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PRESENTED
The Trustees
THE BRITISH MUSEUM.
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NATIONAL ANTARCTIC EXPEDITION
1901-1904
NATURAL HISTORY
VOL. IV.
ZOOLOGY
(VARIOUS INVERTEBRATA)
LONDON
PRINTED BY ORDER OF THE TRUSTEES OF
THE BRITISH MUSEUM
[908
I.-/// A'ig/ifs
SOLD BY LONGMANS AND Co., 39 PATERNOSTER Row, E.G. ; BERNARD QUARITCH, 15 PICCADILLY, W. ;
DULAU AND Co., 37 SOHO SQUARE, \V. ;
AND AT
THE BRITISH MUSEUM (NATURAL HISTORY), CROMWELL ROAD, LONDON, S.W.
PREFACE.
WHEN, in 1901, the Expedition of the S.S. 'Discovery,' under Captain Scott, R.X..
was sent to the Antarctic Regions, the Trustees of the British Museum gave their
assistance to this national enterprise by allowing the cases containing the natural
history specimens which might be obtained by the Expedition to be sent to the
Natural History Museum for unpacking and sorting. They further undertook to
publish a detailed report on the collections so obtained, under the superintendence
of the Director of the Natural History Departments.
Some of the most important collections have been dealt with by naturalists who
were members of the Expedition. Thus, the Mammals* and Birds* are described by
Dr. Edward A. Wilson, the Isopoda and Pycnogonida* by Mr. T. V. Hodgson, and
the Rocks* (in relation to Field Geology) by Mr. H. T. Ferrar. Other groups have
been dealt with by members of the staff of the Natural History Departments of
the British Museum : Mr. Boulenger describes the Fishes* ; Mr. E. A. Smith, tin-
Gastropoda,* Lam*llibranchia,' ;:: and Brachiopoda* ; Mr. Jeffrey Bell, the Echinoderma* ;
Dr. Caiman, the Crustacea Decapoda,* and the Cumacea* ; Mr. Kirkpatrick, the nmi-
calcareous Sponges* ; whilst Dr. G. T. Prior has prepared a petrographical description
of .the Rock-specimens.*
It has been necessary to obtain the assistance of other specialists in order to deal
with the rest of the collections. So far as the latter group of contributors is
concerned, the following is a list of the subject-matters, together with the name of the
naturalist who has undertaken the work in each case :
EMBRYOS *OF SEALS
*ANATOMY OF EMPEROR PENGUIN
TUNICATA .
*CEPHALODISUUS .
*CEPHALOPODA
*NUDIBRAN('HS AND PxEROPODS
POLYZOA ....
*EGGS AND YOUNG OF ASTERIAS
:i; . \MPHIPODA
*S(JHIZOPODA
*NEBALI.E .
*OSTRACODA
*COPEPODA
Di;. MARRETT TIMS.
Mi:. \V. 1'. I'v.'KAKT.
PROF. HERMMAX.
DR. RIDEWOOD.
Di:. HOYLE.
Sn; CHAKI.KS ELIOT, K.C.M.G.
MR. II. W. BURROWS.
PROF. MAC-BRIDE.
MR. A. <>. WALKER.
Mi;. HOLT.
DR. J. THIEI.K.
PROF. BRADY.
I>|;. WoLKKXIiEX.
iv PREFACE.
*CIRRIPEDIA ...... PROF. GRUVEL.
*MYZOSTOMA ...... PROF. v. GRAFF.
*ACARI ....... DR. TROUESSART.
*COLLEMBOLA ...... PROF. CARPENTER.
POLYCH^TA ...... PROF. EHLERS.
*GEPHYRIA ...... MR. A. E. SHIPLEY.
*CH.ETOGNATHA ...... DR. FOWLER.
NEMERTINES ...... PROF. HUBRECHT.
FREE PLATYHELMINTHES .... MR. F. F. LAIDLAW.
*CESTODA ....... MR. A. E. SHIPLEY.
*NEMATODA ...... DR. v. LINSTOW.
*ZOANTHARIA . . . . . .MR. CLUBB.
*ALCYONARIA AND PENNATULIDA . . . PROF. HICKSON.
HYDROMEDUS.E ...... MR. E. T. BROWNE.
*CALCAREOUS SPONGES ..... MR. FREWEN JENKIN.
RADIOLARIA ...... MR. LEWIS H. GOUGH.
* MOSSES ....... M. JULES CARDOT.
LICHENS ....... MR. DARBISHIRE.
*ALG.E (MARINE) ...... MRS. GEPP.
ALG/E (FRESH-WATER) ..... DR. FRITSCH.
*A.LGJE (CALCAREOUS) . ... DR. FOSLIE.
PHYTOPLANKTON ...... DR. LEWIS H. GOUGH.
The work of securing the assistance of these specialists and of distributing the
collections has been performed by Mr. .Jeffrey Bell, of the Zoological Department, who
has also acted as sub-editor of the Zoological and Botanical portions of the reports.
The Keeper of Minerals, Mr. Fletcher, has superintended the reports in the subjects
belonging to his department.
The Director desires to acknowledge the ability and energy which have been
brought to bear on the preparation of the Zoological reports by Mr. Jeffrey Bell.
Owing to his care, the reports have been got ready by the various contributors and
published within a reasonable time after the return of the ' Discovery ' from the
Antarctic Regions. Neither trouble nor expense has been spared in order to
render the illustration and presentation of the Natural History of the Expedition
worthy of the generous efforts both of Captain Scott and his fellow-explorers and
of those who provided the funds for that enterprise.
E. RAY LANKESTER.
October, 1906.
* Have been or are now published. 1/3/1908.
CONTENTS OF VOL IV.
MOLLUSCA.
VII. SOLENOGASTRES. By DR. H. F. NlERSTRASZ . . (13 pp., 2 Pis.)
ARTHROPOD A.
(A) IXSECTA.
APTERA. By G. H. CARPENTER, B.Sc., M.R.I.A. (5pp., 1 PI.)
(B) CRUSTACEA.
VII. SCHIZOPODA. By \Y. ]\1. TATTKKSALL, .M.Sc. . . (42 pp., 8 Pis.)
VIII. COPEPODA. By R. NORRIS WOLFENDEN, M.D. . . (44 pp., 7 Pis.)
ECHINODERMA.
I. ECHINODERMA. By F. JEFFREY BELL, M.A. . . (1(5 pp., 5 Pis.)
II. ECHIXODERM LARV.E. By E. W. MACBBIDK, M.A., F.R.S..
and J. 0. SIMPSON, B.Sr . (9 pp^ i pj.)
MYZOSTOMIDAE.
By DR. RUDOLF RITTER v. STUMMER-TRAUNFKLS (-<; pp., 1 I'D
SIPUNCULOEDEA.
1!\ \\'. V. LAN< HESTKI:. .M.A. . . . (' pp. i
COELENTERA.
IV. ACTINI.P.. By J. A. ('LI;I:I:. M.S.. (12 pp.. :j Pis.)
PORIFERA.
II. TETRAXMNIDA. By R. KIRKPATRK K . . . (56 pp., 19 Pis.)
III.- 'AI.CAI:KA. By ( '. F. JKNKIN. B.A. . (50 pp., 12 Pis.)
'
MOLLUSCA.
VII. SOLENOGASTEES.
By Dr. II. F. NIEKSTRA--/.
(2 Plates.)
THE 'Discovery' expedition brought back one specimen of Soleuogastres from 77 50'
30" S., 165 40' 5" E., 100 fathoms, of which Fig. 1, A gives a representation. The
length is 23 mm., the diameter 2 '25 mm., the index therefore about 10. Proximally
the animal is blunt, slightly broadened, with a slit-like mouth (Fig. 1, B). The distal
part terminates in a dorsal prolongation. On the ventral side the rhomboid opening
of the cloaca is distinctly visible, as also the ventral groove, which runs as far as that
opening (Fig. 1, C). The opening of the ciliated groove (" Flimmcrhohle " of Wiivn)
is also visible. The animal has no lustre. When slightly enlarged, spicules arc seen
crossing each other at obtuse angles. The colour of the animal in alcohol is a yellowish
brown.
After being decalcified in nitric acid (1 per cent, in alcohol 90 per cent.) and
being stained in iron carmalum for twenty-four hours, transverse sections were made
of the proximal and distal portions for the study of the internal organs. The spicules
of the different parts of the body were isolated in Eau de Javelle. They ;uv small,
but strictly uniform. They belong to the true Proneomenia type : hollow, straight, or
more or less curved or S-shaped calcareous tubes, which end either sharply or bluntly
(Fig. 2). In some cases the base is slightly broadened. I have not succeeded in
finding spicules of different shapes along the ventral groove, nor at the proximal and
distal portions. At the proximal end, however, they are somewhat more curved.
The spicules arc arranged indifferent layers upon each other. The whole of the
strong cuticle is pierced by numerous papillae. The greater part of these are oval, pear-
or dub-shapcd, sometimes round, and they show some transparent cells (Fig. 4). They
are placed on strong multiuuclear stalks, the cells of which are more or less fibrous.
Amongst the transparent cells granular supporting cells are found. It is worthy of
note that these papillae are often branched. The side-branch never readies the length
of the stalk, though ending also in a papilla. In most cases these epidermal papillae
do not open on to the exterior, but remain closed. Thielc believes the papillae to be
sensitive, an opinion which I share (13, p. 280). In the stalk a thin thread may
occur, arising from the circular muscular layer. Whether this thread is of a nervous
character or not remains undecided. There is. however, no penetration into the circular
H. F. NIERSTRASZ.
muscular layer by nervous threads. I am also inclined to ascribe to the epidermal
papillae, and more especially to the granular cells amongst the transparent ones, a
secretory function, if not an excretory function as well. The purpose of such a
secretion is doubtful. Perhaps it prevents the cuticle from damage by causing small
particles to adhere to its surface. In any case, it is remarkable that the cuticle of the
Soleuogastres is nearly always covered with a thin layer of mud firmly sticking to it.
The spicules are formed in small cellular accumulations in the epidermis. Such
accumulations frequently occur, and consist of a few transparent cells (Fig. 3, A). In
one of the cells the spicule is formed. This cell often has two nuclei, a fact which may
give rise to the supposition that there are two cells that form the spicule. The
spicules are pushed to the periphery of the cuticle, and are connected with the
epidermis by short stalks. The older spicules lose their connection with these stalks,
but preserve the mother-cells, which surround their base like a cap (Fig. 3, B). In
the case of the oldest spicules, however, this cap has also disappeared. Nowhere did
I meet with a formation of the spicules such as Thiele described in Amphimenia
neapolitana, Thiele (13, p. 246). Thiele is also of the opinion that it is one cell only
that produces a spicule.
The ventral groove is distinct and deep. In it three folds of the epidermis are
found, a large and broad median fold and two small lateral ones. All three run as far
as the opening of the cloaca. Close before the latter the groove and the median fold
broaden considerably. The folds are formed of a single layer of epithelial cells. All
along the median fold the ventral glands open between its cells. These glands (" hintere
Bauchdriise," of Wiren) are strongly developed, especially at the proximal end of the
body, where they entirely surround the digestive tract (Figs. 6-11, vg). They are
conspicuous by their more or less distinct vesicular structure and granular contents,
which stain intensely. At the proximal part of the body, amongst the strongly
developed ventral glands, voluminous anterior glands (" vordere Bauchdriise," of
Wire'n) are also found. They are of a more delicate structure, and stain much less
intensely (Fig. 6, ag). They open out between the epithelial cells which invest
the ciliated cavity ("Flimmerhohle," Fig. 6, /). The latter is large, and divided by
a median dorsal fold. The wall carries strong cilia.
There are two dorso-ternimal sense organs situated on the distal lip behind the
cloaca. Fig. 5 shows three sections through the more proximal of these organs. The
epithelium (ty) shows an evagination (Fig. 5, C) into which the muscular layers and the
connective tissue continue (TO). This evagination separates ventrally from the epidermis
and pierces the thick cuticle (cut. Fig. 5, B). It forms a pedicle (pcd.), upon which a
round knob is resting, the sense organ visible at the surface of the cuticle (Fig. 5, A,
dtn). The external layer of this knob consists of high epithelial cells, amongst which
slender, fine, easily staining cells are found, which I believe to be sense-organs.
The mouth-slit is terminal and vertical. It leads into a spacious mouth-cavity,
provided with numerous ramified cirri, as is the case in other forms. A horseshoe-
SOLENOGASTRES. 3
shaped sense-organ is wanting, though usually present in other species. The digestive
tract opens to the exterior at the same place as the mouth-cavity, with which it has,
however, no communication (cf. Dinomenia, 6, p. 12, PI. 2, Fig. 43). I share Thiele's
opinion, who does not regard the so-called mouth-cavity as a part of the digestive
tract, but considers it an independent ectodermal sense-cavity (17, pp. 308 and 314).
A proper mouth-cavity occurs in the proximal part of the digestive tract, which is
slightly dilated, and the wall of which presents numerous folds. The epithelial cells
of which the wall of this part is composed are very transparent and have large oval
nuclei. The wall is surrounded by a thin circular muscular layer. The folds, as it
appears to me, allow this portion of the digestive tract to extend considerably after
the taking of food. I do not believe them to be sensitive (Figs. G and 7, /'//).
Gradually this portion passes into the following part of the alimentary canal. The
latter is much wider, and likewise provided with a strongly folded wall (Fig. 8, ves).
Its cells are taller than those of the first portion, and have elongated nuclei ; they arc
covered with a cuticle. Consequently these two parts of the digestive tract are easilv
distinguished from one another. More distally, however, between these slender cells
there occur broader ones with more oval nuclei, the cuticle no longer being easily
discerned throughout. Finally the spacious intestine unites \vith the proximal caecum
(Fig. 8, c).
In the second portion of the digestive tract the radula, which is of a very
peculiar structure, is found. In the digestive tract a proximally directed cone appeals
with a strong ventral fold (Fig. 8, en). This cone is beset throughout with radula-
teeth (Fig. 12, c). For its further structure compare figures 12-14 and 8-11. An
epithelial layer appears (), which covers the radula. The ventral wall of the
digestive tract (l>) unites with the ventral wall of the cone (c) ; the epithelial layer
a with the side-walls of the cone. The two lateral walls of the proximal part of the
alimentary canal unite dorsal from the cone. By this complicated process the cone,
which on the outside is beset with small teeth, is divided into two closed pouches ; the
latter are on the inner side closely beset with teeth and are situated outside the
alimentary canal. The dorsal pouch is the radula-sac, the ventral contains adult
radula-teeth and, distally, ends blindly. Consequently the ventral pouch is but
a ventral caecum of the proximal part of the digestive tract, directed distally ;
icrili on the ventral wall are wanting, in the distal part teeth only occur on
the dnr.sil wall. The radular teeth are well developed (Fig. 10). They are
triangular, with a broad base, and more or less strongly curved, often recurved
at the point (Fig. 17, B). Frequently forms occur as represented in Fig. 17, A ;
often other forms are met with, which, however, I consider fragments of whole
teeth. Fig. 17, C shows a tooth as seen from the ventral side : the base
appears to be broader than the curved part. The teeth are placed in irregular
rows, and rest in crypts of the epithelium (Fig. Ifi). A basal membrane is
present. This is remarkable ; Thiele lays stress on the absence of such a membrane
B 2
4 H. F. NIERRTRASZ.
in different forms and doubts its existence (17, pp. 268 and 315). As has been
indicated by me elsewhere, the existence of a basal membrane has to be accepted for
several forms (9). The membrane is stained red with carmalum, consequently its
structure differs from that of the chitinous teeth. Probably it must be considered
as a cuticle, and so its origin is entirely different from that of the radula-teeth, which
are formed in the distal portion of the radula-sac by large odontoblasts. In this
respect Proneomenia Jixcoveri/i resembles those forms, already described by me
elsewhere (9). So we have to do here with a multiserial radula with basal membrane.
Of any double character of the radula, viz., of the existence of a right and a left
half, no trace is to be found. The distal portion of the radula-sac, however, is
divided into two parts, a right and a left one (Figs. 10, 20) ; in both parts radula-
teeth are formed by a thick cushion of odontoblasts. Here also the radula originates
from the coalescence of a right and a left portion.
