 
fishes. Starcross, Devon, on a dab (C. Parker m Mus. 
Normanianum). On Solea vulgaris, Plymouth (Bassetf- 
Smith). Firth of Clyde {T. Scoff). Irish Sea (A. Scoff). 

4. Chondracanthus flurae Kroyer. 
(Plate XL VII, fig. 5; Plate LIII, figs. 10, 11.) 

1863. Chondracanthus flurn> Kroyer. (71) p. 249, pi. xiii. figs. 7 a-d. 
1900. Chondracanthus flune T. Scott. (112) p. 166, pi. vii, tigs. 32-^4. 

Female. — The head and thoracic segments narrow 



172 BRITISH PARASITIC COPEPODA. 

and not so clearly defined as in Ghoudrncauthus solede, 
and together scarcely equal to a third of the entire 
length of the animal. Glenital segment robust, some- 
what flattened, and about twice as wide as the thorax ; 
a distinct transverse constriction dividing it into two 
subequal portions ; the general outline of the segment 
broadly oval, its extreme length being about one 
and a half times greater than the width ; the postero- 
lateral corners of the distal portion produced into 
moderately short, narrow, arcuate, and slightly con- 
vergent lobes, enclosing between them the small 
biarticulated abdomen. 

Antennules moderately robust and indistinctly bi- 
articulated, with one or two subapical setae. Antennas 
short, and armed with small but stout terminal claws. 
Mandibles and other mouth-appendages somewhat 
similar to those of Chondr acanthus cornutu^. Thoracic 
legs rather small and rudimentary. Length about 
5 mm. Egg-strings about the same length. 

Male. — Very small, and somewhat similar in size 
and structure to the male of Ghondracaufhus cornutus. 

Habitat. — Parasitic on the gills and inside the gill- 
covers of long rough dabs {Drepanopsetta platessoides 
Fabr. = Platessa liuuindoides (Bl.)). Firth of Clyde 
{T. Scott). Irish Sea {A. Scott). 

This species of CJiondracanthus has not been observed on 
any other kind of fish. It was also from long* rough dabs 
that Kroyer obtained his specimens of this parasite. 

5. Chondracanthus depressus T. Scott. 
(Plate XLI, figs. 2-4 ; Plate LVI, figs. 1-7.) 

1905. Chondracanthus depressus T. Scott. (116) p. 114, pi. vi, figs. 7-13. 

Female. — Head subquadrangular, nearly as long 
as broad. The first and second thoracic segments 
subequal, very short, and somewhat narrower than 
the cephalic segment, and that segment combined with 
the thorax not more than about a third of the entire 
length of the animal, the other two-thirds comprising 



CHONDRAOANTHnS DEPEESSUS. 173 

the genital segment. This segment fully one and a 
half times as wide as the cephalon and its length only 
about a fourth greater than the width, a distinct 
transverse constriction dividing the segment into two 
nearly equal parts which are considerably flattened ; 
the postero-lateral prolongations moderately short and 
stout, and blunt at the apex, also distinctly con- 
vergent so that the apices are usually contiguous, and 
sometimes overlap each other. Abdomen very short. 

Antennules short and tolerably robust; simple in 
structure ; the distal extremity, which appears to be 
obscurely jointed, bearing scattered apical spinules. 
AntennsB somewhat similar to those of Cliondr acanthus 
cornutus ; the mandibles and maxillipeds also re- 
sembling those of the same species. Thoracic legs 
short, stout, biramose, both rami rudimentary, their 
surface covered with numerous very minute prickles. 
Length about 5 mm. 

Egg-strings short and stout and containing numerous 
ova. 

This species has a general resemblance to Ghondra- 
ccmthus fluTdB from the long rough dab, but is rather 
more flattened. 

'No males have been observed. 

Habitat. — Parasitic on the gills of the flounder 
{PleuTonectes flesns). Firth of Forth and St. 
Andrews Bay '(T. Scott). Irish Sea (A. Scott). 

This Chondr acanthus differs from the other described 
species by the very short thoracic segment, by the genital 
segment being considerably flattened and of a broadly 
quadriform outline, and by the structure of the thoracic 
legs. 

An apparent variety of the species here described, which 
has been observed parasitic on the same kind of fish, differs 
in being more elongated and rather less flattened ; the 
thoracic legs are larger and more rudimentary, and the 
postero-lateral prolongations of the genital segment are 
scarcely convergent. This form, which has been recorded 
as varietv ohlon<jiis, appears to be rare (see PI. XLI, fig. 4 ; 
and PI. LYI, figs. 5-7). 



174 BRITISH PARASITIC COPEPODA. 

6. Chondracanthus limandae Kroyer. 
(Plate XL VII, fig. 6; Plate LVI, figs. 11, 12.) 

1863. Chondracanthus limandx Kroyer. (71) p. 248, pi. xiv, fig. 2, a-h 
1900. Chondracanthiis Ihnandx T. Scott. (112) p. 167, pi. vii, figs. 
38-40. 

Female. — Body tolerably robust ; head large, length 
equal to about two-thirds of the width ; first thoracic 
segment very short, considerably narrower than the 
head, second much larger, nearly as wide as the 
genital segment, and having on each side on the dorsal 
aspect a slightly-elevated rounded knob the diameter 
of which is equal to a third of the width of the seg- 
ment.* Genital segment broad, not greatly elongated, 
and divided by a distinct transverse constriction 
into two nearly equal portions ; the postero-lateral 
corners of the distal portion are produced into mode- 
rately short stout lobes with bluntly-rounded ends. 
Abdomen short, biarticulate, and nearly filling the 
space between the lateral lobes. 

Antennules well developed, basal joint large, end 
one small but fairly well defined, and bearing a few 
small apical spines. Mandibles tolerably large ; other 
mouth-appendages as in Chondracanilms cornutu.^. 
Thoracic legs stout but not greatly elongated. Length 
about 5 mm. Length of egg-strings about the same. 
Colour opaque-white tinged with red. 

Male. — The male of this species does not differ 
greatly in size or structure from that of Chondra- 
canthus cornutits. The cephalothorax is greatly en- 
larged, but the abdomen is small and composed of 
about three segments. Antennse and maxillipeds 
provided with stout terminal claws fitted for grasping. 
Length about 0*65 mm. 

Habitat. — Parasitic on the gills of the dab {Plenro- 
nectes limanda Linn.). This parasite does not appear 
to be very common, and has only been met with on 
the dab. Kroyer obtained his specimens also on the 

* Kroyer says, — " Annulo secundo duobus tuberculis humeralibus praedito 
magnis.*' ' Bidrag til Kvindskab cm Snyltekrebsene,' p. 256 (1863). 



CHONDRACANTHUS LIMAND^. 175 

same kind of fisli. Firth of Clyde and Moray Firth 
(T. Scott). Irish Sea {A. Scott). 

7. Chondracanthus clavatus Bassett- Smith. 
(Plate XL VII, fig. 7; Plate LVI, figs. 14, 15.) 

1896, Chondrocanthus clavatus Bassett-Smith. (6) Ann. and Mag. 
Nat. Hist. (6), vol. 18, p. 13, pi. v, fig. 6. 

1900. Chondracanthus clavatus T. Scott. (112) p. 165, pi. vii, figs. 
35-37. 

1901. Chondracanthus clavatus A. Scott. (107) p. 351. 

Female. — Body claviform, somewhat flattened, 
anterior end narrow, expanded posteriorly. Head 
oval or subcylindrical, rather longer than broad ; a 
median dorsal line, quite distinct in some specimens, 
extending backwards for some distance from the front 
margin. Thorax about as long as the head, but rather 
narrower and not very clearly defined ; the head and 
thorax together equal to about two-fifths of the entire 
length of the animal. Glenital segment clavate, some- 
what flattened, and gradually becoming broader towards 
the posterior end ; postero-lateral prolongations narrow 
and tolerably elongated, about equal in length to half 
the width of the segment, and having a comparatively 
wide space between them. Abdomen very small, indis- 
tinctly biarticulate. 

Antennules short, moderately stout, end joint not 
clearly defined, provided with a few minute apical 
spines. Mandibles and other mouth - appendages 
somewhat similar to those of Ghondracantlms cornatus. 
Thoracic legs small and rudimentary. Egg-strings 
moderately elongated. Length 6*5 mm. 

Male. — Very small ; the cephalothorax is propor- 
tionally greatly enlarged, and the abdomen short and 
segmented. The antennae are armed with short, stout, 
terminal hooks, and the maxillipeds are also similarly 
armed. 

Habitat. — Parasitic on the gills of lemon-soles 
(Pleiironectes microcephalus). Plymouth {Basseft- 
Smith). Firths of Forth and Clyde (71 Scott). Irish 
Sea (A. Scott). 



176 BRITISH PARASITIC COPEPODA. 

8. Chondracanthus nodosus (0. F. Mliller). 
(Plate LII, figs. 1-3 ; Plate XLVIII, figs. 13-10.) 

1777. Lemwa nodosa O. F. Mliller. (85) p. 40, pi. xxxiii, fig. 5. 
1816. Lermea nodosa Lamarck. (72) p. 231. 
1822, Lernentoma nodosa Blaiiiville. (24) p. 125. 
1838. Cliondr acanthus nodosus Krojer. (70) p. 133, pi. iii, fig. 2. 
1840. Clwndracanthus nodosus M. Edwards. (43) p. 503. 
1863. Clwndracanthus nodosus Kriiyer. (71) p. 258. 
1909. Chondracanthus wilUamsoni T. Scott. (118) p. 76, pi. iii. figs. 
8-17. 

Femole. — Body flattened, and of an ovate form when 
seen from above ; cephalon subquadrangular but with 
a shallow rounded projection on each side. A slight 
constriction separating the cephalon from the thorax. 
The genital segment, which is coalescent with the 
thorax, broadly ovate and flattened, and having the 
lateral margins coarsely crenulated or lobate, with six 
rounded but somewhat irregular projections on each 
side, the second and the last three being more promi- 
nent than the others ; the posterior end terminating in 
a narrow median lobe. Abdomen much reduced in 
size, and of a rounded form. 

Antennules small, uniarticulate, and rather rudi- 
mentary in structure. Antennae short and armed with 
strongly-hooked terminal claws. Mandibles similar 
to those of Chondracanthus cornutiis ; both pairs of 
maxillipeds small ; the first pair very similar to those 
of the species named, being composed of a stout basal 
part and a straight spine-like claw finely serrated on 
the inner edge ; the second pair more elongated and 
narrow, and each armed with a short, stout, terminal 
claw and a small rounded process. Both pairs of 
thoracic legs rudimentary and nearly similar in 
structure ; each limb consisting of a short and broad 
basal part, gibbous on both sides, which bears a small 
oblong distal process separated from the basal part by 
a narrow constriction. Egg-strings tolerably elongate 
and stout, and containing many ova. Length about 
7*5 mm. Colour opaque white with a tinge of red. 

Plate LII, fig. 2, represents a young female 



CHONDRACANTHUS NODOSUS. 177 

which, though resembling the adult in having the 
lateral margins coarsely crenulated, differs in being 
proportionally narrower and in the more quad- 
rangular form of the cephalon. 

Male. — The male is very small, the length being 
rather more than half a millimetre. The cephalo- 
thorax is considerably enlarged, and the abdomen 
short and indistinctly segmented. In size and struc- 
ture the male closely resembles the male of Chondra- 
canthus cornukis except that the structure of the 
thoracic part is slightly different. 

Habitat. — Parasitic on Sebastes norvegicus ; usually 
found fixed in the angles formed by the junction of the 
gill-covers with the gill-arches. On specimens of 
Sebastes captured in the Xorth Sea and landed at the 
Fish-Market, Aberdeen, in February 1908 {T. Scott). 

9. Chondracanthus zei De la Roche. 
(Plate XLI, fig. 5; Plate LVI, fig. 13.) 

1811. Cliondracantlms zei De ]a Roche. (41) p. 270, pi. ii, fig. 2. 
1822. Chondracanthus delarochiaua Blainv. (24) p. 422, pi. xxvi, fig. 

1829-1843. Chondracanthus zei Guerin-Meneville. (55) t. 9, fig. 9. 

1831. Chondracanthus zei Burmeister. (26) p. 325. 

1832. Chondracanthus tuberculatiis Nordmann. (89) p. 118. 

1850. Chondracanthus zei Baird. (4) p. 327, pi. xxxv, fig. 1. 

1851. Chondracanthus zei P. J. van Beneden. (11) p. 110, pi. iv, figs. 5-7. 
1892. Chondracanthus zei T. Scott. lOtli Ann. Report Fishery Board 

for Scotland, pt. iii, p. 262. 

1896. Chondracanthus zei Bassett-Smith. (7) p. 162. 

1900. Chondracanthus zeus T. Scott. (112) p. 167, pi. viii, fig. 1. 

1906. Chondracanthus zei Norman & T. Scott. (88) p. 217. 

Female. — Body short, moderately stout, and of a 
somewhat ovate form. Head small, orbicular ; thorax 
very short, narrowed behind the head where it forms 
a kind of neck. The genital segment, with which the 
thorax appears to be more or less coalescent, flattened 
and becoming gradually and considerably wider pos- 
teriorly. Abdomen small and rudimentary. A short 
horn-like process extends outwards from both sides of 
the thoracic segment and there is a trifid process on 

VOL. I. 12 



178 



BIUTISH PAEASITIC COPEPODA. 



its dorsal aspect. Grenital segment furnished with two 
pairs of tridigitate appendages on the under side and 
with three pairs of somewhat similar digitiform pro- 
longations on the lateral aspect, the posterior prolon- 
gations being the larger; these latter appendages 
surround and nearly conceal the short, stout, and 
flattened ovisacs ; several median processes also occur 
on the dorsal aspect of the genital segment. 

Antennules short and moderately expanded. An- 
tennse prehensile and provided with stout terminal 
hooks. Mandibles and other mouth - appendages 




a h 

One of the ovisacs of Chondr acanthus zei. 

similar to those of Chondr acanthus cornutus. Thoracic 
legs indistinctly biramose, and their structure some- 
what rudimentary. Ovisacs, which are short and con- 
tain numerous small ova, compressed laterally so as to 
have a broadly and somewhat obliquely ovate outline ; 
not extending beyond the ends of the prolongations 
described above. Colour white tinged with red. 
Length about 12 mm., but the size somewhat variable. 

Male. — The male of this species does not appear to 
differ greatly from that of Ghondracanthus cornutus in 
size and structure. 

Habitat. — Parasitic on the gills and gill-arches of 
the dory (Zeus faher). Polperro, Cornwall {A. M. 
Normaii). Plymouth (Bassett- Smith). Firths of Forth 



CHONDRACANTHUS ZEI. 179 

and Clyde {1\ Scott). Irish Sea {A. Scott). This 
parasite is of frequent occurrence on the dory, and 
Dr. Bassett- Smith describes it as very common on the 
gills of Zeus faher at Plymouth. 

The ovisacs of Ghondracanthiis zei differ so remarkably 
from those common to most of the species known to us that 
we give here a figure showing a side and an end view, 
enlarged, of one of them. When the parasite is viewed from 
the ventral aspect the ovisacs appear as in figure 6, but the 
side view a can best be seen whem the ovisacs are removed. 
X shows the point of attachment of the ovisac to the genital 
segment. 

10. Chondracanthus lophii Johnston. 
(Plate LIT, fig. 4 ; Plate LVI, figs. 16-18.) 

1836. Chondracanthus lophii Johnston. (67) p. 81, fig. 16. 
1837-1838. Chondracanthus gibbosus Kroyer. (70) (1) p. 252, pi. ii, 
fig. 4. 

1843. Chondracanthus lophii Rathke. (100) p. 116, pi. v, figs. 16-18. 
1847. Chondracanthus gibbosus Thompson. (129) p. 248. 

1850. Lernentoma lophii Baird. (4) p. 330, pi. xxxv, fig. 3. 

1851. Chondracanthus gibbosus P. J. van Beneden. (11) p. 104, pi. ill, 
figs. 10-15. 

1862. Chondracanthus lophii Turner & Wilson. (140) p. 67, pi. iii. 
1877. Chondracanthus gibbosus Yogt. (142) p. 76, pi. v, figs. 1-4 ; pi. 
vi, figs. 1-3. 

1896. Chondracanthus lophii Bassett-Smitli. (7) p. 162. 
1900. Chcytidracanthus lophii T. Scott. (112) p. 167. 

Female, — Body tolerably elongated, narrow in front, 
posterior end moderately broad and flattened. Head 
small, having on each side a horn-shaped appendage 
directed obliquely backwards. Thorax narrow and 
furnished wath one or two short, paired, marginal 
processes directed outwards, and a pair of bifurcated 
moderately long ventral appendages. Grenital seg- 
ment flattened, and about twice as long as broad, 
constricted in the middle so as to form two subequal 
portions, the postero-lateral corners of both of which 
form short prolongations ; this segment also furnished 
with a row of median digitiform processes alternately 
larger and smaller, and directed obliquely backwards. 
Abdomen very short. Egg-strings greatly elongated 
and more or less twisted into loops. 



180 BRITISH PARASITIC COPEPODA. 

Antennules short, conspicuous in front, their basal 
part considerably expanded, but the distal end narrow, 
and minutely setiferous. Antennae short and stout, 
and armed with strono- terminal hook-like claws. 
Mandibles and other mouth-organs somewhat similar to 
those of Ghondracanthus cormitns. Thoracic legs in- 
distinctly biramose, and more or less rudimentary in 
structure. Length about 12 mm. 

Male. — The male of this species, like that of the 
last, has a close resemblance to the male of Ghondra- 
canthus cornutus as indicated by Dr. Baird. It is of a 
pyriform shape, considerably swollen in front but 
tapering towards the distal end, and furnished w^ith 
strongly uncinate foot-jaws. It is usually found 
clino-ino^ to the female, but from its minute size mav 
easily be missed. 

Habitat. — Parasitic usually on inner surface of the 
gill-pouches of angler-fishes {Lophius piscatorius Linn.). 
It is a moderately common species on the angler-fish 
and has been recorded from various parts of the 
British coast. Dublin, 1839; Belfast, 1841 (ir. 
Thompson). Coasts of Devon and Cornwall (G. Parker, 
A. M. Norman, and others). Irish Sea, and various 
parts of the Scottish coasts {T. ^' A. Scott). For a 
detailed account of the anatomical structure of this 
curious species see Turner and Wilson's memoir referred 
to above. The colour of this Ghondracanthus is usually 
opaque white. The egg-strings are of great length, 
usually in screw-form, very slender, and contain 
numerous small ova. Specimens ma}^ often be found 
in considerable numbers in the gill-pouches of both 
large and small anglers. 

11. Ghondracanthus merluccii (Holten). 

(Plate XX, fi^. 10 ; Plate XL VII, fig. 8 ; Plate LIII, 
figs. 16-19.) 

1802. Lernipa merluccii Holten. (63) vol. v, pi. iii. fig. 2. 

1837. Ghondracanthus merluccii Kroyer. (70) p. 278, pi. iii, fig. 9. 

1840. Ghondracanthus merluccii M. Edwards. (43) vol. iii, p. 503. 



CHONDRACANTHUS MERLUCCII. 181 

1892. Chondracanthus merluccii T, Scott. lOth Ann. Rept. Fishery 
Board for Scotland, p. 262. 

1896. Chondracanthus merluccii Bassett-Smith. (7) p. 161. 
1900. Chondracanthus merluccii T. Scott. (112) p. 166. 
1906. Chondracanthus merlucii Brian. (21) p. 97, pi. vii, fig. 4. 
1906. Chondracanthus merluccii Norman & T. Scott. (88) p. 217. 

Female. — Body oblong and flattened. Head some- 
what wedge-shaped, wider behind than in front, the 
front margin rounded, and a more or less distinct 
median line extending from it backwards ; the head 
seen in profile sloping upward and backward, and 
becoming distinctly gibbous behind; provided with 
small, lateral horns — one on each side near the postero- 
lateral corners — Avhicli extend obliquely backwards 
like the barbs on a fishing hook. Thorax short, 
indistinctly bisegmented ; a constriction separating 
the head from the thorax and another separating 
the thorax from the genital segment ; the second seg- 
ment of the thorax wider than the first and nearly as 
w4de as the genital segment; its anterior lateral 
corners produced forwards into small round knobs. 
Genital segment moderately thick, and flattened 
dorsally, a slight constriction dividing it into tw^o 
nearly equal portions ; a pair of tolerably stout and 
elongated processes springing from the ventral aspect 
of the segment just in front of the constriction, and 
extending obliquely backwards so as to reach beyond 
its posterior end; the postero-lateral corners of the 
genital segment also produced backward into elongated 
and moderately stout horn-hke prolongations. Abdo- 
men very small. 

The two pairs of antennae and the mouth-appendages 
similar to those of Oliondrocanthus cormitus; so are also 
the thoracic legs, except that the second pair are con- 
siderably larger than the first. Length to the end of the 
posterior processes about 12 mm. These processes 
about three millimetres in length, being rather shorter 
than the ventral prolongations, which measure about 
four millimetres. Egg-strings about ten millimetres in 
length, moderately thick, and containing numerous ova. 



182 BRITISH PARASITIC COPEPODA. 

Male. — The male of this species is very small, being 
scarcely more than half a millimetre in length ; the 
cephalothorax, as in the male of Chondr acanthus 
eornutiis, is proportionally large ; the abdomen is small 
and composed of two or three segments. Sometimes 
more than one male may be found adhering to a female. 

Habitat. — Parasitic on the hake {Merlucchis vulgaris 
Cuv.). Usually found clinging i^o the roof and sides, 
sometimes on the under side of the tongue and on the 
inside of the gill-covers. Frequently the head is either 
buried in the tissues of the fish or enveloped in mucus. 
This is a tolerably common parasite of the hake, and a 
considerable number of specimens may sometimes be 
obtained from a single fish. Coasts of Devon and 
Cornwall (C. Parker, Lancjlirin, Bassett-Smlth). Irish 
Sea (A. Scott). Firths of Forth and Clyde, Aberdeen, 
&c. {T. Scott). 

12. Chondracanthus ornatus T. Scott. 
(Plate XLI, fig. 6; Plate XLV, fig. 7.) 

1900. Chondracanthus ornatus T. Scott. (112) p. 168 (description 
only). 

1901. Chondracanthus ornatus idem. (113) p. 129, pi. vii, fig. 14 ( ? ). 

1902. Chondracanthus ornatus idem. (114) p. 298, pi. xiii, fig. 34 ( (^). 

Female. — When viewed from above with a general 
outline closely similar to tliat of an equilateral triaugle, 
the bluntly-rounded head forming the apex, and the 
truncated posterior end the base; the front of the 
head indistinctly trilobed, one bluntly-rounded lobe 
being in the centre, and projecting slightly in front of 
the two lateral lobes wdiich are also bhmtly rounded. 
The neck connecting the head w4th the thorax very 
short. Three or four, more or less distinct, tubercles 
along each side of the thorax (forming the sides of the 
triangle), a series of three similar tubercles extending 
along the middle of the dorsum ; the posterior tubercle 
of the middle series standing well up, but each of the 
other two standing at a slightly low^er elevation than 
the one immediately behind. The arrangement and 



CHONDRACANTHUS ORNATUS. 183 

position of these lateral tubercles seems to vary 
sliglitlj in different specimens. Abdomen exceedingly 
small and inconspicuous. Egg-strings of moderate 
length and stoutness, like those of GJiondracanthus 
limandse. 

The size of the more typical of the female specimens 
observed is as follows : — from the forehead to the 
posterior end of the body 5 mm., from the forehead 
to the end of the egg-strings 11*5 mm., the width of the 
genito-thoracic segment at the posterior end about 
equal to the length, so that, as already stated, the 
body of the animal has a general outline closely 
similar to that of an equilateral triangle. 

Male, — The male is very small, measuring scarcely 
half a millimetre in length ; it is of a robust form 
and does not differ very much from the male of 
Ghondr acanthus comutus. 

Habitat. — Parasitic on the gills of spotted dragonets 
(GaUionymus maculatus Bonap.). Firth of Clyde, 1899 
and 1901 ; Moray Firth, 1900. 

Though Chondracanthiis ornatus is of a tolerably large size, 
it is so well concealed that it easily escapes notice, and it is 
only by turning back the gill-covers that the observer can 
be sure whether the parasite is present or not. Hitherto 
this Chondracanthus has only been obtained on the spotted 
dragonet, and frequently it has been found associated with 
another species — the Hsemohaphoides ambiguus (T. Scott), pre- 
viously described. As pointed out in the description of Hsemo- 
haphoides, the two species sometimes occurred on the same 
fish, on opposite sides or on the same side ; occasionally also 
a Chooidr acanthus would be found on one side and a ffsemo- 
haphoides on the other. The Chondr acanthus was usually of 
a purplish colour tinged with brown. About 14 per cent, of 
the fishes examined were found to harbour the Chondra- 
canthiis. 

Genus 37. MEDESICASTE Kroner, 1863. 

Sjn. Clwndracantlius Aiict. ; Leimentoma Baird ; Oralien Bassett-Smith. 

This genus was established for the reception of a 
GJiondracanthus which, though agreeing in some 



184 BRITISH PARASITIC COPEPODA. 

respects with that genus, yet differed because of the 
remarkable arrangement of the cephalic appendages. 
In typical Ghondr acanthus these appendages are 
situated in more or less proximity to each other, but 
in Medesicaste the anterior portion of the cephalo- 
thorax, which has a bulb-like form, and is provided 
with two pairs of antennge; is separated from the 
thorax and the thoracic appendages by a moderately 
long and narrow neck : by this arrangement a con- 
siderable distance intervenes between the two pairs of 
antennae and the mouth and other thoracic organs. 
The posterior part of the body is tolerably large and 
robust, its lateral margins are deeply incised, dividing 
it into two portions : the thoracic, which carries on 
the ventral aspect two pairs of bilobed blunt-pointed 
processes and other appendages; and the genital, 
which is rounded and rather larger than the other, and 
has its postero-lateral corners produced into more or 
less angular lobes. Abdomen small, biarticulated, and 
more or less inclosed between the lateral lobes of the 
genital segment. Egg-strings elongated, claviform. 

Male very small, similar to the male of Cliondra- 
can fines. 

We are unable to find any valid difference between Medesi- 
cas'te Kroyer and Oralien Bassett- Smith. 

'J'he only British species of Medesicaste is that described 
below. 

1. Medesicaste asellinum (Linn.). 
(Plate LII, fig. 6; Plate LVII, figs. 18, 19.) 

1761. Lernsea asellina Linn. Fauna Snec, 2101 ; Syst. Nat., Edit. 12. 
vol. i. pt. ii, p. 1093 (1767). 

1829-1843. ChondracantJms tnglse. Giiei-in-Meneville. (55) Incono- 
grapli., pi. ix, fig. 8. 

1822. Leimentoma h-iglse Blainv. (24) p. 441, pi. Ixii, fig. 12, 

1832. Chondrocanthus trigJx Nordmann. (89) p. 116, pl. ix, figs. 1-4. 

1838. Chmidr acanthus trigliE Kroyer. (70) p. 135, pl. iii, fig. 3. 

1850. Lernentoma asellina Baird. (4) p. 329, pl. xxxv, fig. 4. 

1863. Medesicaste triglarum Kroyer. (71) p. 312, pl. xviii, fig. 1, a-h. 

1869. Chotidr acanthus triglse Steenstrup. (126a) pl. ii, fig. 3. 

1896. Chondracanthus triglse Bassett-Smith. (6) p. 13, pl. iv, fig. 4. 

1899. Oralien aselUnus idem. (8) p. 490. 



MEDESICASTE i^SELLINUM. 185 

1900. Oralien aselUnus T. Scott. (112) p. 163, pi. vii, figs. 16-18. 
1906. Medesicaste triglaruni Brian. (21) p. 94. 

Female. — Head small, subglobose, situated at the 
end of a tolerably long and slender neck, rounded in 
front, and with each side expanded into rounded lobes, 
the whole forming a bulb-like extremity usually in- 
closed in the tissues of the host ; this anterior part 
furnished with two pairs of antennae, which are 
small and have hook-like extremities. The neck, 
where it joins the posterior portion of the body, 
slightly enlarged. This posterior portion of the body 
tolerably large and robust and divided by a deep 
transverse constriction into two subequal parts ; the 
front part appearing to constitute the thorax with its 
appendages, and the lower the genital segment, the 
antero-lateral corners of which extend outwards into 
angular projections ; the lateral margins of this lower 
segment slightly arcuate, converging distally, and 
forming two small postero-lateral lobes, separated from 
each other by a small median sinus, best seen from 
the ventral aspect. Abdomen very small, occupying 
the space between the postero-lateral lobes. 

Both pairs of antennse small, and situated on the 
subglobose cephalic segment as described above. 
Mandibles stout, falcate, and with their margins 
fringed with numerous small teeth as in Chondra- 
cautlms; maxillse and other mouth-appendages also 
somewhat similar to those in Chondracnnthus. Thoracic 
legs two pairs, very rudimentary. Colour pale red. 
Egg-strings nearly as long as the body and containing 
many ova. Length exclusive of egg-strings about 7 mm. 

Male. — Extremely small and similar in structure to 
the male of Gliondracanfhus. 

Habitat. — Parasitic on the gills of Trigla spp., 
Gadus spp. ; also on the plaice (Pleiironectes platessa) 
and other fishes, captured in British waters, frequent. 

A somewhat variable species, and apparently widely 
distributed. 



18() BRITISH PAKASITIC COPEPODA. 

Family vii. Lern^opodid^:. 

Female. — Body in the adult female usually non- 
segmented or indistinctly or incompletely so. Head 
small, thorax not clearly defined, narrow and more or 
less elongated, but sometimes short as in Lerndeopoda 
cluthde. Abdomen usually separated from the thorax 
by a constriction tolerably distinct but seldom if ever 
forming a complete articulation ; the genital segment 
of the abdomen usually enlarged and sometimes pro- 
vided with posterior processes ; the remaining ab- 
dominal segments usually obsolete or rudimentary. 
Appendages of the cephalon and thorax comprising 
antennules and antennae, mandibles, maxillae, and first 
and second maxillipeds, but the thoracic usually obso- 
lete or entirely wanting. The second maxillipeds arm- 
like appendages elongated or short, and modified to 
form organs of attachment, the arms entirely separated 
except at the tip or partly or completely coalescent ; 
united at the distal extremity to a chitinous process 
Avhicli penetrates the tissues of the host, and the 
character of which may vary in the different species. 
Egg-strings generally only moderately elongated and 
the ova small and arranged in a multiseriate manner. 

Male. — Very small and furnished with several ap- 
pendages the structure of which is more or less rudi- 
mentary. Its form varies in the different genera, 
and it is usually found adhering to some part of the 
female — i.e., the rudimentary abdominal segment or 
the cephalothorax. 

Milne Edwards included AncJiorella [Clavella] in his Family 
Lerneopodiens, but Dr. Baird removed this genus and estab- 
lished the Fami^ly Anchorellidae for its reception. 'Jlie dif- 
ference between LerndBopnda and Anchorella is obvious in two 
particulars — in the structure and modification of the second 
maxillipeds and in the form of the pygmy male. The first 
provided Baird with the principal character by which the 
two Families could be distinguished, and might have been 
considered valid except for the occurrence of genera inter- 
mediate in structure, which made it more convenient to include 
them all under the one Family Lernaeopodidse. 



