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THE LIBRARY

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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.

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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 0 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 0 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-
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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
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51. 1899 Giesbrecht, TF. Die Asterocheriden des Golfes.

von Neapel und der Angrenzenden Meeres. —
Abschuitte. (A.u.d. Tit. : Fauna und Flora des
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52. 1842. Goodsir, H. Note sur le developpement des oeufs.

du Caligiis et sur les metamorphoses que ce-
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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-
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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.
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60. 1884 idem. Crustaces rares ou nouveaux des cotes de

France. — Ann. Scienc. Nat., 6 ser., ZooL, Vol.
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61. 1783 Hermann,/. Helminthologische Bemerkungen. —

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61a. 1857 HoBven, /. van der. Note sur les genres Cecrops
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62. 1799 Holten, H. 8. Acta Danica, Vol. V.

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63a. 1897 Horst, R. PhiloHhagoriscus serratns Kr. [Dine-
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64. 1824 Johnston, G. Notice respecting the genus Cali-

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65. 1835 idem. Lernsea uncinata. — Loudon's Mag. of Nat.

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Q>Q. 1835 i*t?em. Pandarusalatus nndlamnse. Loc. cit. VIII,
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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
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68a. 1853 KoUiker, A. Bericht iiber eiiiige im Herbst 1852
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LETEltATURE CHIEFLY EEFEKRED TO. 239

69. 1835 KoUar, V. Beltrage zur Kenntnisslernaeenartigen

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70. 1837-38 Kroyer, H. Om Snyltekrebsene, isaer med

Hensyn til den dansk Fauna. — Naturhistorisk
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71. 1863 idem. Bidrag til Kundskab om Snyltekrebsene. —

Naturhistorisk Tidsskrift, 3 die. Kaekke, II Bind,
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72. J 815-22 Lamarck, G. B. P. de. Histoire naturelle des

Animaux sans Vertebres, Vol. V. (1818.) Seven
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73. 1814 Leach, W. E. Crustaceology. (In) Edinburgh

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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-

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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

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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.
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Albany Museum ; Grahamstown, Cafe Colony, S. Africa,
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Assheton, Richard, M.A., F.L.S., F.Gr.S., F.Z.S. ; Grantchester,

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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

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b LIST OF THE KAY SOCIETY.

Bestow, Charles Horton, F.R.M.S. ; Melford House, 43 Upper

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LIST OF THE RAY SOCIETY. 7

Burr, Malcolm, D.Sc, F.L.S., F.GT.S., F.Z.S., F.E.S.; Castle

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Cambridge Philosophical Library ; New Museums, Gamhridge,

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Campbell, Francis Maule, F.L.S., F.E.S. ; Brynllwydwyn,

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8 LIST OF THE EAY SOCIETY.

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Folkestone Free Public Library and Museum ; Folkestone.

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LIST OF THE RAY SOCIETY. 9

Gascoigne, Major French; Lotherton Hall, Aherford, Leeds.
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W.
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Glasgow University Library; Glasgoiv.
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Halifax Public Library ; Belle View, Halifax.

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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.
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10 LIST OF THE RAY SOCIETY.

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Jackson, B. Daydon, Ph.D., F.L.S., General Secretary of the

Linnean Society; 21 Cautlei/ Avenue, Clapham Common,

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Kiel University Library ; Kiel, Germany.

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King's Inn Library ; Dublin.

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LIST OF THE EAY SOCIETY. 11

Laver, Henry, M.R.C.S. ; 43 Head Street, Colchester.
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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.
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Manchester Public Free Libraries ; Manchester.
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12 LIST OF THE EAY SOCIETY.

Michael, Albert Davidson, F.L.S., F.Z.S.,F.K.M.S., F.R.H.S.,
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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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