The wall of the cone and the dorsal wall of the ventral pouch consist of
epithelial cells with round nuclei, between which numerous extremely slender
supporting cells occur with elongated nuclei (Fig. 16). On the ventral wall of the
ventral pouch and on the dorsal wall of its distal end supporting cells are absent ;
this ventral wall is formed by cylindrical epithelium with oval nuclei, whilst the
constituent cells of the dorsal wall of the distal portion are higher and more
cylindrical. One part of this dorsal wall more especially deserves notice ; the distal
and median part is invaginated and of an entirely different structure. Here very long
and slender cells are found, with pointed nuclei and granular contents. It leaves the
impression that these cells have pushed themselves wedgewise between the epithelial
cells; yet no sharp line can be drawn between these two elements (Fig. 18). Upon
this wedge-shaped part the proximal rows of the radula-teeth are resting ; the basal
membrane, however, fails here. As to the significance of this wedge I am at a loss.
There seems to be a connection between the wedge and the layer of muscular and
connective tissue surrounding the radula-sac, numerous fibrils pushing themselves
between the cells of the wedge. I am bound to mention its existence, having found
in a similar ventral pouch containing the proximal portion of the radula in Proneomenia
weberi, a regeneration either of the cuticle or of the radula (6, p. 4). But in this
new form I do not believe we have to do with a regeneration ; its significance remains
undecided.
Still, in another respect the ventral pouch is of great importance, as the
salivary glands open into it (Figs. 11 and 15 sg). These are tube-shaped, and extend
a long way distally ; they merge separately into the ventral wall of the ventral
pouch.
The connective tissue and the muscles, which are in connection with the radular
apparatus, are strongly developed. Ventrally and laterally from the radula-sac
numerous crystal-clear cells occur ; usually those are small, sometimes, however, of
large size (Fig. 10 <<). They serve as a support for the surrounding connective
SOLENOGASTRES. 5
tissue and muscles. The course of these muscles is difficult to trace ; they surround
the radula-sac and the distal portion of the ventral pouch ; a very broad transverse
muscle takes its course between the radula-sac and the ventral pouch (Fig. 10, ?).
Around the salivary glands a thin muscular layer likewise occurs. I do not know
how to explain the mechanism of the radular movement. No doubt, the diverse
muscles allow of movement in various directions.
A subradular organ, as found by Heath (1) in Proneomenia and Rhopalomenia,
is wanting.
The spacious intestine is formed by the coalescence of the already described
portion of the digestive tract with the dorsal caecum (Fig. 8). I have avoided making
use of the names pharynx and oesophagus, as there cannot be question here of any
sharp separation between the two, even though the proximal part is of a different
structure from the following portion. Thiele mentions a strong fold in which the
salivarv glands open out, and which separates pharynx from oesophagus (13, p. 288).
A similar condition does not exist here. An cesophagus, in the sense Thiele ascribes to
it, fails entirely.
The spacious intestine shows strongly folded walls, with the exception of the
dorsal wall, which remains smooth, except in the median line, where a small fold
occurs. The dorsal wall carries cilia ; the ventral and lateral walls show glandular
epithelium. Lateral caeca are met with as usual ; the transverse sections do not teach
much in regard to their relative situation. The rectum has strong cilia (Fig. 21, r).
The cloaca with its strongly folded wall opens to the exterior through a wide
longitudinal slit. Gills are entirely wanting. The wall of the cloaca is very thick and
consists of several layers of small transparent cells with small nuclei, between which
slender ciliated supporting cells occur with narrow long nuclei, which stain well
(Fig. 19). As it seems to me, the wall of the cloaca does not perform any respiratory
function, it being too thick. A secretory function must not be excluded. Oftentimes
in the cloaca and against its wall little feebly staining globules an- found, which I
believe to be formed by the wall-cells of the cloaca. Yet we must take into
consideration that the cloaca is surrounded by extremely loose connective tissue with
large spaces, filled with blood-corpuscles. This arrangement more especially obtains
in that part of the cloaca which is contiguous with the dorsal lip ; this part is entirely
surrounded by blood-corpuscles, ami. as it is only clothed by a single layer of epithelial
cells, a respiratory function may be ascribed to it. The lateral walls of the ventral
slit, by which the cloaca communicates with the exterior, are coated with ciliated
cylindrical epithelial cells (Fig. 19).
Properly speaking, the praecloacal organ does not open into the cloaca, but into
the ventral slit, as well as both cloacal spicules (Figs. 19, 20). The latter (Kloaken-
spicula of Thiele) are well developed, and consist of small bars of about 75 mm. length.
These bars are probably calcareous. Around the liars a strong envelope is found,
which is not dissolved in nitric acid, but Mains easily with carnialuni. Probably
G H. F. NIERSTRASZ.
this envelope is of cbitinous character (Fig. 24). The two extremities of the envelope
are tube-shaped (Fig. 24, B) ; it is open in the middle portion (Fig. 24, A), which may
have been caused by damage. Around the proper spicule a cellular envelope is found,
consisting of transparent epithelial cells with round nuclei, between which there are
long supporting cells with oblong nuclei (Fig. 24). Between the cellular envelope and
the spicule an almost homogeneous mass occurs, staining pale-red with carmalum, which
I believe to be a secretion of the cells of the envelope, though perfect certainty cannot
be obtained. In this ease the envelope might perform the duty of a poison gland, the
spicule having consequently the power of attack and defence. At the proximal
end of the 'spicules numerous small muscles are attached, which enable the animal
to move the spicules in various directions.
The nervous system does not furnish much worthy of note. The cerebral
ganglion is large (Fig. G, ctj) ; the latero-ventral ganglia are large also (Fig. 7, r/r),
whereas the latero-dorsal ones are very small (Fig. 6, or Dinomenia. The same holds good for Rhopalomenia itself.
This genus was created by Simroth on account of the presence of club-shaped
epidermal papillae (12, p. 229). It is with good reason that Thiele observes, that
the absence of a radula is more typical of. Rhopalomenia than the presence of the said
papillae (13, p. 272). In this sense the genus Rhopalomenia was enlarged by me,
some years ago, to contain new species (6, p. 22).
As to the geographical distribution, nothing confirmative is given. An-tn-
arc xliiiti'r! and //////- //*/*; both forms arc most certainly not closely rclaled
(14, p. 115).
Antarctic forms: gerlachei and discoveryi. Between these two forms there exist
great differences (dimensions, radula, dorsal spicules, buccal gland of _//.r/<7/,7.
structure of the proximal portion of the digestive tract). In dimensions. >/, rfni'/i, i
resembles sluiferi ; for the rest, according to Pelseneer (10, p. 34), these two forms
do not correspond in many respects.
Mediterranean forms : neapolitana, goryonophila, desiderata, riii// /'/, /7,
gerlachei and discoveryi, by the structure of the radula ; on the other hand, a relation
with thulensis is very well possible (radula, salivary glands, abdominal spicules),
though in the latter form cloacal spicules arc wanting.
Indo-pacific forms: webtri. /7\.
Of these forms /lawaiensis is insufficiently known ; only the nervous system has been
accurately studied by Heath (1). IIW>7 and l divide the genus into a number of smaller
c
10 H. F. NIEKSTRASZ.
ones, considering in the first place the exact structure of the radula and salivary
glands. I propose the following division :
FAMILY PRONEOMENIIDAE.
Index at least fi. Cuticle thick, with many layers of spicules, by far the greater part of which is
pointed and hollow. Epidermal papillae present. One or more dorse-terminal sense-organs. Radtila and
salivary glands present. Gills are wanting.
A. Radula multiserial. Salivary glands tube- or bag-shaped.
Proneomenia Hubrecht.
a. Radula with basal membrane. Salivary glands tube-shaped, uniting before opening
out into the digestive tract. Receptacula seminis strongly folded, glandular. The
spicules remain connected with the epidermis by multinuclear stems. Abdominal
spicules present. Index 9-14.
Proneomenia sluiteri Hubr. (4.)
(= lanrji Heuscher),
Barent's Sea, Spitzbergen.
70-160 fathoms.
b. Radula with basal membrane. Salivary glands tube-shaped, opening out separately.
Receptacula seminis vesicular or tube-shaped.
a. With abdominal 'and cloacal spicules. Receptacula seminis vesicular.
One dorso-terminal sense-organ. Index 22-50.
Proneomenia weberi Nierstr. (6, p. 2.)
East-Indian Archipelago, 22-1633 M.
Proneomenia longa Nierstr. (6, p. 7.)
Saleyer, East Indian Archipelago, 1158 M.
/3. Without abdominal, with cloacal spicules. Receptacula seminis vesicular.
Two dorso-terminal sense-organs. Index 10.
Proneomenia discoveryi Nierstr.
77 50' 30" S., 165 40' 5" E. 100 fathoms.
y. Without abdominal or cloaca! spicules. One dorso-terminal sense-organ.
Receptacula seminis vesicular or tube-shaped. Index 9-18 '5.
Proneomenia valdiviae Thiele (16.)
3 7' S., 40 45' 8" E. 748 M.
Proneomenia acuminata Wiren (18, p. 6s.)
West India. 300 fathoms.
Proneomenia hawaiensis Heath (1.)
(? insufficiently known.)
Hawaian Islands.
Proneomenia gerlachei Pels. (10, p. 16.)
70 S., 80 48' W. 500 M.
c. Radula without basal membrane. Salivary glands tube- or bag-shaped, opening out
separately. Receptacula seminis vesicular.
a. With abdominal and cloacal spicules. Salivary glands tube-shaped.
1-3 dorso-terminal sense-organs. Index 6.
Proneomenia vai/ans Kow. a. Mar. (5, p. 29.)
Mediterranean.
/3. With abdominal, without cloacal spicules. Salivary glands bag-shaped.
Receptacula seminis numerous, often lobate. One dorso-terminal sense-
organ. Index 11.
Proneomenia Ihulensis Thiele (14.)
so // is related
tn aglaopheniae. Jf. ////< shows numerous receptacula seminis; cloacal spicules,
however, fail. True abdominal spicules are absent, though around the cloacal opening
spicules occur of special form. Moreover, there only occur two tube-shaped salivary
glands; relation with Proneomenia tlmli'iisix and E/iinn'nin ilix, thoroughly known to us, probably is closely
related to ff/injiitfuiiii'iiiit im/icit. ( >nr knowledge of the stnu'ture of the internal
organs of Rhopalomenia w. ciliated bands. (See text.)
cd. cloacal duct.
cl. cloaca.
rx. cloacal-spicule.
!/ps. ganglion posterius superius with commissure.
i. intestine.
//(. lateral nerve.
n. distal nerves.
p. pericardium.
po. praecloacal organ.
r. rectum.
v. ventricle.
///. ventral nerve.
FIG. 21. Transverse section through a cloacal-spicule. (x 330.)
FIG. 25. Genital products in the pericardium. (See text.) ( x 480.)
FIG. 26. Blood-corpuscles. (See text.) (x 780.)
6.
i
INSECTA APTEBA.
By GEORGE H. CARPENTER, B.Sc. (Loxo.), M.R.I.A.,
Professor of Zoology in the Royal College of Science, Dublin.
(1 Plate.)
THE explorations of the National Antarctic Expedition have established the presence
of a wingless insect of exceptional interest, far south in the Continent of \ ictoria
Land. From Granite Harbour, 77 S. hit. and 162 E. long., on the south-trending
continental coast-line, almost opposite Ross Island, on which stand .Mounts Erelius
and Terror, and about 100 miles N.W. of the AVinter (Quarters of the ' Discovery,' a
jar-full of moss believed to contain Collembola was secured. Examination of this
material has resulted in the detection of half-a-dozen very imperfect specimens of a small
dark-blue spriugtail. Unfortunately, these insects are in a poor state of preservation ;
cither the spirit in which the moss was placed was too strong, or the insects had died
and shrivelled before the moss was collected. The result, however, is that no really
good example of the species can be obtained, and the following descriptions with the
accompanying figures have been made from various fragments. Under these
circumstances the descriptions are necessarily imperfect. It is hoped that the account
will not require correction by the discovery of further and more satisfactory specimens
bv some future expedition, but it will assuredly need supplementing. The remoteness
of the locality, and the difficulty with which the insects must have been obtained,
render it a duty to make the best possible use even of such unpromising material.
At tirst I spent many hours pulling the leaves of the moss _ plants asunder with
the help of a dissecting microscope, and this labour was rewarded by the discovery of
one or two shrivelled specimens, almost useless for study. Afterwards it was found
that the bases of the shoots afforded a better hunting-ground, and several more days'
work resulted in the unearthing of a few specimens in a less shrivelled state. From
these, when cleared in potash and mounted in glycerine jelly, it was possible to make
out various details of structure, and the systematic- position of the insect has bn-n
gradually traced. All the specimens seen arc referable to the same species.
Unfortunately, the general shape of the body can only be surmised, and the
segmentation of thorax and abdomen cannot be clearly distinguished in any specimen.
On the other hand, the delicate sense-organs on the feelers and head, and some details
of the jaws, can lie plainly seen in one or two examples. A large quantity of the
moss still remains unexamined, so that further light may yet be thro\\n on the species
G. E. CARPENTER.
by the collection already in hand ; but I think it unlikely that any specimens more
satisfactory than those here described and figured will be found in it.
This is the second species of Collembola known from the Antarctic Continent
of South Victoria Land. The first, described five years ago (Carpenter, 1902) was
brought home by the ' Southern Cross ' Expedition from Geikie Land, on the shores of
Robertson Bay near Cape Adare, some 380 miles north of Granite Harbour. That
species was an Isotoma, a member of the family Entomobryidae. The present insect
belongs to the family Poduridae, so that the two largest families of Collembola are both
represented on the Antarctic Continent. A summary of the distribution of the known
species of Antarctic and sub-Antarctic Collembola will be found in Wahlgren's recent
paper (1906) on the insects of this order collected by the Swedish Expedition. To the
species that he enumerates have now to be added the insect here described and the two
new species brought from the South Orkney Islands by the ' Scotia ' Expedition
(Carpenter, 1906). The last-named paper contains arguments, from the distribution of
Antarctic springtails, in favour of a former wide extension of the Antarctic Continent.
The present species, having no near allies, does not throw any additional light on such
geographical problems.
COLLEMBOLA.
FAMILY PODURIDAE.
GOMPHIO.CEPHALUS, gen. nov.*
Cuticle very finely granulate. Abdomen with two anal spines. Spring and
catch vestigial. Empodium of foot without appendix (" inferior claw "). Feeler
four-segmented ; sensory setae on the third segment ; a sub-apical, depressed sense-
organ, and a protrusible apical sense-organ on the fourth. On each side of head a
simple post-antennal organ surrounded by a single ovate or cordiform rim ; and a stout
sensory tooth-like spine behind the post-autenual organ. Mandible with molar
surface close to base. Maxillula with apical teeth and a feeble but distinct molar
area.
This genus will probably require a new sub-family (Gomphiocephalinae) to express
its peculiar affinities. On the whole it comes nearest to the Hypogastrurinae in the
recent classification of Borner (1906). But the dentiform sense-organ on the head, the
shortened mandibles, and the mandibuliform maxillulae appear to be absolutely
distinctive characters ; while the feeble granulation of the cuticle, and especially the
simple post-anteunal organ, show an approach to the Anurophorini, a tribe of the
Isotominae which belong to the Eutomobryidae. Indeed the genus Gomphiocephalus
* From yofifjjios, a back-tooth, and KetjtaX!]. I have to thank the Editor, Prof. F. .1. Bell, for kindly proposing
this name to replace my suggestion Odontocephalus, which is pre-occupied. The allusion is to the peculiar
dentiform sense-organ on the head (tig. 1,/J.
I.NSKCTA AI'THK\.
seems ,-i I'odurid, with affinities to (lie Entomobryidae, just ;is the Anurophorinae
\vhidi include that remarkable Antarctic genus Cryptopygus (Willem, I'.HfJ) -arc
I'jutomobryids, with affinities to the I'odtiridae. The presence of such ancient
connecting links on the Antarctic Continent and islands might reasonably have Keen
expected.
I have pleasure in naming the type-species of Gomphiocephalus after Mr.
T. V. Hodgson, of the ' Discovery.'
GOMPHIOCEPHALUS ILOHHSUNI.
Length, 1 nun. Colour dark blue-violet. Feelers rather shorter than head;
proportional length of segments 8 : 11 : 12 : 15. Foot with three long, slender
tenent hairs; daw simple without teeth; empodium vestigial. Anal spines nearly
as long as foot-claw, on short contiguous papillae.
Habitat. In moss (Bryum algens, Cardot *), Granite Harbour, South Victoria Land.
(Types in British Museum.)
The whole insect is of a very deep blue-violet colour. The cuticle is finely
rugose, and bears numerous bristles, many of which, especially on the terminal
abdominal segments, are elongate, but none are feathered. The general aspect of the
species and the form of the foot are suggestive of a \~i-n >/lla.