THYSANOTE. 187 

Genus 38. THYSANOTE Kroyer, 1863. 

Sjn. Brachiella Nordmann (in part). 

Ceplialo thorax short, robust ; genital segment short, 
dilated, and somewhat flattened, the postero-lateral 
corners each provided with two moderately short pro- 
cesses of about equal length ; another pair of rather 
longer submedian appendages springing from the 
ventral aspect and a little in front of the posterior 
margin. Abdomen obsolete or nearly so. 

Both pairs of antennge small and rudimentary. 
Mandibles moderately elongated, and having the inner 
margin near the distal end coarsely and irregularly 
serrate. Maxillae and first maxillipeds somewhat 
similar to those of Brachiella. Second maxillipeds 
free, each with the extremity expanded and obscurely 
trilobed, the tip of the inner lobe of each maxilliped 
being apparently united together. 

Thysanote has a close general resemblance to Brachiella, 
both in the form of the body and in the free second 
maxillipeds ; the structure however of these maxillipeds 
differs, and the genital segment also differs in being fur- 
nished with two pairs of posterior appendages. 

1. Thysanote impudica (Nordmann). 
(Plate XXXIII, fig. 5; Plate XLIX, figs. 8-11.) 

1832. Brachiella impudica Nordmann. (89) vol. ii, p. 92, pi. viii, fig. 1. 

1840. Brachiella impvdica M. Edwards. (43) p. 513. 

1877. Brachiella impudica Yogt. (142) vol. xvi, p. 436. 

1899. Thysanote impudica Bassett-Smith. (8) p. 162. 

1900. Thysanote impudica T. Scott. (112) p. 169, pi. viii, figs. 2-5. 

Female. — Cepbalotliorax short, robust, strongly 
arcuate, and alDout as long as the genital segment. 
Genital segment short, about as broad as long, 
flattened and subquadriform, posterior truncated and 
furnished with three pairs of short appendages, one 
pair on each side and one pair submedian. 

Both pairs of antennjB somewhat rudimentary. Man- 
dibles moderately stout at the distal end, and having 
the inner margin coarsely and irregularly serrate, 



188 BRITISH PARASITIC COPEPODA. 

some of the teetli being alternately larger and smaller. 
Maxillae moderately stout, fnrnislied with two mode- 
rately long, narrow, and spiniferoiis apical processes, 
two small spinules also springing from a siibapical 
rudimentary lobe. The first maxillipeds very short, 
stout, and strongly uncinate. The second maxillipeds 
very short, with the distal end expanded and united 
at the top. Length from the forehead to the end of 
the longer posterior appendages about 6*5 mm. 

Male. — We have not ourselves seen the male of 
Thysanote impudica ; Nordmann gives a figure of it 
and describes it as very minute (about 0*6 mm.). It 
somewhat resembles the male of Brachiella, and its 
appendages also appear to be somewhat similar to 
those of that genus. 

Habitat. — Parasitic on the gills of Trufla sp. and 
Gadus sp. On the gills of sapphirine gurnard {Trigla 
hirundo) captured at Polperro and Starcross {A. M. 
Norman). Plymouth {Bassett- Smith). Irish Sea {A. 
Scott). Moray Firth {T. Scott). Dr. Bassett-Smith 
records this species also from Trigla cucidus and gur- 
nardus ; and M. Nordmann from Gadu.s xglefinus (the 
haddock). 

Genus 39. CHAROPINUS Krdiier, 1863. 

Syn. Lernxa Retzins (in part) ; Lerneopoda Nordmann (in part). 

Female. — Cephalothorax of moderate size, ovate or 
sub-conical, indistinctly segmented, with obscure trans- 
verse constrictions behind the cephalon ; neck short 
or none. Genital segment pyriform or elongate- 
ovate ; abdomen very small or obsolete. Posterior 
appendages two, slender, short, one on each side 
of the abdomen. Antennge short, stout, uncinate. 
Mandibles and first maxillipeds nearly as in Thysanote. 
Second maxillipeds greatly elongated, slender, or 
moderately stout, contiguous or coalescent at the 
apex. Apex enlarged or biramose, and clasping a 
cross bar of chiton, or greatly attenuated and fixed at 



CHAROPINUS. 189 

the tip to a large horizontal chitinic appendage. Egg- 
strings tolerably thick and elongate, and containing 
numerous ova. 

Male. — Very small, somewhat similar in structure 
to the male of Lerncjeojpoda cluthx. 

1. Charopinus dalmanni (Retzius). 
(Plate LV, figs. 1-4; Plate LIV, figs. 12-17.) 

1829. Lemaea dalmanni Retzius. (101a) p. 109, : vol. xxix, p. 6, figs. 5-9, 
(1830). 

1832. Lerneo-poda dalmani Nordmann. (89) p. 138. 
1837. LerniEopoda dalmanni Kroyer. (70) vol. i, p. 264. 

1862, Lerneopoda dalmanni Turner & Wilson. (140) p. 77, pi. iv. 

1863. Charopinus dalmanni Kroyer. (71) p. 280, pi. xiv, figs. 6, a-g. 
1879. Stylopho7'us hypocephalus Hesse. (59) (6) vol. viii, ai-t. 15, p. 31. 
1891. Charopinus dalmanni T. Scott. 9tli Ann. E/epoi*t Fishery Board 

for Scotland, p. 310. 

1900. Charopinus dalmanni idem. (112) p. 169, pi. viii, figs. 6-10. 
1904. Charopinus dalmanni A. Scott. (109) p. 43. 

Female. — Cephalothorax short, somewhat trian- 
gular in outline, and usually bent abruptly downwards 
so as to form a more or less distinct angle with the 
posterior part of the body. There springs from each 
side of the angle thus formed a long, moderately 
slender, and indistinctly annulated appendage which at 
the apex becomes dilated and lunuliform. The lunuli- 
form apices of these elongated appendages, otherwise 
described as the second maxillipeds, though not 
actually coalescent, fit closely together, and clasp a 
cartilaginous or chitinous bar which extends some 
distance on each side of the conjoined apices; this 
complex structure, buried in the tissues of the fish, 
forms a secure anchorage for the parasite. Imme- 
diately in front of the base of each of the second 
maxillipeds there is on each side of the thorax a small 
rounded protuberance termed by Retzius and Kroyer 
" eye-like spots," but they are not supposed to be 
eyes, and their true character seems to be obscure. 
The post erior and genital portion of the body becomes 
gradually and considerably enlarged toward the distal 
end, and the postero-lateral corners form bluntly- 



190 BRITISH PARASITIC COPEPODA. 

rounded knobs. Two moderately long and slender 
appendages spring from the ventral aspect just in 
front of the base of the egg-strings and extend back- 
wards some distance beyond the end of the genital 
segment. Abdomen and caudal rami obsolete. Anten- 
nules stout, short, and apparently three-jointed; 
antennas short with a gibbous basal part, and the 
small end joint provided with a minute terminal claw. 
Mandibles slender, moderately elongated, and with a 
portion of the inner margin of the end joint coarsely 
serrate ; maxillae simple and furnished with a few 
apical setae. No thoracic legs observed. Length 
from the apex of the second maxillipeds to the end of 
the posterior appendages about 47 mm. Colour 
opaque white tinged with red. 

Male. — Very small, scarcely reaching beyond two 
millimetres in length. Somewhat like Lernseopoda 
cliithai in general appearance but larger. Thorax and 
abdomen segmented. Maxillipeds short and stout and 
furnished with strong terminal claws. 

Habitat. — Parasitic in the nasal fossae or spiracles of 
gray (or blue) skates (Baia hatis). Firth of Forth; 
frequent on large Rala hatis brought to the Fish- 
Market at Aberdeen {T. Scott). Firth of Forth, 1862 
{Dr. Wilson). Polperro, Cornwall {A. M. Norman). 
Irish Sea {A. Scott). Usually only one specimen is 
present in a spiracle, but two and sometimes three 
have been observed almost blocking up the spiracle. 

2. Charopinus dubius T. Scott. 
(Plate LV, fig. 5.) 

1900. Charopinus duhkis T. Scott. (112) p. 130, pi. vii, fig. 15. 

Female. — Somewhat resembling Charopinus dalmanni 
Retzius in general appearance and also in some of its 
appendages, but considerably smaller, being little more 
than half the size. 

The principal structural characters by wnich it is 
distinguished from that species are those of the 



CHAKOPINUS DUBIUS. 191 

complex form of the anchoring arrangement of the 
second maxillipeds, which are more slender in propor- 
tion to their length ; their apices appear to be coalesc- 
ent, and are fixed to a moderately large horizontal 
appendage which is cartilaginous and of a brownish 
colour. This appendage, when perfect, is somewhat 
boat-shaped with the extremities slightly turned up ; it 
has a resemblance to the cartilaginous bar clasped by the 
enlarged apices of the second maxillipeds of Charojmms 
dalmaimi, but there the resemblance ends, for in that 
species the ends of the second maxillipeds when freed 
from the cartilaginous bar separate freely, whereas in 
the species under consideration they remain fixed 
together by a small hardened plug as in Lerndeopoch, 
and do not separate freely. The two pairs of antennae, 
the mandibles, and other mouth-organs do not appear to 
differ greatly from those of Gkaropimis dalmanni. The 
ventral appendages at the posterior end of the genital 
segment are also similar. Length of one of the larger 
specimens : maxillipeds 11 mm., cephalothorax 6 mm., 
genital segment 8*5 mm., posterior appendages 4 mm., 
egg-strings 18 mm., and from the chitinic bar to which 
the maxillipeds are fixed to the end of the posterior 
appendages, 26 mm. 

No males of this species have been observed. 

Habitat. — Parasitic on the gills and gill-arches of 
the cuckoo ray (Rata circularis) chiefly, and less fre- 
quently on other skates. Firth of Clyde and Fish-Market, 
Aberdeen {T. Scott), On Raia fiillonica captured off 
Dubh Artach {A. Scott). 

3. Charopinus ramosus Kroyer. 
(Plate LV, figs. 6, 7 ; Plate LIV, figs. 18-22.) 

1863. Charopinus ramosus Kroyer. (71) j). 284, pi. xiv, fig. 5,a-i. 
wot. Charopinus rainosus T. Scott. (113) p. 130, pi. vii, figs. 17-23. 
1904. Charopinus ramosus A. Scott. (109) p. 43. 

Female. — In this species the cephalothorax is not 
deflexed in front of the bases of the second maxilli- 
peds, but is projected forward as in Lernseopoda galei. 



192 BEITISH PARASITIC COPEPODA. 

It has also an ovate or pyriform outline, narrow in 
front. Genital segment elongated, narrow, and only 
slightly enlarged towards the posterior end. 

Antennules short and composed of two joints, a 
stout basal joint and a narrow end one which is 
furnished with a few apical spines, a small bristle also 
springing from near the end of the basal joint. 
Antennae, like those of GharojnniLs duhius, short and 
stout, the outer ramus consisting of a small joint 
articulated to the inner ramus and provided with two 
short unequal spines ; the inner ramus stout with a 
gibbous rounded end, both rami covered with micro- 
scopic prickles. Mandibles narrow, oblong, the apex 
obliquely truncate and coarsely serrated ; maxilla 
small, moderately stout, biarticulate, and furnished 
with three apical and subapical spines and a small 
setiferous process. First maxillipeds three-jointed ; 
second joint with a small setiferous knob near the 
proximal end of the inner margin; end joint slender, 
and armed at the apex with two minute spines and a 
small slightly-curved claw. Second maxillipeds slender 
and about as long as the body, exclusive of the posterior 
appendages, becoming considerably altered at the distal 
end where their inner surfaces are closely adherent, and 
extending on each side into biramose processes which 
penetrate the tissues of the host and form a secure 
anchorage for the parasite. The genital segment 
furnished with two posterior appendages moderately 
short and slightly divaricate. 

Male, — We have not ourselves noticed the male of 
this species, but Kroyer describes and figures it in the 
work quoted above.* It is, though smaller, very like the 
male of Gharojnnus dahnaomi, and is characterized by a 
similar stout proboscis in front of the head and by 
powerful maxillipeds fitted for grasping. ♦ 

Habitat. — Parasitic on the gills and gill-arches of 
Eaia clavata and Baia macidata, Fish-Market, Aber- 

* " Bidrag- til Kvmdskab om Synltekrebsene/' ' Natur. Tidssk./ 3 Raekke, 
2 Bind, p. 287. 



CHAROPINUS RAMOSUS. 193 

deen {T. Scott). Irish Sea (A. Scott). This appears to 
be a moderately rare species in British waters. 



Genus 40. ACHTHERES Nordmann, 1832. 

Female. — Cephalothorax short and broadly ovate, 
separated from the genital segment by a tolerably dis- 
tinct constriction. G-enital segment oval or subglobose, 
and sometimes exhibiting a more or less segmented 
structure. The two pairs of antennae and the mouth- 
appendages not differing much from those of Lernseo- 
'poda ; second maxillipeds short, stout, free, but united 
at the tip to a circular chitinous disk, which is expanded 
in front. Grenital segment usually furnished with two 
small processes at the distal extremity. Egg-strings 
short, dilated or saccate ; ova tolerably large. 

Male. — Very small; first maxillipeds short, very 
robust, strongly uncinate; second maxillipeds elongated, 
stout; terminal claws small. 

Parasitic for the most part on freshwater fishes. 

1. Achtheres percarum Nordmann. 

(Plate L, figs. 6-8 ; Plate LIX, figs. 7, 8; Plate LXIV, 

figs. 1-3.) 

1832. Achtheres percarum Nordmann. (89) p. 63, pi. iv. 
1838. Achtheres percarum Kroyer. (70) ii, p. 143, pi. iii, fig. 6. 
1840. Achtheres percarum M. Edwards. (43) p. 511, pi. xl, fig. 8. 
1861. Achtheres percarum Glaus. (32) p. 287, pis. 23, 24. 
1899. A ether es percarum 'BsL^^Qit-^xniih. (8) p. 498. 
1901. Adheres percarum T. Scott. (113) p. 132. 
1904. Achteres percarum Pelir Gadd. (47) p. 21. 
1906. Achtheres percarum Brian. (21) p. 101. 

Female. — Cephalothorax of moderate size, narrower 
in front than behind. Genital segment sub-globose, 
rather longer than broad, separated from the cephalo- 
thorax by a distinct constriction ; the posterior end of 
the segment slightly angular, bearing two minute pro- 
cesses — one on each side of the median line. Abdomen 
obsolete. Egg-strings short, stout, saccate, scarcely 
equal in length to the genital segment. Ova large. 

VOL. I. 13 



194 BRiTlSlI PAKASITIC COPEl'ODA. 

Antennae and moutli-appendages somewhat similar 
to those of LerndBopoda salmonea. The second maxil- 
lipeds nearly as long as the genital segment, free 
except at the tips where they are united to a small 
obconical chitinous plug. 

"We have not seen the male of Achtheres percarum, 
but it has been described by Nordmann. It is very 
small, measuring little more than a millimetre in 
length. 

Habitat. — Parasitic on the gills of fresh-water perch 
and trout. On the gills of the common trout, Moray 
Firth district (T, Echcard, in 'Museum Normanianum'). 
On the gills of a trout captured by Mr. W. S- Caine 
in Loch Awe, a small fresh-water loch in Assynt, 
Sutherlandshire {T, Scott).* On the gills of a trout 
captured in Loch Tay, Perthshire, by Dr. Williamson, 
to whom we are indebted for the specimen. 

The specimens recorded above agree generally with the 
form described and figured by Nordmann, except that the 
genital segment is not segmented as his drawing shows it to 
be (this drawing is reproduced on PL L, fig. 6, for com- 
parison with Scottish specimens). 

The chitinous plug by which the female Achtheres is fixed to 
its host is considerably and regularly expanded in front, as 
shown by fig. 7, PI. LIX. This form of phig appears to be 
characteristic of the genus, and differs entirely from that of 
the closely allied Lernxopoda salmonea, which is of a broadly 
lingulate form. 

Dr. Pehr Gadd, in his account of the Copepod parasites 
found on the fishes of Finland, describes a species parasitic 
on the fresh-water fish Lucioperca sandra, under the name of 
Achtheres sandrae.f In that species the genital segment is 
not segmented, resembling-, in that respect, the specimens 
described above. 

* " The Invertebrate Fauna of the Inland Waters of Scotland," ' Twelfth 
Annual Keport of the Fishery Board for Scotland/ Part III, p. 287 (1894). 

t " Parasit-Copepoder I Finland," ' Acta Societatis pro fauna et flora 
Fonnica,' vol. xxvi, No. 8, -p. 22, pi. i, fig. 1 (1904). 



LERN^OPODA. 195 

Genus 41. LERNJEOPODA Blainville, 1822. 

Female. — Body generally elongated; cephalotliorax 
short, wider behind than in front, more or less dis- 
tinctly separated from the remaining part of the body, 
and sometimes with a chitinous shield-like dorsum. 
Genital segment elongated, sometimes ovoid or saccate, 
and unsegmented. Abdomen obsolete or nearly so ; 
posterior processes two, short, or none. First maxilli- 
peds near and a little behind the mouth. Second 
maxiUipeds long or moderately shore, slender, wide 
apart, and only united at the tip to a chitinous button 
or plug usually buried in the tissues of the host. 

Male. — Much smaller than the female, somewhat 
elongated and distinctly segmented. Cephalotliorax. 
sometimes provided with a dorsal shield, and usually 
separated from the abdomen by an obvious articulation. 
Abdomen usually segmented. Antennules small, two- 
or three-jointed. Antennge tolerably large. Mouth in 
the form of a short conical siphon. MaxiUipeds large 
and uncinate. 

1. LernsBopoda elongata (Grant). 
(Plate LXI, figs. 5, 6; Plate LVII, figs. 13-17.) 

1827. Lernaea elongata Grant. (53) vol. vii, p. 147, pi. ii, fig. 5. 

1838. LerniBopoda elongata Kroyer. (70) i, pi. 2, fig. 12 ; pi. 3, fig. 3 a. 

1840. Lernxojjoda elongata M. Edwards. (43) p. 515. 

1850. Lerneopoda elongata Baird. (4) p. 333, pi. xxxv, fig. 5. 

1861. Lernxo;poda elongata Stp. & Ltk. (127) p. 422, pi. xv, fig. 37. 

1861. Lernxopoda elongata P. J. van Beneden. (15) p. 154. 

1862. Lerneopoda elongata Turner & Wilson. (140) p. 85. 

1900. Lernxopoda elongata T. Scott. (112) p. 171, pi. viii, figs. 11-15. 

Female. — Cephalothorax distinct, ovate, flattened, 
obtusely and narrowly rounded in front, but becoming 
gradually wider and thicker behind, the posterior 
margin being rounded and somewhat gibbous. Genital 
segment elongate, subcylindrical, slightly depressed, 
about as wide as the cephalothorax but distinctly 
narrower where it joins the head ; distal end truncated 
and provided with two small appendages on the ventral 



196 BRITISH PARASITIC COPEPODA. 

aspect — one on either side of the median line and in 
front of the origin of the egg-strings. Abdomen very 
minute, the genital segment showing no trace of 
segmentation. 

Antennules short, basal joint stout, the others 
narrower. Antennae verj short, stout, and bluntly 
orunded at the end, furnished with a minute branch near 
the distal end (PL LVII, fig. 14). Mandibles small, 
elongate, narrow, and having the inner margin near 
the distal end distinctly serrated. First maxillipeds 
short, stout, armed with moderately strong terminal 
claws. Second maxillipeds long and slender, exceed- 
ing the length of the body; they ''taper abruptly at 
their distal ends and are connected to a small, rounded, 
horny, or chitinous disk."* The extremities of the 
arms, where they join, closely contiguous but not 
coalescent with each other. Length from posterior 
end of the genital segment to the tips of the second 
maxillipeds about 40 mm., the maxillipeds alone 
measuring over 20 mm. 

Male, — We have not seen the male of this species, but 
Steenstrup and Liitken (op. cit., p. 423) give the size as 
about 1 mm., and their figure shows that the cephalo- 
thorax, which is provided with large uncinate maxilli- 
peds, is separated from the abdomen by a not very 
strongly defined segment ; the abdomen is more dilated 
than the cephalothorax and is also unsegmented. 

Habitat. — Parasitic usually on the eyes of the Grreen- 
land shark {Lamna cormthica). Obtained on a Green- 
land shark caught on the English coast in the winter 
of 1848 by Mr. Yarrell, who gave the specimen to Dr. 
Baird (' Entom.,' p. 334). Taken on the smooth 
hound {Miistelus vulgaris) at Polperro {A. M. Norman). 
A specimen was obtained by Mr. H. Dannevig on the 
eye of a shark captured east of Fair Isle between 
Orkney and Shetland in October 1900. Another was 
obtained by Mr. Ingram, Fishery Officer; on a Grreen- 

* Turner and Wilson, " Observations on the Parasitic Crastacea, Chondra- 
canthus Lojphii and Lerneopoda dalmanni," 'Trans. Eoy. See. Edinb./ vol. 
xxiii, Pt. 1, p. 85 (1862). 



LERNiEOPODA ELONGATA. 197 

land shark captured in the North Sea and landed at 
Aberdeen Fish-Market in April 1901. Both specimens 
were examined by us. 

2. Lernaeopoda galei Kroyer. 

(Plate LX, figs. 4-6 ; Plate LVIII, figs. 9-15 ; Plate 
LXIII, fig. 1.) 

1837. Lernaeopoda galei Kroyer. (70) p. 272, pi. iii, fig. 5. 
1840. Lernseopoda galei M. Edwards. (43) p. 516. 

1850. Lerneoiooda galti Baird. (4) p. 334, pi. xxxv, fig. 7. 

1851. Lernseojjoda galei P. J. van Beneden. (11) p. 120, pi. iv. 
1896. Lernasopoda galei Bassett-Smith. (7) p. 163. 

1900. Lernaeopoda galei T. Scott. (112) p. 172, pi. viii, figs. 16-25. 

Female. — Cephalothorax short, flattened, and of an 
oval shape; the dorsum chitinous and forming a shield- 
like covering. Genital segment elongated, subcylin- 
drical, proximal end narrower than the cephalothorax 
but becoming slightly enlarged towards the posterior 
extremity ; distal end provided with two small 
cylindrical processes, one on each side of the minute 
abdomen. Egg - strings narrow, cylindrical, and 
tolerably elongated. 

Antennules short, enlarged at the base; end joints 
three, narrow, and furnished with a few minute apical 
setas. Antennae, mandibles, and maxillae similar to 
those of Lerndeopoda elongata. First maxillipeds short, 
stout, armed with strong terminal claws. Second 
maxillipeds slender, considerably shorter than the body, 
and united at the tip to a chitinous disk or plug inserted 
into the tissues of the host. Length from the forehead 
to the end of the posterior appendages 13 to 14 mm. 
Length of the posterior appendages fully 2 mm., and 
of the second maxillipeds between 4 and 5 mm. 

Male, — The male is small. The cephalothorax is 
nearly equal in length to the genital segment, and the 
form is ovoid; genital segment elongate-ovate, fully 
twice as long as broad, and having at the distal end 
two short reflexed oval appendages, one on each side of 
the nearly obsolete abdomen. Both pairs of maxilli- 



198 BRITISH PARASITIC COPEPODA. 

peds short, stout, and furnished with strong terminal 
claws. 

Habitat. — Found parasitic on various dog- fishes, 
usually adhering to the ventral and anal fins. At 
Polperro on the toper {Galeus vulgaris), and from 
cod {A, M, Norman). At Plymouth "from Galeus 
vulgaris, Mustelus vulgaris, and Acanthias vulgaris '^ 
(Bassett-Smith). Belfast, 1839 (W. Thompson). Off 
Valentia (W. F. Kane). Irish Sea {A. Scott). The 
Firth of Clyde, Moray Firth, Aberdeen {T. Scott). 

3. Lernseopoda cluthae T. Scott. 

(Plate LX, figs. 1-3; Plate LVII, figs. 1-7; Plate 
LVIII, fig. 16.) 

1900. Lemseopoda cluthw T. Scott. (112) p. 173, pi. viii, figs. 27-37. 
1909. Lemseopoda clutlue May E. Bainbridge. (3) p. 49, pi. 10, figs, 

24-27. 

Female. — Cephalothorax small, subtriangular, a dis- 
tinct and narrow neck connecting it with the posterior 
portion of the body, which is somewhat dilated and 
subcylindrical, and exhibits a few pseudo-constrictions. 
Two short processes springing from the distal end of 
the genital segment ; the abdomen, situated between 
them, very small. The two pairs of antennas similar 
to those of the species already described. Mandibles 
small and their biting margins, which are obliquely 
truncated, finely and somewhat irregularly serrated, 
and differing very markedly from the same appendages 
in Lerndeopoda galei ; maxillae small, end joint provided 
with two tolerably long and stout terminal spines. 
The first maxillipeds more slender and rather more 
elongate than those of L. galei. Length, from the 
forehead to the end of the posterior appendages, 
5 mm. ; length of the second maxillipeds nearly 3'5 mm. 

Male. — The male differs considerably from that of 
the species just described {L. galei), particularly in 
the structure of the abdomen and of the caudal rami. 
In the species under consideration the abdomen of the 
male is distinctly segmented, and the caudal rami. 



LERN^OPODA CLUTH^. 199 

wliicli are not divergent, consist of two short slender 
processes instead of being broadly oval or claviform 
and reflexed upon the abdomen. 

Habitat. — Parasitic on the gill-filaments of the 
Fuller's ray {Baia fullonica). Firth of Clyde, May, 
1897 {T. Scott). Northumberland coast {May E, 
Bainhridge) on Raia radiata. 

This species has not been observed on any of the 
dog-fishes, but only on the gill-filaments of the rays 
mentioned. 

P. Olsson lias described a Lern^eopoda (L. longimana) 
from the same species of skate.* The female only is described, 
and it appears from his description and figures to be distinct 
from the species recorded here. 

4. Lernseopoda salmonea (Linn.). 
(Plate LIX, figs. 1-G ; Plate XL VIII, figs. S-12.) 

1761. Lerrnea salmonea Linn. (77) p. 509, No. 2102. 

1780. Lerucea salmonea Cordiner. (36) p. 7, 8, pi. 6, fig. 2. 

1816. Entomoda salmonea Lamarck. (72) p. 233. 

1822. Lerneoiioda salmoyiea Blainville. (24) p. 127. 

1837. LeruiBopoda carpionis (?) Kroyer. (70; (1) pi. ii, fig. 6.. 

1840. Basenistes salmonea M.. Edwards. (43) p. 509, pi. xli, fig. 3. 

1850. Jjerneopoda salmonea Baird. (4) p. 335, pi. xxxv, fig. 6. 

1863. Leru'cBopoda salmonea Kroyer. (71) p. 275, pi. xiv, fig. 3 «-/. 

1900. Lern-ceopoda salmonea T. Scott. (112) p. 173, pi. viii, tig. 26. 

Female. — Cephalothorax, seen from above, sub- 
triangular, small, enlarged behind and distinctly 
separated from the posterior part of the body. The 
genital segment, where it joins the cephalothorax, 
narrowed and forming a kind of neck ; the segment 
then becoming considerably enlarged and ovoid or 
pyriform, the posterior end rounded and provided 
with two minute apical knobs, one on each side of the 
abdomen which is very small. 

The two pairs of antennae and the appendages of 
the thorax somewhat similar to those of Levnseoiooda 
(jalei except that the second maxillipeds are tolerably 
stout and short, and the chitinous plug to which they 

* " Prodr. fauna Copep. parasit. Scand.," ' Acta Universitatis Lundensis/ 
1868 ; * Lunds Univ. Arsskrift/ iii, p. 30, pi. 2, figs. 13-15. 



200 BKITISH PARASITIC COPEPODA. 

are united at the tip is proportionally large. Entire 
length from the posterior end of the genital segment 
to tlie tip of the second maxillipeds about 6 mm. 

The male has not been observed by us. 

Habitat. — Parasitic on the gills of the salmon. 
London Market (IF. Baivd)'. North of Ireland, 1856 
{W. Thompson). Plymouth {B as sett- Smith). On a 
diseased salmon from the Firth of Tay (T. Scott). On 
*' the gills of salmon taken in the Coquet, 1908"^' 
(E. L. Gill). This species was also obtained on the 
gills of salmon by Thomas Edward of Banff.f 

A salmon captured in the river Dee near Aberdeen was 
infested by these parasites : they could be observed adhering 
to the ends of the gill-filaments in considerable numbers, the 
white colour of the parasites showing distinctly against the 
red colour of the gill filaments. (See Plate B, fig. 3). 

5. Lernseopoda bidiscalis W. F. de Visme Kane. 
(Plate LXI, figs. 1, 2; Plate LVII, figs. 8-12.) 

1892. Lernseopoda hidiscalis W. F. de Yisme Kane. (68) (3) vol. ii, 
p. 203, pis. ix, X. 

1900. LerniEopoda hidiscalis T. Scott. (112) p. 172. 

Female. — Cephalothorax, seen from above, oblong 
in shape and rather longer than broad, but in profile 
somewhat pyriform, sloping dorsall}^ upwards and 
backwards, and terminating abruptly behind ; a con- 
striction which is distinct, but which can scarcely be 
described as a "neck," separating the cephalothorax 
from the genital segment. Genital segment short, 
considerably enlarged, and nearly as broad as long ; an 
indistinct median dorsal groove present in some speci- 
mens, the posterior end somewhat truncated, and the 
postero-lateral corners rounded and slightly lobate. 
The genital segment provided with two short, inter- 
mediate, fusiform appendages which spring from the 
ventral aspect near the origin of the egg-strings. 
Abdomen nearly obsolete. 

* Cf. 'Crustacea of Northumberland and Durham/ by Brady and 
Norman (1910). 

t Cf. Smiles, ' Life of a Scottish Naturalist/ 2nd ed. (1877), p. 437. 



LERN.EOrODA BIDISCALIS. 201 

Antennules nearly as in Lernxopoda galei. Antennae 
very short, stout, and prehensile. The mandibles and 
other mouth-organs not differing much from those of 
L. galei. The second maxillipeds very short and 
stout, each terminating in a very large, fleshy, circular 
or ear-shaped disk ; the disks united together by a 
horn-coloured chitinous plug. Colour usually opaque- 
white, more or less tinged with red, sometimes highly 
coloured. Length : forehead to the end of the 
posterior appendages about 7 mm., excluding the 
appendages, 5*5 mm. 

Male. — Though small, proportionally rather larger 
than the male of Lernaeopoda galei, but in its general 
form and in the structure of its appendages having 
a close resemblance to the male of that species. 

Habitat. — Parasitic chiefly on the claspers of male 
specimens of the tope (or toper), Galeus canis. On 
topers captured off Valentia, Ireland {de Vism^e Kane). 
Firth of Clyde (Mr. Duthie, Fishery Officer). Aber- 
deen {T. Scott). Irish Sea (A. Scott). Taken also on 
the smooth hound (Mustelus vidgaris) at Polperro, Corn- 
wall by Mr. W. Laughrin about 1862 {A. M. Norman). 

The dog-fishes on which these parasites were obtained were 
adult males, and they were usually found adhering on, or 
near the ends of, the claspers. We have rarely observed 
them on the claspers of young males. Moreover, as men- 
tioned by the describer of the species, we almost invariably 
found that the ends of the claspers on which the parasites 
occurred were torn and bleeding, but whether the wounds 
were caused directly by the parasites, or through the efforts 
made by the fish to throw off its tormentors, was not very 
clear. Occasionally both claspers had parasites adhering to 
them. 