Feelers. The feelers (figs. 1, -2) have characteristic sense-oriians both on the
third and fourth segments. On the dorsal- aspect of the extremity of the third
segment is a group of four short, sharp spines arising from large circular depressions in
the cuticle, and surrounded by a sinuous ridge (fig. 1, '/) The two outer spines of
this organ are stronger than the two central. At the tip of the fourth .Moment is a
O O L
long tapering sensory bristle (tig. 1, It), and near it is a sub-apical sense-organ (fig. 1, ri. ]'. ."i.
I'
4 G. II. CARPENTER.
The labrum (tig. 8) is narrow and rounded in front, and bears a proximal row of four
and a distal row of three long bristles.
The jaws are in many respects highly remarkable. The mandibles (figs. 3,4) have
four strong teeth at the apex, and a very well-developed molar area, the teeth at its
distal region being especially strong. At the proximal end of the molar area are two
prominent recurved hooks directed dorsalwards. On the outer edge of the mandible is
a strong " shoulder," and just proximal to this a rounded " boss." The large basal
extension of the mandible so characteristic of the Collembola generally is entirely
wanting in the present genus. At first I could not believe this to be so, supposing
that the basal region had somehow got destroyed in the specimens that I was
examining. But on finding the mandibles in all the specimens in exactly the same
condition, I can only conclude that this insect shows a modification of the jaws unique
in the Collembola, As the muscles are usually attached to this missing basal part of
the mandible, fresh specimens of the present genus would be especially welcome
subjects for anatomical research.
The maxillulae (fig. 5) are also very remarkable on account of their likeness to the
mandibles. I have not succeeded in isolating these appendages, but one of the heads is
so transparent that their shape can be easily seen. At its apex the maxillula has three
delicate pointed teeth, while the inner edge of the base carries, not, as is usual, one row
of spines, but four rows of short denticles, forming a molar area analogous to that of
the mandible. I have not succeeded in seeing the tongue (hypopharynx) so that I
cannot state whether the relation of the maxillulae to that structure is of the usual
nature or not. In this genus they are more unmistakably a pair of jaws than in any
Collembolau that I have examined.
Each maxilla (fig. 6) has a " head " of the usual Collembolau form with a two-
toothed galea (//) and three internal lamellae (/) ; on account of their delicate texture
these latter structures are somewhat shrivelled, and the shape given to them in the
figure may be inaccurate. The base of the maxilla (fig. 6, st) can only be traced a
short distance into the head capsule, but I cannot satisfy myself that the cardo,
with its usual associated sclerites, is absent. There is a blunt palp (fig. 6, ]>} carrying
three or four bristles at its tip.
The second maxillae combine, as is usual in the Collembola, to form a transverse
liiliiiiin (fig. 7) with alternating narrower and broader lobes, carrying long bristles.
Legs. The legs (fig. 9) are of the usual Collembolan form. The tibio-tarsus
cariies a simple pointed claw without trace of tunica, and a small rounded empodiuni
without appendix. There are three elongate teneut hairs, without terminal club,
springing from large circular depressions of the cuticle.
Ventral Tube. The ventral tube (fig. 10) on the first abdominal segment is
abrupt on its front and rounded on its hinder face, beset with numerous bristles like
those of the cuticle generally. It appears to be in the undeveloped state characteristic
of the section l<> which this genus belongs.
INSKCTA AI'TKRA. 5
Spring and Catch. .Most of lh<> specimens examined show no trace ot' these
structures, hut in two the appearance drawn in Jig. 11 can he plainly seen. Tin- s|nin-
(fig. I 1, .) is clearly in a vestigial condition, its niannliriiini hcinu; merged in the ventral
region of the fourth ahdoniiual segment, while the combined dens and miicio terminate
in a blunt, rounded knoh.
The catch (retinaculum) is represented by a pair of .small rugose prominences
(fig. 11 /) on the third abdominal segment.
Anal Spines. The anal spines are nearly as long as the foot-daw, slightly and
evenly curved (figs. 11, 12). They are situated on short, contiguous papillae. In all
the specimens examined the spines are slightly asymmetrical (fig. 12). bur this is
probably due to the position of one having been slightly shifted.
linn',
linn;.
I'.ioi;.
I'.ni2.
REFERENCES.
('. BORXER. Das System der Collembolen ncbst Beschreibong nener Collembolen dee Hamburger
Datnrb.iBtoriflcb.en Museums. 2 IViheft zum Jnln-h. ., xxvi., I'.ini;, pp. 1 7:;- Is:;, pis. 1-2.
E. WAIIIJIUKX. Antarktische und Bubantarktiflche Collembollen jrcsamnielt von dcT sclnvc(li>rln-ii
Siidpolarexpedition. Wix*e/ti'/t. Enjclm. ilrr s'/vv/yy/y/^/-c.;. ."). Kight maxillula. showing teeth and molar area.
Fid. (!. Left maxilla ; .v. siipes : //, galea : /, lamellae ; //. palp.
Fir,. 7. Second pair of maxillae forming labium.
Fiii. s. Labnuii.
FlG. 0. Right hind leg, showing claw, vestigial empndiuni, and leiieni hairs mi foot.
Fin. lo.. -Ventral tube on first aMi'initial seiiiin/nt, seen from right side.
Fn:. 11. Part of bind abdominal segments, showing anal spines, and vestigial spring (.) and raidi
lateral view.
Fn:. 12. Tip of abdomen, witli papilhe and anal spines ; postern-ventral view.
All figures are magnified Coo diainet'T>.
I. 2
.
Antarctic (Discover.
ii'" jsoni.
north, >c.
C E U S T A C E A.
VII.-SCHIZOPODA.
By AV. M. TATTERSALL, M.Sc.
(8 Plates.)
TIIK colleeiion of ' Discovery ' Schixopoda is a large one from the point of vir\v of the
ninnber of specimens it contains, considerably over ten thousand, including larva-,
having been collected. By 1'ar the majority of the-e -peciinens. nearly ten thousand.
however, belong to one species, and the total number of species reaches thirteen. The
collection is in a generally excellent state of preservation, though many specimens have
undoubtedlv suffered from being frozen after coming out of the water and from the
subsequent thawing before preservation.
In the preliminary notice of this collection (Holt and Tattcrsall, 1906(1)*) ten
species were noted, of which five were absolutely new, two were only known under
manuscript names given to them by Dr. Hansen from the collections of the ' Brlgira
Antarctic Expedition, while the remaining three were already described forms.
Since the publication of that notice further material has come to hand, collected
on the homeward journey of the 'Discovery.' It contained two species of En/ >li. Sars), but which Dr. Hansen has kindly informed me is distinct,
and will be described by him in a forthcoming paper. A re-examination of the two
specimens which in the preliminary note were referred to .l///x/.-- maxima,
Ihmseii (MS.), has shown that they represent in reality two very closely allied
species, the second of which will also be described by Dr. llansen in a future work.
Previous to the recent activity in South Polar exploration only three expeditious
to the Antarctic had brought back zoological material from which Schizopoda were
recorded. Dana (185:2) in his great work <>n Crustacea records two species from
Antarctic waters (/>., south of lat. '',() S. ) vix., Eitj>/tv //-.///./ and /-jn'o/'ln tuitr s .
W. M. TATTERSALL.
JVIr. Hodgson (1902) described two species as new, Euphausia glacialis and E. australis,
collected by the ' Southern ( 'ross ' South Polar Expedition. Both species arc
synonymous with E. xuperbti, Dana, so that previous to 1903 only four Antarctic,
species of Schizopoda were known, viz., Eucopia aust rails, Dana, Euphausia superba,
1 )ann, Thysanoessa macrura, G. 0. Sars, and Pseudomma x/trxi, Will.-Suhm.
Of the recent expeditions to the South Pole, which number seven, the results of
the Schizopoda of the French Antarctic Expedition have been published in full, and of
the ' Valdivia ' Expedition in part only. M. Coutiere (1906) notes from the French
Antarctic collections, Enphaasia superba, Dana, E. similis, G. 0. Sars, Thysanoessa
macrura, G. 0. Sars, and Antarctomysis maxima (Hansen, MS.), a species also recorded
in the preliminary note on the present collection.
Preliminary descriptions have been published of two Antarctic Mysidse collected
by the. ' Valdivia ' (Illig, 190G), Dactylerythrops arcuala and Echinomysis chum. The
first of these is synonymous with a species Dactylamblyops hodr/soni, described below.
This completes the bibliography as regards purely Antarctic Schizopoda, though a
number of sub-Antarctic species are known.
Ten of the thirteen species collected by the ' Discovery ' were taken in Antarctic
waters, and when, as we have seen above, the total previously recorded species number
seven, it will be recognised that the present collection has added considerably to our
knowledge of South Polar species of this group.
The most abundant species in the collection is a small Euphausia, E. crystal-
lorophias, H. and T., which evidently has its head-quarters under the ice, since all the
specimens were collected from ice-holes at Winter Quarters, and none were met with in
the open sea. On the other hand, the dominant species of the collections from open
Antarctic waters are Eupliauxia xuperba, Dana, and Thysanoessa macrura, G. 0. Sars.
The abundant material of the first-named species has afforded opportunity for
some observations on the sexual characters and life history of the species, with the'
result that four other species, hitherto regarded as distinct from E. superba, must now
be allocated to its synonymy, having been founded either on characters which are
sexual and not specific," or else from immature specimens.
The scarcity of fully grown males of E. superba seems worthy of note. This fact
seems to be established by the results of the various collections of which we' have
knowledge, since, so far as I am aware, the only one recorded is Sars' ' Challenger '
specimen. M. Coutiere (1906), it is true, notes that male specimens were more
numerous than females in the collection he examined, but it is equally clear that none
were fully grown, since the largest measured only 45 mm., and was in the stage pre-
vious to the last moult into completely grown specimens. There are no fully-grown
males in the ' Discovery ' collections, but to judge from the development of the copulatory
apparatus on the first pleopods, many of them must be sexually mature at any rate.
Thysanoessa macrura, G. 0. Sars, too, seems never to have been previously met
with in full-grown condition, and but very few of the ' Discovery ' specimens can be
8CHIZOPODA.
said to have reached that state. Still, they afford material for a re-description and
figures of the species in the adult state, with some notes on the growth change-.
The Mvsidacea consist of eleven specimens, referable to six species. The most
interesting of these species is Hansenomysis inifnn 4 fii-,i, an Antarctic representative of
a genus hitherto known from hut three specimens from Arctic and Itoival waters.
The chief interest of the collection lies in the evidence which it may afford as to
the similarity or dissimilarity of the fauna at the two poles. There are no species
common to the fauna of both polar regions in the collection : but, on the other hand,
all the genera save one, Antarctomysis, are represented in northern waters bv species
which are quite distinct from their southern allies.
Exploration of the bottom fauna of the, deep waters of the nlolic. especially in
tropical and sub-tropical regions, is as yet only in its infancy, and it is therefore
extremely probable that what are now known to be bipolar genera and species will in
future be found to be cosmopolitan in their distribution. The8chizopoda were long
thought to have in Lophogaster fj/jiicux a stock instance of a bipolar form, but tin-
gaps in its distribution have been almost completely filled up as a result of recent deep-
sea work, and, with the exception of the tropical Atlantic, its range is known to be
complete from Norway to the < 'ape.
Two bipolar species of Mysidse are known, however Boreomysis scyphops, C,. < ).
Sars, from Arctic waters, and Lat. 50 S. , near the Crozet Islands, and Ani/>/>/t>jix
crozetti, from the seas of Ureeuland and Ian Mayeu in the north and the Crozct Islands
in the south. They are not known from the intermediate waters.
Of the genera of Antarctic Schixopoda, EH-II/KIHX/H, Tlii/xini<>; : xel\-
related in structure to the northern species Michtheimysis mixta (Lilljeboig), a species
inhabiting chiefly the colder waters of the northern hemisphere. The two genera are
separated only in the characters of the male pleopods. which arc more primitive in the
Antarctic form, and neither genus is likelv to be found to have a distribution which
extends very far from the poles they frequent.
In the preparation of this report 1 have received miicli valuable help from manv
sources. The authorities of the British .Museum kindly allowed me to examine and
dissect two specimens from the Challenger' collections in their charge. To 1 'r.
(',-ilman, of the British Museum, I have been much indebted for information on manv
points connected with the Challenger' material, and he has. also, at my reqiu-.-t.
furnished me with drawings of various species. Mr. Iv \V. 1,. HO|I kindly examined
4 W. M. TATTERSALL.
the British Museum material, and provided me with valuable notes as a result, which
I have freely used in this report. The Eev. T. R. R. Stebbing, F.R.S., placed two of
the type specimens of Euphausia vu/l< ////'/// at my disposal, with full permission to
dissect them if found to tie necessary, while Prof. D'Arcy W. Thompson, C.B.,
allowed me to examine and retain for some time a small collection of Antarctic
Schizopoda from the collections of the Museum at University College, Dundee. I am
specially indebted to Dr. Han sen, of Copenhagen, for the most generous and valuable
assistance. Not only did he kindly confirm or modify my determinations of the
more subtle species of Euphausia, but he placed at my disposal his notes and drawings
of the male copulatory apparatus of the species of this genus, of which he has made
a special study. Without his help I should have failed to recognise that the two
specimens of Antarctomysis belonged to two distinct species, while I should have
recorded as E. liict'tix the new Antarctic species which Dr. Haiisen will himself describe.
For all this assistance, I wish to express my best thanks.
ORDER EUPHAUSIACEA.
FAMILY EUPHAUSIID.E.
SUB-FAMILY EUPHAUSIN.E, Holt and Tattersall.
GENUS EUPHAUSIA, Dana.
EUPHAUSIA SUPERBA.
(Plate L, Figs. 1-12.)
Kii.li/iin/xi/i xiijiir/nf, Dana, is.'c'.
K/ijifiii.ii.xia st/perlni, G. 0. Sars, INS:-; and iss.V
Ei/pliaiiaia nuirrfiyi, G-. 0. Sars, 1883 and 18s:..
Kiipl/anxifi an/af/i/ni, G. 0. Sars, 18X3 and INN:>.
Enphaiinfa t/lan'ulix, Hodgson, 1902.
Kiipliai/^in ni/sf ralis, Hodgson, I'.lni'.
Eii]}lian.$i/i Kiiprla, Holt and Tattersall. I'.ioi; (1).
EnpIi/tnxi/1 xii/>/'rl/ri, Coutim-, I'.iftf!.
L<>c<>n carcinophaga, 3. 1. 02, eleven specimens, 43-47 mm.
Lat. 72' 5' S., long. 172 23' E. ? 10. 1. 02, eleven specimens, 15-47 mm.
Lat. 72 10' 33" S., long. 172 26' 2" E., 11. 1. 02, ninety specimens, 35-48 mm.
From River Koettlitx, 2. 1. 0: : J, sixty-nine specimens, 13-27 mm.
Lat. 70' 29' 27" S., long. 168 51' 46" E., 26. 2. 04. live specimens, 45-47 mm.
None were taken at Winter (Quarters from the holes dug in the ice.
The synonymy given here, whereby five suppo.-ed distinct species of En)>lm*i) Small blunt .>///. on t/i>- outer h- I'll tli,' outer //*?///// <>f tin- mil' nmil >/- <>/ t/m j><'iinltin> the three shown by Sar> being frequently noticed. In all
VOL. IV. K
6 AV. M. TATTERSALL.
other respects the 'Discovery' material arid the 'Challenger' types are in perfect
agreement, and the facts noted above establish the identity of the females here
referred to E. superba, with the species described by Sars as E. murrayi. It now remains
to show that the differences between E. murrayi and E. superba are only sexual.
Male. Under the name E. superba Sars has described and figured this sex
adequately. The only point in which his description is deficient is the structure and
armature of the telsou. He figures no dorsal spiuules on the telson, and both describes
and figures the apex as slightly produced and obtusely pointed. Examination of Sars'
type shows that the apex of the telson is clearly broken, so that Sars' figure is in this
respect entirely imaginary. In the present material the apex of the telson is much
produced and acutely pointed, and the number of dorsal spinules is usually three pairs,
but may be four or two, placed as in Sars' figure of the telson of E. murrayi. One pair
of spinules still remains in Sars' type of E. superba, but the others had probably been
broken off (or obsolete ?).
The most conspicuous difference between E. superba and E. murrayi, as described
by Sars, is the presence in the latter and absence in the former of a lateral denticle on
the carapace. But both Sars' E. murrayi were females, and his single specimen of
E. superba a male, In all the females in the present collection, the largest of which is
47 mm. in length, the spine on the lateral margin is large and prominent, and even in a
female, 50 mm. in length, in the collection from University College, Dundee, the spine
is equally well-developed. I have figured the spine of the latter specimen on PI. I.,
Fig. 10. In male specimens, on the other hand, only those which are less than 42 mm.
in length have the spine well-developed (cf. PI. I., Fig. 12, taken from a male, 39 mm.
in length). In males above 42 mm. up to 47 mm. in length the lateral spine on the
carapace is nearly obsolete and persists only as a blunt protuberance (cf. PL I,
Fig. 11, taken from a specimen 45 mm. long, and also Coutiere (1906), PI. II., Fig. 22,
taken from a male of the same size). The ' Discovery ' collection contains no male
specimens exceeding 47 mm. in length, but the ' Challenger ' type measures 48 mm.