The specimens observed by us had usually the front 
portion of the head of a bright red colour, and sometimes 
there were blotches of the same colour on other parts of the 
body ; frequently also we found one, and more rarely two, 
males adhering to the female. 

Among the more prominent features by which this Ler- 
naeopod is distinguished is the comparatively large size of 
the nearly circular disks which terminate the short second 



202 BRITISH PARASITIC COPEPODA. 

maxillipeds ; the whole animal is also short and robust, and 
very different from the more elegant L. galei which is also 
sometimes met with at the base of the claspers. 



6. Lernseopoda similis sp. nov. 
(Plate LXI, figs. 3, 4.) 

Female, — Somewhat similar in general appearance 
to Lernseopoda cluthse, but the head tolerably elongated ; 
the second maxillipeds short, free except at the ex- 
tremities, where they become united to a chitinous, 
horn-coloured button which penetrates the tissues of 
the host. Genital segment elongate-pyriform, widest 
posteriorly. Abdomen nearly obsolete, furnished with 
two tolerably long digitiform processes. Egg-strings 
long, narrow-cylindrical ; ova small, numerous. 

Mouth-appendages apparently somewhat similar to 
those of Lernaeopoda chithm. Colour yellowish grey. 
Length to the end of the posterior processes about 
8 mm. 

Habitat. — Parasitic on the gills of the blue or grey 
skate {Eaia hatiH) captured in the Firth of Clyde at 
Stations I and II in December 1899. 

This species may be distinguished from L. cluthse, which 
it somewhat resembles, by the short second maxillipeds, 
and by the form of the head. 

7. ? Lernaeopoda lampri sp. nov. c^ . 
(Plate XXXIII, fig. G; Plate XLIX, figs. 12-16.) 

1901. ? Lermeojwda sp. c?, T. Scott. (113) p. 136, pi. viii, figs. 4-10. 

" A few male specimens of what appears to be a 
species of Lernaeopoda were obtained on the gills of an 
opah or king-fish, Lampris pelar/icus {Gunn,) = Lampris 
luna, Grmel.), forwarded from Shetland to the Fishery 
Board's Laboratory at Bay of Ni gg, 10th October 
1900 ; no females were observed. . . . The body 
of this parasite of the king-fish is comparatively 
slender ; it tapers more or less gradually towards the 



? LERNiEOPODA LAMPRI. 203 

posterior end. The antennules are small and four- 
jointed, the penultimate joint being shorter than the 
others. The antennse are moderately robust except 
the last two joints, which are small, and the end joint 
is very feebly clawed. The mandibles are small, but 
somewhat similar in form and armature to those of 
Gharopinus dalmanni. The maxillge are also somewhat 
similar to those of that species, except that the ends 
are furnished with two instead of three spiniform 
appendages. The first and second maxillipeds are 
very robust and armed with short but powerful 
hooked terminal claws. 

" The form of the various appendages shows a close 
relationship with the Lernasopodidae, closer perhaps 
with Gharopinus than with Lerndeopoda.'' 

The male specimens, of which the above is a short descrip- 
tion, have been doubtfully referred to the genus Lernseopodaj 
but as they are considerably larger than the males of any 
species of that genus the females of which are known to us, 
they probably do not represent Lernseo'poda at all, but if so, it 
is a genus closely related to it, as indicated by their form and 
anatomical details. One of the males figured in the Fishery 
Board's Report mentioned above measured 4'3 mm. If the 
size of the female is of about the same proportion as we find 
it to be in some of the otlier species of the Lerna3opoda, it 
should reach to about 16 or 18 mm. in leno-tli. 



Genus 42. BRACHIELLA Guvier, 1817. 

Nearly related to Lernseopoda, but the female differ- 
ing in having the cephalothorax usually more elon- 
gated and without a shield-like dorsum, the distance 
from the base of the second maxillipeds also being 
usually greater. The second maxillipeds separate 
except at the tip as in Lenidepoda, but usually shorter, 
rarely elongated. Genital segment short and robust, 
rarely elongate, and furnished with one, sometimes 
with two pairs of posterior appendages. 

Male very minute ; the cephalothorax and abdomen 
of nearly equal thickness and not very distinctly 



204 BRITISH PARASITIC COPEPODA. 

defined. Both pairs of maxillipeds large and un- 
cinate. 

The male shows a closer relationship Avith the male of 
Clavella than with that of Lernxopoda. 

1. Brachiella thynni Cuvier. 
(Plate LXIV, figs. 4-6.) 

1817. Brachiella thynni Ciiv. (37) p. 287, pi. xv, fig. 5. 

1829-1843. Brachiella thynni Guerin-Meneville. (55) j)l. ix, fig. 6 a-c. 

1832. Brachiella thynni Nordmann. (89) jd. 90. 

1840. Brachiella thynni M. Edwards. (43) i). 512. 

1851. Brachiella thynni P. J. van Beneden. (11) p. 128. 

1861. Brachiella thynni Stp. & Liitk. (127) p. 420, pi. xv, fig. 36. 

1877. Brachiella thynni C. Vogt. (142) p. 426. 

1896. Brachiella thynni Bassett-Smith. (7) p. 162. 

1906. Brachiella thynni Norman & T. Scott. (88) p. 219. 

Female. — Ceplialothorax slender, flexuose, fully as 
long as the genital segment, and separated from it by 
a more or less distinct articulation or constriction. 
Genital segment narrow at its union with the cephalo- 
tliorax but becoming gradually enlarged towards the 
posterior end, where the width is equal to about half 
the length ; the posterior end abruptly truncated and 
furnished with two pairs of elongated and moderately 
slender appendages, one pair springing in front of and 
the other behind the base of the egg-strings, which 
thus issue from between them, each having at its 
base an appendage both in front and behind ; the 
appendages on the ventral aspect scarcely so elongated 
as the others, and about as long as the genital seg- 
ment. That segment also somewhat flattened and 
with several transverse constrictions more or less indis- 
tinct, imparting to it an obscure lobate appearance. 

Antennules small, three- join ted. Antennae stout, 
moderately large, and furnished with a few apical 
spines. Mandibles narrow, elongated, inner margin 
near the distal end coarsely toothed, the teeth being 
to some extent alternately larger and smaller. Maxillae 
small, furnished with two or three moderately stout 
apical setae and one or two minute submarginal 



BRACHIELLA THYNNI. 205 

spines. First maxiilipeds sliorfc, stout, and strongly 
uncinate. The second maxiilipeds, which are equal 
to about two- thirds of the length of the genital seg- 
ment, slender and free except at the tip, where thej 
are joined to a reddish-brown coloured chitinous plug. 
Length from the forehead to the end of the posterior 
appendages of a full-grown specimen about 32 mm. ; 
length of cephalo thorax about 14 mm., of the genital 
segment about 9 mm., and of the posterior appendages 
9 mm. 

Male. — Very small, being about a millimetre in 
length; having a closer resemblance to the male of 
Ler7ideopoda than to that of a true Brachiella, 

Habitat. — Parasitic on the gills of the tunny (Orcy- 
nus thynnus), and other nearly related fishes. Four 
specimens were obtained by Dr. Bassett-Smith on a 
tunny captured at Plymouth. 

The female of this species is somewhat similar to B. rostrata 
in general appearance, but the cephalothorax is distinctly 
longer, and the posterior appendages are four in number and 
much more elongated. 

2. Brachiella rostrata Kroyer. 

(Plate XLV, fig. 8 ; Plate LXII, fig. 3 ; Plate LXIII, 

figs. 2-8.) 

1837. Brachiella rostrata Kroyer. (70) Rekka 1, p. 207, pi. ii, fig. 1. 

1840. Brachiella rostrata M. Edwards. (43) vol. iii, p. 415. 

1863. Brachiella rostrata Kroyer. (71) p. 290, pi. xviii, figs. 8, a-i. 

1877. Brachiella rostrata C. Yogt. (142) p. 426. 

1900. Brachiella rostrata T. Scott. (112) p. 174, pi. viii, figs. 38, 39. 

Female. — Cephalothorax elongated, flexuose or 
vermiform, and separated from the genital segment 
by a more or less clearly-defined constriction. Genital 
segment narrow, subcylindrical, fully three times 
longer than broad and equal to about one and one- 
third times the length of the cephalothorax ; posterior 
end subtruncated and furnished with two subapical 
appendages. Abdomen obsolete or nearly so. The 
egg-strings, which are tolerably elongated, arising 
immediately outside of the subapical appendages. 



206 BRITISH PARASITIC COPEPODA. 

Antennules small, apparently four- jointed, but the 
articulation between the first two joints indistinct. 
Antennae and mouth-orf^ans somewhat similar to those 
of BrarJiiella thyniil. The second maxillipeds spring- 
ing from the lower half of the cephalo thorax and 
reaching to about the proximal end of it ; free except 
at the tip, where they are jointed to a horn-coloured 
chitinous plug. Length from the extremity of the 
cephalothorax to the end of the genital segment 
about 15 mm., and 17 mm. to the tip of the posterior 
appendages. 

Male. — The male is nearly twice as long as broad 
and measures about 2 mm. in length; it is compara- 
tively robust, the cephalothoracic appendages are 
somewhat crowded together at the proximal end, and 
there is no distinct division between the cephalothorax 
and abdomen. The antennae (both pairs), mandibles, 
and maxillae are nearly as in the female ; but both 
pairs of maxillipeds, though short, are furnished with 
strong terminal claws. Abdominal appendages very 
small. 

Habitat. — Parasitic on the gills of halibut {Hippo- 
glossus vulgaris). On large halibut captured in the 
North Sea, and landed at the Fish-Market, Aberdeen. 

Kroyer records two forms, apparently distinct, under B. 
rostrata, one from the Hippoglossiis here mentioned and the 
other — a shorter form — from Hippoglossus pinguis. This 
form we have not seen. 

8. Brachiella insidiosa Heller. 

(Plate XLVIII, fig. 18; Plate LXII, fi^. 2; Plate 
LXIII, figs. 9-16.) 

1865. Brachiella insidiosa Heller, (58) p. 239, pi. xxiv, fig. 1. 

1896. Brachiella insidiosa Bassett-Smith. (6) vol xviii, p. 14, pi. vi, 
fig. 2. 

1900. Brachiella insidiosa T. Scott. (112) p. 175, pi. viii, figs. 40, 41. 

1906. Brachiella insidiosa Brian. (21) p. 104, pi. viii, figs. 1 and 4. 

Female. — Body tolerably robust. Cephalothorax 
short, flexuose, and vermiform. Genital segment 



BRACHIELLA INSIDIOSA. 207 

subcylindrical, flattened, about two and a half times 
longer than broad ; the width increasing slightly 
towards the posterior end, which is truncated and 
provided with two pairs of appendages. The two 
lateral appendages moderately elongated and slender, 
but the intermediate pair short, and springing from 
each side of the very small abdomen. 

Antennules short and stout, and each composed of 
about three joints; antennse robust, and somewhat 
similar to those of Lernxopoda galei. Mandibles 
narrow at the base but wider towards the posterior 
end, and armed with irregular coarse teeth on the 
obliquely-truncated distal extremity. Maxillae small, 
simple, and provided with a few apical spines. First 
maxillipeds robust, and strongly uncinate ; second 
maxillipeds short, and having the appearance of being 
a prolongation of the cephalothorax. Length from 
the extremity of the cephalothorax to the end of the 
longer posterior appendages about 14 mm. Egg- 
strings long and slender. 

Male. — Very small, stout, about twice longer than 
broad and similar in structure to the male of Brachiella 
rostrata ; the antennse and both pairs of maxillipeds 
strongly uncinate. 

Habitat. — Parasitic on the gill-rays of the hake 
(Merhiccius vulgaris), moderately common. Plymouth 
{Bassett- Smith). Firths of Forth and Clyde, and at 
Aberdeen Fish-Market {T. Scott). Irish Sea {A. Scott). 

4. Brachiella merluccii Bassett-Smith. 
(Plate LXII, figs. 4, 5; Plate LXIII, figs. 17-20.) 

1896. Brachiella merluccii Bassett-Smitli. (7) vol. iv, p. 163. 
1896. Brachiella merluccii idem. (6) (6) vol. xviii, p. 14, pL vi, fig. 1. 
1900. Brachiella merluccii T. Scott. (112) p. 175, pi. viii, fig. 42. 
1906. Brachiella merlucii Brian. (21) p. 107, pi. viii, fig. 3. 

Female. — This species of Brachiella differs from 
most of the others by its bizarre appearance. Cephalo- 
thorax not very clearly defined from the genital 
portion of the body ; bending round and forward at 



208 BIUTISH PARASITIC COPEPODA. 

nearly a right angle to the dorsum, so that the animal 
appears to be very short and dilated when viewed 
from above. G-enital segment considerably enlarged 
and furnished with two pairs of appendages ; one 
pair moderately short prolongations of the postero- 
lateral corners of the segment, the other pair tolerably 
elongated, springing from the ventral surface in front 
of the egg-strings, extending backward, curving round, 
and partly inclosing them. 

Antennules stout and composed of three joints ; 
the mandibles and maxilla, as well as the second pair 
of antennae, somewhat similar to those of Brachiella 
insidiosa ; the first maxillipeds, however, scarcely so 
robust, but furnished with more powerful terminal 
claws. The second maxillipeds very short and 
enclosed together within a gelatinous envelope : 
though thus enclosed, not coalescent, but may be seen 
through the semi-transparent envelope extending 
alongside each other to where at the apex they are 
united to a hard chitinous plug. Length about 8 mm. 

Male. — We have not observed the male of this 
species, but Dr. Bassett-Smith describes it and states 
that the cephalothorax is large, and distinct from 
the posterior portion of the body, this portion being 
divided into five indistinct segments, and furnished at 
the posterior end with a pair of short two-jointed 
processes with pointed ends. The two pairs of an- 
tennae, and the various other thoracic appendages^ 
apparently similar to those of the male of Brachiella 
insidiosa. 

Habitat. — Parasitic on the gill-rakers of the hake 
{Merlucciusvidgaris), and not, like Brachiella insidiosa^ 
attached to the gill-rays. Plymouth (Bassett-Smith). 
Firth of Forth and Aberdeen Fish-Market {T. Scott). 

Brachiella merliiccii does not appear to be so common as the 
species just alluded to; and the general structure of the 
female, and especially tlie partial union of the second maxilli- 
peds, show, as pointed out by Dr. Bassett-Smith, a tolerably 
close relationship with Clavella {Anchorella) . 



BRACHIEfXA TEIGL^. 209 

5. Brachiella triglae Claus. 
(Plate LXII, fig. 1 ; Plate LXIII, figs. 21-25.) 

I860. Brachiella trigliE Claus. (30a) Wiirzburger naturwiss. Zeitsclir., 
vol. i. p. 32, pi. i, fig. 6. 

1877. Ancliorella trigliE Kiirz. (71a) Zeitschr. f. wiss. Zool., vol. xxix, 
p. 404, pi. XXV, figs. 13-15. 

1896. Brachiella trighe Bassett-Smith. (7) p. 163. 

1901. Brachiella triglse T. Scott. (113) p. 133, pi. vii, figs. 24-29. 

Female. — Small but moderately stout; cephalo- 
tliorax moderately short and flexuose ; somewhat 
vermiform, and about as long as the genital segment. 
G-enital segment considerably expanded, the width 
being rather greater than the length ; lateral margins 
more or less irregularly lobate, and the postero-lateral 
produced into bluntly-rounded lobes ; also provided 
with two short posterior appendages, situated one on 
each side of the nearly obsolete abdomen. 

Antennules short, tapering, and composed of four 
joints, end joint rather longer than the preceding one. 
Antennse moderately short and stout, the end joint 
with a small terminal spike ; outer ramus very short, 
biarticulated, and bearing two or three minute apical 
spines. Mandibles slender and obliquely truncated, 
the truncated margin being rather coarsely serrated ; 
maxillge moderately stout, bearing three elongated 
apical spines, and two others at the end of a small 
lateral process. First maxillipeds short, robust, and 
provided w4th small terminal claws. Second maxilli- 
peds very short and stout, free except at the tips, 
where they are joined to a chitinous horn-coloured 
disk. Length about 4*5 mm., varying shghtly in 
different specimens. 

Male. — We have not observed the male of this 
species, but Dr. Bassett-Smith has figured one which 
apparently does not differ much from the males of 
other species of Brachiella. 

Hahitat. — Parasitic on the gills and gill-arches of 
Trigla spp. On Trigla gurnardns, cuculiis, and hirundo 
at Plymouth {Bassefi- Smith). On Trigla lineata, Firths 
of Forth and Clyde (7'. Scott). It has also been 

VOL. T. 14 



210 BEITISH PARASITIC COPEPODA. 

obtained on the gill-arches of Trigla lineata in the 
Adriatic. 

6. Brachiella ovalis (Kroyer). 
(Plate LX, fig. 7 ; Plate LXIII, figs. 26-30.) 

1837. Anchorella ovalis Kroyer. (70) p. 289, pi. iii, fig. 6. 
1870. Anchorella ovalis P. J. van Beneden. (16) p. 31, pi. ii, fig. 8. 
1901. Brachiella ovalis T. Scott. (113) p. 133. pi. vii, figs. 30-35. 
1904. Brachiella ovalis A. Scott. (109) No. 12, p. 44. 

Female. — Cephalothorax stout, moderately short, 
but rather longer than the genital segment. Genital 
segment subpyriform, considerably expanded pos- 
teriorly and narrowed at the proximal end to nearly 
the width of the cephalothorax ; the posterior margin 
sloping backward slightly, and equally from both 
sides, and forming an obtuse angle at the base of the 
abdomen ; a short subniedian appendage occurring on 
each side of the abdomen midway between it and 
origin of the subglobular egg-strings, these append- 
ages usually being more or less concealed by the egg- 
strings. Abdomen very small. 

Antennules apparently composed of four joints, the 
first dilated and the others small, the third being 
much shorter than either the second or fourth. 
Antennae tolerably large ; end joint narrowly rounded 
at the apex and covered with minute bristles, the 
outer ramus consisting of two small subequal joints 
bearing three very small apical spines. Mandibles 
moderately stout and slightly expanded towards 
the distal end, the inner margin at this end being 
armed with a series of somewhat coarse teeth, 
alternately larger and smaller. Maxillae and first 
maxillipeds somewhat similar to those of Brachiella 
triglde. Second maxillipeds very short, stout, and re- 
curved, free except at the distal end, where they are 
united to a chitinous plug fixed in the gill-arches of 
the host. Length about 8*5 mm. 

AVe have been unable to obtain the male of this species. 

Habitat. — Parasitic on the oi-ill-arches of Trigla 



BRACHIELLA OVALTS. 211 

gurnardus. Moray Firth, in April and May 1900 (T. 
Scott). Irish Sea (A. Scott). Apparently more fre- 
quent on young fishes than on adults. Both Kroyer 
and P. J. van Beneden record this BrachieUa from 
Trigla gurnardus. 

7. BrachieUa bispinosa Nordmann. 
(Plate L, fig. 9 ; Plate LXIV, fig. 9.) 

1832. Bracliiella hispinosa JN'ordmann. (89) p. 94, pi, viii, figs. 4-7. 

1840. BrachieUa bispinosa M. Edwards. (43) p. 513. 

1877. Bracliiella bispinosa C. Yogt. (142) p. 456. 

1901. BrachieUa bispinosa T. Scott. (113) p. 132. 

1906. BrachieUa bispinosa Norman & T. Scott. (88) p. 220. 

Female. — Cephalothorax narrow, rather shorter than 
the genital segment and usually more or less doubled 
back upon it. Grenital segment seen from above 
oblong or somewhat lyre-shaped, nearly twice longer 
than broad ; posterior end broadly rounded, not 
angular, and provided with two small submedian 
spiniform appendages. 

Antennules short, slender, and composed of three 
or four joints. First maxillipeds strongly uncinate. 
Second maxillipeds scarcely reaching to the end of 
the cephalothorax, free but united at the apex to a 
round chitinous knob. Length about 8 mm. 

We have not seen the male of this species, but 
Dr. Bassett-Smith states that it resembles the male of 
Thysanote imjmdica (Xordmanu).* 

Habitat. — Parasitic on the gill-arches of Trigla 
spp. Plymouth, on Trigla cuculus, Trigla gurnardus, 
and Trigla lyra (Bassett-Smith). Moray Firth, on 
Trigla sp. {T. Edivard : 'Museum Xormanianum '). 

8. Bracliiella pastinaca P. J. van Beneden. 
(Plate LXIV, fig. 8.) 

1851. BrachieUa pastinaca P. J. van Beneden. (11) p. 118, pi. 4, figs. 8. 9. 

1877. BrachieUa pastinaca Kurz. (71a) Zeitsclir. f. wiss. Zool., vol. 
xxix, pis. 25, 26, 27, figs. 2, 3, 36, 45. 

1880. BrachieUa pastinaca A. Yalle. (141a) Boll. Soc. Adriat. Sc. 
Nat., vol. iv, fasc. i, p. 77. 

1904. BrachieUa pastinaca T. Scott. (115) p. 278. 

* ' Joiirn. Mar. Biol. Assoc' (n.s.), vol. iv, No. 2 (February 1896), p. 163. 



212 BRITISH PARASITIC COPEPODA. 

1906. Brachiella pastinacx Brian. (21) p. 103. 

1909. 5rac^^eZZa^as/^l^aca May E. Bainbridge. (3) p. 50, pi. 8, figs. 
6, 7 ; pi. 9, fig. 8. 

Female, — Ceplialotliorax tolerably enlarged and 
some what gibbous behind on the dorsal aspect, con- 
nected with the genital segment by a narrow neck. 
Genital segment oblong, robust, and at the distal end 
bearing two slender submedian appendages nearly half 
as long as the segment. Abdomen obsolete or nearly so. 

Antennules moderately slender and four-jointed ; 
antennae stout, composed of two joints, and fur- 
nished with a small rudimentary outer ramus. 
Mandibles tolerably slender and armed with several 
coarse teeth at the distal end of the inner margin. 
The first maxillipeds tolerably large and strongly 
uncinate. The second maxillipeds nearly as long 
as the cephalothorax, free except at the tips where 
they are united to a horn-coloured chitinous ping. 
Length about 7*5 mm. 

Habitat. — Parasitic in the nasal fossae of Trigon 
pastinaca and also of the piked dog-fish [Acanthi as 
tmlgaris). In the nasal fossas of a Trir/ou captured in 
the Dornoch Firth in October, 1903 (T. Scott). In 
the spiracle of a spiked dog-fish captured off the 
coast of Xorthuml)erland {Maij E. Bahibridrje).^ 

9. Brachiella parkeri G. M. Thomson. 
(Plate LXIV, fig. 7.) 

1889. Brachiella parheri Thomson. (132) p. 374, pi. 10, figs. 18-23. 
1909. Brachiella parkeri May E. Bainbridge. (3) p. 52, pi. 9, figs, 
16, 17 ; pi. 10, figs. 18-23. 

Female. — Cephalothorax moderately stout, deflected 
so as usually to be nearly at right angles to the genital 
segment, and equal to about two-thirds the length of it. 
Genital segment of an ovate form, moderately dilated, 
and provided with two elongated and rather slender 
distal appendages, one on each side of the papilliform 
abdomen. Antennules moderately elongated, slender, 
and composed of two articulations; antennae some- 

* Miss May Evelina Bainbridge, now the Hon. Mrs. Henn Collins. 



BRACHIELLA PAEKEKI. 213 

what similar in structure to those of Brachiella fas- 
tinaca. Eostrum somewhat prominent ; mandibles 
slender, their inner margins armed at the distal end 
with about four tolerably large teeth and with smaller 
teeth between. Maxillaa of the usual type, resembling 
those of Brachiella triglse. First maxillipeds stout 
but somewhat rudimentary and only feebly uncinate ; 
second maxillipeds long and slender, fully twice the 
length of the cephalothorax, and entirely free except 
at the tip, where they are united to a chitinous horn- 
coloured disk. 

Habitat. — Obtained on the gills of a long-nosed 
skate (Baia oxyrhynchus) captured off the Northumber- 
land coast in May 1908 {May E. Bainbridge). 

We have not met with this species, hut Miss Bainbridge, in her 
paper referred to above, gives a careful description of it. Her 
description and drawings agree very well with those of Gr. M. 
Thomson. The following measurements of the specimen 
examined are given by Miss Bainbridge : — 

Length from top of arms to end of abdomen 9*2 mm. 
„ from end of head to extremity of 

attachment .... 19*0 „ 
„ of arms ..... 14*5 „ 
„ of body from bend of neck to end 

of abdomen .... S'O ,, 



„ of head to bend of neck 
„ of process . 
Width of neck 

,, of genital segment 



5-3 
4-0 
1-5 
5-2 



Genus 43. CLAVELLA Olmi, 1815. 

Syn. Schisfui'us Oken (1815), a name preoccupied by Rndolphi (1809) 
for a genus of Vermes. Ancliorella Cuv, (1817). Lerneomijzon Blainville 

(1822). 

Female. — Body usually short and somewhat swollen. 
Head small, situated at the end of a long flexuous 
neck which is usually more or less deflexed. Both 
pairs of antennae small and somewhat rudimentary; 
the mandibles, maxilljB, and first maxillipeds resem- 
bling those in Lernmopoda, and, together with the two 



214 BRITISH PARASrilC COPEPODA. 

pairs of antennae, placed at the end of the elongated 
neck, or cephalothorax. The second maxillipeds, by 
which the parasite fixes itself to its host, situated at 
the base of the cephalothorax, and coalescent so that 
they appear to consist of a single organ; this organ is 
usually short, sometimes very small and papilliform, 
and united at the tip to a horn-coloured chitinous plug 
which penetrates the tissues of the fish. Genital seg- 
ment usually unprovided with terminal or lateral pro- 
cesses, but if present they are generally very small. 
Abdomen obsolete or nearly so. Egg-strings moderately 
elongated. 

Male. — The male is very small and proportionally 
shorter than the male of .Bnichiella, which it more 
nearly resembles than that of the genus Lernseopoda ; 
the dorsum, however, is more boldly arched, and the 
height is usually greater than the length. The append- 
ages of the cephalon and thorax are somewhat similar 
to those of the male of Bracldelln. 

The genus comprises a large number of species, and those 
described below have been obtained on British fishes. 

1. Clavella uncinata (0. F. Midler). 
(Plate LXV, figs. 2, 3, 6 ; Plate LXVI, figs. 21-23.) 

1777. Lermea uncinata O. F. Miiller. (85) p. 120, pi. xxxiii, fig. 2. 

1815. Schisturus iincinatus Oken. (91) part iii, p. 183. 

1815. Clavella uncinata idem. (91) pai-t iii, p. (?) 357. 

1817. Anchorella uncinata Cuv. (37) Icon, du Regne Anim., pi. 9, 
fig. 5. 

1822. Lerneomyzon uncinatum Blainville. (24) Joiira. de Physique, 
vol. 95, p. 438. 

1832. Anchorella uncinata Nordmann. (89) p. 102, pi. 8, figs. 8-12 ; 
pi. 10, figs. 1-5. 

1850. Anchorella uncinata Baird. (4) p. 337, pi. xxxv, fig. 9. 

1900. Anchorella uncinata T. Scott. (112) p. 177, pi. viii, figs. 43, 44. 

1906. Clavella uncinata Brian. (21) p. 110. 

1909. Anchorella uncinata May E. Bainbridge. (3) p. 57, pi. ii, figs. 
43-46. 

Female. — Cephalothorax moderately slender and 
about as long as the genital segment. Genital segment 
ovate, slightly flattened, and moderately elongated. 



CLAVELLA UNCINATA. 215 

being fully twice as long as broad. Abdomen small 
but quite distinct. 

Antennules three- jointed, tlie proximal joint large 
but tapering towards the distal end ; other joints small 
and subequal but the end one rather the larger. Man- 
dibles small, biting part serrated on the inner edge. 
Maxillae small, expanded towards the distal end, with 
the outer margin somewhat gibbous ; each bearing a 
couple of terminal spines and also a small subterminal 
lobe with two spinules at its apex. The first maxilli- 
peds rather diminutive in size but provided with 
moderately stout terminal claws. The second maxilli- 
peds completely coalescent, very short, expanded at 
the apex, and fixed to a nipple-like plug which pene- 
trates the tissues of the fish. Egg-strings tolerably 
slender and fully twice as long as the genital segment. 
Length of the specimen represented by figure 2, 
PI. LXV, exclusive of the cephalothorax, about 
6'5 mm., but the size is somewhat variable. 

Male. — The male is very small, and, viewed laterally, 
the height is equal to about one and a half times the 
length; the dorsum tapers upwards and assumes a 
somewhat conical outline, but with the apex boldly 
rounded. Both pairs of maxillipeds short but tolerably 
stout and strongly uncinate ; other cephalothoracic 
appendages small and somewhat rudimentary. Length 
about 1 mm. 

Habitat. — Found parasitic on the whiting {Gadus 
merlangus) and other Gadoids. This tolerably common 
species has been recorded from various places round 
the British coasts. Larne and Dublin {W. Thompson). 
Polperro and Falmouth {A. M. Norman). Plymouth 
{B ass ett- Smith). Irish Sea {A. Scott), and also from 
various Scottish localities. 

A young Clavella uncinaua with two males adhering to it 
is represented by fig. 6 on PL LXV. In this specimen 
the genital segment is much less rohust, and the mouth- 
appendages are more rudimentary. 



21(3 BRITISH PAKASITIC COPEPODA. 

2. Clavella rugosa (Kiojer). 
(Plate LXV, figs. 4, 5 ; Plate LXVI, figs. 1-7.) 

1837. Anchorella rugosa Kroyer. .(70) pi. 1. vol. i, p. 294, pi. iii, fig. 6. 

1850. Anchorella rugosa Baird. (4) p. 338, ^\. xxxv, fig. 8. 

1851. Anchorella rugosa P. J. van Benecleii. (11) p. 114, pi. vi, fis:j. 7. 
1900. Anchorella rugosa T. Scott. (112) p. 176, pi. viii, figs. 45-48. 

Female. — Ceplialothorax tolerably elongated and 
vermiform, about one and a half times longer than the 
genital segment ; nsnally more or less recurved and 
reaching considerably beyond the distal end of the 
segment. Genital segment stout, rugose, quadriform, 
and somewhat flattened, length and width about equal. 
Abdomen obsolete or nearly so. 

Antennules composed of three joints, the first 
moderately large and stout, the other two smaller 
but the end joint rather longer than the preceding one ; 
antenna stout and somewhat similar in structure to 
those of BracMella. Mandibles small, slender, and 
coarsely dentate near the distal end of the inner 
margin. Maxillae furnished with three moderately 
stout terminal spines, and two smaller ones on a some- 
what rudimentary subterminal lobe. First maxillipeds 
short, stout, and strongly uncinate. Second maxilli- 
peds rudimentary, being reduced to a small button-like 
projection at the anterior end of the genital segment, 
which is fixed to a horn-coloured chitinous plug that 
penetrates the tissues of the fish. Egg-strings toler- 
ably stout and elongated, more than twice the length 
of the genital segment, and containing numerous ova. 
Length exclusive of the ceplialothorax about 4 mm., 
but the size is slightly variable. 