It is well preserved and shows no trace of the lateral spine at all. Obviously, then, the
absence of a spine is a sexual character confined to absolutely full-grown males only.
The remaining differences between E. superba and E. murrayi given by Sars are
as follows :
(l) E. superba has the antenuules considerably more robust than in E. murrayi
and the lobe from the second joint almost -obsolete. This difference is, I think, a
purely sexual one, affording a parallel instance to that seen in the northern species,
Nyctiphanes couch). PI. I., Figs. 1 and ~2 are taken respectively from male and
female specimens of the same size, viz.. 45 mm., and from the same bottle. They
indicate, clearly, the difference in relative stoutness in the two sexes, and that of the
male shows the lobe from the second antenuular joint in an intermediate stage of
reduction between that of the female and that shown by Sars in his figure of the male
E. xiiperliti, 48 mm. in length.
SCHIZOPODA. 7
(J) In A'. xtijii-rfxi the rostrum is shorter and blunter tlian in A'. ninrriii/i. and lias
the margins less deeply concave. This, again, is dearly shown to lie a sexual
difference in PI. I., Figs. 1 and 2. The rostrum of the male figured (Fig. 2) is shorter
than that of the female, but is still rather more acute than in Bars' figure of K. .v//y/et. \veeu the two sexes is brought out by the figures here
given, namely, the reduction in the male of the spine 011 the outer distal corner of the
basal joint of the auteuuular peduncle. It is not visible in dorsal view, being hidden
by the slightly projecting anterior margin of the joint, but it still persists as a small
blunt protuberance. In the female, on the contrary, it is well-developed, distinctly
visible in dorsal view, and acutely pointed throughout life.
A fourth distinction shown in the figures, the absence in the female of the curved
setse on the dorsal surface of the basal joint of the antennules, is due to the accident
that in the female from which the figure was taken, these setse had become broken off.
They are, in reality, present, and equally developed in both sexes.
The above detailed description proves, I think, dearly, that E. >/////// and
E. murrayi are the adult male and female, respectively, of one species which must I/ear
the name E. superba Dana.
I also give (Plate I., Figs. 5-9), figures of the mouth organs and endopods of the
first two thoracic limbs, to show two characters in which E. >////./// differs from all
other Euphausia yet described. The first of these points is the narrow and elongate
form of the terminal joint of the mandibular palp, with its peculiar armature of four or
five terminal strong plumose setse. In all the other species of the genus (with the
exception of E. untarctica, Sars, and E. i/lncial!*, Hodgson), the terminal joint of the
mandibular palp is much shorter and stouter. In the two exceptions just mentioned
the mandibular palp is figured by Sars and Hodgson respectively, almost exactly as
here given for E. superba. This fact first suggested to me that these (wo species were
only developmental stages of E. superba, a suggestion fully borne out by the evidence
derived from a study of the present collection. The second distinctive character of the
appendages is found in the great length of the setaj arming the joints of the thoracic
limbs. They are very much longer than in any other species of the genus, and with
the character of the mandibular palp serve for recognition of E. supei/m \\\ aiiv sta^v
in its development.
E/i/i/iit/ixiit Kiij><-rlni is the giant of the genus, and the only one of Dana's original
four species which is now retained by Hanseu (1905 (2)), the other three having been
cancelled by that author as unrecognisable.
Some Notes n ///* development <>f A'.
These notes were made chiefly with a view to confirming the suspicion, aroused
by the similarity in mouth organs, that E/i/>/i///>.///. ami must
therefore lie regarded as synonymous with that species.
It should be mentioned that Hodgson's types of E. australis differ in no \vav from
E. sujn'rfni (females), except in being considerably damaged.
EUPHAUSIA CRYSTALLOROPHIAS.
(Plate II., Figs. 1-10 ; Plate IV., Fig. 10.)
i-fi/vtal/uni/i/iiitf:, Holland T;itti/rs:ill, I'.MMi (1).
of O'y/^/v.x :
Winter (Quarters.
26. 1. 02-8. 3. 02, 1 specimen, 24 mm.
,, ,, 216 specimens, larval.
No. 3 Hole, 52 specimens, larval to 25 mm.
No. 4 Hole, 4572 specimens, larval to 32 mm.
No. G Hole, 13 specimens, larval to 25 mm.
No. 8 Hole, 4642 specimens, larval to 32 nun.
No. 13 Hole, 50 specimens, larval.
No specimens were captured either on the outward or homeward journey.
Form, moderately robust.
( 'it/'uj'iii-t' (Plate II., Figs. I and 2), with a prominent, rather long and acute
spine on its lateral margins, a little anterior to the middle, and just above the
insertion of the second thoracic limb ; antero-lateral angles terminating in an
acute spine ; anterior margins inflated above the eyestalks and produced into a long
acute rostrum extending to the visual part of the eye and about half-way along the
basal joint of the anteuuular peduncle; there is a faint gastro-hepatic groove and
a distinct keel runs forward medio-dorsally from the latter into the rostrum.
Pli'nn (Plate II., Fig. 1) without ridges or dorsal spines ; none of the epimeral
plates much produced ; sixth segment about one and a half times as long as the
fifth ; preanal spine well developed and usually simple, but in large examples bifid.
A//'.y (Plate II., Fig. 1) globose and rather large ; greatest diameter of the
cornea exceeding half the length of the last pleon segment ; pigment black.
Aittt'iunilai- /ifi/in/i-li- (Plate II., Fig. 2), with the basal joint as long as the second
and third joints combined and much wider; no lobe or lappet; a row of about
twelve long curved plumose setae set cu a ridge on the distal pan of its length;
a short stout spine on the outer distal corner, which is more or less concealed by
the numerous setse which arm the outer half of the anterior margin and the Ji-lal
10
W. M. TATTERSALL.
lialf of the exterior margin; a bunch of coupling seta? on the inner distal corner;
second joint slightly longer than the third and without a lobe, its anterior margin
a little oblique.
Antennal peduncle about equal in length to the basal two joints of the antennular
peduncle, the third joint only very slightly shorter than the second.
Antennal scale reaching the centre of the third joint of the antennular peduncle,
about three times as long as broad, outer margin entire and terminating in a spine,
apex broadly rounded ; spine on the outer corner of the basal joint long and slender,
extending one-third of the way along the scale, plumose at least on the
proximal part.
The mouth parts (Plate II., Figs. 3, 4, 5) are figured for comparison with those
of other species. They do not appear to present any striking peculiarities.
Firxt thoracic limb (Plate II., Fig. 6), has the penultimate joint of the endopod
longer than either the preceding or ultimate joints ; the latter has the lower margin
armed with a row of short fine setae in addition to the longer ones at the apex.
Second thoracic limb (Plate II., Fig. 7), with the terminal joint armed with a row of
three (sometimes four) short, rather stout and curved spines on the inner face.
The remaining thoracic limbs have the penultimate joint in all cases longer than
the ultimate and slightly longer than the antepenultimate. The following table
gives the lengths of the joints of the first six thoracic limbs in millimetres and the
total length of the limbs from a specimen 27 mm. long.
Thoracic
limb.
Lengths of the joints in min
Total length
of linih in mm.
1234
5 G
1
50
I'll
I -50
77
1-00
66
5 :, (.
2
50
1-22
1-77
I'll
1-22
50
6'32
3
66
1-50
1-88
1-22
1-28
72
7 21;
4
66
1 83
2-05
1-28
1-83
83
7'98
5
61
2-00
2-11
1-00
1 ' 05
66
7-43
<;
5;-,
2 '(HI 2' Illl
72
S3
44
6-54
First pleopod of the male (Plate IV., Fig. 10) with both movable processes on the
inner plate of the eudopod shorter than the plate itself; distal process feebly curved,
bifid at the tip ; proximal process expanded at the tip into two lobes not in the
same plane, the outer lobe the larger, and wider than long, the inner lobe but little
expanded ; uncinus of the inner plate of the endopod without secondary spiuule.
Telson about one and a half times as long as the last segment of the pleon ;
apex acutely pointed ; sub-apical spines extending for half their length beyond the
apex of the telsou and hearing a few minute spinules on their inner margins ; dorsal
denticles usually in two pairs, the first aliout half-way towards, tin- second at the
base of, the sub-apical spines.
Uropixh reaching to the level of the insertion of the sub-apical spines, (lie inner
very slightly longer than the inner, with a prominent denticle at its outer extremity.
Length of the largest adult specimens of both sexes, 32 mm.
Eupliiiuxin crystollorophios approaches most nearly among the species uf the
genus to E. 'x'nniti*, G. 0. Sars, but differs (1) in the different shape of the rostral
projection, (2) in the shape of the epimeral plates of the fourth and fifth se^mem.-
of the pleou, (3) in the absence of antenuular lobes and lappets.
From E. spti-inli'iift. G. < >. Sars (E. Inci'iis, Hausen) the present species is dis-
tinguished by the greater length of the rostrum and by the absence of antenuular
lobes and leaflets, the types of E. splendens, G. 0. Sars, being possessed of a small
but distinct antenuular lobe. E. crystallorophios is an enormously abundant species
under the ice, some ten thousand specimens having been taken. None, however,
were met with in any other locality except Winter Quarters.
Larva; of E. CRYSTALLOROPHIAS.
The collection contains individuals in all stages of development from the
M<-liiiiniil>rms to the adult condition.
The Calyptopi* larvze (Plate II., Fig. 8) first appear at the beginning of January
and continue in the tow-nettings till nearly the end of February. The 1 1 of the
carapace is very obtusely pointed in front and has the margins quite smooth. There
is no posterior median spine on the carapace, while the telson has the apical margin
lightly emarginate. I can see the beginning of the lateral spine of the carapace at
this stage. The largest Calyptopis larva measures .".<) mm.
The Furi-Hiii stages (Plate II., Km. '.0 first appear during the last week of Fehruarv
and are abundant all through March. Thev cease after the (\\-^\ week in April.
The emaruinatioii of the apex of the telson is most marked during this stage and
serves readily to connect it with the early CW/// "'"/"''' larva-. The .-pine on the
lateral margin of the carapace is now well developed. The size of the Furciim larva-
Is from 4 - 5 mm. to 8 mm.
The Cyrtopin larva- (Plate II.. Kig. LO) first occur about the last week in March,
and late post-larval stage- are still to be had at the beginning of August. The size of
this stage is from 8 to II mm. The rostral projection is now an acutely pointed
triangular plate, but the sides are still bin little concave. The final shape of the
rostrum is not assumed till the animal is in all other respects like the adult.
At a size of 11 mm. the telson assumes its adult shape, but the pair of spines
immediately anterior to the sub-apical spines are -till lon^ ami plumose. Tliev
linallv become reduced to adult size when a length of I:; mm. i> readied. At this
12 W. M. TATTEKSALL.
latter size the species has all the adult characters, except perhaps the rostrum, which
has the margins hardly as concave as fully adult specimens. Examples of 13 mm.
in length are to be met with in January, and so were presumedly larv;u of the
preceding season, from which it would appear that the species takes at least one
year, and very probably longer, to reach the final adult size of 32 mm.
EUPHAUSIA TRIACANTHA.
(Plate IV., Figs. 1-3.)
!:'n.jiJiausia triacantha. Holt and Tattersall, 100(5 (1).
Locality of capture -.Lai. 66 52' 09" S., long. 178 08' 15" E., 2030 fathoms;
one specimen, immature male, 23 mm.
Carapace (Plate IV., Fig. 1), with a single lateral denticle posterior to the centre
of the lower margin of the carapace ; antero-lateral margins somewhat inflated over
the eyestalks, and then produced into a long and very acute rostrum, which extends
beyond the eyes and almost to the distal end of the basal joint of the antennular
peduncle ; a faint keel is present on the carapace behind the rostrum.
Picon (Plate IV., Fig. l) with the posterior dorsal margin of the terga
of the third, fourth, and fifth segments produced into rather long, slender, very acute
and slightly curved median spines ; sixth segment rather long, nearly twice as long-
as the fifth segment without the spine.
Eyes somewhat damaged in the single specimen, but apparently rather small,
pyriform in shape.
Antennular peduncle (Plate IV., Fig. 2) bearing on the inner distal corner of
the basal joint a well-developed bifid leaflet, the lappets of the leaflet of about equal
size ; outer corner of the basal joint rounded and adorned with numerous rather long
plumose setae ; a row of six curved plumose setae on the dorsal surface of the basal
joint ; second joint with a simple acutely spiuiform lappet arising from the median
anterior margin : third joint slightly narrower and shorter than the second.
Antennal peduncle shorter than the scale, the third joint a little shorter than
the second.
Antennal scale reaching very slightly beyond the distal extremity of the second
joint of antenuular peduncle, broadly oval in shape, about three times as long as
broad, apex broadly and obtusely rounded, spine at the distal end of the outer margin
small but distinct ; spine on the outer distal corner of the basal joint long, slender
and smooth.
First pleopods of the male (Plate IV., Fig. 3) obviously not fully metamorphosed,
since both the proximal and distal movable processes on the endopod are small
and simple, and the uncinus on the middle lobe is without a secondary spinule.
Telson with the portion between and posterior to the sub-apical spines acutely
produced and smooth; sub-apical spines extending beyond the apex of the telsun.
SCHIZOPODA. 13
smooth ; dorsal denticles in two pairs, the first situated at about two-thirds of the
distance from the base of the telson to the insertion of the sub-apical spines, second
pair just above the spines.
I 'nijiiiils sub-equal in length, rather slender, extending to the level of the
insertion of the sub-apical spines of the telson.
Preanal spine small and simple.
A fuller description of this species is not possible, since the single specimen is
in bad condition and dissection was not desirable.
The species belongs to that group of the genus with a posterior median dorsal
spine on the third segment of the pleon, and is distinguished from the other members
of the group by having an equally developed spine on the fourth and fifth segments
of the pleon in addition. It presents no very near kinship with any described
species of the genus, and from the depth at which it was captured is probably a deep-
water form.
E U P H A U S I A V A L L E X T I N I.
(Plate IV., Figs. 4-6.)
Eitphavsia splendens (pars), G. 0. Sars, 1885.
En/ihai/tnxitt ./(/fWc//.v by Sars. They were from the second of the localities
given by Sars on p. 82 of his " ' Challenger ' Report," viz., " October 21, 1875, South
Pacific." It became at once apparent on examination that one of these specimens
did not agree with Sars' description, since the antennule was furnished with a large
evenly rounded lappet on the basal joint, very conspicuous in lateral view.* Further
examination showed that it probably, indeed almost certainly, belongs to the present
species. It is true that 1 could not see the spine on the third pleon segment, but
the specimen is in very poor condition, and if, as I suspect to be the case, the
spine has been broken off, the scar would be difficult to detect.
I give (Plate IV., Fig. 5) an outline sketch of the rostrum and the basal joint
* Examination of the ' Challenger ' types of E. splcndens shows that this species possesses a small
antonnular lobe, but it is nothing like so well developed and conspicuous as in F.. fulli-ntini (are Hansen
(1900 (2) ), Holt and Tattersall U906 ^ an a below, p. 14.)
VOL. IV. [,'
14 AV. M. TATTERSALL.
of the antennule of the 'Challenger' specimen and (Plate IV., Fig. 4) a sketch of
the antennule of one of the ' Discovery ' examples for comparison with those given
by Stebbing (1900). This shows clearly, in my opinion, that all three specimens
belong to one species, and that the absence of the spine on the third pleou segment
of the ' Challenger ' example is the result of accident or possibly an abnormality.
The peculiar shape of the antennular lobe is practically the same in the ' Discovery '
and ' Challenger ' individuals, and only differs from Stebbing's types in degree, a
result of more complete growth.
The rostrum of E. vallentini is very like that of E. splendens, G. 0. Sars, but is
slightly longer, and the angle formed by its margins a little more acute. Sars may
have been misled by the resemblance between the rostra of the two species, which
caused him to overlook the marked differences which exist in the antennulfe. Dr.
Hansen has seen the ' Challenger ' specimen referred to, and agrees with my
interpretation of its specific identity.