Male. — The male is very small and short, the height 
being equal to fully one and a half times the length ; 
the general outline, seen from the side, is subconical. 
Both pairs of maxillipeds, though short, are furnished 
with stout terminal claws. 

Hahitat. — Parasitic on the gills and gill-covers of 
cat-fishes (Aiiarrhlrhas hijms). On cat-fishes captured 



CLAVELLA liUGOSA. 217 

in the Firths of Forth and Clyde, off Aberdeen, and in 
the Moray Firth (T. Scott). Off the coast of Northum- 
berland and Durham (May E. Bamhriclge), Larne, 
Ireland {W. Thonq^soii). 

3. Clavella dubia T. & A. Scott. 
(Plate LXV, %. 7; Plate LXVI, figs. 8-11.) 

1900. Anchorella rugosa var. T. Scott. (112) p. 177, pi. viii, fig. 52. 

Female. — Generally resembling the female of C. itn- 
cinatus, but having the cephalothorax rather shorter, 
and the second maxillipeds or fixative organ scarcely 
so prominent; the genital segment also being more 
robust. 

Antennules short and three-jointed, the first joint 
robust, but tapering quickly towards the distal end ; 
the second small, and the end one about twice as long 
as the preceding joint, and very sparingly setiferous 
at the apex. Mandibles tolerably stout, somewhat 
expanded near the middle, and the biting part of the 
inner margin provided with about four coarse teeth 
and two or three smaller ones ; the first two teeth 
nearly close together, a slight gap in which is a small 
denticle dividing these two from the next one, but a 
considerable distance separating it from the end tooth, 
and the intervening space occupied by tw^o denticles. 
Maxillge moderately elongated, and provided w^ith two 
narrow^ terminal lobes ending in tolerably long spines. 
First maxillipeds short, moderately robust, and pos- 
sessing fairly stout terminal claws. 

Habitat. — Found adhering to a haddock (Gadus 
deglefinus) from the North Sea. 

This form was at first regarded as a variety of Clavella 
riigosa, but further examination reveals differences both in its 
form and structure which, though showing some relationship 
wdth its two nearest allies, C. rugosa and C. uncinata, cannot 
be reconciled with either. We therefore prefer to describe it 
under a distinct name. 



218 BKITISH PAKASITIO COPEPODA. 

4. Clavella emarginata Kroyer. 
(Plate LXIX, %. 7 ; Plate LXVI, fig. 17-20.) 

1837. Anchorella emarginata Kroyer. (70) pi. i, vol. i, p. 287, pi. iii, fig. vi. 
1851. Anchorella emarginata P. J. van Beneden. (11) p. 113, pi. vi, fig. 4. 
1877. Anchorella emarginata Kurz. (71a) Zeitschr. f. wiss. Zool., vol. 
xxix, p. 398, pi. XXV, fig. 8. 

1900. Anchorella emarginata T. Scott. (112) p. 176, pi. viii, figs. 49-51. 
1906. Clavella emarginata Brian. (21) p. 109, pi. x, fig. 3. 

Female. — Cephalothorax very long and slender, 
twice as long as the genital segment ; head small and 
separated from the long neck hj a slight constriction. 
Genital segment subquadriform, considerably dilated, 
and somewhat truncated at the posterior end. Ab- 
domen obsolete or nearly so. Egg-strings short and 
tolerably thick. 

Antennules short, composed of three articulations. 
Mandibles small and armed with only moderately 
large teeth; maxillas narrow, provided with three 
apical spines and with two small ones on a rudimen- 
tary branch near the middle of the joint. First maxil- 
lipeds moderately small and uncinate. Second maxil- 
lipeds moderately thick, very short, and not completely 
united except at the tip, where they are joined together 
to a chitinous plug. Length. — The specimen repre- 
sented by the drawing measures, exclusive of the 
cephalothorax, about 2*5 mm., while the cephalo- 
thorax alone measures 3*5 mm. 

We have not seen the male of this species. 

Habitat. — Parasitic on tlie gill-arches of the alhs 
shad {Clnjjea alosa) and of the twaite shad (Cliijyea 
finfa). V\ jmovith o\\ Glnpea {Bas^eM- Smith). Dunbar, 
Firth of Forth, on Clwpea finta (T. Scott). 

This species is readily distinguished by the long, slender 
cephalothorax, the dilated genital segment^ and by the 
imperfectly united second maxillipeds. 

5. Clavella scombri Kurz. 
(Plate LXVII, figs. 1-3 ; Plate LXVIIT, figs. 1-7.) 

1877. Anchorella scombri Kurz. (71a) Op. cit., p. 403, pi. xxv, figs. 12, 
35,41. 



CLAVELLA SCOMBRI. 219 

1901. Aachorella scomhri T. Scott. (113) p. 135, pi. viii, fig. 3. 
1906. Anchorella scomberi A. Scott. (110) p. 53, pi. vii. 
1906. Clavella scombri Brian. (21) p. 116, pi. x, fig. 6. 

Female. — Ceplialothorax extremely elongated, flex- 
uose, vermiform, and nearly tliree times as long as 
the genital segment, the proximal half moderately 
stout, the distal portion rather more slender. More- 
over, the proximal portion of the cephalothorax bends 
gently downwards, but near the middle there is a 
somewhat abrupt change of curvature in the opposite 
direction, which causes the cephalothorax to assume 
a geniculated or sigmoid appearance. Genital seg- 
ment of a subglobular form, rather longer than broad, 
and small in comparison w^ith the cephalothorax, also 
appearing to be unprovided with distal appendages 
of any kind. Abdomen rudimentary or obsolete. 
Egg-strings small and saccate. 

Antennules small, two-jointed, and furnished with a 
few small apical setas. Antennae short, stout, and 
somewhat rudimentary ; outer ramus very small. 
Mandibles and other mouth-organs similar to those of 
Clavella emarijinata. Second maxillipeds entirely 
coalescent, extremely short, and united at the apex 
to a small horn-coloured chitinous plug. Length, ex- 
clusive of the cephalothorax, about 2*5 mm. Length 
of cephalothorax fully 6 mm. 

Male. — Extremely small, and its form that which 
appears to be characteristic of the males of this genus, 
being very short and with the dorsum elevated so that 
the height is distinctly greater than the length. Both 
pairs of maxillipeds short, stout, and strongly uncinate. 

Hahitat. — Parasitic on the gill-arches of mackerel 
{Scomber scomhrus). Found on mackerel captured 
in the neighbourhood of Aberdeen in August, 1900 
{T. Scott). On mackerel captured in the Irish Sea 
off Walney Island, and in Carnarvon Bay in 1905 
{A. Scott). Recorded by Dr. Ed. Grraeffe as parasitic 
on mackerel captured in the Gulf of Trieste.* 

* 'Arbeiten dcr zoolog. Institut zii Wien,' T. xiii. Heft 1, p. 17 (1900). 



220 BRITISH PAHASITIC COPEPODA. 

6. Clavella brevicollis M. Edwards. 
(Plate LXV, fig. 1 ; Plate LXVI, figs. 12-16.) 

1840. Anchorella brevicollis M. Edwards. (43) vol. iii, p. 518. 

1877. Anchorella brevicollis C. Yogt. (142) p. 432. 

1901. Anchoi'dla brevicollis T. Scott. (113) p. 135, pi. viii, figs. 11-16. 

Female. — Resembling Clavella uncinata in some 
respects but differing considerably in the following 
particulars : cephalothorax comparatively much shorter 
and stouter, and appearing rather to be a prolonga- 
tion of the posterior portion of the body forwards, 
the only limitation being a slight constriction imme- 
diately behind the second maxillipeds. Genital seg- 
ment subglobular, rather longer than broad, and 
without posterior appendages. Abdomen small, dis- 
tinct. Antennules short, two-jointed, basal joint large, 
end joint small and furnished with a few apical 
setEe. Antennae simple, rudimentary, but stout and 
apparently uniarticulate. Mandibles slender, their 
armature consisting of a few comparatively large 
teeth with intermediate smaller ones. Maxillae mode- 
rately stout, provided with two subapical processes 
bearing short terminal spines, and two minute lateral 
spines on the same side as the processes. The first 
maxillipeds large, with strong terminal claws. The 
second maxillipeds entirely coalescent, very short and 
stout, and terminating in a chitinous knob. Length 
about 4*5 mm. 

Male not observed. 

Habitat. — Parasitic near the base of the anal fin of 
a haddock captured in the Firth of Forth in January 
1896. P. J. van Beneden records this species as 
parasitic on the body (not on the gills) of haddocks ; 
so also does M. Edwards. 

7. Clavella stellata (Kroyer). 
(Plate LXV, figs. 8, 9.) 

1837-1838. Anchorella stellata Kroyer. (70) pi. i, vol. ii, p. 142, pi. iii, 
fig. 5. 

1877. Anchorella stellata C. Yogt. (142) vol. xiv, p. 432. 



CLAVELLA 8TELLATA. 221 

1900. Anchorella stellata T. Scott. (112) p. 178. 

1901. Anchorella stellata idem. (113) p. 134, pi. viii, figs. 1, 2. 

1909. Anchorella stellata May E. Bainbridge. (3) p. 57, pi. ii, figs. 
38-42. 

Female. — Cephalo thorax tolerably elongated and 
slender, and appearing to be but a prolongation of 
tlie coalescent second maxillipeds, having the head- 
appendages at the end of it. Genital segment short, 
moderately stout, ovoid in shape, and joined to the 
cephalothorax by a narrow neck. Abdomen rudi- 
mentary, appearing as a slightly produced middle 
portion of the posterior end of the genital segment ; 
on each side of the abdomen a small tubercle, which, 
with the rudimentary abdomen, imparts a slightly 
trilobed appearance to the end of the segment. 

The two pairs of antennae, the mandibles, and other 
mouth-organs are apparently similar to those of 
Glavella uncinata. The whole animal including the 
maxillipeds appeared to be, in the living state, 
surrounded by semi-transparent gelatinous matter 
through which some of the structures could be plainly 
seen ; the two second maxillipeds were visible within the 
gelatinous investment, extending alongside each other 
to where they joined the chitinous disk by which the 
parasite was anchored to its host. All the specimens 
observed were fixed to scales, and the chitinous fixture, 
after piercing the outer surface of the scale, spread 
out into an extremely thin and dark horn-coloured 
circular disk ; the disk was ornamented all round with 
pellucid, oval markings arranged at more or less 
regular intervals and in the manner of radii which 
did not quite extend to the circumference of the disk. 
The parasite could not easily be removed without 
also removing the scale to which it was attached or 
by severing the attachment where it penetrated the 
scale. The specimen represented by the drawing 
(PI. LXV, fig. 8) measured from the forehead to the 
apex of the second maxillipeds about 6 mm., and the 
genital segment about 4 mm. in length by fully 2 mm. 
in thickness. 



222 BRITISH PARASITIC COPEPODA. 

No males have been observed. 

Habitat. — Parasitic on the skin of the hake {Merliic- 
cius vulgar is). The specimens were found adhering 
to scales near the base of the pectoral and ventral 
fins of a hake captured in the Firth of Clyde in 
December 1899. The whole parasite was enveloped 
in some soft, nearly transparent, jelly-like matter, and 
at first sight had the appearance of a small roundish 
mass of mucus, and thus easily escaped notice : when 
preserved, the mucus assumed a whitish colour. Miss 
Bainbridge (op. cit.) described some specimens found 
on the skin of a hake, " purchased at Sheringham, 
Norfolk." 

8. Clavella paradoxa P. J. van Beneden. 
(Plate LXVII, figs. 4-6 ; Plate LXVIIT, figs. 8-14.) 

1851. Anchorella paradoxa P. J. van Beneden. (11) vol. xvi, p. 117, 
pi. vi, fig. 1. 

1896. Anchorella paradoxa Bassett-Smith. (6) p. 15, pi. v, fig. 2. 
1906. Anchorella paradoxa A. Scott. (110) p. 53, pi. vii. 

Female. — Cephalothorax tolerably elongated, usually 
deflected backward upon the genital segment, and 
extending somewhat beyond its distal end. Genital 
segment subquadriform, the postero-lateral corners 
prolonged backward and slightly outward in the form 
of narrow subtriangular processes, their width at the 
base being equal to about half the length and with the 
apex bluntly pointed; genital segment exclusive of 
the lateral prolongations about as broad as long. 
Abdomen intermediate between the processes, com- 
paratively rather longer than broad and bluntly 
rounded at the end. Three shallow knobs at the 
anterior end of the genital segment, on the dorsal 
aspect, one on each of the flatly-rounded corners, and 
one intermediate and somewhat behind the others ; 
two similar but smaller knobs also present on the fore- 
head as shown in the figure (PI. LXVII, fig. 4). 

Antennules nearly as in Clavella scombri, but the 
antennae, which are three-jointed, are scarcely so 



CLAVELLA PARADOXA. .223 

robust. Mandibles somewhat similar to those of that 
species, but the maxillae are slender and provided with 
three digitate terminal processes. First maxillipeds 
similar to those of Glavella scombri ; second maxillipeds 
very short, coalescent, and fixed to a terminal chitinous 
plug. Length, exclusive of the cephalothorax, about 
4 mm. 

Male. — Very small, nearly globular in shape, and 
somewhat similar in structure and appendages to the 
males of other species of Glavella.^ 

Habitat. — Parasitic on the gill-filaments of mackerel 
(Scomber scombrus). Plymouth (Bassett- Smith). Irish 
Sea {A. Scott). 

9. Clavella quadrata Bassett-Smith. 
(Plate LXVIII, fig. 15.) 

1896. Anclwrella quadrata Bassett-Smith. (6) p. 15, pi. iv, fig. 5. 

1899. Anclwrella quadrata idem. (8) p. 504. 

1906. Anchorella quadrata Norman & T. Scott. (88) p. 221. 

Female. — Cephalothorax much longer than the 
genital segment. Grenital segment almost quadri- 
lateral in outline. Abdomen of extraordinary size for 
animals of this genus; somewhat club-shaped, and 
equal to about two-thirds the length of the genital 
segment. Egg-strings small and broadl}^ ovate. First 
pair of maxillipeds placed close to the mouth, well 
developed ; second pair short, opposite to the abdomen 
and at the base of the neck, thick and completely 
united, terminating in the organ of adhesion, which 
has the form of a cup with a long pedicel. 

This species is much like Anchorella falax Heller, in 
form, except for the great size of the abdomen. 

The male has not been observed. 

Habitat. — Parasitic on the gill-arches of Callionymus 
Jyra, at Plymouth ; only a few sjDecimens were obtained 
{B assett- Smith) . 

* One of the authors, when first recording the male of C.paracloxa, states 
that he could only find one pair of maxillipeds (see ' Trans. Biol. Soc. Liver- 
pool,' vol. XX, p. 53, 1906). This was evidently due to the second pair having 
been accidentally destroyed : he has since found that there are two pairs, 
as in other males of the same irenus. 



224 BRITISH PARASITIC COPEPODA. 

The above is quoted from Dr. Bassett-Smitli's description 
of this species, as we have not ourselves observed it. 

10. Clavella canthari (Heller). 
(Plate LXIX, figs. 1-3.) 

1863. (?) Anchorella pagelli Kroyer. (71) p. 295, pi. xvi, fig. 3. 

1865. Anchorella canthari Heller. (58) p. 212, pi. xxiv, fig. 6. 

1877. Anchorella pagelli C. Yogt. (142) p. 432. 

1880. Anchorella canthari Richiardi. (104) p. 152. 

1906. Clavella macrotrachelus Bnan. (21) p. 116. 

1910. Lerneomyzon canthari T. R. R. Stebbing. (125) p. 562. 

Female. — Small and tolerably robust, but the cephalo- 
thorax is ratlier slender and considerably longer than 
tlie genital segment, being equal to fully one and a 
half times the length of that segment ; the proximal 
end of the cephalothorax bluntly rounded and on each 
side bearing a small but fairly distinct lobe ; a minute 
process arising from between the two lobes, and termi- 
nating in a fascicle of chitinous bristles which penetrates 
the tissue of the gill filament and then gradually be- 
comes separated and forms a brush-like appendage. 
Genital segment moderately stout and subcylindrical, 
the width equal to rather more than half the length. 
A small gibbous projection at the proximal end of the 
segment, as Aaewed from the side, separated from the 
segment by a fairly distinct constriction, and forming 
the base of the cephalothorax. The distal end of the 
genital segment truncated and obscurely trilobate, but 
only the middle lobe fairly distinct ; the lateral lobes 
indistinct and scarcely produced, but the middle one 
fairly prominent and thickly covered with minute 
hairs. Egg-strings moderately elongated. Length. — 
The following measurements are taken from a fairly 
typical specimen : — 

Genital segment . . . about 1'8 mm. 

Cephalothorax . . . about 2*5 mm. 
Egg-strings . . . about 3*0 mm. 

Male. — Small, somewhat similar to the male of C. 
(data, Brian. 

Habitat. — Parasitic on the o-iH-filaments of the black 



CLAVELLA CANTHARI. 225 

sea-bream {Gantharus lineatuH) and the common sea- 
bream (Pagelhcs centrodonhis). On Gantharus lineatus, 
captured in the North Sea in November 1910, as well 
as on a few subsequent occasions. Kroyer obtained 
what appears to be the same species on Fagellus 
centrodontus. 

The characters by which this species seems to be dis- 
tinguished are the proportional length of the cephalothorax, 
the small lobe on each side of its promixal end, the peculiar 
appendage by which the species is anchored to the gill 
filament, and the densely ciliated central lobe of the posterior 
margin of the genital segment. Some of the specimens 
examined by us have exhibited slight differences, but they 
all agree in the more important characters mentioned above. 

11. Clavella alata Brian. 
(Plate LXIX, figs. 4-6.) 

1906. Clavella alata Brian. (21) p. 114, pi. iii, fig. 5 ; pi. xx, figs. 5 and 6. 

Female. — Short and stout. Cephalothorax reaching 
only to about, or slightly beyond, the end of the genital 
segment; at the base of the cephalothorax a small 
horn-like process projecting outwards on each side. 
The second maxillipeds apparently completely coalesc- 
ent, very short, tapering slightly towards the tip, 
which is provided with a rounded cartilaginous knob. 
Genital segment short and stout, subglobose or pyri- 
form, greatest width about equal to the length, pos- 
terior end subtruncate or broadly rounded ; the middle 
portion, however, slightly produced, and on each side of 
this slightly produced part is the attachment of the short 
and tolerably stout ovisacs. The antennules, antennas, 
and several mouth-appendages apparently not differing 
greatly from those of other species of the genus. 
Antennules short, moderately stout, indistinctly seg- 
mented, and bearing a few apical setae ; mandibles 
very slender, but the first maxillipeds tolerabl}^ en- 
larged and uncinate. The entire length, exclusive of 
the egg-strings, is about 3 mm., while the egg-strings 
are about 2 mm. in length. Colour yellowish. 

VOL. I. 15 



226 BRITISH PARASITIC COPEPODA. 

The males, which are very small, measure only about 
0*3 mm., and are provided with large uncinate maxil- 
lipeds. Males, however, do not appear to be very 
common. 

Habitat. — Parasitic on the greater fork - beard 
{Phycis hlennoides (Briin.)). It is usually found adher- 
ing to the gill-arches and not to the filaments of the 
gills. 

One of the more obvious characters of this species seems to 
be the small but quite distinct horn-like processes at the base 
of the cephalothorax, which can be seen — one on each side — 
extending outwards at about right angles to the cephalo- 
thorax. 

12. Clavella lophii (M. Edwards). 
(Plate LI, fig. 7.) 

1840. P Brachiella lophii M. Edwards. (42) vol. iii, p. 514, pi. xli, fig. 4. 
1877. ? Brachiella lophii Vogt. (142) vol. vi, p. 426. 

Female. — A small and robust species. Cephalo- 
thorax moderately short, subcylindrical or subconical, 
stout but less robust in front. Grenital segment some- 
what dilated, widest posteriorly, postero-lateral corners 
rounded. Abdomen very short. Egg-strings short, 
saccate, rather longer than the genital segment, and 
containing tolerably large ova. 

Mouth-appendages apparently somewhat similar to 
those of G. riigosa. The second maxillipeds very short 
and coalescent, and terminating in a small, horn- 
coloured, chitinous button. ^Colour of the specimen 
yellowish. Length about three and a half millimetres. 

Male. — Very small and of the usual Clavella 
{Ancliorella) type; short, and with the dorsum boldly 
arched. 

Habitat. — Found adhering to the gills of an angler- 
fish {LopJiiiis piscatorius) ; locality uncertain. 

We are indebted to Mr. W. M. Tattersall, M.Sc, Keeper 
of the Manchester Museum, for the privilege of describing 
this species. The drawing is made from the mounted specimen. 



AEGULUS. 227 



Genus ARGULUS Milller, 1785. 

Body flattened, ceplialothorax scutiform; cephalon 
and first thoracic segment coalescent ; other segments 
free. Abdomen small, comparatively narrow, and 
ending in two equal lobes, unsegmented. Antennae 
two pairs, small, first pair tolerably stout and termi- 
nating in a small hook. Mouth siphon-like, enclosing 
a flexible, sharp-pointed, sting-like process furnished 
with a poison-gland. Two pairs of maxillipeds, the 
first pair transformed into sucking-discs; posterior 
pair prehensile, their integument rough with minute 
prickles, and having a tridentate plate on the inferior 
aspect of the basal joint. Swimming-legs four pairs, 
biramose, and furnished with plumose hairs. Females 
without external egg-strings. Eyes conspicuous. 

1. Argulus foliaceus (Linn.). 
(Plate LXXII, figs. 1-10.) 

1758. Monoculus foliaceus Linn. Systema Naturae, lOtk edit., vol. i, 
p. 634. 

1762. Binoculus gasterostei Geofroy. Insectes de Paris, vol. ii, p. 661. 

1785. Argulus foliaceus Miiller. Entomost., p. 123. 

1785. Argulus charon idem. Ibidem, pi. 20, figs. 1, 2. 

1793. Monoculus arguluso Fabr. Entom. Syst., vol. ii, p. 489. 

1798. Monoculus gyHni Ciivier. Tab. element. Hist. Nat., p. 45. 

1802. Ozolus gasterostei Latreille. Hist. Nat. Crust, et Ins., vol. iv, 
p. 128, pi. 29, fig. 4. 

1814. Argulus m-gulus Leach. Edin. Encyclop., vol. vii, p. 388. 

1839. Argulus foliaceus Thompson. Ann. and Mag. Nat. Hist., vol. v, 
p. 221. 

1845. Argulus foliaceus Yogt. Nouv. Mem. Soc. Helvet.,vol. vii, pi. i, 
fig. 2. 

1850. Argulus foliaceus Baird. Entom., p. 255, pi. xxxi, figs. 1, 2 a-l. 

Carapace of a rounded oval shape, rather longer 
than broad, and with the front margin slightly pro- 
jecting forward ; posterior sinus narrow and extending 
forward for about two-fifths of the carapace. Abdo- 
men small, rather longer than broad, and about one- 
fourth the length of the cephalothoracic shield ; lobes 
hluntly rounded at the end, separated by a sinus 
extending forward to about half the length of the 



228 BRITISH PARASITIC COPEPODA. 

abdomen ; inner edges proximally contiguous and 
somewhat divaricate at the distal extremity. An- 
tennae normal. Sucking-discs of moderate size. Pos- 
terior maxillipeds tolerably large and stout, their 
basal plates somewhat expanded posteriorly ; the distal 
margin of the plates with three short and subequal 
tooth-like projections. Swimming-legs tolerably elon- 
gated. Eyes conspicuous. A black mark on each 
side near the base of the abdomen. Carapace of a 
greenish colour and ornamented on both sides with 
darker coloured ramifications. Size of the female 
about 6 or 7 mm. in length. Male somewhat like the 
female, but smaller. 

Hahifat, — Parasitic on various fresh-water fishes, as 
carp, sticklebacks, trout, pike {Esox hiciits), greyling 
{Thymallus vitlgaris), and others. The species appears 
to be generally distributed throughout the British 
Islands. Some years ago it was very common on 
greyling in the upper waters of the Clyde, but after a 
few days' heavy rain Avhich flooded the river the ArgnJus 
had all disappeared. This species has also been obtained 
on a fish captured in the Faroe Channel by G. H. 
Fowler. The specimen is now in the British Museum, 
and we are indebted to Dr. W. T. Caiman for drawing 
our attention to it and for the loan of it for exami- 
nation. 

There appears to he some uncertainty as to the relationship 
between the Argnlidae and the species usually recognized as 
Copepoda. Dr. Baird includes the Family Argulidae in. the 
same Tribe — Peltocephala — with the Caligidge, tlie Pandarid^e, 
and the Cecropid^.^ In 1866 Gerstaecker, revising liis 
previous classification, replaces the Argulidae under the 
Branchiopoda.t Clans in 1875 proposed making the Argulida) 
a second suborder of the Branchiura, under the Order Cope- 
poda. J On the other hand, the Rev. T. 11. R. Stebbing, in 
'A History of Crustacea/ places the carp-lice (Argiilu.s) in 
the suborder Branchiura of tlie Order Branchiopoda, and 

* * Entomostraca,' p. 15. 

t C/._Bronn's 'Klassen unci Ordnungen des Thierreichs/ vol. v, p. 16. 

X " tJber die Entwickelung, Organisation, imd Systematische Stellung^ 

der Arguliden," ' Zeitsch. f. wissensch. Zool./ vol. xxv, pp. 217-2S4, pis. 



ARGULUS FOLIACEUS. * 229 

not with the Copepoda."^ Dr. Bassett-Smith, in his ^System- 
atic description of the parasitic Copepoda found on Fishes/ 
also excludes the Family Argulidae from tlie Order Cope- 
poda.t And lastly, in the first of an important series of 
* Memoirs on North American parasitic Copepods/ by Charles 
Branch Wilson, that author makes the Branchiura a suborder 
of the Copepoda, with Argulidas as the only Family. J 

There is doubtless a close relationship between the Argulidas 
and the other parasitic groups enumerated in this volume; 
but apparently, as indicated, there is also a considerable 
divergence of opinion as to whether Arguhts and its allies 
should be recognized as a suborder of the Copepoda. We 
have, however, had few opportunities for studying these 
interesting forms, as there is only a single British species 
known to us, and therefore we do not propose to enter further 
into a discussion of this question. 



Tripaphylus musteli (P. J. van Beneden). pp. 160- 

161. 

Amended description of male. 

Our description of the male of this copepod parasite was 
copied, to some extent, from van Beneden^s work, as we had 
not then seen the male ourselves. We discovered a male 
attached to a female after the description was in type, and 
some changes appear to be necessary to bring the description 
more in line with the views now generally accepted regarding 
the names of the appendages, but it was too late to do this in 
the proper place. This description, taken from our own 
specimen, should read as follows : — 

Male. — The male of this species has the body divided into 
two unequal portions ; the anterior portion is large and carries 
the whole of the appendages — viz. one pair of antennules, one 
pair of antennae, one pair of mandibles, one pair of maxillae, 
and two pairs of maxillipeds; the posterior portion is much 
more narrow^ and rounded and is terminated by short stout 
furca. The antennules, antennee, mandibles, and maxillae are 

* ' A History of Crustacea : Recent Malacostraca.' The International 
Scientific Series, vol. Ixxiv (1893), pp. 10 and 49. 

t "A systematic description of the Copepoda found on fishes, with an 
enumeration of the known species," ' Proc. Zool. Soc. Lond.,' April, 1899. 

t " North American Parasitic Copepods of the Family Argulidse," ' Proc. 
United States National Museum,' vol. xxv (1902), pp. 635-742, pis. viii- 



230 



BRITISH PARASITIC COPEPODA. 



somewhat rudimentary. The two paiis of maxillipeds are 
comparatively well developed and form prehensile appendages 
similar to those found in the males of many of the sedentary 
female fish-parasites. The appendages described as feet by 
van Beneden, and also in the second paragraph on p. 161, 
are the first and second maxillipeds. No appendages corre- 
sponding to feet in the more highly organized parasitic 
Copepoda appear to be present. 



List of the Fishes on which the Parasitic Copepoda 
described in this volume were obtained. 



Acipenser sturio Linn. 
Agonus cataphractus Linn. 
Alopias vulpes [Gmelin) . 
Anarrhichas lupus Linn. . 
Bothus maximus (Linn.) . 
Bothus rhombus [Linn.) 
Brosmius brosme Cuv. 
Callionymus lyra Linn. 
Callionymus maculatus Linn. . 
Cantharus lineatus (Mont.) 
Car ch arias glaucus (Linn.) 
Clupea alosa Linn. 
Clupea finta Cuv. 
Clupea sprat tus Linn. 
Conger niger Risso . 
Corregonus pollan Thomps. 
Crenilabrus melops (Linn.) 
Ctenolabrus rupestris (Linn.) 
Cyclopterus lumpus Linn. . 
Cyprinus carpio Linn. 
Drepanopsetta platessoides 

(Fahr.) 
Engraulis encrasicholus Linn. . 
Esox lucius Linn. 
Gadus aeglefinus Linn. 
Gadus callarius Linn. 
Gadus luscus Linn. . 
Gadus merlangus Linn. 
Gadus minutus Linn. . 
Gadus pollachi us Li7i7t. 



Sturgeon. 

Pogge. 

Thrasher Shark. 

Cat-fish; Wolf-fish. 

I'urbot. 

Brill. 

Torsk or Tusk. 

Gemmeous Dragonet. 

Spotted Dragonet. 

Black Bream. 

Blue Shark. 

Allis Shad. 

Twaite Shad. 

Sprat. 

Conger. 

Pollan. 

Connor; Goldsinny. 

Jago's Goldsinny. 

Lumpsucker. 

Carp. 

Long Eough Dab. 

Anchovy. 

Fresh-water Pike. 

Haddock. 

Codfish. 

Bib ; Whiting Pout. 

Whiting. 

Poor Cod. 

Pollack. 



LIST OP THE FISHES. 



231 



Gadus virens Linn. . ... 
Galeorliinus galeus {Linn.) 

(Galeus canis Bonap.) 
Gastei'osteus aculeatus Linn. 
Gastraea spinachia {Linn.) 
Gobius miiiutus Gmelin 
Hippoglossus vulgaris Flem. 
Labrax lupus Guv. Sf Val. . 
Labrus bergylta Ascan. 
Labrus mixtus Linn. . 
Lainna coruubica Cuv. 
Lampris pelagicus {Gun.) . 
Leuciscus rutilus {Linn.) . 
Lophius piscatorius Linn. . 
Lu varus iinperiab's Rafin . 
Macrurus coelorhynchus Risso . 
Merluccius vulgaris {Cuv.) 
Molva vulgaris {Flem.) 
Mugil chelo Guv. . . . 

Mullus barbatus {Linn.) 
Mustellus vulgaris Mull. ^ Henle. 
Onos cimbrius {Linn.) 
Onos mustelus {Linn.) . . . 

Onos tricirratus {Briln.) 
Orcynus thyunus {Linn.) . 
Orthagoriscus mola {Linn.) 
Pagellus centrodontus {De la 

Roche) 
Perca fluviatilis Linn. 
Pleuronectes cynoglossus Linn. . 
Pleurnoectes flesus Linn. . 
Pleuronectes limanda Linn. 
Pleuronectes microcephalus Don. 
Pleuronectes platessa Linn. 
Pholis gunnellus {Linn.) 
Phycis blennoides {Briin.) 
Raia batis Linn. 
Raia circular is Gouch 
JRaia clavata Linn. 
Raia fullonica Linn. . 
Eaia maculata Mont. . 
Raia oxyrhynchus Linn. 
Rliina squatina {Linn.) 
Salmo fario Linn. 
Sal mo salar Linn. 



Coalfish. 
Tope or Toper. 

Three-spined Stickleback. 

Fifteen-spined Stickleback, 

Speckled Goby. 

Halibut. 

Basse. 

Ballan Wrasse ; Bergylt. 

Striped Wrasse. 

Porbeagle. 

Opah; King-fish. 

Roach. 

Angler or Angler-fish. 

Grenadier-fish. 

Hake. 

Ling. 

Grey Mullet. 

Surmullet ; Red Mullet. 

Smooth-hound. 

Four-bearded Rockling. 

Five-bearded Rockling. 

Three-bearded Rockling. 

Tunny. 

Short Sunfish. 

Sea-bream. 

Perch. 

Pole-dab ; Witch-sole. 

Flounder. 

Dab. 

Lemon- sole. 

Plaice. 

Gunnel; Butter-fish. 

Greater Fork-beard. 

Grey or Blue Skate. 

Sandy Ray. 

Thornback Skate. 

Shagreen Ray. 

Homelyn Ray. 

Long-nosed Skate. 

Angel-fish. 

Trout. 

Salmon. 



232 



BRITISH PARASITIC COPEPODA. 



Salmo trutfca Linn, . 
Sciaena umbra (Lacep.) 
Scomber scombrus Linn. 
Scylliorhinus canicula {Linn.) 

Scy Ilium catulus Cuv. 
Scymnus borealis Flem, 
Sebastes norvegicus {Ascan.) 
Solea vulgaris Quensel 
Squalus acanthias Linn. 

Thymallus vulgaris Nilss. 
Trigla cuculus Linn. . 

(Trigla pini Bloch.) 
Trigla gurnardus Linn. 
Trigla lineata Gmelin 
Trigla lucerna Linn. . 

Trigla lyra Linn. 
Trygon pastiiiaca (Ldnn.) 
Xiphias gladius Linn. 
Zeugopterus punctatus [Blainv.) 
Zeus faber lAnn. 



Salmon trout. 

Maigre. 

Mackerel. 

Kough-hound; Lesser Spot- 
ted Dog-fish. 

Nurse or Nurse-hound. 

Greenland Shark. 

Bergylt; Norway Haddock. 

Black Sole. 

Spur - Dog ; Piked Dog- 
fish. 

Greyling. 

Red Gurnard. 

Grey Gurnard. 
Streaked Gurnard. 
Sapphirine Gurnard; Yel- 
low Gurnard. 
Piper; Crowner. 
Sting-ray. 
Sword-fish. 
Muller's Topknot. 
Dory ; John Dory ; Doree. 



^o^e.-— The scientific names of* the fishes in this list are, with a few- 
exceptions, from 'A History of Scandinavian Fishes,' by B. Fries, 
C. U. Ekstrom, and C. Sundevall, 2nd edit., revised by Prof. F. A. 
Smitt (1893-95). 



(233) 



LITERATURE 

CHIEFLY KEFKKEED TO IN THE PRECEDING PAGES. 

1. 1794 Abildgaard, P. C. Beskrivelse over tvende nye 
MonocuU Liini. (Callgiis crassus et ohlongus.) — 
Skrift. Naturliist. Selsk. Kiobenhavii, III, 2, pp. 
46-54. 

:2. 1865 Agassiz, A. Sur un Pennella, parasite de VOrtha- 
goriscus mola. — Illustrated Catalogue of the 
Museum of Comparat. Zoology, Cambridge, p. 87. 

3. 1909 Bainbridge, May E. (Hon. Mrs. Henn Collins). 

Notes on some parasitic Copepoda with a de- 
scription of a new species of Chondr acanthus 
{C. inflatus). — Trans. Linn. Soc. London, ZooL, 
Ser. 2, Vol. XL part 3. 

4. 1850 Baird, Dr, W. The Natural History of the British 

Entomostraca. — Ray Society. London. 

5. 1783 Barbut, /. The genera Vermium exemplified by 

various specimens of the animals contained in 
the orders of the Intestina et Mollusca Linnsei. 
Drawn from Nature. London. 

6. 1896 Bassett-Smith, Dr. P. W. Notes on the Para- 

sitic Copepoda of fish obtained at Plymouth 
with descriptions of new species. — Ann. and 
Mag. Nat. Hist. (6), Vol. XVIII, pp. 8-16, with 4 
plates. 

7. 1896 idem. List of Parasitic Copepoda of fish obtained 

at Plymouth. — Journ. Mar. Biol. Assoc, N.S., 
Vol. IV, pp. 153-163. 

8. 1899 idem. A Systematic Description of Parasitic Cope- 

poda found on fishes, with an enumeration of 
the known species. — Proc. Zool. Soc. London, 
1899, Pt. IT, pp. 438-507, with one plate. 

9. 1765 Baster, Job, Opuscula subseciva. Vol. II, p. 138. 

Harlemi. 
10. 1905 Baudouin, M, Les parasites de la Sardine. — Rev. 
Scient. Paris, No. 23, 10 Juin, pp. 715-722, with 
4 text-figs. 



234 



BRITISH PARASITIC COPKPODA. 



11. 1851 Bene den, P. ,7. van. Recherches siir quelques 

Crustaces inferieurs. — Aiiti. Scieiic. Nat., 3 Sei\ 
ZooL, Vol. XVI, pp. 71-131, pis. 2-6. 

12. 1851 idem. Note snr uii Crustace parasite nouveau,. 

avec renumeratjon des especes de cette classe 
qu'on observe sur les poissoiis du littoral de 
Belgique.— Bull. Acad. Roy. de Belg., T. XVIII, 
pt. 1, pp. 286-290 avec pi. 

13. 1852 idem. Note sur un nouveau genre de Crustace 

parasite de la famille des Peltocepliales. — Bull. 

Acad. Roy. de Belg., T. XIX, pt. 3, pp. 462-467, 

avec pi. 
13a. 1852 idem,. Note sur quelques parasites d'un poissou 

rare sur nos cotes (le maigre d'Europe, Sciwna. 

aquila Cuv.).— Bull. Acad. Roy. de Belg., T. XIX,. 

pt. 3, pp. 98-109 avec pi. 
13b. 1853 idem. Note sur un nouveau genre de Crustace- 

parasite, Eudactylina [E. acuta). — Bull. Acad. 

Roy. de Belg., T. XX, pt. 1, pp. 235-239. 
13c. 1853 idem. Notice sur un genre nouveau de la tribu des- 

Caligiens (genre Kroyeria van Ben.). — Bull. Acad. 

Roy. de Belg., T. XX, pt. 1, pp. 23-30 avec pi. 

14. 1854 idem. Notice sur un nouveau genre de Siphono- 

stome (genre Congericola). — Bull. Acad. Roy. de- 
Belg., T. XXI, pt. 2, pp. 583-589. 
14a. 1857 idem. Sur un nouveau Dinemoure provenant du 
Scimiiiis glacialiii. — Bull. Acad. Rov. de Belg., (2) 
T. I, pt. 1, pp. 226-235 avec pi. 

15. 1861 idem. Recherches sur les Crustaces du littorale de 

Belgique. Mem. Acad. Roy. de Belg., T. XXXIII, 
n. 3, 174 pp., 21 pis. 

16. 1870 idem. Les poissons des cotes de Belgique, leurs 

parasites et leurs commensaux. — Mem. Acad. 
Roy. de Belg., T. XXXVIII, n. 4, 100 pp., 6 pis. 

17. 1864 Bergsoe, F. P/u7i6'/i^%sa;*27/ii«Stp.,Monograpliisk 

Fremstittet.-Naturliist.Tidkssrift,3Raek., 3 Bind^ 
pp. 87-130, tab. 13, Kjobenhavn; {oppure Ann. 
Scienc. Natur., 5 ser. Zoologie, Vol, III, p. 213, 
pi. I, 1865). 

18. 1898 Brian, A. Note preliminaire sur les Copepodes 

parasites des poissons. — Bull. Instit Oceanogr., 
No. 110, p. 19, with text-figs. (Records ' Pen- 
nella Jilosa ' from Sunfish, &c.). 



LITEllATUHE CHIEFLY REFERRED TO. 235 

19. 1899 Brian, A. (contd.). Di alcuni Crostacei parassiti 

dei pesci dell'Isola d'Elba.. — Atti Soc. Lig. 
Sc. Nat. e Geogr., Vol. X, pp. 1-10, with text- 
figs. 

20. 1899 idem. Diphyllogaster thompsoni n. gen. e sp. di 

Caligidae della Dicerohatis' giornse Griintli. — Atti 
8oc. Lig. Sc. Nat. e Geogr., Vol. X, pp. 1-11, 
con. tav. 

21. 1906 idem. Copepodi Parassiti dei Pesci d'ltalia. Con 

21 Tavole. Genova. 
21a. 1908 idem. Note preliminaire sur les Copepodes para- 
sites des poissons provenantdes Campagnes Scien- 
tifiques de S.A.S. le Prince Albert P^ de Monaco 
ou deposes dans les collections du Musee Oceano- 
graphique. — Ball. Instit. Oceanogr. No. 110. 

22. 1816 Blainville, H. M. D. de. Prodromus d'une Nou- 

velle Classification du Regno Animal. Paris. 

23. 1816 idem. Article Lernasa in Diet. d. Scienc. Natur., 

Vol. XXVI, p. 112, ff., 1816 (e 1823). 

24. 1822 idem. Memoires sur les Lernees. — Journ. de Phy- 

sique, T. 95, pp. 372-380, et pp. 437-447, avec 
Ipl. 

25. 1883 Brady, G. S. Report on the Copepoda (including 

Copepoda parasitic on fishes). — Voyage of H.M.S. 
' Challenger,^ ZooL, Vol. VII, Part XXIII. 

26. 27. 1833 Burmeister, H. Beschreibung einiger neuen 

oder wenig bekannten Sclimarotzerkrebse, nebst 
allgemeinen Betrachtungen iiber die Gruppe, 
welcher sie angehoren. — Nova Acta Acad. 
Leopold.-Carolin., XVII, 1, pp. 269-336, mit 3 
Taf. 

28. 1870 idem. Bomolochus helones, occurrence in Mediter- 

ranean and German Sea. — Hartmann's Archiv» 
Anat. Physiol. (1870), pp. 116-163. 

29. 1892 Canu, E. Les Copepodes du Boulonnais, parasites 

des poissons. Annales de la Statione Aquicola 
de Boulogne-sur-mer (1892). 

29a. 1884 Cams, /. V. Prodromus faunae mediterraneae. 
Pars I, pp. 345-378. Stuttgart. 

30. 1858 Claus, C. Ueber den Bau und die Entwickelungs- 

geschichte parasitischer Crustaceen, pp. 1-34^ 
2 plates. Cassel. 



236 BRITISH PARASITIC COPKPODA. 

30a. 1860 Claus, C. (coutd.). Zur Morphologie der Cope- 
poden. 1. Eine Uemmuiigsbildung von Cyclops. 
2. Ueber den Ban von Nicothoe. 3. Ueber die 
Leibesgliederuiig und die Mundwerkzeuge der 
Sclimarotzerkrebse. — Wiirzb. Naturw. Zeitschr. 
Bd.I, pp. 20-36/JaF. 1. 

31. 1861 idem. Uebei* die Familie der Lerniien : Lern^ocera 

gohina. — Wiirzb. Naturw. Zeitsclir., Bd. II, pp. 
JO-22, Taf. 1. 

32. 1861. idem. Ueber den Ban und die Entwickelung von 

Achieves percarum. Zeitschr. f. wiss. Zool., Bd. 
XI, pp. 287-308, Taf. 23 u. 24. 

33. 1864 idem. Beitrage zur Kenntniss der Sclimarotzer- 

krebse. — Zeitschr. f. wiss. Zool., Bd. XIV, pp. 365- 
382, T. 33-36. 

34. 1868 idem. Beobachtungen iiber Lerndeocera, Peniculus, 

und LeriLfea. Ein Beitrag zur Naturgeschichte 
der Lern«en, mit 4 Taf. — Schriften der Gesell- 
scliaft zur Beforderungder yesammten Naturwiss. 
zu Marburg, 2 Supplement-Heft. 

35. 1875 idem. Neue Beitrage zur Kenntniss parasitischer 

Copepoden nebst Bemerkungen iiber das System 
derselben. — Zeitschr. f. wiss. Zool., Bd. XXV, 4. 
mit 3 Taf. Leipzig. 

36. 1780 Cordiner, Rev. Charles. Lernasa salmonea Linn. — 

Antiquities and Scenery of the North of Scotland, 
pp. 7-8, pi. VI, fig. 2. 
36a. 1865 Gornalia, E. Sulla Lophoura rdwardsii di Kol- 
liker. Osservazioni zoologiche e anatomiche. — 
Atti Soc. Ital. Sc. Nat., Vol. IX, pp. 1-10, tav. 1. 

36b. 1912 Cuenot, M. L. Contributions a la faune du 
bassin d'Arcachon : description d'Argulus arcas- 
sonensis, nov. sp. — Bull, de la Station Biol. d'Arca- 
chon, i4c, pp. 117-127, pis. 1 & 2. (Extrait.) 

37. 1817 Cuvier, G. L. Regno Animal. Premiere edit. 

Tom. Ill, pp. 60-66. Paris. 

38. 1829 idem. Regne Animal. Nouvelle edit. Tom. IV, 

pp. 189-202. Paris. 

39. 1838 Dana, J. D. (see Pickering and Dana). 

40. 1852 idem. United States Exploring Expedition during 

the years 1838-1842, under the command of 
Charles Wilkes. U.S.N. XIII, Crustacea. (Atlas 
Crustacea, Philadelphia 1855.) 



LITERATURE CHIEFLY REFERRED TO. 237 

41. 1811 De la Roche, F. Sur deuxanimaux vivants surles 

branchies des poissoiis. — Noiiv. Bull. Soc. Philo- 
mat., II, pp. 270-272. 

42. 1833 Edwards, H. Milne. Memofre sur Torganisation de 

la bouche chez les Ciustaces suceurs. — Ann. 
Scienc. Natur , Yol. XXIX, pp. 78-86. 

43. 1840 idem. Histoire naturelle des crustaces, comprenant 

Fanatomie, la physiologie, et la classification de 
ces animaux. Paris. 

44. 1763 Ellis, /. Pliilo.<. Trans.^Yol. 53, p. 433.— Penwaht?^ 

filosa and sagitta. 

45. 1909 Elwes, ^. F. The Sunfish and its parasites.— Journ. 

Torquay Nat. Hist. Soc, Yol. I, No. I, pp. 17-20. 

46. 1906 Evans, TF. Lsemargusmuricatusivov[i^^w.n^&la.. — 

Ann. Scott. Nat. Hist. (Jany.). Zoological notes. 

47. 1904 Gadd, Pehr. Parasit-Copepoder i Finland. — Acta. 

Soc. pro Fauna et Flora Fennica, XXYI, No. 8. 

48. 1853-54 Gerstaecker, A. D. Ueber eine neue und eine 

weniger gekannte Siplionostomen-Gattung, und 
Besclireibimg zvveier neuer Siphon ostomen-Gat- 
tuugen. — Wiegniann^s Arcliiv f. Naturg., XIX,. 
1, pp. 58-70, Taf. 3 u. 4; XX, pp. 185-195, Taf. 7.. 

49. 1870-71 idem. In Dr. H. G-. Bronn's Klassen und Ord- 

nungen des Thier-Reichs, Y. Band, Grliederf ussier. . 
Lieferung 11-16. Leipzig u. Heidelberg. 

50. 1889 Giard, J.. Suv Va^ssocmtion de PenneJ la or thagoris-ci,. 

Peicival Wright, et de Conchoderma virgatum 
Spengl.— Le Naturali^te, (2) III, No. 50, p. 82. 

51. 1899 Giesbrecht, TF. Die Asterocheriden des Golfes. 

von Neapel und der Angrenzenden Meeres. — 
Abschuitte. (A.u.d. Tit. : Fauna und Flora des 
Golfes von Neapel, 25 Monogr.) 

52. 1842. Goodsir, H. Note sur le developpement des oeufs. 

du Caligiis et sur les metamorphoses que ce- 
Crustace eprouve. — Ann. Scienc. Natur., 2 ser., 
ZooL, Yol. XYIII, pp. 181-184. 

53. 1827 Grant, E. E. On the structure and characters of 

Lenixa elongata from the Arctic Seas. — Edin- 
burgh Journ. of Science, YII, pp. 147-154. 

54. 1828 idem. Froriep's Notizen, Yol. XIX, p. 18. 

55. 1829-1843 Guerin-MeneviUe, F. E. Iconographie du 

Eegne animal. Crustaces. (Text, 1844.) 



238 BRITISH PAKASITIC COPEPODA. 

56. 1879 Heider, C. Die Gattnng Lemanthropus. — Arbeit. 

ZooLInstit.Wien.n. Triest., T. II, Hft. 3, pp. 269- 
368, mit 5 Taf.— Audi separ.; Wien, Holder, 1879. 

57. 1904 Hofer, B. Handbucli der Fisclikrankheiten. — 

Verlag der allg.. Fischerei-Zeitung. Miinchen. 

58. 1865 Heller, C. Crustaceen der Novara-Expedition. 

59. 1879 Hesse, E. Description des Crustaces rares ou 

nouveaux des cotes de France. — Ann. Scienc. 
Nat. 6 ser., Zool., Vol. VIII, art. 29. 

60. 1884 idem. Crustaces rares ou nouveaux des cotes de 

France. — Ann. Scienc. Nat., 6 ser., ZooL, Vol. 
XVI, art. 34, pp. 1-18, pis. 12-14. 

61. 1783 Hermann,/. Helminthologische Bemerkungen. — 

Naturforscher, n. 19. 

61a. 1857 HoBven, /. van der. Note sur les genres Cecrops 
et Lemargus. — Memoires d'Entoniol. de la Soc. 
Entom. des Pays-bas, I, pp. 67-87. 

62. 1799 Holten, H. 8. Acta Danica, Vol. V. 

63. 1802 idem. Lernsea merlucii og Exocosti, to nye Arter. 

— Skrift. Naturhist. Selsk. Kjobenhavn, V, 2, pp. 
135-137. 

63a. 1897 Horst, R. PhiloHhagoriscus serratns Kr. [Dine- 
matura serrata Kr.) With 1 pi. Notes Leyden 
Mus., Vol. 19, N. J. Note XIV, pp. 137-144. 

64. 1824 Johnston, G. Notice respecting the genus Cali- 

gus of Leach. — Edinburgh Philosoph. Journ. X, 
pp. 292-294. 

65. 1835 idem. Lernsea uncinata. — Loudon's Mag. of Nat. 

Hist., VIII, pp. 565-566. 
Q>Q. 1835 i*t?em. Pandarusalatus nndlamnse. Loc. cit. VIII, 
pp. 202-205. 

67. 1836 idem. Chondracanthus lophii, nov. spec. Loc. cit. 

IX, pp. 81-83. 

68. 1892. Kane, W. V. de Vismes. On a new species of 

Lerndeopoda (hidiscalis) from the West Coast of 
Ireland, and Polperro, Cornwall. — With 2 pis. 
Proc. Koy. Irish Acad. (3) Vol. 2, n. 2, pp. 203- 
211. Abstr. ; Journ. Hoy. Micr. Soc. London, 
1892, Pt. 4, p. 480. 

68a. 1853 KoUiker, A. Bericht iiber eiiiige im Herbst 1852 
in Messina angestellte vergleichend anatornische 
Untersuchungen. Gatt. Lophoura. — Zeitschr. f. 
Wiss. Zool. IV, p. 359. 



LETEltATURE CHIEFLY EEFEKRED TO. 239 

69. 1835 KoUar, V. Beltrage zur Kenntnisslernaeenartigen 

Crustaceen. Annal. des Wiener Museums fur 
Natiirgeschiclite, I, 1, pp. 72-92, mit 2 Taf. 
69a. 1877 Koren, /., and Danielssen, D. C. En ny art af 
slaegten Pennella. — Faun. litt. Norweg. 3 Hft., 
pp. 157-163. 

70. 1837-38 Kroyer, H. Om Snyltekrebsene, isaer med 

Hensyn til den dansk Fauna. — Naturhistorisk 
Tidsskrift, Vol. I. pp. 172-208, pp. 252-304, pp. 
476-505, pp. 605-628. tab. 2-3 u. 5-6— II. pp. 
8-52, pp. 131-157, tab 1 u. 3. Kjobenhavn. 

71. 1863 idem. Bidrag til Kundskab om Snyltekrebsene. — 

Naturhistorisk Tidsskrift, 3 die. Kaekke, II Bind, 
pp. 75-426, tab. 1-18. — separat ; Kjobenhavn. 
71a. 1877 Kurz, W. Studien iiber die Familie der Lernaeo- 
podiden. — Zeitschr. f. wiss. Zool. Bd. XXIX. 

72. J 815-22 Lamarck, G. B. P. de. Histoire naturelle des 

Animaux sans Vertebres, Vol. V. (1818.) Seven 
species of copepod parasites described. 

73. 1814 Leach, W. E. Crustaceology. (In) Edinburgh 

Encyclopedia. 

74. 1816 idem. Supplement. — Annulosa. loc. cit. 

75. 1819 idem. Article Entomostraca in Diet. d. Scienc. Natur. 

76. 1746 Linnaeus, C. Fauna Suecica. 1st Edition. Lug- 

duni Batavorum. 

77. 1761 ideyn. loc. cit. 2nd Edition. Stockholmiae. 

78. 1766, 67, 68 idem. Systema Naturte. 12th Edition. 

79. 1899 Lonnberg, E. Bidrag til Kannedomen om fritt 

lefvande Caligider. — Verh. Biol. Ver. Stock- 
holm, Bd. I, pp. 148-158. 

79a. 1860 Lubbock, /. On some oceanic Entomostraca col- 
lected by Capt. Toynbee. — Trans. Linn. Soc. 
London, XXIII, pp. 173-192, pi. 29. (Describes 
Bacidus elongatus recognized by Mrazek 35 years 
later to be a larval Pennella.) 

SO. 1890 Malard, A. E. Catalogue des Poissons (with their 
parasites). — Bull. Soc. Philomathique, Paris, 8* 
ser. Vol. 2, No. 2. 

^1. 1835-39 Mayer, A i^./.C. Analekton f ur vergleichenden 

Anatoniie. Bonn. 
S2. 1863 M'lntOSh, IF. C. Notes on the food and parasites 

of the Sahno salar Linn, of the Tay. — Journ. 

Linn. Soc. London, Zool., Vol. VII, pp. 145-154. 



240 BRITISH PARASITIC COPEPODA. 

83. 1880 Miers, E. J. On a small collection of Crustacea made- 

by Edmund Wliymper, Esq., clnefly in the N. 
Greenland seas, &c. — Journ. Linn. Soc. London, 
Zool., Yol. XY, pp. 59-73 (records Caligus curtus,. 
Diiiematura feiox, and Lern8eoj)oda elongata). 

84. 1895 Mrazek, AL Ueber Baculus Lubb. und Hessella 

Brady, ein Beitrag zur Anatomic der Lernaeiden. 
Mit 2 Taf. und 2 Holzclm.— Sitzgsber. K. Bohm. 
Ges. Wiss., matli.-nat. CI. XLIY. Ausz. vom.. 
Yerf. Zool. Centrabl. 3 Ihg. n. 7, pp. 237-238. 

85. 1777 Miiller, 0. F. Zoologia Danica. 

SQ. 1785 idem. Entomostraca, sen Insecta testacea quae in 
aquis Daniae et Norvegias reperit. descripsit, et 
iconibus illustravit. LipsiaB et Hafni£e (Leipzig 
and Copenhagen). 

87. 1903 Norman, A. M. New generic names for some 

Entomostraca and Cirripedia. — Ann. and Mag. 
Nat. Hist., (7) XI, pp. 367-3G9. 
87a. 1910 Norman, A. M., and Brady, G. 8. The Crus- 
tacea of Northumberland and Durham. — Trans. 
Nat. Hist. Soc. Northumbeilaiid, Dnrham, and 
Newcastle-upon-Tyne. N.S., Yol. Ill, part 2. 

88. 1906 Norman, A. M., and Scott, T. Crustacea of 

Devon and Cornwall. London. 

89. 1832 Nordmann, J. Mikrographisclie Beitrage zur 

Naturgeschichte der wirbellosen Thiere, 2 Hefte. 
Berlin. 

90. 1864 idem. Neue Beitrage zur Kenntniss parasitischer 

Copepodeu. Erster Beitrag. — Bull, de la Soc. 
des Natur. de Moscou, XXXYII, 2, pp. 461-520, 
Taf. 5-8. 

91. 1815 Oken, L. Lehrbuch der Naturgeschichte. Th. 

Ill, p. 184, ff., und p. 857, IT. [Schisturus replaced 
by Clavella as the first name was occupied by 
Rudolphe in 1809 for a genus of Yernies). 

92. 1868 Olsson, P. Prodromus faunae Copepodorum para- 

sitantium Scandinaviae. Lunds Univ. Arsskrift.,. 
Tom. Y. 

93. 1828 Otto, A. W. Beschreibung einiger neuen, in den 

jahren 1818 und 1819 im mittellandischen Meere 
gefundener Crustaceen. — Nova Acta Acad. Leo- 
pold. Carolin., Yol. XIY, Pt. 1, pp. 351-354-, Mit 
drei Kupfertafelen. 



LITERATUKE CHIEFLY REFERRED TO. 241 

94. 1861 Pagenstecher, Dr. A. H. Ther sites gasterostei, 

eine Gattung parasitischer Crustaceen. — Weig- 
mann's Arohiv. f. Naturgesch. XXVII, 1, pp. 
118-126, Taf. 6. 

95. 1905 Pearson, /. A list of the Marine Copepoda of 

Ireland. I, Littoral forms and Fish Parasites. — 
Fish. Ireland Sci. Invest. 1904, iii. 

96. 1838 Pickering, Ch., and Dana, /. D. Description of 

a species of Caligus {C. americanus). — Silliman's 
Americ. Joiirn. of Science, XXXIV, pp. 225-266, 
pi. 3-5. Read before the Yale Nat. Hist. Soc, 
Feb. 20, 1838. 

97. 1902 Poche, F. B. Bemerkungen zu der Arbeit des 

Herrn Bassett-Smith 'A Systematic Description 
of Parasitic Copepoda found on Fishes, with an 
Enumeration of the known Species ^ — Zool. 
Anzeig., 26 Bd. n. 685, pp. 8-20. 

97a. 1884 RB.th.hlin, Rich. Annotated List of the described 
species of parasitic Copepoda (Siphonostoma) 
from American Waters contained in the United 
States National Museum. — Proc. U. S. National 
Museum, Vol. 7, pp. 483-492. 

98. 1887 ide7n. Descriptions of (4) new species of parasitic 

Copepods belonging to the genera Trehius, 
Ptrissopus, and Lernanthroj>iis . — Proc. U. S. 
National Museum, Vol. 10, pp. 559-571. 

99. 1887 idem. Descriptions of parasitic Copepoda belong- 

ing to the genera Pandariis and Ghrondr acanthus. 
With 7 plates.— Xoc. cit., Vol. 9, pp. 310-324. 
Abstr. in Journ. Roy. Microsc. Soc. 1887, pt. 3, 
p. 395 (4n. sp.). 

99a. 1839 Rathke, S. Bemerkungen iiber den Bau de& 
Diclielesthium stiirionis und der Lernseopoda stel- 
lata. — Nova Acta Acad. Leopold.-Carolin., Vol. 
XIX, 1, pp. 125-168, mit 1 plate. 

100,101. 1843 idem. Beitrag zur Fauna Norwegeus. — Nova 
Acta Acad. Leopold.-Carolin., Vol. XX, 1, pp. 
1-264, mit 12 Taf. 

VOL. r. 16 



242 BRITISH PARASITIC COPEPODA. 

101a. 1829-31 Retzius, A. Beskrifning ofver en ny Skandi- 
navisk Lernsea fran Nordsjon, kallad Lernsea 
Dalmanni. — Kongl. Vetensk. Akad. Handling, 
• Stockholm, pp. 109-119, c. tab. oppure : Be- 
schreibung einerneuen Scandinavischen Lernsea 
aus dem Nordsee, Lern^a Dalmanni gennant. 
Froriep's Notizen, Bd. XXIX, p. 6, figs. 5-9, 
1830. Idem in Isis, 1831, p. 1345, Taf. IX. 

102. 1877 Richiardi, S. Descrizione di due specie nuove 

di Lernseenicus (L. neglectus, L. vorax) con 
osserv. intorno a questo ed ai gen. Lernseocera 
Bl. e Ler7ifP,onema M. Edw. — Atti Soc. Tosc. Sc. 
Nat. Pisa, Yol. Ill, fasc. 1. 

103. 1878 idem. Tripapliylus mnsteli. — Processi verbali, 

Soc. Tosc.^Sc. Nat. Pisa, XX. 

104. 1880 idem. Catalogo sistematico dei crostacei che 

vivona sul corpo degli animali aquatici. — Cata- 
logo Sez. Ital. Esposiz. Internaz. di Pesca, 
Berlino, 1880 — Firenze — anche in Pisa tip. 
Vannucchi 1880 (8 pp.). For Clavella can- 
thari, &c. 

105. 1826 Risso, A. Histoire naturelle des productions de 

I'Europe meridionale, Yol. Y, p. 175 Paris. 

Refers to Otto's Caligus minimus, &c. 

105a. 1886 Sars, G, 0. The Norwegian North Atlantic Ex- 
pedition 1876-1878, Zoology — Crustacea, Part 
II, p. 80. (Records Anchorella uncinata and 
HsemohapJies cyclojiterinus,) 

106. 1850 Salter, Jawe^. Descriiption oi Lerneonema Bairdii. 

—Ann. and Mag. Nat. Hist., (2) YI, pp. 85-87, 
pi. vii B. 

107. 1901 Scott, A. Some additions to the Fauna of Liver- 

pool Bay. — Trans. Biol. Soc. of Liverpool, 
Yol. XY, pp. 342-853, with 2 pis. 

108. 1901 idem. Lepeophtheirus and Lernsea. — Liver}30ol 

Marine Biology Committee Memoirs, No. YI, 
54 pp., with 5 plates. 

109. 1904 idem. Some parasites found on fishes in the Irish 

Sea. — Trans. Biol. Soc. Liverpool, YoL XYIII, 
pp. 119-125. 

110. 1906 idem. Faunistic Notes. — Loc. cit., Yol. XX, pp. 

191-201, with pis. ii-vii. 



LITERATURE CHIEFLY REFERRED TO. 243 

111. 1907 Scott, A. (contd.). Faunistic Notes. — Loc, cit., 

Vol. XXI, pp. 191-198, with pis. i-v. 

Ilia. 1893 Scott, T. Additions to the Fauna of the Firth 
of Forth. Party. — 11th Annual Report Fishery 
Board for Scotland, Part III, p. 212, pi. Y, figs. 
1-13. 

111b. 1894 idem. Additions to the Fauna of the Firth of 
Forth. Part VI.— 12th Annual Eeport loc. 
cit. p. 233. 