One of the 'Discovery' E. vallentini is a male, but unfortunately the copulatory
apparatus on the first pleopods is considerably damaged, so that I am obliged to refer
to the ' Challenger ' example, which is likewise an adult male, for a description and
figure of this apparatus (Plate IV., Fig. 6). The figure represents the inner lobe of
the endopodite of the first pleopod of the male. This inner lobe bears internally two
movable processes, the inner and more distal of which is feebly curved, slightly over-
reaching the inner lobe and bifid at the tip. The external and more proximal of the
two processes has the distal extremity greatly expanded, the expansion very much
broader than long, oblique, and divided into two lobes, the more distal of which is the
larger. On the under side of the expansion of the proximal process as viewed in the
figure there is a small spine-like process. The inner lobe of the endopodite itself bears
a strongly curved uucinus with a small secondary spine near the tip.
Distribution. Southern Pacific, between New Zealand and Chili (' Challenger ') ;
Falkland Islands (Stebbing].
EUPHAUSIA, sp.
Locality f capture. Lat. 57 25' 30" S., long. 151 43' E., nineteen specimens,
10-18 mm.
On first looking over these specimens I identified them with Euphausia splemlens,
G. 0. Sars (1885), a species which Hansen (1905 (2)) considers to be different from
E. splendens, Dana, and which he has re-named E. lucens. Hansen, in the same
paper, notes that E. splendens, G. 0. Sars, has the first joint of the anteunular
peduncle without a leaflet, but distally produced above. Holt and Tattersall (1906
(1)) have confirmed this statement by an examination of Sars' type specimens of E.
splendent, in which they found that in the female type the lobe is quite conspicuous
both in lateral and dorsal view ; while in the male type, which is considerably smaller
than the female, the lobe is less developed, but still easily seen in lateral view. Sars
SCHIZOPODA.
15
was therefore in error when he .described the antennular peduncle of his E, splendens
as " more particularly distinguished by the total absence of any dorsal leaflet or lobe."
Moreover, it is apparent from what has already been written above in dealing with
Euphausia vallentini that Sars confused at least two distinct species under the name
E. splendens. It was subsequent to the publication of the preliminary notice of the
' Discovery ' collection that the present specimens came to hand. I therefore appealed
to Dr. Caiman for further information with regard to the 'Challenger' .species, and he
very kindly sent me a sketch of the dorsal aspect of the anterior end <>f both types.
From these sketches and Sars' description in the ' Challenger' report I identified the
'Discovery' specimens as Euphausia splendens, G. 0. Sars = E. lucens, II. J. Ilansen.
Wishing, however, to have confirmation of my identification, I submitted the
specimens to Dr. Hansen, who at first was inclined to agree with me that they
belonged to E. xj>I< //< //*, G. 0. Sars. I may, perhaps, lie allowed to ijuote Dr.
Ilansen's remarks. They read as follows: " E. lucens (splendens). I have specimens
from the southern Atlantic and the southern Pacific, and in all these the leaflet from
first antennular joint is easily seen, triangular, but not acuminate, with the end often
a little obtuse. In the material from the Swedish Antarctic expedition I have a large
number of specimens which differ only from the Copenhagen specimens in the feature
that the antennular leaflet is extremely small (visible as a very small triangular
distally obtuse plate when seen from in front . . . . )or rudimentary, but I find it
necessary to consider this difference only as a variation " (Hansen, in litt.}. Then,
after some remarks in which he noted that my specimens agree with the latter
condition, he concludes by saying that he considers them to belong to the more
Antarctic variety of E. lucens. In a later communication Dr. Hansen kindly informed
me that, after an elaborate study of the copulatory organs on the first pleopods of the
males of the genus Euphausia, he had found that these two varieties were readily
distinguishable in the characters of the male pleopods, and that he proposed to
consider them as two species. At the same time he was good enough to send me
sketches of the first pleopods of both species for comparison with my own specimens.
The largest 'Discovery' specimen is a male 18 mm. in length, and as far as I can
judge, it appears to be quite adult. The copulatory apparatus on the first pleopods
agrees exactly with the sketch which Dr. Hansen sent me of the same apparatus in his
Antarctic form. It would therefore appear that these specimens belong to Ilansen's
new Antarctic species. I have not attempted to give a detailed description with
figures of this form, since it is quite evident that an accurate diagnosis can only be
drawn up from a close study of this species and the true E. />/<>//* side by side, and
a careful comparison, character by character. There are no specimens of the true
E. lucens in the ' Discovery ' collection, so I leave the descriptions of the two species
to Dr. Hansen, who has abundant material for the purpose.
I may mention here that some specimens of an 7v >;> .-/'" (labelled E. xpli'ml, us,
G. 0. Sars) in the small collection of Antarctic Schizopods kindly lent me by Prof.
^1 F 2
16 W. M. TATTERSALL.
D'Arcy W. Thompson, from the collections of the University College, Dundee, appear
to belong to this species. They were collected in the Antarctic Ocean, the exact
locality being uncertain, but it is believed to be in the neighbourhood of the South
Shetland Islands.
E U P H A TJ S I A, Sp. ?, JUV.
(PL IV., Figs. 7-9.)
Localities of captures : Lat. 49 40' S., long. 172 18' 30" W., five specimens,
immature, 8-9 mm.
Lat. 58 49' 45" S., long. 154 48' W., three specimens, immature, 10 mm.
The specimens from the above two localities all belong to the same species. They
were submitted to Dr. H. J. Hansen of Copenhagen, who agreed with my suggestion
that they were too young for absolute specific determination. I give here only a brief
description, pointing out a few of the characteristic features.
Carapace with a prominent slender denticle on lateral margins just over the base
of the third thoracic limbs; antero-lateral margins- slightly undulate, only partially
concealing the eyestalks and produced into a long, narrowly acute rostrum (Fig. 7)
extending almost to the anterior end of the eye and about half-way along the basal
joint of the autennules.
Pleon having the third segment provided dorsally on the median posterior margin
of the tergum with a slender spine (Fig. 9) ; sixth segment long and slender, about
twice as long as the fifth.
Antennular peduncle (Figs. 7 and 8), with a minute bluntly pointed simple lobe
on the inner distal corner of the basal joint ; a thin oblique lamella-like ridge running
across the third joint from the inner proximal to the outer distal corner and partly
continued down the inner side of the second joint.
Antennal scale reaching to about half-way along the terminal joint of the
antennular peduncle.
Telson having the portion beyond the sub-apical spines produced into an acute
apex with smooth margins ; two pairs of spinules present.
Uropods reaching to the level of the insertion of the sub-apical spines.
This species belongs to that section of the genus provided with a spine on the
dorsal surface of the third segment of the pleou. Among members of this section it
approaches most nearly to E. gibboides, Ortmann (1893), but Dr. Hansen has kindly
pointed out to me that it differs from that species in the much greater length of the
rostrum and its different shape.
The smallest of the specimens, i.e. all under 9 mm. in length, have the spine on
the third pleon segment still undeveloped, only the two largest ones, 9 5 mm. and
10 mm. in length, showing it fully formed. We have here slight evidence as to the
stage in development at which this spine appears. None of the specimens present any
larval characters in the form of the telson or antennules. It would appear, then, that
SCIII/.oi'ODA. 17
the spine, at any rate in this species, developes late in life, only after the final adult
form is reached.
The species was taken on the homeward voyage of the ' Discovery,' in the
extreme southern part of the Pacific Ocean between New Zealand and Cape Horn.
SUB-FAMILY NEMATOSCELIN.E, Holt and Tattersall.
GENUS THYSANOESSA, Brandt.
THYSANOESSA MACRURA.
(PI. III., Figs. 1-12).
Thysanoessa macrura, G. 0. Sars, 1883 ; id. (1885) ; Ortmanu, 1893 ; Stebbing, 1900 ; Holt and
Tattersall, 190(5 (l) ; Coutierc, 19m;.
Localities of captures :
Winter Quarters.
No. 4 Hole, 47 specimens, 7-18 mm.
No. 8 Hole, 40 8-21 mm.
No. 12 Hole, 2 7 and 20 mm.
From River Koettlitz, 2. 1. 03, 6 specimens, 9-14 mm.
Outward or Homeward Journey.
Lat. 61 46' S., long. 141 12' K, 16. 11. 01, 18 specimens, 14-20 mm.
Lat. 57 25' 30" S., long. 151 43' E., 20. 11. 01, 35 specimens, 12-22 mm.
Lat. 54 1' 15" S., long. 170 49' K, 27. 12. 01, 1 specimen, 6 mm.
Lat. 61 13' 30" S., long. 173 33' K, 31. 12. 01, 30 specimens, 12-19 mm.
Lat. 66 52' 9" S., long. 178 8' 15" K, 3. 1. 02, 3 specimens, 8-9 mm.
Lat. 70 29' 27" S., long. 168 51' 46" E., 26. 2. 04, 1 specimen, 28 mm.
Lat. 49 40' S., long. 172 18' 30" W., 12. 6. 04, 2 specimens, 8-12 mm.
Lat. 58 49' 45" S., long. 154 48' W., 24. 6. 04, 4 specimens, 10 mm.
Lat. 59 34' 30" S., long. 106 28' 12" W., 28. 6. 04, 3 specimens, 7 mm.
Lat, 55 44' S., long. 95 43' 30" W., 1. 7. 04, 3 specimens, 6-8 mm.
Form. (Fig. 1) of the body rather slender.
Carapace (Fig. 1) with a single rather long slender spine on the lower lateral
margin posterior to the middle, just above the origin of the sixth thoracic limb ;
antero-lateral corners acute and somewhat produced ; anterior margins very concave
and produced forwards into a long, slender, acute rostrum which reaches beyond the
eyes and far beyond the middle of the basal joint of the autenuules ; there is a low
keel on the anterior part of the carapace behind the rostrum, and a very faint
gastro-hepatic groove.
Pleon (Fig. l) rather elongate, narrow and attenuate; segments unarmed; sixth
segment equal to or slightly less than the combined lengths of the preceding two ;
18 TV. M. TATTERSALL.
preanal spine well developed, provided with au external strong tooth and a comb-like
row of finer teeth up to twelve in number.
Antennular peduncle (Fig. 2) rather long and slender, considerably longer than half
the carapace ; basal joint rather flattened, considerably wider than the remaining two
joints, bearing on its outer distal corner a rather long and slender spine, its anterior
margin somewhat overlapping the basal part of the second joint, fringed with
numerous setae and exhibiting near the inner side a shallow fissure : terminal two
joints very long and slender, subequal in length, their combined lengths slightly
exceeding that of the basal joint.
Antenna! peduncle (Fig. 3) very long and slender, nearly as long as the scale,
third joint less than half as long as the second.
Antenna! scale (Fig. 3) extending to about midway along the terminal joint of
the anteunular peduncle, almost five times as long as broad, outer margin terminating
in a well-marked spine, inner margin sloping away obliquely towards the apex ; spine
on the basal joint moderately long, slender and smooth.
Mouth parts (Figs. 4-6) do not offer any marked differences from those of
Thysanoessa gregaria as figured by Sars (1885, PL XXII).
Second thoracic limbs (Fig. 8), with the endopod remarkably long and well
developed, in full-grown specimens equalling nearly three-quarters of the total length of
the body from the eyes to the telson ; meral joint the longest, extending beyond the
tip of the antennular peduncle ; carpal joint a little shorter than the meral and more
slender, nearly four times the length of the propodal joint, armed with four slender
spiniform setae on the outer margin at the distal extremity, and five similar setse on
the inner distal margin ; propodal joint with five long spiniform setse on the outer
and six on the inner margin ; terminal joint small, tipped with six spiniform setse.
The remaining thoracic limbs (figs. 7, 9, 10) not differing greatly from the same
limbs in T. gregaria.
Copulatary apparatus (Fig. 11) on the first pleopod of the male exhibits a
structure very similar to that figured by Sars for T. gregaria, except that the distal
extremities of the two movable processes on the inner lobe do not seem to be serrate.
Telson slender, apex acutely produced and smooth ; sub-apical spines smooth ;
dorsal denticles two pairs, the first pair just anterior to the centre pf the telson, the
second pair a little anterior to the insertion of the sub-apical spines.
Uropocl.s exceedingly slender, inner one reaching the apex of the telson, outer
slightly shorter.
Length of the largest specimen, 28 mm.
I have thought it well to describe and re-figure this species, since Sars' original
description was taken from admittedly young forms, and I cannot find that mature
specimens have ever been described and figured. The changes that take place
during growth affect principally the rostrum, the second thoracic (or elongate) limb
and the preanal spine.
XMIIZOPODA. 19
The rostrum is figured correctly by Sars from a specimen, 13 mm. iu length.
At that stage iu development the rostrum is an acutely triangular projection, the
sides of the triangle very nearly straight and but little concave. As growth proceeds,
however, the sides of the triangular plate become deeply concave and the rostrum
assumes the form of a long narrow acutely spiuiform projection, as shown in Fig. 2.
This gives it something of the form as seen in T. grei/aria, but it is much longer and
more slender than in that species.
Sars describes and figures the preanal spine in his 13 mm. specimen as armed
with only two teeth. This is correctly stated for a specimen that size, but increase
in size is also accompanied by an increase in the number of teeth in the preanal spine,
till in a specimen 28 mm. iu length I found the teeth to number twelve in addition
to the strong external one. The teeth are, moreover, fine and comb-like, and the
whole spine closely resembles that figured by Sars for T. gregaria. I have noticed
all intermediate stages in the present material. This clearly undermines the value of
the preanal spine as a specific character, since the number of teeth is dependent first of
all upon age, while individual variation must also be taken considerably into account.
The most considerable changes due to growth are, however, exhibited by the
second thoracic limbs. Sars describes them as much smaller than in T. y/vfopis,
and Furcilia, larvae.
All these are apparently stages of one species. The largest Calyptopis larva
measures 3 mm. and is without a posterior median spine on the carapace. The
anterior margin of the hood is smooth ; there is a lateral spine on the carapace, and the
apex of the telson is very lightly convex. The largest Furcilia measures 4 5 mm.
Lat. 61 13' 30" S., long. 173 33' E., 31. 12. 01, one Furcilm larva, 5 mm.,
apparently the same species as the following larvse.
Lat. 63 4' 24" S., long. 175 47' 57" E., 1. 1. 02 ; numerous Furcilia larvse from
4 mm. to 5*25 mm. in length, and apparently the same species as the preceding larva.
Wood Bay, 22. 2. 04, numerous small Mctmnniplii and ('////. /ia larva.
9 mm. in length, which, from its size, is probably Euphnuxi specimens of any of
VOL. IV. ( '
22 W. M. TATTERSALL.
the genera of the family before him, but drew up his definition from the descriptions
and figures of Willemoes-Suhm (1875). He includes in the family the single type
genus, Petalophthalmus, Will.-Suhm, with two species P. armtger, Will.-Suhm, and
P. willemoesii, a new species which he founds for the reception of the female ascribed
by Suhm to P. armiger.
Faxon and Hansen, however, have since pointed out that the female specimen
described by AVill.-Suhm is in reality a Boreomysis, probably B. scyphops, G. 0. Sars,
so that if this latter view of its identity be adopted, P. willemoesii becomes a synonym
of B. scyphops.
Czerniavsky's definition of the family is inadequate, inasmuch as no reference is
made therein to the remarkable characters of the carapace, and the first and second
thoracic limbs, while the importance which is given to the supposed characters of the
exopods of the thoracic limbs is exaggerated, the difference in development as
compared with those of the Mysidse being very slight.
The definition given by Holt and Tattersall, 1906 (2), may, therefore, be adopted
with some slight alteration in the characters ascribed to the eyes, rendered necessary
by recent discoveries.
Examination of British specimens of Hansenomysis fyllse (Hanseu, 1887) has
revealed the fact that eyes are, in reality, present in this species. They resemble those
described below for the Antarctic species, except that the lappets are much reduced
and almost obsolete.
Further, in a new species of Petalophthalmus, P. oculatus, recently defined by
Illig (1906), the eyes are described, as well developed, with the cornea bright brown in
colour and distinctly facetted.
In the amended definition of the family, therefore, the description of the eyes
would read : Eyes (first cephalic appendages) small, either imperfectly developed as
lamellar or spiniform organs, without visual elements, or furnished with a distinct
cornea in which visual elements are clearly defined and functional.
GENUS HANSENOMYSIS, Stebbing.
Arctomysis, Hansen, 1887 (HOW Czerniavsky, 1883).
Hansenomysis, Stebbing, 1893.
Hansenomysis, Holt and Tattersall, 190C (1 and 2) ; Tattersall, 1907.
The name Arctomysis, given to this genus by Hansen (1887), having been already
used by Czerniavsky (1883) for an entirely different form, was changed to
Hansenomysis by Stebbing (1893). Arctomysis Czerniavsky is itself a synonym of
Boreomysis G. 0. Sars.