111c. 1897 idem. The Marine Fishes and Invertebrates of 
LochFyne. — 15th Annual Report loc. cit. p. 155. 

llld. 1898 idem. On the distribution of the pelagic Inverte- 
brate Fauna of the Firth of Forth and its 
vicinity during the seven years from 1889 to 
1895, both inclusive. — 16th Annual Report loc. 
cit. p. 177. 

112. 1900 idem. Notes on some Crustacean Parasites of 

Fishes. — 18th Annual Report loc. cit. pp. 144- 
188, pis. V-VIII. 

113. 1901 idem. Notes on some Parasites of Fishes. — 19th 

Annual Report loc. cit. pp. 120-151, pis. VII- 
VIII. 

114. 1902 idem. Notes on some Parasites of Fishes. — 20th 

Annual Report loc. cit. pp. 288-299, pis. XII, 
XIII. 

115. 1904 idem. On some Parasites of Fishes new to the 

Scottish Marine Fauna. — 22nd Annual Report 
loc. cit, pp. 275-278, pi. XVII. 

116. 1905 idem. Observations on some Parasites of Fishes 

new or rare in Scottish Seas. — 23rd Annual 
Report loc. cit. pp. 108-115, pis. V-VI. 

117. 1907 idem. Some additional notes on Copepoda from 

the Scottish Seas. — 25th Annual Report loc. cit, 
pp. 216-218, pi. XV. 

118. 1909 idem. Some notes on Fish Parasites. — 26th 

Annual Report loc. cit. pp. lZ-11 , pi. III. 

119. 1900 idem. On Copepods living in the nostrils of Fishes. 

—Ann. Scott. Nat. Hist. pp. 153-155 (July 1900). 

120. 1901 idem. Glavella Za6raci5, van Ben., a Copepod new 

to Britain. — Ann. Scott. Nat. Hist. pp. 110-121 
(April, 1901). 



244 EEITISH PARASITIC COPEPODA. 

121. 1883 Smith, S. I. Eeview of the marine Crustacea of 

Labrador. — Proc. U. S. National Museum, pp. 
223-232. 

121a. 1899 Stebbing, T. R. R. Genus Sphyrion Cuvier.— 
Rep. Marine Biolog. Cape Town, 1898, p. 897. 

122. 1900 idem. South African Crustacea, Part I. — Cape of 

Good Hope Department of Agriculture, Marine 
Investigations in South Africa. — Records 8phy- 
rion IssvigatuSy Guerin-Meneville. 

123. 1900 idem. On Crustacea brought by Dr. Willey from 

the South Seas. — A. Willey's Zoological Results, 
Pt. V (Cambridge Univ. Press), Dec. 1900. 

124. 1905 South African Crustacea, Part iii. — Marine Investi- 

gations in South Africa, Vol. IV. 

125. 1910 idem. General Catalogue of South African Crus- 

tacea. (Contains also descriptions of new 
species.) — Annals of the South African Museum, 
Vol. VI, Part IV. 

126. 1862 Steenstrup, /. J. S. PMIichthys xij)hife, en ny 

snylter hos Svaerdfisken. — Overs. Kongel. 
Danske Vidensk. Selsk. Forhandl. 1861, pp. 
295-305, pi. 2. 
126a. 1869 idem. Cm Lesteira, Silenium, og Pegesimallus, 
tre af Prof. H. Kroyer opstillede Slaegter af 
Snyltekrebs. — Overs. Kongel. Danske Vidensk. 
Selsk. Porhandl. 1869. 

127. 1861 Steenstrup, J, J. S., and Liitken, C. Bidrag til 

Kundskab om det aabne Havs Synltekrebs og 
Lerna^er, etc. — Kongel. Danske Vidensk. Selsk. 
Skrifter, 5te Raekke, naturh. og math. Afdeling, 
Bd. V. Kjobenhavn. 

128. 1844 Thompson, jr. Report on the Fauna of Ireland, 

div. Invertebrata. — Report of the British Asso- 
ciation for 1843. 

129. 1847 idem. Additions to the Fauna of Ireland. {Lepe- 

oplitheirus stromi, etc.). — Ann. and Mag. Nat. 
Hist., Ser. I, Vol. XX, p. 248. 

130. 1856 idem. Natural History of Ireland, Vol. IV, 

p. 407. London. 

131. 1885 Thomson, G. M. Parasitic Crustacea.— New 

Zealand Journ. of Science, Vol. 2, p. 455; New 
Crustacea, loc. cit. p. 576. 



LITERATURE CHIEFLY REFERRED TO. 245 

132. 1889 Thomson, G. M. [contd.) Parasitic Copepoda of 

New Zealand, with descriptions of new species. 
— Trans. New Zealand Instit., Vol. 22 (n.s. 
Vol. 5), pp. 353-376 (13 n. sp.), with 5 plates. 

133. 1890 idem. A new parasitic Copepod. — Trans. New 

Zealand Instit., Vol. 23 (n.s. Vol. 6), pp. 227- 
229, with 1 plate. 

134. 1889 Thompson, I. 0. Third Report on the Copepoda 

of Liverpool Bay. — Proc. Biol. Soc. Liverpool, 
Vol. Ill, pp. 181-19], pi. viii. 

135. 1893 idem. Revised Report on the Copepoda of Liver- 

pool Bay. — Trans. Biol. Soc. Liverpool, Vol. VII, 
pp. 175-230, pis. xv-xxxv. 

136. 1897 idem. Further Report upon the Free- Swimming 

Copepoda of the West Coast of Ireland. — Trans. 
Biol. Soc. Liverpool, Vol. XI, pp. 127-131. 
Refers to the swimming* habits of G aligns. 

137. 1900 Thompson, I. C, and Scott, A. Some recent 

additions to the Copepoda of Liverpool Bay. — 
Trans. Biol. Soc. Liverpool, Vol. XIV, pp. 139- 
144, pi. viii. 

138. 190S idem. Supplementary Report VII. On the Cope- 

poda. — Report to the Government of Ceylon on 
the Pearl Oyster fisheries of the Gulf of Manaar, 
Part I, pp. 227-307, with 20 plates. Published 
by the Royal Society, London. 

139. 1905 Turner, Sir W. On Fennella halasnopterai : a 

Crustacean, parasitic on a ^nner Whole Balsenop- 
tera musculus. — Trans. Roy. Soc. Edinburgh, 
Vol. XLI, Part II, No. 18, pp. 409-434, with 4 
plates. 

140. 1862 Turner, Sir W., and Wilson, E. S. Observations 

on Parasitic Crustacea Ghondr acanthus lophii 
and Lernasopoda dalmanni. — Trans. Roy. Soc. 
Edinburgh, Vol. XXIII, part I, pp. 67-87, pi. 4. 

141. 1807 Turton, W. British Fauna, Vol. I. 

141a. 1880 Valle, A. Delia. Crostacei parassiti dei Pesci 
del mare Adriatico. — Boll. Soc. Adriat. Sc. Nat., 
Vol. VI, pp. 55-90. 

142. 1877 Vogt, G. Kecherches cotieres faites a Roscoff ; 

Crustaces parasites des Poissons. Avec 6 pis. 
Geneve. H. Georg. 1877, p. 104 — oppure : Arch. 
Zool. Exper. et Gen. VI, 1877, pp. 385-456. 



246 BRITISH PARASITIC COPEPODA. 

143. 1879 YoGT, C. [contd,), Crustaces parasites des Poissons. 

— Memoires de Tlnstitut National Genevois, Yol. 
XIII. (Two Memoirs — separate copies) pp."l- 
104, 6 plates. 

143a. 1902 Wilson, Ch. Branch. North American Parasitic 
Copepods of the Family Argulidas, with a Biblio- 
graphy of the Group and a Systematic Review 
of all the known Species. — Proc. U.S. National 
Museum, Yol. XXY, pp. 635-742, with plates 
YIII-XXYII. 

143b. 1904 idem. Anew species oiArgulus, with a more com- 
plete account of two species already described. 
—Proc. U. S. National Museum, Yol. XXVII, 
pp. 627-655. 

144. 1905 idem. New species of Parasitic Copepods from 

the Massachusetts Coast. — Proc. Biol. Soc. 
Washington, Yol. 18, pp. 127-131. 

145. 1905 idem. North American Parasitic Copepods belong- 

ing to the Family Caligida3. Part 1. The 
Caliginae. — Proc. U. S. National Museum, Yol. 
XXYIII, pp. 479-672, with plates Y-XXIX. 

146. 1906 idem. Supplementary Report XXXIY. On some 

Parasitio Copepoda. — Report to the Government 
of Ceylon on the Pearl Oyster Fisheries of the 
Gulf of Manaar, Pt. Y, pp. 189-210, with 5 
plates. Published by the Royal Society, London. 

147. \^01 idem. North American Parasitic Copepods belong- 

ing to the Family Caligidai. Part 2, The Trebinae 
and Euryphorina?. — Proc. U.S. National Museum, 
Yol. XXXI, pp. 669-720, with plates XY-XX. 

148. 1907 idem. North American Parasitic Copepods be- 

longing to the Family Caligidas. Parts 3 and 4, 
A Revision of the Pandarinae and the Cecro- 
pinae. — Proc. U. S. National Museum, Yol. 
XXXIII, pp. 323-490, with plates XYII-XLIII. 
148a. 1907 idem. Additional Notes on the development of 
the Argulid^e, with description of a new 
species. — Proc. U. S. National Museum, Yol. 
XXXII, pp. 411-424, with plates XXIX-XXXII. 

149. 1908 idem. North American Parasitic Copepods: A 

list of those found upon the fishes of the Pacific 
Coast, with descriptions of new genera and spe- 
cies. — Proc. U. S. National Museum, Yol. XXXY, 
pp. 431-481, with plates LXYI-LXXXIII. 



LITERATURE CHIEFLY REFERRED TO, 247 

149a. 1911 WiLSO-^, C. B. {contd.). North American Parasitic 
Copepods belonging to the Family Ergasilidae. 
— Proc. U.S. iSational Museum, Vol. 39, pp. 
263-400, with plates 41-60. 

149b. 1911 idem. North American Parasitic Copepods. — 
Part 9. the Lernasopudae. — Proc. U. S. ^'ational 
Museum, Vol. 39, pp. 189-226, with plates 29-36. 

149c. 1911 idem. North American Parasitic Copepods. 
Descriptions of new genera and species. — Proc. 
U. S. National Museum, Vol, 39, pp. 625-634, 
with plates 65-68. 

149d. 1912 idem. Descriptions of new species of Parasitic 
Copepods in the collections of the United States 
National Museum. — Proc. U.S. NationalMuseum, 
Vol. 42, pp. 233-243, with plates 30-34. 

149e. 1912 idem. Parasitic Copepods from Nanaimo, British 
Columbia, including eight species new to science. 
— Contributions to Canadian Biology being 
studies from the Marine Biological Stations of 
Canada 1906—1910, pp. 85-102, with plates 
III-IX. 

150. 1870 Wright, Edw. Perceval. On a new species of the 
genus Pennella. — Ann. and Mag. Nat. Hist., 
Ser. 4, Vol. V, p. 45, pi. 1. 



(248) 



INDEX. 

{Syiionynis in italics,) 



A. 

ahhreviatus, Caligus 

acanthii, Eudactylina 

Achieves . 

Achtheres 

Adheres . 

acuta, Eudactylina 

ieglifini, C aligns 

alata, Clavella . 

alata, Dinemoura 

alatus, Pandarus 

ambiguus, Hiemobaphoides 

amhiguus, Hxmobaphus 

ambiguus, Nogagus 

ambiguus, Nogaus 

Anchorella 

angustulus, Nogagus 

annulatus, Chondracanthus 

Anops 

Antbosoma 

Argulus . 

argulus, Argulus 

argulus, Monoculus 

ArnsBUs . 

asellina, Lernxa 

asellina, Lernentoma 

asellinum, Medesicaste . 

asellinus, Oralien 

asterodermi, Liitkenia . 



B. 

hairdi, Lernseonema 
balsenopterse, Penella 
Basanistes 
bicolor, Pandarus 
hicuspidatus, Caligus 
bidiscalis, Lemseopoda 



PAGE 

50 
128 
193 
193 
193 
126 

45 
225 

89 

89 
148 
148 

78 

78 
213 

95 
169 
168 
1U8 
227 
227 
227 

83 
184 
184 
184 
184 

76 



156 
153 
199 
95 
45 
200 



Binoculus 

bispinosa, Brachiella 

Bomolocbus 

hoscii, Pandarus 

Bracbiella 

Brachiella 

brachyptera, Elytrophora 

hranchialis, Caligus 

branchial! s, Lernaea 

brevicaudatus, Caligus . 

brevicollis, Anchorella . 

brevicollis, Clavella 

hrevipedes, Caligus 

brevipedes, Pseudocaligus 



Caligid^ 
Caligoida 
Caligus . 
Caligus . 

canthari, Anchorella 
canthari, Clavella 
canthari, Lerneomyzon 
carpionis, Lerneopoda 
caudatus, Trebius 
Cecrops . 
Cecropsina 
centrodonti, Caligus 
charon, Argulus 
Charopinus 

CnONDRACANTHIDiE 

Chondracanthus 
Chondracanthus . 
clavatus, Chondracanth 
Clavella . 
Clavella . 
cluthae, Clavella 



INDEX. 



249 



cluthae, Hatschekia 
clnthse, Lernaeopoda 
coleoptrata, Dinematura 
<joleoptratus, Echthrogaleus 
€ongericola 

cornigera, Hatschekia . 
cornuta, Anops . 
cornuta, Entomoda 
cornuta, Lernaea 
cornuta, Lernentoma 
cornuta, Phyllophora 
cornutus, Chondracanthus 
cormdus, Fhyllophorus . 
cornutus, Phyllothreus 
crass um, Anthosoma 
jcrassus, Caligus . 
cvirtus, Caligus . 
Cyclopoida 

cyclopterina, Hsemobaphes 
■cyclopterina, Lernsea 
cyclopterina, Lernaeocera 
Cycnus . 

cyprinacea, Lernsea 
cyprinacea, Lernaeocera 
cyprinacea, Lerneocera . 

J). 
dalmanni, Charopinus . 
dalmanni, Lernaea . 
dalmanni, Lernaeopoda . 
dalmanni, Lerneopoda . 
delarochianus, Chondracanthus 
Demoleus 

depressus, Chondracanthus 
diaphanus, Caligus 
diaphanus, Caligus 
Dichelestiidae 
Dichelestium 
Binematura 
Dinemoura 
dubia, Clavella 
dubius, Charopinus 

E. 

Echthrogaleus . 
edwardsi, Lophoura 
edwardsi, Lophura 



PAGE 




PAGE 


. 117 


edwardsi, Rebelula 


. 162 


. 198 


elongata, Dinematura 


. 86 


89 


elongata, Lernaea 


. 195 


. 89 


elongata, Lernaeopoda 


. 195 


. 124 


elongata, Lerneopoda 


. 195 


. 118 


elongatus, Caligus 


. 48 


. 168 


Elytrophora 


82 


. 168 


emarginata, Anchorella 


. 218 


. 168 


emarginata, Clavella 


. 218 


. 168 


encrasicola, Lernaea 


. 158 


. 92 


encrasicola, Lernaeenicus . 158 


. 168 


encrasicola, Lerneonema 


. 158 


92 


Entomoda 


. 168 


. 92 


Ergasilid^ 


33 


108 


Ergasilus 


34 


. 108 


esocina, Lernaea . 


. 154 


. 45 


esocina, Lernaeocera 


. 154 


33 


Eudactylina 


. 125 


. 147 






. 147 


F. 




. 147 


fibrosa, Pennella 


. 151 


. 124 


filosa, Pennela . 


. 151 


154 


filosa, Pennella . 


. 150 


154 


fissifrons, Pandarus 


95 


, 154 


flura?, Chondracanthus 


. 171 




foliaceus, Argulus 


. 227 




foliaceus, Monoculus 


. 227 


189 






189 


G. 




189 


galei, Lernaeopoda 


. 197 


189 


galei, Lerneopoda 


. 197 


3 177 


g aster ostei, Binoculus 


. 227 


79 


gasterostei, Ergasilus 


42 


172 


gasterostei, Ozolus 


. 227 


60 


gasterostei, Ther sites 


42 


45 


gasterostei, Thersitina 


42 


105 


gihhosus, Chondracanthti 


s . 179 


105 


glabra, Cecropsina 


. 76 


5,103 


gracilis, Caligus 


69 


85 


gracilis, Leopeophtheirus 


. 69 


217 


grandis, Nogagus 


. 79 


190 


gurnardi, Caligus 


. 52 




gyrini, Monoculus 


. 227 


88 


H. 




162 


Haemobaphes 


. 147 


162 


Haemohaphes 


. 148 



260 



BRITISH PARASITIC COPEPODA. 







PAGE 




PAGE 


Haemobaphoides 


. 148 


Lerneopoda 


. 188 


Hatschekia 


. 112 


Lesteira . 


. 164 


hippoglossi, Caligus 


. 67 


Lestes 


164 


hippoglossi, Clavella 


. 112 


limandsB, Chondracanthus 


174 


hippoglossi, Hatschekia 


. 112 


lineata, Kroyeria 


121 


hippoglossi, Lepeophtheirus 


. 67 


lineatum, Lonchidium . 


121 


hypocephalus, Stylophorus 


. 189 


Lonchidium 


. 120 






lophii, Brachiella 


226 


I. 




lophii, Chondracanthus 


179 


imhricatus, Caligus 


. 108 


lophii, Clavella . 


226 


impudica, Brachiella 


. 187 


Lophoura 


162 


impudica, Thysanote . 


. 187 


Lophura . 


162 


inflatus, Chondracanthu» 


. 169 


lumpi, Lernsea . 


146 


inominatus, Lepeophtheirus 


. 73 


lumpi, Lesteira . 


164 


insidiosa, Brachiella . 


. 206 


lumpi, Lestes 


164 


insolens, Eudactylina . 


. 135 


lumpi, Sphyrion 


164 


isonyx, Caligus . 


60 


lusci, Lernsea 


144 






lutkeni, Echthrogaleus 


90 


K. 




lutkeni, Nogagus 


90 


Kroyeria 


. 120 


Liitkenia 


75 


kroyeri, Lernanthropus 


. 110 


M. 

macroirachelus, Clavella 




L. 




224 


labracis, Caligus 


. 53 


Medesicaste 


183 


lahracis, Clavella 


. 115 


merluccii, Brachiella . 


207 


labracis, Hatschekia . 


. 115 


merluccii, Chondracanthus 


180 


Lsemargus 


. 100 


rtierlucii, Brachiella 


207 


Isevirajse, Chondracanthus 


. 169 


merlucii, Chondracanthus 


181 


lamnae, Dinemoura 


. 86 


merlucii, Lernsea 


180 


lamtice, Pandarus 


86 


minimus, Caligus 


46 


lampri, ? Lernaeopoda . 


. 202 


minuta, Eudactylina 


133 


latreillii, Cecrops 


98 


minuta, Lernsea 


145 


latus, Nogagus . 


95 


minutus, Caligus 


46 


Lepeophtheirus 


. 64 


monilaris, Lerneonema . 


156 


Lernsea . 


. 142 


Monoculus 


227 


Lernsea . 




. 184 


miilleri, Caligus 


45 


Lernaeenicus 




. 156 


rmilli, Clavella . 


114 


LBBNiEID^ 




. 141 


mulli, Hatschekia 


114 


Lernseocera 




. 154 


muricata, Orthagoriseicola 


lUO 


Lern^oidea 




. 141 


muricatus, Laemargus 


100 


Lernaeopoda 




. 195 


musteli, Lernseenicus 


160 


LERNiEOPODIDA; 




. 186 


musteli, Lerneonema 


160 


Lernanthropus 




. 110 


musteli, Tripaphylus 


160 


Lernentoma 




167, 183 






Lerneocera 




. 154 


N. 




Lerneomyzon 




. 213 


nanus, Ergasilus 


34 


Lerneonema 




. 159 


nodosa, Lernxa . 


176 



INDEX. 



251 





PAGE 




PAGE 


nodosa, Lernentoma 


. 176 


Phyllophora 


. 92 


nodosus, Chondracanthus 


. 176 


Phyllothreus . 


92 


Nogagus . 


77 


poUachii, Lepeophtheirus . 73 


Nogaus . 


77 


pollachius, Lepeophtheirus . 73 


nordmanni, Caligus 


. 66 


producta, Dinematura 


. 86 


nordmanni, Lepeophtheirus 


66 


producta, Dinemoura 


86 






productus, Caligus 


. 86 


0. 




productus, Nogagus 


. 86 


oblongum, Dichelestium 


. 106 


Pseudocaligus . 


. • 61 


ohlongus, Caligus 


. 106 


pygmsea, Hatschekia 


. 119 


obscurus, Caligus 


70 






obscurus, Lepeophtheirus 


70 


Q. 




obscurus, Lepeophtheirus 


67 


quadrata, Anchorella 


. 223 


obscurus, Lepeophtheirus 


69 


quadrata, Clavella 


. 223 


onosi, Bomolochus 


39 






Oralien . 


183 


E. 




ornatus, Chondracanthus 


182 


ramosus, Charopinus 


. 191 


orthagorisci, Pennella . 


150 


rapax, Caligus . 


48 


Orthagoriscicola 


100 


Rebelula 


. 162 


oralis, Anchorella 


210 


rhombi, Lepeophtheirus 


69 


ovalis, Brachiella 


210 


rostrata, Brachiella 


. 205 


Ozolus 


227 


rugosa, Anchorella 


. 216 


P. 

pagelli, Anchorella 




rugosa, Clavella 


. 216 


224 


S. 




pallida, Congericola 


124 


salnionea, Basenistes 


. 199 


pallidus, Cycnus 


124 


salmonea, Entomoda 


. 199 


Pandarus 


94 


salmonea, Lernsea 


. 199 


paradoxa, Anchorella . 


222 


salmonea, Lernaeopoda 


. 199 


paradoxa, Clavella 


222 


salmonea, Lerneopoda 


. 199 


paradoxus, Caligus 


79 


salmonis, Caligus 


71 


paradoxus, Demoleus . 


79 


salmonis, Lepeophtheiri 


IS . 71 


parkeri, Brachiella 


212 


Schisturus 


. 213 


pastinaca, Brachiella . 


211 


Sciaenophilus 


. 62 


pastinacse, Brachiella 


212 


scomberi, Anchorella 


. 219 


pectoralis, Caligus 


64 


scomberi, Caligus 


. 57 


pectoralis, Lepeophtheirus 


64 


scombri, Anchorella 


. 218 


pectoralis, Lernsea 


64 


scombri, Clavella 


. 218 


pelamydis, Caligus 


57 


serrata, Dinematura 


. 104 


Pennela 


151 


serratus, Philorthragori 


scus . 104 


Pennella 


149 


similis, Eudactylina 


. 130 


percarum, Achteres 


193 


similis, Lernaeopoda 


. 202 


percarum, Achtheres . 


193 


smithii, Anthosoma 


. 108 


percarum, Adheres 


193 


solese, Bomolochus 


. 36 


Philichthyid^ 


137 


solese, Chondracanthus 


. 170 


Philichthys . 


138 


Sphyrion 


. 164 


Philorthragoriscus 


103 


spratta, Lernsea . 


. 156 



252 



BRITISH PAHASITIO OOPEPODA. 





PAGE 




PAGE 


spratta, Lernseenicus 


156 


triglae, Chondracanthus . 


184 


spratta, Lermeonema 


156 


triglae, Lernentoma 


184 


sprattse, Lernseenicus . 


156 


triglarum, Medesicaste . 


185 


steMata, Anchorella 


220 


Tripaphylus 


159 


stellata, Clavella 


220 


tuherculatus, Chondracanthus 


177 


stromi, Lepeophtheirus . 


71 






stromii, Lepeophtheirus . 


71 


U. 




sturionis, Caligus 


74 


uncinata, Anchorella 


214 


sturionis, Dichelestium . 


106 


uncinata, Clavella 


. 214 


sturionis, Lepeophtheirus 


74 


uncinata, Lernaea 


. 214 


Stylophorus 


189 


uncinatum, Lerneomyzon 


. 214 


T. 




uncinatus, Schisturus . 


. 214 


tenuis, Scisenophilus 


63 


W. 




tenuis, Scienophilus 


63 


williamsoni, Chondracanthus 


. 176 


Thersites 


41 






Thersitina 


. 41 


X. 




thompsoni, Lepeophtheirus 


. 69 


xiphiae, Philichthys 


. 138 


thynni, Arnseus . 


83 






thynni, Brachiella 


204 


Z. 




Thysanote 


187 


zei, Caligus , 


. 54 


Trebius . 


81 


zei, Chondracanthus . 


. 177 


triglae, Anchorella 


209 


zeugopteri, Bomolochus 


. 40 


triglsB, Brachiella 


209 


zeus, Chondracanthus 


. 177 



Errata. 

p. 8, line 8, for siphon read chitin filament. 

„ 31, Genus 28, for cyclopterinus read cyclopterina. 

„ „ (jrenus S3, for Tbypaphylus read Tripaphylus. 

„ 136, lines 13 and 14, for Eudactylin aacuta read Eudactylina acuta. 

„ 137, line 17, for Kroyeria cineata read Kroyeria lineata. 

„ 199, Synonymy, line 6, for Basenistes read Basanistes. 



EXPLANATIONS OF PLATES A AND B, 



(254) 



Plate A. 

Figs. 

1. Ventral surface of the pectoral fin of a flounder [Pleicro- 

nectes jiesus Linn.), with thirty-two adult specimens of 
Lepeophtheirus pectoralis (Miiller) adhering to it. 
(x3.) 

2. Head of a whiting {Gadus merlangus Linn.), with part of 

the operculum removed to shew an adult female Lernsea 
hranchialis Linn, adhering to the gills. ( x 2.) 

3. Part of the gills of a flounder {Pleuronectes Jiesus Linn.), 

shewing the '^Cyclopoid" stage of Lernsea hranchialis 
attached to the ends of the gill-filaments. ( x 2J.) 

{Reproduced from photographs,) 



PLATE A 




(256) 



Plate B. 

Figs. 

1. Part of the gills of a dory {Zeus faher Linn.), shewing an 

adult female Ghondr acanthus zei De la Roche in situ. 
(x24.) 

2. Adult female of Chondracanthus zei removed from its 

host, (x 2.) 

3. Gill of a salmon {Salmo salar Linn.), shewing adult 

female specimens of Lernasofoda salmonea (Gisler) 
attached to the ends of the gill-filaments. The salmon 
was captured in the river Dee, near Aberdeen; the 
parasites adhered to the gill-filaments in considerable 
numbers, their white colour shewing distinctly against 
the red colour of the gills. (Nat. size.) 

{Reproduced from photographs.) 



PLATE B. 




RAY SOCIETY 

INSTITUTED 1844 

FOR THE PUBLICATION OF WORKS ON 
NATURAL HISTORY 

ANNUAL SUBSCRIPTION ONE GUINEA 



LIST OF THE SOCIETY 

FOR THE YEAR 1913 



COERECTED TO 28th FEBEUARY, 1913 



OFFICERS AND COUNCIL. 

1912-13. 



Ipresttrent. 

The Et. Hon. LORD AVEBURY, P.C., D.C.L., LL.D., For.Sec.R.A., 
F.R.S., F.S.A., F.L.S., etc. 

S. F. FARMER, M.A., Sc.D., F.R.S., F.Z.S. 

A. D. MICHAEL, F.L.S., F.Z.S., F.R.M.S. 

R. F. SCHARFF, Ph.D., B.Sc, F.L.S., F.Z.S. 



€onncii, 

Robert Adkin, F.E.S. I G. B. Longstaff, M.D., F.E.S. 

Rev. C. R. N. Burrows, F.E.S. I J. W. S. Meiklbjohn, M.D., F.L.S. 

T. A. Chapman, M.D., F.Z.S., F.E.S. ' James Murray, F.Z.S., F.R.M.S. 



Rev. Alfred Fuller, M.A., F.E.S. 

A. E. GiBBS, F.L.S., F.Z.S., F.E.S. 

B. Daydon Jackson, Ph.D., F.L.S. 
Albert H. Jones, Treas.E.S. 



A.W.Oke,B.A.,LL.M.,F.L.S.,F.G.S. 
Charles Oldham, F.Z.S., M.B.O.U. 
C. D. Soar, F.L,S., F.R.M.S. 
W. Mark Webb, F.L.S., F.R.M.S. 



dbrcasitrer. 

F. DuCANE GODMAN, D.C.L., F.R.S., F.L.S., F.G.S., F.Z.S., F.E.S. 

JOHN HOPKINSON, F.L.S., F.G.S., F.Z.S., Assoc.Inst.C.E.; 
Weetwood, Watford. 



LIST OF THE EAY SOCIETY. 



Aberdeen University Library ; King's College, Aberdeen, 

Adelaide Public Library; Adelaide, S. Australia. 

Adkin, Robert, F.E.S. ; Wellfield, 4 Lingards Road, Lewisham, 

S.E. 
Adlard, Robert Evan ; Bartholomew Close, E.G. 
Advocates^ Library; Edinhnrgh. 

Albany Museum ; Grahamstown, Cafe Colony, S. Africa, 
American Museum of Natural History ; New York, U.S.A. 
Armstrong College; Newcastle-tipon-Tyne. 

Assheton, Richard, M.A., F.L.S., F.Gr.S., F.Z.S. ; Grantchester, 

Cambridge. 
Athenaeum Club ; Pall Mall, S.W. 
Australian Museum ; Sydney, New South Wales. 
Avebury, The Right Hon. Lord, PC, D.C.L., LL.D., F.R.S., 

F.S.A., F.L.S., F.a.S., F.Z.S., F.E.S., President ; 15 

Lombard Street, E.C. ; 48 Groscenor Street, W. ; and 

High Elms, Farnborough, Kent. 

Baer, Joseph, & Co. ; 6 Hochstrasse, Frankfort, Germany. 

Bagshaw, Walter, F.R.M.S. ; Moorfield, BIrhenshaw, Bradford. 

Barraud, Philip J., F.E.S. ; Bushey Heath, Watford. 

Barrow-in-Furness Public Library ; Barrow-in-Furness. 

Battersea Public Library; Lavender Hill, S.W. 

Belfast Library ; Donegal Square, Belfast. 

Bentley, Richard, F.S.A., F.L.S., F.R.Met.Soc, F.R.G.S., 

F.S.S. ; The Mere, Upton, Slough. 
Berens, A. A., M.A. ; 50 Eaton Square, S.W. 
Bergens Museums Bibliotek ; Bergen, Noricay. 
Berlin Royal Library ; Berlin. 
Berlin Royal Zoological Museum ; Berlin. 
Berne Natural History Museum ; Berne, Switzerland. 



b LIST OF THE KAY SOCIETY. 

Bestow, Charles Horton, F.R.M.S. ; Melford House, 43 Upper 

Clapton Road J London , N.E. 
Bethune-Baker, George T., F.L.S., F.E.S.; 19 Clarendon 

Road, Edghaston, Birmingham. 

Bibliotheque Nationale ; Paris. 

Binks, Mrs. I. ; 9 Burton Street, Wakefield. 

Birmingliam Free Libraries; Birmingham. 

Birmingham Natural History and Philosophical Society ; 
Avehury House, 55 Newhall Street, Birmingham. 