Of the other three genera belonging to the Petalophthalmidse Petalophthalmus,
Ceratomysis and Scolophthalmus, Hansenomysis comes nearest to the last. Both
agree in having the first thoracic limbs devoid of exopods and lacking the internal
lamelliform moral lobe, and in the presence of well-developed exopods to the second
SCHIZOPODA. 23
thoracic limbs. Whereas, however, in Scolophthalmus the rostrum is prominent and
the eyes are modified into sharp spiniform organs, Hansenomysis has the rostrum
obsolete and the eyes more or less lean 1 ike.
Males of this genus have not yet been noted, but specimens of that sex of the
northern species, H. fyllse (Hansen, 1887), have come into my hands. Detailed
examination and description are reserved for a future occasion, but it may be
mentioned here that, besides having the pleopods biramous, males also have the basal
portion of the inner flagellum of the antennule considerably thickened and adorned
with rings of setae.
HANSENOMYSIS ANTARCTICA.
(PI. V., Figs. 1-19.)
ffanseiiomysis antarctica, Holt and Tattersall, 1900 (1).
Locality of capture. Off Coulman Island, 100 fathoms, two specimens, females,
20 mm.
Form (Fig. l) compact, rather slender, tapering considerably towards the
posterior end.
Carapace (Figs. 1 and 3) short, sub-membranous, leaving the last two thoracic
segments completely exposed, and part of a third visible behind its posterior
emargination ; anterior border produced, but very slightly, into a broadly and evenly-
rounded but somewhat strongly upturned rostrum ; antero-lateral angles evenly
rounded and extending forwards as much as the rostrum ; cervical sulcus well marked
and rather deep, the posterior margin bounded by a conspicuous and rather sharp ridge
formed by the carapace. Behind the cervical sulcus is a shield-shaped dorsal area,
indicating the attachment of the carapace to the thorax, behind, and on either side of
which the wings of the carapace are free. A slight ridge runs from the antero-lateral
angles, first downwards and then -posteriorly, to meet the cervical sulcus, while a
shallow groove runs forward on each side from the dorsal shield-shaped area, thus
marking off a hepatic area, on which is a prominent forwardly-directed spine with a
broad base. A shallow depression follows the base of the rostrum, and merges on
either side into the groove formed by the ridge from the antero-lateral angles. A
small blunt and rounded spine is present on the gastric aiv;i.
Picon (Fig. 1)9 mm. in length, a little longer than the thorax, which measures 8 nun.
from the eyes to the posterior margin of the last free segment ; segments cylindrical,
postero-lateral inferior margins not at all produced as cpimera ; first segment arcuate
in dorsal contour, its anterior margin slightly raised above the level of that of the last
thoracic segment, its posterior margin broadly produced, so as to partly cover the
second segment, the whole forming a sort of " cap " over the junction of the thorax and
pleon ; second to fifth segments sub-equal in length and succeediogly narrower ; sixth
segment narrower than any of the preceding ones, and nearly twice as long.
a 2
24 AY. M. TATTERSALL.
(Figs. 1, 2 and 3) small, united at their base into a thick flattened pad, from
the anterior part of which proceed two thin, short, sub-triangular, slightly-diverging
lappets, which do not reach the middle of the basal joint of the antennular peduncle ;
visual elements entirely absent.
Antennular peduncle (Fig. 2) short and stout, its three joints sub-equal in length
and quadrangular in outline ; basal joint, with a single long seta on its internal distal
corner, and a more or less continuous submarginal row of setae across the anterior
dorsal region ; second joint with about seven long stout plumose setse on its inner
margin and two or three long setae on the outer distal corner ; third joint with about
eleven long stout plumose setse on the inner margin. On the dorsal surface of the
basal joint, partly concealed by the eye in dorsal view, is an organ of rather
problematical function (Figs. 2, 4 and 5). It appears to consist of a shallow depression
bounded by a raised ridge marked with pigment, and overhung by a membranous flap,
which apparently rises from its posterior border. The flap only imperfectly covers the
depression. In the preliminary notice of this collection it was suggested that this
organ might prove to be auditory in function, but under moderately high powers of
the microscope no otoliths or even sensory hairs could be distinguished in the
shallow pit.
Antcnnal peduncle longer than the antennular, and considerably more slender ;
distal joint shorter than the preceding.
Antennal scale (Fig. 2) lanceolate in shape, about three-and-a-half times as long as
broad, apex evenly rounded, the whole of the inner margin and distal third of the
outer margin setose ; proximal two-thirds of the outer margin devoid of setae, but
armed with eleven strong spines, the proximal one of which is the shortest, and is
situated at the end of the proximal quarter of the outer margin, the spines increasing
in size distally ; spine on the outer distal corner of the basal joint short, but
prominent.
Mandibles (Fig. 6) with the cutting edge prominent and molar process well-
developed and rather long ; between the cutting edge and the molar process is a
single spine-like seta, and in the left mandible a lacinin mobilis in addition; palp
(Fig. 7) rather long and powerfully developed, terminal joint shorter than the
penultimate, both joints armed on both outer and inner margin with numerous long
and rather stout setse.
First maxilla (Fig. 8) having the outer lobe much larger than the inner, and
armed at its apex with about thirteen spines, behind which is a row of five plumose
setae ; inner lobe armed at its tip with four long plumose setae.
Second maxilla (Fig. 9) consisting of the usual three lobes, a two-jointed palp
and outer setiferous plate, the setae arming the appendage being numerous and
rather strong.
First thoracic limb (Fig. 10) short and stout, devoid of exopod, but with well-
developed epipod ; third joint small ; fourth joint with numerous setre and a row of
snuzopom. 25
six short stout spines on its inner margin ; fifth joint with three, sixth j'>iut two, and
seventh joint four rather long, strong plumose spines on their inner margins as well
as numerous setae.
Second thomcic //////<* (Fig. 11) with the endopods longer and rather more slender
than the first, exopods well developed ; fourth or meral joint produced internally into
a large setiferous lamelliform lobe nearly as long as the iiftli joint ; the latter, the
longest joint of the limb, longer than the combined length of the sixth and seventh
joints, its outer margin armed with a single seta, the distal half of the inner margin
slightly excavate with a row of eight short, closely-set plumose spines and a single
long plumose seta on the emarginate portion, and a few long simple seta', set widely
apart, on the proximal portion of the inner margin ; sixth joint longer than the
seventh, its outer margin armed with a few long setae, the proximal portion of the
inner margin bearing a row of about nine short closely-set plumose spines and a single
long plumose seta, the distal portion of the inner margin with a few long simple seta) ;
seventh joint small, armed with numerous long and rather stout simple set;e.
Third in jiff It thoracic limbs (Fig. 12) with the endopods feeble, long and slender;
sixth joint slightly longer and more slender than the fifth ; seventh joint very small,
forming with two strong spines a very minute chela, densely clothed with short fine
set;e ; the rest of the endopod armed with a few short scattered sct.-r.
Sixth to I'ii/hlh thoracic //infix (Fig. 13) with the endopods slightly longer and
stouter than those of the three preceding pairs ; sixth joint shorter than the fifth ;
seventh joint small and bearing a long slightly-curved nail, the junction between the
nail and seventh joint being indicated by a seta on the inner margin ; rest of the
endopod feebly armed with short setae.
Exopods of the second to eighth thoracic limbs well-developed ; basal joint long
and rather narrow, the outer distal corner rounded : rlagelliforin part composed of from
ten to thirteen joints.
Incubatory lamellse, seven pairs, situated on the second to eighth thoracic limbs.
Pleopods (Figs. 14 to 18) in the female uniramous, the first pair small, succeeding
pairs increasing in size to the fifth pair, which are slightly longer than the sixth
segment of the pleon ; first four pairs one-jointed; fifth pair two-jointed, I he second
joint longer than the first ; all the pleopods bearing long >((;> at the apex.
T< l*nn (Fig. 19) rather massive, longer and a little wider than the last segment
of the pleon, dorsally grooved, oblong in shape, slightly wider at the apex than at the
base, its margins lightly arcuate ; apex truncate or very lightly emarginate, bearing
a single median spine with six or seven long spines on either side ; lateral margins
armed with from twenty-five to thirty fairly long spines arranged more or less
in series.
Iiunr nrojtotl* broken in both specimens.
Outer uropud* (Fig. 19) nearly twice as long as the sixth segment of the pleon,
two-jointed, the terminal joint about one-seventh as long as the basal ; outer margin
26 W. II. TATTERSALL.
of the basal joint without setae, but armed with twenty-one stout spines increasing in
size posteriorly.
Length of adult and ovigerous female, 20 mm. from the eyes to the tip of the
telson.
Colour of preserved specimens light brown, with a broad band of dark brown
pigment across the dorsal surface of the first segment of the pleon and scattered
patches of dark pigment on the lateral parts of the carapace, basal joints of the
anteunules arid antennas and the basal membranous pad of the eyes.
One of the specimens has young, considerably advaiiced in development, in the
incubatory lamellae.
In both specimens the telson is considerably damaged, and the description and
figures have been drawn up from both specimens and fragments of the telsons found
along with them. This fact must be borne in mind in dealing with specimens of this
species which may be found by future expeditions. It was a matter of considerable
surprise and no little interest to find in this collection two specimens belonging to
a genus hitherto known only from a single specimen from Greenland and two taken
off the cost of Ireland.
There can be no doubt that H. antarctica is co-generic with //. fyllse (Hansen,
1887). All the distinctive characters of generic importance in the mouth parts and
thoracic limbs of the latter are reproduced in H. antarctica down to the minutest
detail. The points of difference between the two species are, however, sufficiently
well-marked and numerous enough to justify specific separation. They may be pointed
out as follows :
H. antarctica is in general build a more robust and less fragile species than
H. fylb*.
Antennae. In H. fyllse the terminal joint of the peduncle is longer than the
penultimate, whereas in H. antarctica the reverse obtains.
Antennal scale. In H. fyllse the outer margin bears only five spines, between
which are numerous setae. In H. antarctica, on the other hand, there are eleven spines
on the outer margin and no setae between them.
Telson. The telson in Hansen's type-specimen was broken, but so much of
it as remained showed that the armature consisted of both spines and setae. In
H. antarctica the telson is armed with spines only, which are probably more numerous
than in //. fyllse. The shape of the telson in both species is also somewhat divergent.
That of //. antarctica recalls rather markedly the telson of Petalophthalmus armiger as
figured by Sars in the ' Challenger ' Report.
Outer uropods. //. fyllce has the outer uropods armed with both spines and
setae, whereas in //. antarctica there are spines only present. The spines in H. fi/llce
number six, while in H. antarctica there are twenty-one.
Pleopods. The type specimen of H. fyllce had only one pleopod remaining. This
was one of the third pair, and is described by Hansen as biarticulate. Presumably,
8CHIZOPODA. 27
therefore, the fourth and fifth pleopods will likewise be found to be biarticulate when
perfect specimens are examined. In //. unfuri-tica only the fifth pleopods are
biarticulate, the remaining pairs consisting of a single joint only.
The eyes in the genus are described for the first time. They are remarkable
chiefly for their small size and degenerate structure, for the complete absence of visual
elements, and the subservience, either entirely or in great part, of ophthalmic functions
to those of probably a tactile nature.
The cap-like form of the tergum of the first segment of the pleon recalls the
somewhat similar form of the third pleon segment in many Carida, and suggests that
the posterior part of the body is capable of great ventral flexure. The ' Discovery '
expedition is to be congratulated on the finding of this species, by far the most
interesting Schizopod in the collection.
FAMILY MYSID^E.
SUB-FAMILY LEPTOMYSIX.E, Norman.
GENUS PSEUDOMMA, G. 0. Sars.
PSEUDOMMA BELGICjE.
(Plate VI, Figs. 1-8.)
Pseudomma belgicse, Holt and Tattersall, 1006 (1).
Locality of capture. Lat. 78 25' 40" S., long. 185 39' 6" E., 300 fathoms, one
specimen, immature female, 23 mm.
Fi.inn (Fig. l) compact and moderately stoutly built,
Carapace (Fig. 1) large, less than half the total length of the body, covering
laterally all the segments of the thorax, but dorsally exposing the last one behind its
posterior emargination ; its anterior margin very slightly produced into a blunt, very
broadly rounded rostrum ; antero-lateral angles rounded; cervical sulcus well marked.
Pleon (Fig. 1), excluding the telson, about half the total length of the body from
the eye to the tip of the telson ; first four segments subequal in length and slightly
longer than the fifth ; sixth segment twice as long as the fifth.
Eye plates (Fig. 1) contiguous, exhibiting only a very slight anterior median
cleft ; each plate subquadrangular or rhomboidal in shape, nearly twice as broad as
long, antero-lateral angles rounded, anterior margin nearly straight, no serrations or
armature of any kind ; no pigment present in preserved specimens ; corneal lenses
absent, but the ramifications of the optic nerve are clearly visible in dorsal view.
Antennular peduncle (Fig. l) short and stout, not extending beyond half the
length of the autennal scale ; basal joint almost entirely covered by the ocular laminae,
a few plumose setse on each anterior corner ; second joint very short, more than twice
as broad as long, a few short seta; on the outer distal corner, inner margin with a few
longer plumose sehe ; third joint longer than either of the other two and .-lightly
W. M. TATTEKSALL.
narrower, rectangular in shape, outer margin unarmed, inner margin beset with a few
plumose setae.
Antenna! peduncle (Fig. 1) equal in length to the autennular, but more slender;
last two joints subequal in length.
Antennal scale (Fig. l) slightly longer than the last segment of the pleon and
twice as long as the antennular peduncle, about three and a half times as long as
broad, outer margin entire and terminating in a very strong spine, beyond which the
apex of the scale is but slightly produced ; spine on the basal joint short and acute.
Mmitli, parts (Figs. 2, 3, 4, and 5) exhibit no salient points of difference from
those of the type species of the genus P. roseuin, G. 0. Sars.
First tun I xi'ivi/il tlii'i-iiclc limbs (Figs. 6 and 7) agreeing in the main with those
figured by Sars for P. roseum, but a dactylus is distinctly visible among the dense
mass of plumose setfe arming the terminal joints, though it is smaller and shorter
than the terminal joint ; exopods having the outer distal corner of the basal joint
round, though produced, and the flagelliform part composed of ten to twelve joints.
Telson (Fig. 8) slightly shorter than the last segment of the pleou, tapering to
an evenly rounded apex which is one-third as wide as the base ; apex armed with a
median pair of plumose setae and four pairs of strong smooth spines, the innermost
and largest of which equals one-sixth of the telson in length ; distal half of the lateral
margins armed with five shorter spines.
Inner uropods half as long again as the telson, armed with a single long spine
in the region of the inner posterior corner of the otocyst.
Outer uropods about twice as long as the telson.
Length of an immature female, 23 mm. By an error the specimen was described
in the preliminary report as adult. This is scarcely correct, since the incubatory
lamella; are still only about half developed, so that the adult female probably reaches
to nearly 30 mm. The specimen is badly mutilated, the third to the eighth thoracic
limbs being entirely absent.
P. beljicce is far and away the largest species of the genus yet described, none
of the other ten known species exceeding 15 mm., whereas adult specimens of this
species must reach to nearly 30 mm. It is most nearly related to P. sarsi Will.-
Suhm, described by Sars (1885) from the ' Challenger ' collections for specimens taken
at Kerguelen Island. Besides the great difference in size (P. sarsi measures only
14 mm., adult specimens), the only other conspicuous difference is in the ocular
lamiuse. In P. sarsi the antero-lateral angles of the eye-plates are serrate, whereas
in P. In'lijica 1 they are quite smooth. Minor differences in the shape of the antennal
scale and telson may also be noted. The antennal scale in P. sarsi has the spine
terminating the outer margin less strong than in P. belgicce, while the apex of the
scale is more produced. The telson in P. sarsi has the apex more truncate than
P. fn'ft/ii'tr and the lateral margins, according to Sars, bear eight short spines.
Mr. Holt, however, who has kindly examined the types of P. xarsi in the British
SCIIIZOI'ODA. '-".I
Museum, informs me that only five spines are present on the lateral margins, so that
the armature of the telson of P. fmrni approximates closelv to that of P. belgicce.
The only other species of Pseudomma having smooth ocular lamina- is
P. australe, G. 0. Sar* (1885), from Bass Straits, Australia. The vastly different
form of the antennal scale in the latter, however, abundantly distinguishes it from
/'. fi't'/icce.