Bloomfield, The Rev. E. N., M.A., F.E.S. ; Guestling Rectory, 

Hastings. 
Board of Education, Secondary Branch; Science Library, 

South Kensington, S.W. 
Bodleian Library ; Oxford. 

Bootle Free Library ; Oriel Road, Bootle, Liverpool. 
Bostock, E. D., F.E.S. ; Holly House, Stone. 
Boston Public Library ; Boston, Mass., U.S.A. 
Boston Society of Natural History ; Berkeley Street, Boston, 

Mass., U.S.A. 
Bowles, Edward Augustus, M.A., F.L.S., F.E.S. ; Myddelton 

House, Waltham Cross. 
Bowman, John Herbert ; Greenham Common, Newbury. 
Bradford Natural History and Microscopical Society ; Church 

Institute, North Parade, Bradford. 
Breslau University Library ; Breslau, Germany. 
Briggs, C. A., F.E.S. ; Rock House, Lynmouth, Devon. 
Briggs, T. H., M.A., F.E.S. ; Rock House, Lynmouth, Devon. 

Brighton and Hove Natural History Society ; Public Library, 

Brighton. 
Bristol Municipal Public Libraries ; Bristol. 
Bristol Museum and Art Gallery ; Queeri's Road, Bristol. 
Bristol Naturalists' Society ; 20 Berkeley Square, Bristol. 
British Museum; Bloomsbury, W.C. 
Brokenshire, Fred. R. ; 7 Hitlsboro' Avenue, Pennsylvania, 

Exeter. 
Bromley Naturalists' Society; 92 London Road, Bromley, 

Kent. 
Buchan-Hepburn, Sir Archibald, Bart., F.E.S. ; Smeaton- 

Hepbiirn, Prestonkirk, N.B. 



LIST OF THE RAY SOCIETY. 7 

Burr, Malcolm, D.Sc, F.L.S., F.GT.S., F.Z.S., F.E.S.; Castle 

Hill House J Dover. 
Burrows, The Rev. C. R. N., F.E.S.; Muching Vicarage, 

Stanfo7'd-le-Hope, Essex. 

Cambridge Philosophical Library ; New Museums, Gamhridge, 

Cambridge University Library ; Cambridge. 

Campbell, Francis Maule, F.L.S., F.E.S. ; Brynllwydwyn, 

Machynlleth. 

Canterbury, Philosophical Institution of; Christchurch, New 

Zealand. 
Cardiff Free Libraries ; Cardiff. 
Chapman, Thomas Algernon, M.D., F.Z.S., F.E.S. ; Betula, 

Reigate. 
Cheltenham Natural Science Society ; Public Library and 

Museum, Cheltenham. 
Chester Society of Natural Science ; Grosvenor Museum, 

Chester. 

Chicago University Library ; Chicago, III., U.S.A. 

Christiania, University of ; Christiania, Norway. 

Church, Sir William Selby, Bart., K.C.B., M.B. ; 130 Harley 

Street, W. 
Cleland, John, M.D., D.Sc, LL.D., F.R.S. ; Drumclog, Crew- 

heme. 
Colgan, Nathaniel, M.R.I. A. ; 15 Breffni Terrace, Sandy cove, 

Co. Dublin. 
Collins, The Hon. Mrs. Henn, B.Sc, F.L.S.; Colins Farm, 

Durrington, Salisbury. 
Congress, Library of ; Washington, D.C., U.S.A. 
Copenhagen University Library ; Copenhagen. 
Cornell University Library ; Ithaca, New York, U.S.A. 
Cornwall, Royal Institution of ; Truro. 
Cotton, John, M.R.C.S., F.E.S. ; Simonswood, Prescot Road, 

St. Helens. 
Cox, Benjamia Cornell ; Gilston, Largoward, Fifeshire. 
Crawford, W. C. ; 1 Lockharton Gardens, Colinton Road, 

Edinburgh. 
Crawshay, Lionel Rutledge, M.A.(Oxon) ; Marine Biological 

Laboratory, Plymouth. 
Croydon Public Libraries ; Town Hall, Croydon, 



8 LIST OF THE EAY SOCIETY. 

Dames, Felix L. ; Steglitz-Berlin. 

Davies, Arthur Ellson, Ph.D., F.L.S., F.C.S.; Ttceedhanh, 

West Savile Road, Edinburgh. 
Derby Free Library and Museum ; Wardwich, Derby. 
Detroit Public Library; Detroit, Mich., U.S.A. 
Dilks, Arthur C, B.Sc; Tardebigge, Bromsgrove. 
Doesburgh, S. C. van; Breestraat, Leiden, Holland. 
Downing College ; Cambridge. 

Drinkwater, J. W., F.R.M.S. ; >S'^. Margarets, WalUngton, 
Surrey. 

Dublin, Royal, Society ; Leinster House, Dublin. 

East Kent Natural History Society ; Medical Hall, Canterbury. 
Eastwood, John E., F.E.S. ; Enton Lodge, Witley, Godalming. 
Eddy, James Ray ; The Grange, Carleton, Shipton. 
Edinburgh Public Library ; Edinburgh. 
Edinburgh, Royal Society of; Edinburgh. 
Edinburgh University Library ; Edinburgh. 
Elliott, John, M.D., F.R.C.P. ; 24 Nicholas Street, Chester. 
Elphinstone, Sir Howard W., Bart., M.A., F.L.S., F.R.G.S. ; 

2 Stone Buildings, Lincoln's Lin, W.C., and Struan, 

Wimbledon Parh, S.W. 
England, Royal College of Surgeons of ; Lincoln's Lnn Fields, 

W.C. 

Fielding, Clement, M.P.S., Ph.C. ; Clover Hill, Halifax. 

FitzGerald, The Rev. H. Purefoy, M.A., F.L.S., F.C.S. ; 
Lidwells, Goudhurst, Kent. 

Fletcher, Thomas Bainbrigge, R.N., F.L.S., F.Z.S., F.E.S. ; 
Agricultural Research Lnstitute, Pusa, Bihar, India. 

Fletcher, W. H. B., M.A., F.Z.S., F.E.S.; Aldwich Manor, 
Bognor. 

Folkestone Free Public Library and Museum ; Folkestone. 

France, Institut de ; Paris. 

Friedlander & Sohn; 11 Carlstrasse, Berlin, N.W. 6. 

Friend, The Rev. Hilderic, F.R.M.S.; Wilmot Road, Swadlin- 
cote, Burton-ujpon- Trent. 

Fuller, The Rev. Alfred, M.A., F.E.S.; The Lodge, Sydenham 
Hill, S.E. 

Garnett, Frank W., M.R.C.V.S.; Dalegarth, Windermere. 



LIST OF THE RAY SOCIETY. 9 

Gascoigne, Major French; Lotherton Hall, Aherford, Leeds. 
Geological Society of London; Burlington House, Piccadilly, 

W. 
Gerold & Co. ; Vienna. 
Gibbs, Arthur Ernest, F.L.S., F.Z.S., F.E.S., F.R.H.S.; 

Kitchener's Meads, St. Albans. 
Gibson, Miss; Hill House, Saffron Walden. 
Giles, Harry M., F.E.S. ; Zoological Society, South Perth, 

Western Australia. 
Glasgow Natural History Society ; 207 Bath Street, Glasgow. 
Glasgow, Royal Philosophical Society of; 207 Bath Street, 

Glasgoiv. 
Glasgow University Library; Glasgoiv. 
Godman, F. DuCane, D.C.L., F.R.S., F.L.S., F.G.S., F.Z.S., 

F.E.S., F.R.H.S., Teeasurer; 45 Po7it Street, Belgrave 

Square, S.W. : and South Lodge, Horsham. 
Gooding, Henry Cornish; Ipswich Street, StowmarJcet. 
Gottingen University Library ; Gottingen, Germany. 
Great Britain, Royal Institution of ; Albemarle Street, W. 
Green, E. Ernest, F.E.S. ; Mote Hall, Bearsted, Maidstone. 
Grosvenor Public Library ; Buffalo, N.Y., U.S.A. 
Guille-Alles Library; Guernseij. 
Guitel, Frederic, Professeur de Zoologie a la Faculte des 

Sciences; Universlte de Rennes, France. 
Gurney, Robert, F.Z.S. ; Ligham Old Hall, Stalham. 

Haileybury College ; Hertford. 

Halifax Public Library ; Belle View, Halifax. 

Hamburg, Naturhistoriche Museum zu; Steinthorwall, Ham- 
burg, Germany. 

Hardy, Alfred Douglas, F.L.S., F.R.M.S.; State Forests De- 
partment, Melbourne, Victoria, Australia. 

Harley, John, M.D., F.R.C.P., F.L.S. ; Beedings, Ptdhorough, 
Sussex. 

Harmer, Sidney F., Sc.D., F.R.S., F.Z.S. , Keeper of Zoology. 
British Museum, Vice-President; British Museum (Nat, 
Hist.) Cromivell Road, S.W., and 58 Albemarle Road, 
Beckenham. 

Harvard Museum of Comparative Zoology ; Cambridge, 
Mass., U.S.A. 



10 LIST OF THE RAY SOCIETY. 

Hastings and St. Leonard's Natural History Society; The 

Museicvi, Hastings. 
Heidelberg University Library ; Heidelberg, Germany. 
Hertfordshire County Museum; Hatfield Road, St. Albans. 
Hertfordshire Natural History Society and Field Club ; Upton 

House, Watford. 
Hewitt, David Basil, F.R.C.S.I. ; Grove Mount, Davenham. 
Hope, Gr. P. ; Havering Grange, Romford. 
Hopkinson, John, F.L.S., F.G.S., F.Z.S., F.R.M.S., F.R.Met. 

Soc, Assoc. Inst.C.E., Secretary; Weetwood, Watford. 
Huddersfield Naturalist and Photographic Society ; The 

Technical College, Huddersfield. 
Hull Public Libraries ; Hull. 

niinois University Library ; Urbana, III., U.S.A. 
India, Geological Survey of; Calcutta. 
Ireland, National Library of ; Kildare Street, Dublin. 
Irish, Royal, Academy; 19 Dawson Street, Dublin. 
Irving, John, M.D. ; Inagh Mount, 15 Filey Road, Scar- 
borough. 

Jackson, B. Daydon, Ph.D., F.L.S., General Secretary of the 

Linnean Society; 21 Cautlei/ Avenue, Clapham Common, 

S.W. 
James, Robert Detiley, F.A.I. (Natal), F.R.M.S.; EarVs 

Croome, Durban, Natal {P.O. Box 212). 
John Crerer Library ; Chicago, HI., U.S.A. 
Jones, Albert H., F.E.S. ; Shrublands, Eltham. 
Jones, William Llewellyn, F.R.M.S. ; Manley Knoll, Helsby, 

Warrington. 
Junk, W. ; 201 Kurfurstendamm, Berlin, W. 15. 

Kappel, August Wilhelm, F.L.S., F.E.S., Librarian of the 
Linnean Society ; Creeveroe, Golder's Green, N. W. 

Kenrick, Sir G. H., F.E.S. ; Whetstone, Somerset Road, 
Edgbaston, Birmingham. 

Kiel University Library ; Kiel, Germany. 

Kilmarnock Public Library and Museum ; Kilmarnock, N.B. 

King's Inn Library ; Dublin. 

Knight, H. H. ; The Lodge, All Saints' Road, Cheltenham. 



LIST OF THE EAY SOCIETY. 11 

Laver, Henry, M.R.C.S. ; 43 Head Street, Colchester. 
Lebour, Miss Marie Y., M.Sc. ; Radcliffe House, Corhridge,. 

Northumberland. 

Leeds Philosophical and Literary Society ; The Museum, Leeds. 

Leeds Public Free Libraries ; Leeds. 

Leeds University Library ; Leeds. 

Leicester Free Public Library ; Wellinr/ton Street, Leicester. 

Lewis, John Spedan, F.Z.S. ; Spedan Tower, West Heath, 

Hamf stead, N.W., and 278-288 Oxford Street, W. 
Linnean Society of London ; Burlington House, Piccadilly, W, 
Liverpool Athenieum ; Liverpool. 
Liverpool Free Public Libraries ; Licerpool. 
Liverpool Microscopical Society; Royal Listitution, Liverpool. 

Liverpool School of Tropical Medicine ; Exchange Buildings, 
Liverpool. 

London Library; 12 St. James's Square, S.W. 

Longstaif, George Blundell, M.A., M.D., F.C.S., F.S.S. ; 

Highlands, Putney Heath, S.W., and Twitchen, Morthoe,. 

Devon. 

Los Angeles Public Library ; California, U.S.A. 
Lyon Universite Bibliotheque ; Lyons, France. 

McLitosh, W. Carmichael, M.D., LL.D., F.R.S.L.&E., F.L.S., 

Professor of JSTatural History, University of St. Andrews ; 

2 Ahhotsford Crescent, St. Andrews, N.B. 
McMillan, William Singer, F.L.S.; Ardenholm, Maghull, 

Liverpool. 
Magdalen College ; Oxford. 
Manchester Literary and Philosophical Society; 36 George 

Street, Manchester. 

Manchester Microscopical Society ; 397 Cheetham Hill Road, 
Manchester. 

Manchester Public Free Libraries ; Manchester. 
Marlborough College Natural History Society; Marlhorougli . 
Massey, W. H. ; Twijford, Berks. 
Meiklejohn, John William S., M.D., F.L.S. ; 105 Holland 

Road, Kensington, W. 
Melbourne Public Library ; Alelhourne, Australia. 
Mennell, Henry Tuke, F.L.S. ; 10 St. Dunstan's Buildings, 

Great Toiver Street, E.C. 



12 LIST OF THE EAY SOCIETY. 

Michael, Albert Davidson, F.L.S., F.Z.S.,F.K.M.S., F.R.H.S., 
Vice-President ; The Warren, Studland, Wareham. 

Middlesbrough Free Libraries ; Middleshrough, 

Mitchell Library; 21 Miller Street, Glasgow. 

Moore, Henry; 12 Whiston Grove, Rotherham. 

Morey, Frank, F.L.S.; Wolverton, Cai'ishroohe Road, Newport, 
Isle of Wight. 

Morgan, Ralph ; 9 Clifton Hill, Exeter. 

Moulton, John C, F.L.S., F.E.S. ; Kuching, Sarawak, Borneo. 

Munich Royal Library ; Munich, Germany. 

Museum d'Histoire Naturelle ; Paris. 

Nebraska University Library ; Lincoln, Neh., U.S.A. 
Newcastle-upon-Tyne Literary and Philosophical Society ; 

Westgate Road, Newcastle-upon-Tyne. 
Newcastle-upon-Tyne Public Library; Newcastle-upon-Tyne, 
New South Wales, Public Library ol; Sydney, N.S.W. 
New South Wales, Royal Society of ; Sydney, N.S.W. 
Newstead, Robert, M.Sc.,F.R.S., A.L.S.,F.E.S.,Hon.F.R.H.S., 

Dutton Memorial Professor of Entomology; School of 

Tropical Medicine, The University, Liverpool. 
New York Public Library; New Yorh, U.S.A. 
Nicholson, Charles; Lansdowne House, Morley Street, Bradford. 
Niemeyer, Dr. Max, Halle a Saale 1, Germany. 
Noble, Sir Andrew, Bart., K.C.B., D.Sc, D.C.L., F.R.S., F.C.S., 

F.R.A.S. ; Jesmond Dene House, Newcastle-upon-Tyne. 
Norfolk and Norwich Library ; Norwich. 
Norman, TheRev.A.Merle,M.A.,D.C.L,LL.D.,F.R.S.,F.L.S., 

Hon. Canon of Durham ; The Red House, Berkhamsted. 
Northumberland, Durham, and Newcastle-upon-Tyne Natural 

History Society ; Hancock Museum, Newcastle-upon-Tyne. 
North Staffordshire Field Club ; Stone. 
Norwich Free Library; Norwich. 
Nottingham Free Public Libraries; Nottingham. 

OKamura, Prof. K. ; 4 Nichone, Shin-oganamachi, Ushigome, 
Tokyo, Japan. 

Oke, Alfred William, B.A., LL.M., F.L.S., F.G.S. ; 32 Den- 
mark Villas, Hove, Brighton. 



LIST OF THE RAY SOCIETY. 13 

Oldham, Charles, F.Z.S., M.B.O.U., Hon. Sec. Herts Nat. 

Hist. Soc. ; Kelvin, Berhhamsted. 
Ontario Agricultural College ; Guelph, Canada. 
Otago, University of ; Dunedin, New Zealand. 
Owens College (Christie Library) ; Manchester. 

Pack-Beresford, Denis K., D.L., B.A., M.R.I.A. ; Fenagh, 

Bagenalstown, Go. Carlow, Ireland. 
Paisley Philosophical Institution ; 3 County Place, Paisley. 
Peabody Institute ; Baltimore, Maryland, U.S.A. 
Perthshire Society of Natural Science; Tay Street, Perth. 
Philadelphia Academy of Natural Sciences; Philadelphia, 

Pa., U.S.A. 
Pickard-Cambridge, The Rev. O., M.A., F.R.S.; Bloxworth 

Rectory, Wareham. 
Plowman, T. ; Nystuen Lodge, Bycnllah Park, Enfield. 
Plymouth Institution ; Athenseuw , Plymonth. 
Portsmouth Free Public Library ; Town Hall, Portsmouth. 
Poulton, Edward B., M.A., D.Sc, LL.D., F.R.S., F.L.S., 

F.G-.S., F.Z.S., Hope Professor of Zoology, University of 

Oxford ; Wykeham House, Oxford. 
Preston Free Public Libraries and Museum; Preston. 

Quaritch, Bernard; 11 Grafton Street, Bond Street, W. 
Queen's University ; Belfafit. 

Quekett Microscopical Club ; 20 Hanover Square, W. 
Quelch, William Paul, F.R.B.S.; 65 Shakespeare Road, Han- 
well, W. 

Radcliffe Library ; Museum, Oxford. 
Rashleigh, Evelyn William; Stoheton, Saltash. 
Robertson, James Alexander; Lune View, Fleetwood. 
Rotherham Naturalists' Society; 6 Whiston Grove, Rotherham. 
Rothschild, The Hon. Lionel Walter, Ph.D., F.R.S., F.L.S., 

F.Z.S., Pres. Herts Nat. Hist. Soc. ; Tring Park, Tring. 
Royal Academy of Sciences ; Amsterdam. 
Royal Academy of Sciences ; Stockholm, Sweden. 
Royal College of Science ; Dublin. 
Royal Microscopical Society ; 20 Hanover Square, W. 
Royal Society; Burlington House, Piccadilly, W. 



14 LIST OF THE EAY SOCIETY. 

St. Albans Public Library ; 8t. Alhans. 

St. Andrews University Library ; >S'^. Andrew si, N.B. 

St. Catharine^s College ; Cambridge. 

Salisbury Microscopical Society; 14 Wyndham Terrace, 

Salishury. 
Scbarff, Robert Francis, Ph.D., B.Sc, F.L.S., F.Z.S., Yice- 

Peksident; Knockranny, Bray, co. Wichlow. 
Schmidle, Prof. W. ; Villa Hansagarteii, Konatang, Baden, 

Germany. 
Schmidt, Max, Ph.D. ; 95 iv Eppendorfer, Landdrasse, Ham- 
burg. 
Scott, Thomas, LL.D., F.L.S. ; 2 Bevanha Terrace, Aberdeen. 
Scottish, Royal, Museum ; Edinburgh. 
Scourfield, David Joseph, F.Z.S., F.R.M.S. ; 63 Queen's Road, 

Leytonstone, N.B. 
Sears, R. S. Wilson ; 1 Lisson Grove, Marylebone, N.W. 
Selborne Society ; 42 Bloomsbury Square, W.C. 
Sheffield Literary and Philosophical Society ; Leopold Street, 

Sheffield. 
Sidney-Sussex College ; Cambridge. 
Sion College Library; Victoria Embankment, E.C. 
Soar, Charles David, F.L.S., F.R.M.S. ; 37 Dryburgh Road, 

Putney, S.W. 
Somersetshire ArchaBological and Natural History Society; 

The Castle, Taunton. 
Southport Free Library ; Southport. 
Spicer, Henry, B.A., F.L.S., F.Gr.S.; 14 Aberdeen Park, 

Highbury, N. 

Sprague, T. B., M.A., LL.D. ; 29 Buckingham Terrace, 
Edinburgh. 

Stazione Zoologica ; Naples. 

Stechert, Gr. E. ; 2 Star Yard, Carey Street, W.C. 

Stoke Newington Public Libraries ; Church Street, N. 

Storey, J. E. ; 26 Grosvenor Road, Mlialley Range, Man- 
chester. 

Sunderland Library and Literary Society ; Fawcett Street, 
Sunderland. 

Taverner, Henry, F.R.M.S. ; 319 Seven Sisters' Road, Finsbury 
Park, N. 



LIST OF THE BAY SOCIETY. 15 

Tindall, William B.; 39 St. Mary Street, and 1315 Traders 

Banh Buildings, Toronto, Canada. 
Toronto, University of ; Toronto, Canada. 
Torquay Natural History Society ; Museum, Bahhacomhe Road, 

Torquay. 
Trinity College ; Cambridge. 
Trinity College ; Dublin. 

Trondhjem, Royal Library of; Trondhjem, Norway. 
Tupman, Lt.-Col. George Lyon, F.G.S., F.Z.S., F.R.A.S., 

F.R.M.S. ; Hillfoot Obse7'vatory, College Road, Harrow, 
Turner, Cliarles ; 20 Minster Road, Crichlewood, 'N.W. 

University College ; Cork. 

University College, London; Gower Street, W.C. 

Upsala Royal University Library; Upsala, Sweden. 

Victoria Institute ; Worcester. 

Wailes, George Herbert, F.L.S. ; Vancouver, B.C. (c/o G. 

Wailes & Co., 386 Euston Road, N.W.) 
Wales, National Library of ; Aberystwyth. 
Walker, Alfred 0., F.L.S., F.Z.S. ; Ulcombe Place, Maidstone. 
Walsingham, The Right Hon. Lord, M.A., LL.D., F.R.S., 

F.L.S., F.Z.S., F.E.S., M.B.O.U.; Merton Hall, Thetford. 
Warrington Municipal Museum ; Warrington. 
Webb, Wilfred Mark, F.L.S., F.R.M.S., Hon. Sec. Selborne 

Society ; Odstoch, Hanwell, W., and 42 Bloomsbury 

Square, W.C. 
Weg, Max; 1 Leplaystrass, Leipzig. 
Weigel, Oswald ; 1 Konigstrasse, Leipzig. 
Wesley, E. F., A.K.C.; 28 Essex Street, Strand, W.C. 
West Kent Natural History Society ; 42 Shooter's Hill Road, 

Blackheath, S.E. 

West, William, F.L.S. ; 26 Woodville Terrace, Bradford. 
Whittle, F. G. ; 3 Marine Avenue, Southend-on-Sea. 
Williamson, William; 9 Plewlands Terrace, Edinburgh. 
Wilson, H. Maclean, M.D., B.Sc, Chief Inspector, West 

Riding of Yorkshire Rivers Board ; Wakefield. 
Wilson, Joseph, F.R.M.S. ; Hillside, Avon Road, Upper 

Walthamstoiv, N.E. 



16 LIST OF THE RAY SOCIETY. 

Wood, J. H., M.B.; Tarrington, Ledbury. 
Worssam, Cecil ; 29 Ashhurnham Road, Bedford. 

Yale University Library; New Haven, U.S.A. 
Yorkshire Philosophical Society ; Museum, York. 

Zoological Society of London ; Regent's Park, N.W. 

The Advocate's Library, Edinburgh ; the Bodleian Library, Oxford ; the 
British Museum ; Cambridge University Library ; I'rinity College, Dublin ; 
and the National Library of Wales ; entered in the List, receive the 
Society's publications in accordance with the Copyright Act. 



GEOGRAPHICAL LIST OF THE SOCIETY. 

{Beyond the limits of the Loyidon Postal District,) 



GREAT BRITAIN AND IRELAND. 

ElSraLAND. 

Bedfordshire. 

Bedford — Worssa^m, C. 

Berkshire. 

Newbury — Bowman, J. H. Twyford — Massey, W. H. 

Buckinghamshire. 

Slough — Bentlej, E. 

Cambridgeshire. 

Cambridge — Assheton, E. Cambridge — Sidney- Sussex Col- 

Downing College. lege. 

Philosophical Library. Trinity College. 

St. Catherine's College. University Library. 

Cheshire. 
Chester — Elliott, Dr. J. Davenham. — Hewitt, D. B. 
Society of Natural Science. Helsby — Jones, W. L. 

Cornwall. 
Saltash — Eashleigh, E. W. Truro — Eoyal Institution of 

Cornwall. 

Derbyshire. 
Berby — Free Library and Museum. 

Devonshire. 
Exeter — Brokenshire, F. E. Plymouth — Crawshay, L. E. 

Morgan, E. Plymouth Institution. 

Lynmouth — Briggs, C. A. Torquay — Natural History So- 

Briggs, T. H. ciety. 

Mortehoe — Longstaff, Dr. G. B. 

Dorsetshire. 
Wareham — Michael, A. D. Wareham — Pickard- Cambridge, 

Eev. O. P. 

Durham. 
Sunderland — Library and Literary Society. 



18 



GEOGRAPHICAL LIST OF THE SOCIETY. 



Essex. 

Colchester — Laver, H. Southend — Whittle, F. Gr. 

Romford — Hope, Gr. Stanford - le - Hope — Burrows, 

Saffron Walden — Gibson, Miss. Rev. C. E. N. 



Bristol — Museum. 

Naturalists' Society. 

Public Libraries. 



Gloucestershire. 

Cheltenham — Knight, H. H. 



Natural Science Society. 



Hampshire. 
Newport, I. W. — Morey, F. Portsmouth— Free Public Library. 

Herefordshire. 
Ledbury — Wood, J. H. 

Hertfordshire. 



Berkhamsted — Norman, Canon 
A.M. 

Oldham, C. 

Hertford — Haileybury College. 
St. Alhans—Q^ihhs, A. E. 
Hertfordshire County 



Museum. 



St. Albans. — Public Library. 
rr2«.(7— Eothschild, Hon. W. 
Waltham Cross — Bowles, E. A. 
Watford — Barraud, P. J. 

Hertfordshire Natural 

History Society. 
Hopkinson, J. 



Kent. 



Bechenham — Harmer, Dr. S. F. 
Bro7nley — Naturalists' Society. 
Canterbury — East Kent Natural 

History Society. 
Dover — Burr, Dr. M. 
Eltham — Jones, A. H. 



Farnborough — Avebury , Lord . 
Folkestone — Free Public Library 

and Museum. 
Goudhurst — Fitzgerald, Eev. H, 
Maidstone — Green, E. E. 
Walker, A. O. 



Lancashire. 



B arrow-in- Ftii-ness — Public Lib- 
rary. 

Fleetwood — Eobertson, J. A. 

Liverpool — Bootle Free Library. 

Free Public Libraries. 

Liverpool Athenaeum. 

Macmillan, W. S. 

Microscopical Society. 

Newstead, Prof. E. 

School of Tropical Medi- 
cine. 



Manchester — Literary and Philo- 
sophical Society. 

Microscopical Society. 

— Owens College. 

Public Free Libraries. 

— Storey, J. E. 

Preston — Free Public Libraries. 
St. Helens — Cotton, J. 
Soutliport — Free Library. 
Warrington — Municipal Mu- 
seum. 



GEOaRAPHICAL LIST OF THE SOCIETY. 19 

Leicestershire. 
Leicester — Free Public Library. 

Middlesex. 
Enfield — Plowman, T. Harrow — Tiipman, Lt.-Col. G.L. 

Norfolk. 
Norwich — Free Library. Stalham — Gurney, R. 

Norfolk and Norwich Thetford — Walsingham, Lord. 

Library. 

Northumberland. 
Corhridge—ljoboviY, Miss M. Y. Newcastle-upon-Tyne — Noble, 
Newcastle - upon - Tyne — Arm- Sir A. 

strong College. Northumberland Natu- 

Literary and Philosophi- ral History Society. 

cal Society. Public Library. 

Nottinghamshire. 
Nottingham — Free Public Libraries. 

Oxfordshire. 
Oxford — Bodleian Library. Oxford — Poulton, Prof. E. B. 
Magdalen College. Eadcliffe Library. 

Somersetshire. 

Crewherne — Cleland, Dr. J. Taunton — Somersetshire Natural 

History Society. 

Staffordshire. 
Burton - upon - Trent — Friend, Stone — North Staffordshire 

Eev. H. Field Club. 

8tone — Bostock, E. D. 

Suffolk. 

Stowmarhet — Grooding, H. C. 

Surrey. 

Croydon — Public Libraries. Beigate — Chapman, Dr. T. A. 

Godalming —'Ea.stwood, J. E. Wallington — Drinkwater, J. W. 

Sussex. 
Bognor — Fletcher, W. H. B. Hastings — Natural History So- 

Brighton — Natural History So- ciety. 

ciety. Horsham — Godman, Dr. F. D. 

Oke, A. W. Pulhorough — Harley, Dr. J. 

Hastings — Bloomfield, Rev. E . N. 

Warwickshire. 
Birmingham — Bethune - Baker, Birmingham — Kenrick, Sir G-. H. 

G. T. Natural History and 

Free Libraries. Philosophical Society. 



20 GEOGRAPHICAL LIST OF THE SOCIETY. 

Westmoreland. 

Windermere — Gamett, F. W. 

Wiltshire. 

Marlborough — College Natural Salisbury — Collins, Hon. Mrs. H. 

History Society. Microscopical Society. 

Worcestershire. 
Bromsgrove — Dilks, A. C. Worcester — Victoria Institute. 

Yorkshire. 
Bradford — Bagshaw, W. Leeds — University Library. 



Natural History and Middlesbrough — Free Libraries. 

Microscopical Society. Motherham — Moore, H. 

Nicholson, C. Naturalists' Society. 

West, W. Scarborough — Irving, Dr. J. 

Halifax — Fielding, C. Sheffield — Literary and Philo- 

Public Library. sophical Society. 

Huddersfield — Naturalist and Shiptoyi — Eddy, J. R. 

Photographic Society. Wakefield — Binks, Mrs. I. 

Hull—FnhliQ Libraries. Wilson, Dr. H. M. 

Leeds — Gascoigne, Major F. York — Yorkshire Philosophical 



Philosophical Society. Society. 

Public Free Libraries. 



WALES. 

Cardiganshire. 

Aberystwyth — National Library of Wales. 

Glamorganshire. 

Cardiff — Free Libraries. 

Montgomeryshire. 

Machynlleth — Campbell, F. M. 

SCOTLAND. 

Aberdeenshire. 

Aberdeen — Scott, Dr. J. Aberdeen — University Library. 

Ayrshire. 

Kilmarnock — Public Library and Museum. 

Edinburghshire. 

Edinburgh — Advocates' Library. Edinburgh — Royal Scottish Mu- 

Crawford, W. C. seum. 

Davies, Dr. A. E. Sprague, Dr. T. B. 

Public Library. • University Library. 

Royal Society. ■ Williamson, W. 

Fipeshire. 
Largoward — Cox, B. C. St. Andrews — University Li- 

St. Andrews — Mcintosh, Prof. brary. 
W. C. 



GEOGEAPHICAL LIST OF THE SOCIETY. 21 

Haddingtonshire. 
Frestonldrh — Buchan-Hepburn, Sir A. 