Besides the single ' Discovery ' specimen, this species is also known from the
' Belgica' collections, and has been described by Hansen in ^\IS. under the name which
is here used. It is possible that the mutilated specimen noted by Bars (1885, p. I'.n )
from 1675 fathoms in the Antarctic Ocean may have belonged to this species rather
than to P. air*!. Sars notes that it was much larger than the latter.
( !f)ius I).M i vi.AMDLVors. Holt and Tattersall.
/ti/r/i/litmd/i/iijix. Holt and TattiTsall, I'.iml (1).
/>,i,-li///r//t/ii-x, Illiir, I'.HK;, linn Holt and Tattursall, l'.M5.
? Amlli/ops (///), Ohliu, i:i"l.
Dactylamblyops, Tattersall, I'.lnT.
This genus was established for the reception of the single rather mutilated specimen
of D. /iiiili/xi'/i! in the present collection. Since the publication of the preliminary
notice of the 'Discovery 1 Schizopoda, however, two closelv allied species have Keen
discovered off the west coast of Ireland (Tattersall, 1907). A clearer idea of the
exact relationships of the genus has thus been gained, and while the species referred
thereto appear, in the present state of our knowledge, to form a natural group, it is
undeniably very nearly allied to Dactylerytkrops, Holt and Tattersall (1905), to
which genus, indeed, the present species was referred by Illig (1906).
The definition of the genus given by Holt and Tattersall, 1900 (1), may therefore
be amended as follows :
DACTYLAMBLYOPS, Holt and Tattt-rsall.
Characters generally as in Airiblyops, G. 0. Sars, except :
/v/.'.v placed close together, but not contiguous, more or less pyriform in shape,
furnished with distinct and definite peduncles ; visual elements imperfectly developed.
numerous, reaching to the surface of the eye, and probably directly functional as
organs of sight; outer distal corner rounded, and not produced into a digitifonn
process ; a short blunt process always present on the inner and upper surface.
Second thoracic limb* with the eiidopods not noticeably short, but well developed,
and considerably longer than the endopods of the first thoracic limbs.
Telson not very long, triangular in shape, the distal parts of its margins armed
with more or fewer spines ; median set?e absent.
Type species, D. hodgsoni, Holt and Tattersall.
VOL. IV. II
30 W. M. TATTEBSALL.
The absence of median setaj from the apex of the telson is not necessarily
of generic importance, since the genera Pseudomma and Dactylerythrops both contain
species in some of which these setae are present, and others in which they are
wanting. As, however, all three species at present referred to this genus are without
median apical setae, it is convenient to retain this character in the generic definition.
DACTYLAMBLYOP.S HODGSONI.
(PI. VI., Figs. 9- 16.)
Dactylamblyops hocljsoni. Holt and Tattersall, 1906 (1).
Durtylerythrops areiiata, Illig, 1000.
Locality of capture : Lat. 66 52' 09" S., long. 178 08' 15" E., 2030 fathoms,
one specimen, male, 13 mm.
The single specimen in the collection is considerably damaged, the antenuules,
autennal scales, and the third to the eighth thoracic limbs being missing. A complete
description is therefore not possible, but it is hoped that as many of the characters as can
be made out with certainty will suffice for future recognition of the species in collections.
Carapace submembrauaceous, covering all the thoracic segments except the last
one, anterior margin produced into a blunt, broadly but evenly rounded rostrum
projecting between the eyes ; cervical sulcus well marked ; antero-lateral angles rounded.
Picon slightly longer than the carapace ; first five segments subequal in length ;
sixth nearly twice as long as the fifth.
Eyes (Fig. 9) small, placed- on definite peduncles, not in any way contiguous,
pyriform in shape, external angle evenly rounded, a short digitate process arising from
the inner dorsal face ; visual elements imperfectly developed, apparently represented
by numerous minute granular bodies with a refractive centre ; a large opaque gaugliouic
mass, probably the optic nerve, visible in the peduncle, from which a nerve fibre
proceeds to the cornea.
Antennal peduncle short, composed of three subequal quadrangular joints.
Antennal scale broken on both sides, but there does not appear to be a spine on
the outer corner of the basal joint.
Mouth parts (Figs. 10-13) not differing markedly from those figured by Sars for
AmUyops abbreviata (1870-79).
First thoracic limbs (Fig. 14) with the eudopod substantially of the same form as
in A. abbreviata.
Secund thoracic limbs (Fig. 15) of essentially the same structure as in A. abbreviata,
but with the endopod apparently much longer, being nearly twice as long as the
endopod of the first thoracic limbs.
Genital appendix to the last thoracic limbs of the male terminating in two lobes,
the larger of which bears six long setre, the smaller one being devoid of seta;, but
apparently having a covering of very fine hairs.
SCIIIZOPODA. :! I
in the male agreeing in all points with those of males of tin- genus
Arriblyops.
7i-/.\-nn (Fig. 1G) not unite as long as the last segment of the pleon, triangular in
shape, tapering evenly to a narrowly rounded apex, nearly twice as long as broad a1
its base; distal half of each lateral margin armed with nineteen spines, increasing in
length towards the apex, the terminal ones about one-tenth of the total length of the
telson ; median setae absent.
f nyW.v broken on both sides, but the inner one possesses a single strong .-pini-
on the ventral surface at the inner posterior angle of the otocyst.
Length of the single specimen, an apparently adult male, 1:1 mm.
There can be little doubt, I think, that Dactylerythmps nn'mitn, Illig (190G), is
the same species as the present one. Minor differences, it is true, are to lie IK it iced.
For instance, the visual elements of the eye in Illig's species are represented as larger
and less numerous than in D. ln>(l/jsj'. it
is of quite peculiar form, with an outer equatorial membranous ridge, while in
D. goniops the eye is quadrangular rather than pyrilbrm in shape. Otherwise the
four species are rather closely allied and form quite a distinct generic group to
themselves.
D. hodgsoni is at present only known from very deep water in the Antarctic
Ocean. Illig's specimens were collected over a depth of 4000-5000 metres, while the
present specimen was dredged in 2030 fathoms (,-,i. :;7o<) metres).
ii 2
W. M. TATTERSALL.
SUB-FAMILY MYSIDETIN^E, Holt and Tattersall.
GENUS MYSIDETES, Holt and Tattersall.
? Mi/sMiysis, G. 0. Sars, 1883 aud 1885, -non G. 0. Sars, 1804.
Mysideis (pars), Holt and Tattersall, 1905, non G. 0. Sars, LSI;;).
Myxidttes, Holt and Tattersall, 190(5 (1) and (2).
Jfi't), lanceolate in shape; between four and five times as
long as broad ; setose all round ; a minute second joint at the apex ; a spine on the
outer distal corner of the basal joint.
^fllndinl^'s (Fig. 4) with a well-developed molar process and cutting edge ; palp
(Fig. 5) with the second joint somewhat expanded and armed with lung seta: on
both margins ; third joint not much expanded, a row of strong plumose setoe on
the lower edge, and two very strong simple spine-like seta; at the tip.
Second iiuu'illni' (Fig. 7) with the setiferous expansion of the basal joint well
developed.
Endopods of the first thoracic limbs (Fig. 8) of about the same build as in the
genus Mysidopsis, but seven-jointed ; masticatory lobe well developed : inner margins
of the proximal four joints armed with numerous plumose setae ; sixth joint bearing
a well-developed nail and beset with numerous plumose setae.
Eiidopods of the second thoracic limbs (Fig. 9) very similar to those of Mysidupsis ;
longer than the first ; sixth joint armed with numerous plumose seta:, but in the
specimen dissected I was unable to detect a nail, though it may have been
broken oti.
34 W. M. TATTERSALL.
Endopods of the third thoracic limbs (Fig. 10) with the merus longer than the
tarsus; latter composed of six joints; nail well developed and Linger than the last
joint of the tarsus.
Endopods of the remaining thoracic limbs become successively longer and more
slender from the fourth to the eighth ; the increase in length takes place chiefly in
the ischial joint ; the number of joints in the tarsus of the endopods also increases
in the more posterior limbs ; in one specimen there were six joints in the tarsus of
the third limb, six in the tarsus of the fourth, ten in the tarsus of the seventh, ami
twelve in the tarsus of the last limb.
Genital appendix (Fig. 11) on the last thoracic limb of the male exceedingly
long and slender, equal in length to the first three joints of the limb to which it is
attached.
Pleopods (Fig. 1'2) similar in both sexes, consisting of a single ramus bearing
proximally and externally a rather large process tipped with setse.
Tdson (Fig. 13) a little longer than the last segment of the pleon, and more
than twice as long as broad at its base ; cleft at the apex for nearly a quarter of its
length, cleft rather wide, its margins armed with about eighteen teeth on each side ;
the apex of each lobe of the cleft armed with a pair of spines, the inner one the
shorter ; lateral margins armed throughout their whole length with about seventy
spines, which become arranged in series towards the apex.
Inner wopods slightly longer than the telson, with a row of moderately slender
and long spines on its inner ventral margin, varying in number from twenty-six to
twenty-eight, and extending from the otocyst to near the apex ; spines not arranged in
series, but increasing in size distally. In some specimens the spines extend further
down the uropod than in others.
Outer uropods about half as long again as the inner.
Length of an adult female with embryos in the brood pouch, 21 mm. ; of an
apparently adult male, 25 mm. A second female with embryos in the brood pouch
measured 23 mm.
It is not a little interesting that this genus should have been discovered almost
simultaneously in the northern and southern hemispheres, M. farrani, Holt and
Tattersall (1906 (2)) having just been described when the 'Discovery' collections
came to hand.
M. posthon is a more stoutly built form than M. farrani, and is further
distinguished from the latter in the following characters :
Antennules. M. farrani has not the outer corner of the basal joint of the
peduncle produced nearly as much as in M. post/ton.
Thoracic limbs. The tarsus of the endopods in M. posthon is composed of six to
twelve joints, while in M. farrani there are only four.
(.< rii i tttl appendix to the last thoracic limb of the male is much longer and more
slender in Hf. posthon than in ]\f. farrani.
SOHIZOPODA.
Pltopods. The lateral lobe is less developed in M. farrani than in M. [
Telson. In M. farrani the cleft is armed with only about thirteen spines.
whereas in M. postlion there are about thirty-six. In the former, moreover, the lateral
margins of the telson are armed with not more than twenty-six spines, not arranged in
series, and situated only on the distal two-thirds of the margin. In 3[. posthon the
lateral margins are armed throughout the entire length with about seventy spines,
arranged, at any rate, distally in series. The whole telsoii in ^^. farrani is more
slender than in M. posthmi.
The spines on the inner nropods of M. posthon appear to be somewhat longer than
in M. farrani I have already expressed the opinion that the genus Metamysidella of
Illig is .synonymous with Mysidetes. The type species of the former, ^f. kerguelensis,
Illig (190G), is, however, a much smaller species than M. /><>*f/i<'ii. measuring onlv
10 mm. in length. It is otherwise closely allied to the latter, but differs in having
the antennular peduncle almost equal in length to the antennal scale and in the details
of the armature of the telson.
If .l///.v/rAyw.v incisn, G. O. Sars (1885), should in future be found referable t<> the
genus Mysidetes, as seems probable, it differs from the present species in size, in
having fewer joints in the tarsus of the thoracic limbs, and in the details of (he
armature of the telson.
I should mention here that I do not attach too great an importance to i In-
difference in size between M. kerguelensis and Mysidopsis incisa as compared with
JA poxtlton as a specific character, for I have found both males and females of
M. farrani quite sexually mature at 15 mm. (judging from the characters of the
antennular brush in the male and the incubatory lamella- in the female), while the
species, fully grown, reaches to 28 mm. in total length.
SUB-FAMILY ^
OKNUS ANTAKCTO.MYSIS, ('mitiere.
.l/y.s/.s, Holt ami TattLTsall, I'.H'i; (1).
Antarctomysis, Cnntir-rc, I'.ioi;.
This genus has been recently established by Coutiere for the reception of the
species briefly noted as J ///*/* ii/ii.r!in
Mysis (sens, stricto). The genus ///////////.< /'.v has the fifth pair of pleopods in the male
liiramoiis and natatorv. but the third pair are only imperfectly liirannnis. the outer
ramus being very minute and single-jointed, whereas in . l;//i//r/.i//,_//.v/'x the third pair
resemble the fifth in having both rami multiarticulate and setose.
36 AV. M. TATTBRSALL.
1 became aware, only after the plates illustrating this report had been printed,
that the two specimens of Antarctomysis in the ' Discovery ' collections, referred in the
preliminary note to one species A. maxima, in reality belong to two distinct but very
closely allied species. On my appealing to Dr. Hausen, he very kindly sent me some
notes and sketches of .4. maxima, and a second species of the genus discovered by him
in a collection which he is engaged in working out. These notes and drawings placed
the matter beyond doubt, the larger of my two specimens clearly belonging to Hansen's
second species. I note the species here, and give the points of distinction, but leave a
full description and name to Dr. Hansen. The drawing on PI. VIII. , Fig. 1, was taken
from the real A. maxima, but the remaining figures on the plate represent the
appendages of the second species, which, at the time, I took to be A. maxima also.
They will probably be of use, however, in illustrating how closely allied the two species
are when they are compared with the figures given by Coutiere (1906) of the true
A. maxima.
ANTARCTOMYSIS MAXIMA.
(PI. VIII., Fig. 1.)
Mysis maxima (pars), Holt and Tattersall, 190G (1).
Antarctomysis maxima, Coutiere, 1!)06.
Locality of capture. AVinter quarters, 5. 6. 02, D-uet hole, 56 fathoms, one
specimen, immature male, 33 mm.
Coutiere (1906) has recently described this species in great detail from mature
examples collected by the French Antarctic Expedition. I have practically nothing to
add to his description, but since no figure of the entire animal was given by him, my
drawing on PL VIII., Fig. 1 may be useful.
Coutiere does not mention the spines arming the inner ventral edge of the inner
uropod. They extend from the posterior inner corner of the otocyst to the extreme
tip of the uropod, and posteriorly, at least, are arranged in series of twos, threes and
occasionally fours. The species would appear to be circumpolar in distribution, since,
besides the single specimen in the ' Discovery ' collection, it has been taken by the
French, Swedish and Belgian Antarctic expeditions ; by the two former, in considerable
numbers.
ANTARCTOMYSIS sp.
(PL VIII. , Figs. 2-12.)
M>jsis maxima (pars), Holt ami Tattersall, I'.KiG (1).
Locality of capture : Lat. 78 25' 40" S., long. 185 39' 6" E., 300 fathoms, one
specimen, immature male, 40 mm.
This species is so closely allied to A maxima that I only became aware that it
was distinct when too late to properly illustrate it. The figures 2-12 on Plate VIII.
were taken from the appendages of this specimen.
DA. .".7
The species will lie fully described and named by Dr. Hanson, so here 1 will
merely note the points of distinction between it and A. maxima.
(1) Eye. In .1. maxima (I'l. A' 111., tig. I) the eye is largo and the visual elements
occupy a large part of -the outer side of the eye-stalk, so that in dorsal view the inner
eye-stalk proper is much longer than the outer, and in external lateral view verv
little of the latter is visible. In the present form the eye is smaller and narrower than
in A. maxima, the visual elements occupy the terminal part of the eye-stalk only, so
that the inner and outer margins of the latter are snboqiial in length, and in external
lateral view practically the whole of the eye-stalk is visible.
(2) Rostrum. In A. maxima the angle contained by the antero-lateral margins
of the carapace which form the rostrum is equal to or slightly greater than a right
angle, so that in lateral view the antero-lateral margins are not very oblique. The
tip of the rostrum is produced into a very small spine.
In the new species the angle of the rostrum is considerably less than a riglit
angle, so that the antero-lateral margins of the carapace in lateral view are very
oblique. The apex of the rostrum is bluntly rounded.
(3) Aiifi'i/i/ii. In A. maxima the basal joint of the antenna, from which the
antennal scale and peduncle arise, bears two spines vcntrallv. one at each of the outer
and inner distal corners. In the new form, only the one on the outer distal corner is
present, the inner corner being rounded.
(4) In ^1. maxima the tarsus of the third to the eighth thoracic limbs is seven
to eight-jointed (excluding the nail); in the present species the tarsus is six to
seven-jointed, so that the two distal joints before the nail are proportionately longer
than in A. maxima (cf. PL VIII., Fig. 8, with Coutiere (1900), I'l. I.. Fig. I 1).
In other characters the t\ve species are practically identical.
38 AV. M. TATTER8ALL.
LIST OF AUTHORITIES QUOTED.
COUTIERE, H.. 1906. Expedition Charcot. Crustaces Schizopodes et Decapodes, Paris.
CZKRNIAVSKY, W., 1882-3. Monographia Mysidaruui imprimis Impcrii Rossici, fasc. i.-iii., St. Petersburg.