Lanarkshire. 
Glasgow — Mitchell Library. Glasgow — Eoyal Philosophical 



Natural History Society. Society. 

University Library. 

Perthshire. 
Perth — Perthshire Society of Natural Science. 

Eenfrewshire. 
Paisley — Philosophical Institution. 

lEELAND. 

Antrim. 
Belfast — Belfast Library. Belfast — Queen's University. 

Carlow. 
Bagenalstown — Pack-Beresford, D. E. 

Cork, 
Corlc — University College. 

Dublin. 
Dublin — King's Inn Library. Dublin — Eoyal Dublin Society. 

National Library of Ire- Eoyal Irish Academy. 

land. Trinity College. 

Eoyal College of Science. Sandy cove — Colgan, N. 

WiCKLOW. 

5m2/— Scharff, Dr. E. F. 

CHANNEL ISLANDS. 

Guernsey — Guille-Alles Library. 

BRITISH DOMINIONS. 

AFEICA, SOUTH. 

Cape Colony. 
GraJiamstown — Albany Museum. 

Natal. 
Durban — James, E. D. 

ASIA. 

Borneo. 

Sarawak — Moulton, J. C. 



22 GEOGRAPHICAL LIST OF THE SOCIETY. 

AUSTRALIA. 

New South Wales. 
Sydney — Australian Museum. Sydney — Royal Society of New 

Public Library of New South Wales. 

South Wales. 

South Australia. 

Adelaide — Public Library. 

Victoria. 

Melhoiirne — Hardy, A. D. Melbourne — Public Library. 

Western Australia. 

So7dh Per^A— Giles, H. M. 

CANADA. 

British Columbia. 

Vancouver — Wailes, G-. H. 

Ontario. 

Guelph — Ontario Agricultural Toronto — Tindall, W. B. 

College. University. 

INDIA. 
Bihar — Fletcher, T. B. Calcutta— Geological Survey of 

India. 

NEW ZEALAND. 
Christ church — Philosophical In- Bunedin — University of Otago. 
stitution of Canterbury. 

FOREIGN COUNTRIES. 

AMERICA, UNITED STATES OF. 
California. 

Los Angeles — Public Library. 

Columbia, District of. 

Washington — Library of Congress. 

Illinois. 

Chicago — John Crerer Library. TJrhana — Illinois University Lib- 

University Library. rary. 

Maryland. 

Baltimore — Peabody Institute. 

Massachusetts. 

Boston — Public Library. Ca7nhridge — Harvard Museum of 

Society of Natural His- Comparative Zoology. 

tory. 



GEOGRAPHICAL LIST OF THE SOCIETY. 23 

Michigan. 

Detroit — Public Librtary. 

Nebraska. 

Lincoln — Nebraska Universitj. 

New Haven. 

Yale — University Library. 

New York. 

Buffalo — G-rosvenor Public Lib- New York — American Museum 

rary. of Natural History. 
Ithaca — Cornell University. Public Library. 

Pennsylvania. 
Philadelphia — Academy of Natural Sciences. 

AUSTEIA. 
Vienna — Grerold & Co. 

DENMAEK. 

Copenhagen — University Library. 

FEANCE. 

Lyons — Lyon Universitc Biblio- Paris — Museum d'Histoire Na- 

theque. turelle. 
Paris — Bibliotheque Nationale. Bennes — Guitel, Prof. F. 
Institut de France. 

GEEMANY. 

Baden — Sclimidle, Prof. W. Halle a Saale — Niemeyer, Dr. M. 

Berlin — Dames, F. L. Hamburg — Naturliistoriche Mu- 



Friedlilnder & Solin. seum. 



Junk, W. Schmidt, Dr. M. 

Eoyal Library. Heidelberg — University Library. 

Eoyal Zoological Kiel — University Library. 

Museum. Leipzig — Weg, M. 

Breslau — University Library. Weigel, 0. 

Frankfort — Baer & Co. Munich — Eoyal Library. 
Gottingen — University Library. 

HOLLAND. 

Leiden — Doesburgli, S. C, van. 

ITALY. 

Naples — Stazione Zoologica. 

JAPAN. 

Tokyo — OKamura, Prof. K. 



24 GEOGEAPHICAL LIST OF THE SOCIETY. 

NETHEELANDS. 
Amsterdam — Koyal Academy of Sciences. 

NOEWAY. 

Bergeii — Museums Bibliotek. Trondhj em— 'Royal Library. 

Christiana — University . 

SWEDEN. 

StocJcholm — Eoyal Academy of ZJ^saZa— University Library. 
g [Sciences. 

SWITZEELAND. 
Berne — Natural History Museum. 



LIST OF THE PUBLICATIONS 



OF THE 



RAY SOCIETY. 



For the First Year, 1844. 

1. Keports on tlie Progress of Zoology and Botany, 1841, 
1842. viii + 496 + xx pp. 8vo. 1845. 

The State of Zoology in Europe, as regards the Vertebrata. By 
C. L. Bonaparte. Transl. by H. E. Strickland, pp. 1-44. 

The Progress of Zoology in 1842. Transl. from the German by W. B. 
Macdonald. pp. 1-348. 

The Progress of Physiological Botany in 1841. By H. F. Link. 
Transl. by Edwin Lankester. pp. 1-104. Index, pp. i-xx. 

2. A Monograph of the British Nudibranchiate MoIIusca. 
By Joshua Alder and Albany Hancock. Part I. x + 20 pp., 
10 plates. Folio. 1845. 

3. Memorials of John Kay, consisting of his Life by Dr. 
Derham ; . . . with his Itineraries, etc. Edited by Edwin 
Lankester. xii + 220 pp. 8vo. 1846. 

For the Second Year, 1845. 

4. On the Alternation of Generations. By J. J. S. Steen- 
STRUP. Transl. from the German version of C. H. Lorenzen 
by George Busk, viii + 132 pp., 3 plates. 8vo. 1845. 

5. A Monograph of the British Nudibranchiate Mollusca. 
By Joshua Alder and Albany Hancock. Part II. iv -f 34 pp., 
13 plates. Folio. 1846. 

6. Reports and Papers on Botany, viii + 494 pp., 7 plates. 
.8vo. 1846. 

On the Morphology of the Coniferse. By J. J. Zuccarini. Transl. 
by George Busk. pp. 1-54, 441-444, pis. i-v. 

Eeport on Botanical Geography, 1842. By A. Grisebach. Transl. 
by W. B. Macdonald. pp. 55-212. 

Memoir on the Nuclei, Formation, and Growth of Vegetable Cells. 
By Carl Nageli. Transl. by Arthur Henfret. pp. 213-292, 445- 
459, pis. vi, vii. 

Eeport on Physiological Botany, 1842 and 1843. By H. F. Link. 
Transl. by J. Hudson, pp. 293-440. Index pp. 461-494. 



ii PUBLICATIONS OF THE EAY SOCIETY. 

For the Third Year, 184G. 

7. Outlines of the Geography of Plants. By F. J. F. 
Meyen. Transl. by Margaret Jobnston. x + 422 pp., 1 
plate. 8vo. 1846. 

8. The Organization of Trilobites. By Hermann Burmeister. 
Edited by Thomas Bell and Edward Forbes, xii + 136 pp., 
6 plates. Folio. 1846. 

9. A Monograph of the British Nudibranchiate Mollusca. 
By Joshua Alder and Albany Hancock. Part III. iv + 30 
pp., 12 plates. Folio. 1847. 

For the Fourth Year, 1847. 

10. Elements of Physiophilosophy. By Lorenz Oken. From 
the German by Alfred Tulk. xx + 6QQ pp. 8vo. 1847. 

11. Keports on Zoology for 1843, 1844. Transl. from the 
German by George Bosk, Alfred Tulk, and A. H. Haliday. 
viii + 596 pp. 8vo. 1847. 

Progress of Zoology in 1843. pp. 1-232. 

1844. pp. 233-564. 
Index, pp. 565-596. 

12. A Monograph of the British Naked-eyed Medusae. By 
Edward Forbes, viii + 104 pp., 13 plates. Folio. 1848. 

For the Fifth Year, 1848. 

13. Bibliographia Zoologia3 et Geologiae. A General Cata- 
logue of all Books, Tracts, and Memoirs on Zoology and 
Geology. By Louis Agassiz. Edited by H. E. Strickland. 
Vol. I. Periodicals, and A-B. xxvi + 506 pp. 8vo. 
1848. 

14. The Correspondence of John Ray. Edited by Edwin 
Lankester. xvi 4- 502 pp., 2 plates. 8vo. 1848. 

15. A Monograph of the British Nudibranchiate Mollusca. 
By Joshua Alder and Albany Hancock. Part IV. iv + 
28 pp., 12 plates. Folio. 1848. 

For the Sixth Year, 1849. 

16. Eeports and Papers on Botany. Edited by Arthur 
Henfrey. viii + 514 pp., 3 plates. 8vo. 1849. 

On the Structure of the Palm-stem. By H. von Mohl. pp. 1-92, 
495, pi. i. 

Ou the Nuclei, Formation, and Growth of Vegetable Cells. By Carl 
Nageli. Part II. pp. 93-158, 495-502, pis. ii, iii. 

On the Utricular Structures in the Contents of Cells. By Carl 
Naqeli. pp. 159-190. 

Report on Physiological Botany for 1844 and 1845. By H. T. Link. 
pp. 191-314. 



PUBLICATIONS OF THE HAY SOCIETY. 3 

Report on Geographical Botany for 1844. By A. Grisbbach. pp. 
315-414. 

Eeport on Geographical and Systematic Botany for 1845. By A. 
Grisebach. pp. 415-494. 

17. Tlie Natural History of the British Entomostraca. By 
W. Baied. viii 4- 364 pp., 36 plates. 8vo. 1850. 

For the Seventh Year, 1850. 

18. Bibliographia Zoologize et Geologise. A General Cata- 
logue of all Books, Tracts, and Memoirs on Zoology and 
Geology. By Louis Agassiz. Edited by H. E. Strickland. 
Vol. 11. C-F. iv + 492 pp. 8vo. 1850. 

19. A Monograph of the British Nudibranchiate Mollusca. 
By Joshua Aldee and Albany Hancock. Part V. iv -f 62 
pp., 16 plates. Folio. 1851. 

For the Eighth Year, 1851. 

20. The British Species of Angiocarpous Lichens, elucidated 
by their Sporidia. By the Rev. W. A. Leighton. vi -f 102 
pp., 30 plates. 8vo. 1851. 

21. A Monograph of the Sub-class Cirripedia. By Chaeles 
Daewin. The Lepadidse, or Pedunculated Cirripedes. xii + 
400 pp., 10 plates. 8vo. 1851. 

For the Ninth Year, 1852. 

22. Bibliographia Zoologiae et Geologiae. A General Cata- 
logue of all Books, Tracts, and Memoirs on Zoology and 
Geology, By Louis Agassiz. Edited by H. E. Strickland. 
Vol. IIL G-M. vi-t-658pp. 8vo. 1852. 

23. A Monograph of the British Nudibranchiate Mollusca. 
By Joshua Aldee and Albany Hancock. Part VI. iv -f 62 
pp., 12 plates. Folio. 1852. 

For the Tenth Year, 1853. 

24. Botanical and Physiological Memoirs. Edited by Arthur 
Henfrey. xvi + 568 pp., 6 plates. 8vo. 1853. 

The Phenomenon of Rejuvenescence in Nature. By Alexander 
Braun. pp. vii-xxvi, 1-342, pis. i-v. 

The Animal Natvire of Diatomeae. By G. Menighini. pp. 343-514. 

The Natural History of Protococcus pluvialis. By Ferdinand Cohn. 
pp. 515-564, pi. vi. 

25. A Monograph of the Sub-class Cirripedia. By Charles 
Darwin. The Balanidse, the Verrucidse, etc. viii + 684 pp., 
30 plates. Svo. 1854. 



4 PUBLICATIONS OF THE RAY SOCIETY. 

For the Eleventh Year, 1854. 

26. Bibliograpliia Zoologiae et Geologise. A General Cata- 
logue of all Books, Tracts, and Memoirs on Zoology and 
Geology. By Louis Agassiz. Edited by H. E. Strickland. 
Vol. IV. N-Z. vi + 604 pp. 8vo. 1854. 

For the Twelfth Year, 1855. 

27. A Monograph of the British Nudibranchiate Mollusca. 
By Joshua Alder and Albany Hancock. Part VII. vi -f 54 
4- 40 + xlvi pp., 9 plates. Folio. 1855. 

For the Thirteenth Year, 1856. 

28. A Monograph of the Fresh-water Polyzoa, including all 
the known species, both British and Foreign. By George 
James Allman. viii + 122 -\- 22 pp., 11 plates. Folio. 1856 

[1857]. 

For the Fourteenth Year, 1857. 

29. The Kecent Foraminif era of Great Britain. By William 
Crawford Williamson, xx + 108 pp., 7 plates. Folio. 
1858. 

For the Fifteenth Year, 1858. 

30. Tlie Oceanic Hydrozoa ; a Description of the Calyco- 
phorida) and Physophoridae. . . . By Thomas Henry Huxley. 
X 4- 144 + 24 pp., 12 plates. Folio. 1859. 

For the Sixteenth Year^ 1859. 

31. A History of the Spiders of Great Britain and Ireland. 
By John Blackwall. [Part I.] vi + 174 + 24 pp., 12 
plates. Folio. 1861. 

For the Seventeenth Year, 1860. 

32. Introduction to the Study of the Foraminifera. By 
William B. Carpenter, assisted by William K. Parker and 
T. Rupert Jones, xxii + 320 + 44 pp., 22 plates. Folio. 
1862. 

For the Eighteenth Year, 1861. 

33. On the Germination, Development, and Fructification of 
the Higher Cryptogamia, and on the Fructification of the 
Coniferae. By Wilhelm Hofmeister. Transl. by Frederick 
CuRREY. xviii + 506 pp., 65 plates. 8vo. 1862. 



PUBLICATIONS OF THE RAY SOCEETY. 5 

For the Nineteenth Year, 1862. 

34. A History of the Spiders of Grreat Britain and Ireland. 
By John Blackwall. Part II. iv + 210 (175-384) + 34 
pp., 17 plates. Folio. 1864. 

For the Twentieth Year, 1863. 

35. The Reptiles of British India. By Albert C. L. Gr. 
GiJNTHEE. xxviii + 452 pp., 26 plates. Folio. 1864. 

For the Twenty-first Year, 1864. 

36. A Monograph of the British Spongiadas. By J. S. 
BowERBANK. Vol. I. XX + 290 pp., 37 plates. 8vo. 1864. 

For the Twenty-second Year, 1865. 

37. The British Hemiptera. Yol. I. Hemiptera-Heteroptera. 
By John William Douglas and John Scott, xii + 628 + 
42 pp., 21 plates. 8vo. 1865. 

38. A Monograph of the British Spongiada?. By J. S. 
BowERBANK. Yol. II. XX + 388 pp. 8vo. 1866. 

For the Ticenty-third Year, 1866. 

39. The Miscellaneous Botanical Works of Robert Buown. 
[Edited by John J. Bennett.] Yol. I, containing I, G-eo- 
graphico-botanical, and II, Structural and Physiological 
Memoirs, viii + 612 pp. 8vo. 1866. 

40. Recent Memoirs on the Cetacea. Edited by William 
Henry Flower, xii + 312 pp., 6 plates. Folio. 1866. 

I. On the Greenland Kig-ht- Whale. By D. F. Eschricht and 
J. Eeinhardt. pp. 1-150, pis. i-vi. 

II. On the Species of the Grenus Orca inhabiting the Northern Seas. 
By D. F. Eschricht. pp. 151-188. 

III. Pseudorca crassidens, a Cetacean hitherto unknown in the Danish 
Fanna. By J. Eeinhardt. pp. 189-218. 

IV. Synopsis of the Cretaceous Mammalia of Scandinavia (Norway 
and Sweden). By W. Lilljeborg. pp. 219-310. 

41. NiTZSCH^s Pterylography, translated from the German^ 
Edited by Phjlip Lutley Sclatek. xii + 182 pp., 10 plates. 
Folio. 1867. 

For the Twenty -fourth Year, 1867. 

42. A Monograph on the Structure and Development of 
the Shoulder-girdle and Sternum in the Yertebrata. By 
W. Kitchen Parke u. xii + 240 + 60 pp., 30 plates. Folio. 
1868. 



6 PUBLICATIONS OF THE RAY SOCIETY. 

43. The Miscellaneous Botanical Works of Robert Brown. 
[Edited by John J. Bennett.] Vol. II, containing III, 
JSystematic Memoirs, and IV, Contributions to Systematic 
Works, viii + 780 pp. 8vo. 1868. 

For the Twenty-fifth Year, 1868. 

44. The Miscellaneous Botanical Works of Egbert Bkown. 
[Edited by John J. Bennett.] Vol. III. Atlas of Plates. 
iv + 16 pp., 38 plates. Folio. 1868. 

45. Vegetable Teratology, an Account of the Principal Devia- 
tions from the Usual Structure of Plants. By Maxwell 
T. Masters. With numerous illustrations by E. M.Williams. 
xxxviii + 534 pages. 8vo. 1869. 

For the Twenty-sixth Year, 1869. 

46. A Monograph of the Gymnoblastic or Tubularian 
Hydroids. By George James Allman. Part I, the Hydroida 
in General, xxii -f 154 + 24 pp., 12 plates. Folio. 1871. 

For the Twenty-seveiith Year, 1870. 

47. A Monograph of the Gymnoblastic or Tubularian 
Hydroids. By George James Allman. Conclusion of Part I, 
and Part II, containing descriptions of the Gene?a and Species 
of the Gymnoblastea. iv + 2 (xxiii, xxiv) + 296 (155-450) 
+ 22 pp., 11 plates (xiii-xxiii). Folio. 1872. 

For the Twenty-eighth Year, 1871. 

48. Monograph of the Collcmbola and Thysanura. By Sir 
John Lubbock, x + 276 pp., 78 plates. 8vo. 1873. 

For 'the Ttcenty-ninth Year, 1872. 

49. A Monograph of the British Annelids. By W. C. 
McIntosh. Part I. The Nemertians. xiv + 96 + 20 pp., 
10 plates. Folio. 1873. 

For the Thirtieth Year, 1873. 

50. A Monograph of the British Annelids. By W. C. 
McIntosh. Part I continued. The Nemertians. iv + 122 
(97-214, 213 a-d) + 26 pp., 13 plates (xi-xxiii). Folio. 
1874. 

For the Thirty-first Year, 1874. 

51. A Monograph of the British Spongiadae. By J. S. 
Bowerbank. Vol. III. xxviii + 368 pp., 92 plates. 8vo, 
1874. 



PUBLICATIONS OP THE RAY SOCIETY. 7 

For the Thirty -second Year, 1875. 

52. A Monograph of the British Aphides. By Geoege 
BowDLER Bdckton. Vol. I. X + 194 + 78 pp., 42 plates 
(A-C, i-xxxviii, iv bis). 8\^o. 1876. 

For the Thirty-third Year, 1876. 

53. A Monograph of the Free and Semi-parasitic Copepoda 
of the British Islands. By Gr. Stewardson Brady. Vol. I. 
iv + 148 + 72 pp., 36 plates (1-33, 10a, 24a, 24b). 8vo. 

1878. 

For the Thirty -fourth Year, 1877. 

54. A Monograph of the British Aphides. By George 
Bowdler Buckton. Vol. II. iv + 176 + 100 pp., 50 plates 
(xxxix-lxxxvi, li bis, Ixix bis) . 8vo. 1879. 

For the Thirty-fifth Year, 1878. 

55. A Monograph of the Free and Semi-parasitic Copepoda 
of the British Islands. By G. Stewardson Brady. Vol. II. 
iv + 182 + 98 pp., 49 plates (34-82). 8vo. 1880. 

For the Thirty-sixth Year, 1879. 

56. A Monograph of the Free and Semi-parasitic Copepoda 
of the British Islands. By G. Stewardson Brady. Vol. III. 
iv -{- 84 + 22 pp., 11 plates (83-93). 8vo. 1880. 

57. A Monograph of the British Spongiadse. By the late 
J. S. Bowerbank. Vol. IV (Supplementary). Edited, with 
additions, by the Eev. A. M. Norman, xviii + 250 + 34 pp., 
17 plates. 8vo. 1882. 

For the Thirty-seventh Year, 1880. 

58. A Monograph of the British Aphides. By George 
Bowdler Buckton. Vol. III. vi + 142 -|- 56 pp., 28 plates 
(Ixxxvii-cxiv). 8vo. 1881. 

For the Thirty-eighth Year, 1881. 

59. A Monograph of the British Phytophagous Hymenop- 
tera. By Peter Cameron. Vol. I. viii -f 340 + 42 pp., 21 
plates. 8vo. 1882. 

For the Thirty-ninth Year, 1882. 

60. A Monograph of the British Aphides. By George 
Bowdler Buckton. Vol. IV. x -f- 228 + 62 pp., 27 plates 
D-I, cxiv bis, cxv-cxxxiv). 8vo. 1883. 



8 PUBLICATIONS OF THE EAY SOCIETY. 

Fur the Fortieth Year, 1883. 

61. British Oribati da?. Bv Albert D. Michael. Yol.I. xii 
+ 336 + 62 pp., 31 plates *(i-xxiv, A-G). 8vo. 1884. 

For the Forty-first Year, 1884. 

62. A Monograph of the British Phytophagous Hymen- 
optera. By Petkr Cameron. Vol. II. vi + 234 + 54 pp., 
27 plates. 8vo. 1885. 

For the Forty-second Year, 1885. 

63. The Larvse of the British Butterflies and Moths. By 
the late William Buckler. Vol. I. (The Butterflies.) Edited 
by H. T. Stainton. xvi + 202 + 34 pp., 17 plates. 8vo. 
1886. 

For the Forty-third Year, 1886. 

64. The Larvae of the British Butterflies and Moths. By the 
late William Buckler. A^ol. II. (The Sphinges or Hawk- 
moths and part of tlie Bonibyces.) Edited by H. T. Stainton, 
xii + 172 + 36 pp., 18 plates (xviii-xxxv). 8vo. 1887. 

For the Forty-fourth Year, 1887. 

65. British Oribatidte. By Albert D. Michael. Vol. 11, 
xii + 322 (337-658) + 62 pp., 31 plates (xxv-liv, xlviiA), 
8vo. 1888. 

For the Forty-fifth Year, 1888. 

66. The Larv« of the British Butterflies and Moths. By 
the late William Buckler. A^ol. III. (The concluding por- 
tion of the Bombyces.) Edited by H. T. Stainton. xvi + 
80 + 36 pp., 18 plates (xxxvi-liii). 8vo. 1889. 

For the Forty-sixth Year, 1889. 

67. A Monograph of the British Phytophagous Hymen- 
optera. By Peter Cameron. Vol. III. vi + 274 + 34 pp., 
17 plates. 8vo. 1890. 

For the Forty-seventh Year, 1890. 

68. The Larva? of the British Butterflies and Moths. By 
the late William Buckler. Vol. IV. (The first portion of 
the Noctu^.) Edited by H. T. Stainton. xii + 116 + 32 
pp., 16 plates (liv-lxix). 8vo. 1891. 



PUBLICATIONS OF THE EAY SOCIETY. 9 

For the Forty-eighth Year, 1891. 

69. The LarvaB of the British Butterflies and Moths. Bj 
the late William Buckler. Vol. Y. (The second portion of 
the Noctuay.) Edited (in part) by the late H. T. Stainton. 
xii + 90 + 34 pp., 17 plates (Ixx-lxxxvi). 8vo. 1893. 

For the Forty-ninth Year, 1892. 

70. A Monograph of the British Phytophagous Hymen- 
optera. By Peter Cameron. Vol. IV. vi + 248 + 38 pp., 
19 plates. 8vo. 1893. 

For the Fiftieth Year, 1893. 

71. The Larvas of the British Butterflies and Moths. By 
the late William Buckler. Vol. VI. (The third and con- 
cluding portion of the Noctuae.) Edited by Gko. T. Porritt. 
xii + 142 + 38 pp., 19 plates (Ixxxvii-cv) . 8vo. 1895. 

For the Fifty-first Year, 1894. 

72. The Larvae of the British Butterflies and Moths. By 
the late William Buckler. Vol. VII. (The first portion of 
the Geometrae.) Edited by Geo. T. Porritt. xvi + 176 + 
44 pp., 22 plates (cvi-cxxvii). 8vo. 1897. 

For the Fifty-second Year, 1895. 

73. The Larvas of the British Butterflies and Moths. By 
tne late William Buckler. Vol. VIII. (The concluding 
portion of the Geometras.) Edited by Geo. T. Porritt. 
xii + 120 + 70 pp., 20 plates (cxxviii-cxlvii) . 8vo. 1899. 

For the Fifty-third Year, 1896. 

74. The Tailless Batrachians of Europe. By G. A. 
Boulenger. Part I. viii + 210 pp., 10 plates. 8vo. 1897. 

For the Fifty-fourth Year, 1897. 

75. The Tailless Batrachians of Europe. By G. A. 
Boulenger. Part II. ii + 166 (211-376) pp., 14 plates 
(xi-xxiv). 8vo. 1898. 

For the Fifty-fifth Year, 1898. 

76. A Monograph of the British Annelids. By William 
Carmighael McIntosh. Part II. Polychaeta. Amphinomidae 
to SigalionidjB. x + 228 (215-442) + 40 pp., 20 plates 
(xxiv-xlii, xxviA). Folio. 1900. 



10 PUBLICATIONS OF THE EAY SOCIETX". 

Fo)' the Fifty-sixth Year, 1899. 

77. The Larvas of the British Butterflies and Moths. By the 
late William Bucklee. Vol. IX. (The Deltoides, Pyrales, 
Crambites, Tortrices, Tinese, and Pterophori.) Edited by 
G-Eo. T. PoRRiTT. xviii + 420 + 34 pp., 17 plates (cxlviii- 
clxiv). 8vo. 1901. 

For the Fifty-seventh Year, 1900. 

78. Monograph of tlie Coccidae of the British Isles. By 
Egbert Newstead. Vol. I. xii -h 220 + 78 pp., 39 plates 
(a-e, i-xxxiv). 8vo. 1901. 

For the Fifty-eighth Year, 1901. 

79. British Tyroglyphidas. By Albert D. Michael. Vol. I. 
xvi + 294 + 44 pp., 22 plates (a-c, i-xix). 8vo. 1901. 

For the Fifty-ninth Year, 1902. 

80. Monograph of the Coccidae of the iBritish Isles. By 
EoBEiiT Newstead. VoI. II. viii + 270 + 84 pp., 42 plates 
(f, xxxv-lxxv). 8vo. 1903. 

For the Sixtieth Year, 1903. 

81. British Tyroglyphidae. By Albert D. Michael. Vol.11, 
xii + 184 + 40 pp., 20 plates (xx-xxxix). 8vo. 1903. 

For the Sixty-first Year, 1904. 

82. A Monograph of the British Desmidiacea3. By W. and 
G. S. West. Vol. I. xxxvi + 224 + 64 pp., 32 plates. 
8vo. 1904. 

83. British Tiinicata. By the late Joshua Alder and the 
late Albany Hancock. Edited by John Hopkinson. Vol. I. 
With a History of the Work by Canon A. M. Norman. 
xvi + 146 + 42 pp., 20 plates, and frontispiece. 8vo. 
1905. 

For the Sixty-second Year, 1905. 

84. A Monograph of the British Desmidiaceae. By W. and 
G-. S. West. Vol. II. x + 206 + 64 pp., 32 plates. 8vo. 
1905. 

85. The British Freshwater Rhizopoda and Heliozoa. By 
James Cash, assisted by John Hopkinson. Vol. I. ^J'he 
Rhizopoda, Part I. x + 150 -f 32 pp., 16 plates. 8vo. 1905. 



PUBLICATIONS OF THE RAY SOCIETY. 11 

For the Sixty-third Year, 1906. 

86. The British Tunicata. By the late Joshua Alder and the 
late Albany Hancock. Edited by John Hopkinson. Vol. 

II. With Lives of the Authors by Canon Norman and 
Dennis Embleton. xxviii + 164 + 62 pp., 30 plates (xxi-1), 
and frontispiece. 8vo. 1907. 

For the Sixty -fourth Year, 1907. 

87. A Monograph of the British Annelids. By William 
Carmichael McIntosh. Yol. II, Parti. Polychaeta. Neph- 
thydidae to Syllida3. viii + 232 + 46 pp., 22 plates (xliii-1, 
Ivli-lxx). Folio. 1908. 

For the Sixty-fifth Year, 1908. 

88. A Monograph of the British Desmidiacese. By W. and 
G. S. West. Vol. HI. xvi + 274 + 62 pp., 31 plates (Ixv- 
xcv). 8vo. 1908. 

89. The British Freshwater Ehizopoda and Heliozoa. By 
the late James Cash, assisted by John Hopkinson. Vol. II. 
The Rhizopoda, Part II. xviii + 168 + 82 pp., 16 plates 
(xvii-xxxii), and frontispiece. 8vo. 1909. 

For the Sivty-sixlh Year, 1909. 

90. The British Nudibranchiate Mollusca. By the late 
Joshua Alder and the late Albany Hancock. Part 8 (supple- 
mentary). Text by Sir Charles Eliot, viii +198+18 pp., 
8 plates. Folio. 1910. 

For the Sixty -seventh Year, 1910. 

91. A Monograph of the British Annelids. By William 
Carmichael McIntosh. Vol. II, Part 2. Polychaeta. Syllid^ 
to Ariciidte. vii + 292 (233-524) + 46 pp., 23 plates (li-lvi, 
Ixxi-lxxxvii). Folio. 1910. 

For the Sixty-eighth Year, 1911. 

92. A Monograph of the British Desmidiaceae, By W. and 
G. S. West. Vol. IV. xW + 194 + 66 pp., 33 plates 
(xcvi-cxxviii). 8vo. 1912. 

93. The British Tunicata. By the late Joshua Alder and 
the late Albany Hancock. Edited by John Hopkinson. Vol. 

III. xii + 114 + 34 pp., 16 plates (li-lxvi), and frontispiece. 
8vo. 1912. 



12 PUBLICATIONS OF THE EAY SOCIETY. 

For the Sixty-ninth Year, 1912. 

94. A Bibliograpliy of the Tuiiicata. By John Hopkinson. 
xii + 288 pp. 8vo. 1913. 

95. The British Parasitic Copepoda. By Dr. Thomas Scott 
and Andeew Scott. Vol. I (Copepoda parasitic on Fishes, 
Part I), xii + 256 pp., 2 plates. 8vo. 1913. 



In Course of Publication. 

The British Desmidiaceae. By W. West and Prof. G. S. 
West. 

The British Freslivvater Rhizopoda and Heliozoa. By 
the late James Cash and (alter Vol. II) G. H. Wailes. 

The British Marine Annelids. By Prof. W. C. McIntosh. 

The British Parasitic Copepoda. By Dr. Thomas Scott 
and Andrew Scott. (Vol. II for 1913.) 



Preparing for Pitblication . 

The British Centipedes and Millepedes. By W. M. Wejbb. 

The British Earthworms. By the Rev. Hilderic Friend. 
' The British Hydrachnida). By C. D. Soar and W. 
Williamson. 

The British Ixodoidea. By W. F. Cooper and L. E. Robinson. 

The Earwigs of the World. By Dr. Malcolm Burr. 

March, 1913. 



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