DANA, J. D., 1852. United States Exploring Expedition, Crustacea.
HANSEN, H. J., 1887. " Malacostraca marina Groenlandiae occidentalis." Vid. Medd. naturh. Foren.
Kjobenavn.
HANSEN, H. J., 1905 (1). " Prelim. Report Schizopoda 'Princess Alice,' 1904." Bull. Mus. Ocean.
Monaco, no. 30.
HANSEN, H. J., 1905 (2). "Further notes on the Scbizopoda." Bull. Mus. Ocean. Monaco, no. 42.
HODKSON, T. V., 1902. Schizopoda in " Report collections Natural History ' Southern Cross.' " Loudon.
HOLT, E. W. L., and TATTERSALL, W. M., 1905. " Schizopod. Crust. N. E. Atlantic Slope." Report
Sea and Inland Fisheries, Ireland, 1902-3, Pt. ii., Scientific Investigations, Appendix no. iv.
HOLT, E. "W. L., and TATTERSALL, "W. M., 190G (1). "Prelim, notice Schizopoda 'Discovery.'" Ann.
and Mag. Xat. Hist. ser. 7, vol. xvii.
HOLT, E. W. L., and TATTERSALL, W. M., 190G (2). "Schizopod. Crust. N. E. Atlantic Slope. Supple-
ment." Fisheries, Ireland, Sci. Invest., 1904, v.
ILLIG, G., 190fi. "Bericht ii. die neuen Schizopoden-Gattuug und ArtenderDeutschenTief see-Expedition,
1898-1899." Zoologischer Anzeiger, Bd. xxx., no. 7.
NORMAN, A. M., 1902. "Notes on the Natural History of East Finmark." Ann. and Mag. Nat. Hist.,
ser. 7, vol. x.
OHLIN, A., 1901. "Arctic Crustacea." Bihang Kongl. Sven. Yet.-Akad. Hand!., Bd. 27, Afd. iv.
ORTJLANN, A. E., 1893. Decapoden und Schizopoden der Plankton-Expedition. Ergeb. Plankton Erf. der
Humboldt-Stiftung, Bd. ii. G, b.
SARS, G. 0., 1SIU. Beretning om en i Sommeren 1863 foretageu Zoologisk Reise i Christiania Stift.
SARS, G. 0., 1869. Undersogelser over Christianiafjordens Dybvaudsfauna. Christiania.
SARS, G. 0., 1870-79. Carcin. Bidrag til Norges Fauna. I. Monog. Norges Mysider.
SARS, G. 0., 1883. "Prelim, notices Schizopoda ' Challenger.'" Forhandl. Vidensk. Selsk. Christiania,
no. 7.
SARS, G. 0., 1885. Report on the Schizopoda collected by H.M.S. 'Challenger.' Zool. 'Challenger'
Exped., Pt. xxxvii., vol. xiii.
STEBBING, T. R. R., 1893. History of Crustacea. London.
STEBBIXI;, T. R. R., 1900. " On some Crustaceans from the Falkland Islands." Proc. Zool. Soc., London.
TATTERSALL, W. M., 19n7. "Prelim, diagnoses of six new Mysidat from the West Coast of Ireland."
Ann. and Mag. Nat. Hist., ser. 7, vol. xix.
WiLLEMOES-Sumi, R., 1875. "Some Atlantic Crustacea from the ' Challenger ' expedition." Trans.
Linn. Soc., London, ser. ii., vol. i.
ZIMMER, C., 190-4. Arktische Schizopoden. Fauna Arctica, ii.
ZODIEK, C., 1905. "Biologische Notizen iiber Schizopoden." Verhandl. d. Deutsch. Zool. Gesellschaft.
39
EXPLANATION OF THE PLATES.
PLATE I.
Euphausia superba.
FIG. 1. Male, 4;"i mm., dorsal view of anterior end x 10.
Fn:. 2. Female, 45 mm., doi&al view of anterior end x 10.
Fin. :;. " GlfiriiiHx '' stage, dorsal view of anterior end x 15.
Fin. 4. Late Cyrtopia stage, dorsal view of anterior end x 30.
Fin. 5.- Maudilmlar palp x 1:!.
FIG. d. Second maxilla X 1:1.
FIG. 7. First maxilla x l:i.
FII;. *. First thoracic limb, endopod x '.'.
FIG. '.i. Sirond thoracic limb, endopod X '..
Fin. 10. Lateral s| line on carapace of female, 50 mm. x :!.
Fn;. 11. Lateral spine on carapace of male, 4.1 mm. x :.
Fin-. 12. Lateral spine on carapace of male, :!'.) nun. X 30.
PLATE II.
FIG. 1. Male, lateral view of entire animal x U.
FIG. 2. Female, dorsal view of anterior end x l">.
FIG. 3. Manclibular palp x :H>.
Fin. 4. First maxilla x 30.
FIG. 5. Second maxilla x 30.
FIG. 6. First thoracic limb x 20.
FIG. 7. Second thoracic limb, eudopod x -".
Fin. 8. Calyptopis larva x 4n.
FIG. 9. Furcilia larva x 20.
FIG. 10. Cyrtopia larva x 2i>.
I'l.ATE III.
ma? runt.
FIG. 1. Female, 28 mm., lateral view x G.
Fn;. 2. Female, dorsal view of anterior end x li>.
Fin. 3. Antenna! scale and peduncle x K>.
FIG. 4. Mandibiilar palp x :<>.
PlG. 5. First maxilla x 30.
Fn:. (1. Second maxilla x 80.
FIG. 7. First thoracic limb, endopod x '>.
Fin. 8. Second thoracic limb, endopod x 1">.
FIG. 9. Seventh thoracic limb, endopod x :'".
Fm. in. -Rudimentary eighth thoracic limb x f>.
l-'in. 12. Endopod of the second jilenpod "f the male x t'.n.
I -2
40 W. M. TATTERSALL.
PLATE IV.
FIG. 1. Male, lateral view x 0.
FIG. 2. Male, dorsal view of anterior end x 20.
FIG. 3. Inner lobe of the endopod of the first pleopods of the male to show copulatory apparatus X 80.
Euphausia vaU?iitini.
FIG. 4. Outline of antennular peduncle of ' Discovery ' specimen x 20.
FIG. 5. Outline of rostrum and basal joint of antennular peduncle of 'Challenger' specimen x :!0.
FIG. G. Inner lobe of the endopod of the first pleopods of the male in ' Challenger*' specimen, to show
copulatory apparatus X 80.
Euphausia sp, juv.
FIG. 7. Dorsal view of anterior end of specimen, 10 mm. x 00.
FIG. 8. Lateral view of antennular peduncle of the same specimen x GO.
FIG. 9. Spine on the third segment of the pleon of the same specimen x 2<>.
Euphausia orystalloroph iii*.
FIG. 10. Inner lobe of the endopod of the first pleopods of the male, to show copulatory apparatus X 80.
PLATE V.
iit antarctica.
FIG. 1. Female, dorsal view x 10.
FIG. 2. Enlarged view of anterior end x 23.
FIG. 3. Side view of anterior end x 23.
FIG. 4. Dorsal view of peculiar antennular organ x 70.
FIG. 5. Side view of same x 70.
FIG. 0. Mandible x 20.
FIG. 7. Mandibular palp x 20.
FIG. 8. First maxilla X 20.
FIG. 9. Second maxilla x 20.
FIG. 10. First thoracic limb x 20.
FIG. 11. Second thoracic limb, endopod X 10.
FIG. 12. Third thoracic limb, endopod x 10.
FIG. 13. Sixth thoracic limb, endopod x 10.
FIG. 14. First pleopod x 20.
FIG. 15. Second pleopod x 20.
FIG. 16. Third pleopod x 20.
FIG. 17. Fourth pleopod x 20.
FIG. 18. Fifth pleopod x 20.
FIG. 19. Telson and uropods x 20.
PLATE VI.
Pseudomma lelyicae.
FIG. 1. Female, dorsal view x 10.
FIG. 2. Mandible x 30.
FIG. 3. Mandibular palp x 30.
FIG. 4. First maxilla x 3ii.
FIG. 5. Second maxilla x 30.
FIG. 6. First thoracic limb, endopod x 13.
FIG. 7. Second thoracic limb, endopod X 13.
FIG. 8. Telson X 20.
srilIZOPODA. II
D(Ktylain /i///iij/.i
FIG. i). Eye, external lateral view x i'".
FIG. 10. Mandible x I".
FIG. 11. Mandibular palp x 4<>.
FIG. li'. First maxilla x 4i>.
FIG. 1:1. .Second maxilla X In.
FIG. 14. First thoracic limb, endopod X -'7-
FIG. 15. Second thoracic limb, endopod X :.'7.
FIG. 111. Telson x 20.
PLATE A 71.
FIG. 1. Female, dorsal view x 13.
FIG. 2. Antennnlar peduncle x is.
FIG. 3. Antenna! peduncle with antennal scale x is.
FIG. 4. Mandible X IS.
FIG. .">. Mandibular palp x is.
FIG. C. First maxilla X IN.
IMG. 7. Second maxilla x is.
FIG. s. First thoracic limb, endopod x is.
FIG. 9. Second thoracic limb, endopod x IN.
FIG. ID. Third thoracic limb, endopod x IN.
FIG. 11. Genital appendix of male x IN.
FIG. 12. First pleopod x 24.
FIG. 13. Telson X IS.
PLATE VIII.
iiitt.riiiin.
FIG. 1. Immature male, dorsal view x (!.
FIG. 2. Mandible X 2<>.
FIG. :!. -.Mandibular palp X 2n.
FIG. 4. First maxilla x 20.
FIB. ii. Second maxilla x L'H.
FIG. 6. First thoracic limb, endopod x s.
FIG. 7. Second thoracic limb, endopod x s.
FIG. 8. Third thoracic limb x s.
FIG. 9. Second pleopod of the male (immature) x 15.
FIG. 10. Third pleopod of the male (immature) x 15.
KIG. 11. Fourth pleopod of the male (immature) x 15.
Fio. 12. Fifth plcopod of the male (immature) x 15.
INDEX OF GENEPvA AND SPECIES.
Amblyops, 29.
Amlly/ijix iililu-fi'iata, 30.
Ainbli/ops rrozetti, 3.
Aiifturtmnyxin, 3, 35.
Antarctomysis maxima, 2, 36.
Arctomysis, 22.
Boreomysis, 22.
Boreomysis scypJwps, 3, 22.
Geratomysis, 22.
Dactylamblyops, 3, 29.
Dactylamblyops ymuops, 31.
Dactylamblyops hodgsoni, '2, 30.
Dactylamblyops sarsi, 31.
Dactylamblyops ihaumatops, 31.
Dactyleryfhrops, 29.
Dactyleryfhrops arcuata, 2. 30.
Echinomysix cfnmi, 2.
Eitcopia, 3.
Eucopia awl rails. 1, 2, 3.
Euphausia, 4.
EupJiausia sp., 14, Hi.
Euphausia anfarctica, 1, 4.
Euphausia australis, 1, 4, .">.
Euphausia crystallorophias, 2, 9.
Euphausia yibboides, 10.
Eiiplimisia glacialis, 1 , 4, 8.
Euphausia luceiis, 1, 11, 14.
Eiiplmusia murrayi, 1, 4, C.
Euphausia. similis, 2,11.
Euphausia splendens, 1, 11, 13, 14.
Euphausia svperba, 1, 2, 4.
Euphausia triaca/itka, 12.
Euphausia wUentini, 13, 14.
Hansenomysis, 3, 22.
Hannfiiomysis antan-tifii, 3, 23.
Hansenomysis fyllse, 22, 2fi.
Hemimysis, 35.
Heteromyxis, 32.
Lophog aster typicus, 3.
Metamysidella, 32.
Metamysidella Icerguelmsis, 35.
Michtheimysis mi.i'ta, 3.
Mysideis, 32.
Mysiddla, 32.
Mysiiktes, 3, 32.
Mysidetesfarrani, 34.
Mysidetes posthon, 33.
Mysidopsis, 32.
Mysiilopsis ini'isa, 32, 35.
J///N/.S-, 35.
My sis maxima, 1, 35, 36.
Nyctiphanes couchi, fi.
Petalophthalmus, 22.
Pi'talophthalmus armiger, 22.
Petalophthalmus omlatus, 22.
Petalophthalmus tvillemoesii, 22.
Pseudomma, 3, 27, 30.
Pseudomma australe, 29.
Pseudomma belyicae, 27.
Pseudomma roseum, 28.
Pseudomma sarsi, 1, 2, 28.
Scolophthalmus, 22.
Thysanoessa, 3, 17.
Thysanoessa gregaria, 18.
Thysanoessa macrura, 1, 2, 17.
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CRUSTACEA.
VIIL-COPEPODA.*
BY R. NORRIS WOLFENDEN, M.D. (Cantab.), F.L.S., F.Z.S.
(7 Plates.)
PREFATORY NOTE.
THE collection handed to me for examination and report thereon was contained in 163
bottles, the contents of a few of which were in such bad preservation that they were
practically worthless for the purpose of identification. The collection of individual
species is not a large one, though there were great quantities of the more common
species. But few absolutely new forms were found ; these comprised a new genus
(Paralabidocera] and seven new species (Euckceta similis, Stephus ant /////*. Fitr<>< tlit nntiirfti<-ii, and Gaetanus
mttarcticus). As such of these copepods as may be considered Antarctic were
collected within a small area, I have not deemed it necessary to occupy space by the
repetition of individual captures, which would be monotonous and of no particular
interest. The drawings have been made by Miss Marion Lees.
The signs used in the following pages are B 1 and B 2 for first and second basals :
Ri and Re for endopodite and exopodite ; Si for inner marginal and N<- for outer
marginal spine (or bristle) ; Li for inner and L<- for outer lobe ; Th for thoraric
somite. As they were first used in (liesbrecht's great work, and have been subse-
quently often employed by others as abbreviations, the author has thought no
excuse necessary for their use here, in order to avoid the constant repetition of the
words " exopodite " and " endopodite," etc.
I.
UNTIL the expedition of the ' Belgica ' there existed no records of the collection of
Copepoda south of Kerguelen, except those of Dr. Brady, which referred to the
' Challenger' collections made from the south of Kergucleu to the pack-ice at 66" 'J9' S.
The collections made by the ' Discovery,' the ' Belgica,' and the ' Gauss ' form a
most important contribution to the plauktonology of this southern region, and the
* Owing to the author's absence from England he was unable to see the " revise " of this Report. ED.
2 II. NO H HIS WOLFEXDEX.
results of any one expedition cannot properly be appreciated without reference to the
others.
The 'Belgica' collections were made S. and S.E. of Peter I. Island, between (59 48'
and 71 18' S., and 81 19' and 92 22' W., between April 21st and December 6th, 1898,
by means of nets lowered through holes in the pack-ice to a depth of 0-500
metres.
The 'Discovery' collections were made by lowering and raising a vertically
actuated net through holes cut in the ice, while the ship was in Winter Quarters.*
The ' Gauss ' collections were made from the South of Kerguelen to the winter
station in Gauss Bay, Kaiser Wilhelm II. Land, and were of very extensive character,
and as the collections were further made throughout the Atlantic traverse of the ship,
they afford an opportunity for the comparison of the purely Antarctic fauna with that
of the Southern Ocean.
In considering the question of the distribution of the Copepoda of the southern-
most area of the Atlantic (the Antarctic region) it is convenient to consider the results
of these expeditions together, since any conclusions drawn from the results of the
' Discovery ' alone would be incomplete and even misleading. The ' Belgica ' collections
have been reported upon by Dr. Giesbrecht (" Eesultats du Voyage du S.Y. ' Belgica '
en 1897-1898-1899"; Rapports Scientifiques, 1902), and the ' Gauss ' collections are still
under examination, and I only now refer to the results of my examination of that
collection in so far as they assist the elucidation of the ' Discovery ' results.
From the results of the three expeditions (' North American,' ' Challenger ' and
' Vettor Pisaui') which, previously to the 'Belgica,' had collected in the Southern
Ocean as far south a* the pack-ice. Giesbrecht accepts seventeen species as
correct, after rejecting a number of species as "uugeniigend beschriebenen und nicht
zuverlassig geuug bestimmten "),y c/r., Ai'tuliii* iirnmtus (50 S.), Calanus jmmarchiehus
(52), Caltinus patagoniensis (47), C. propiiiquus (64 37'), simittimm (52), Centropages
Iifcie/tiiiftix (52), ClrtiiHiiciilniiu* arcuicornis (53), Clytemnestra yeuf<-tlaf2"). Caiulacin i-nrta (oO"| and truncata (64- 37'), EiiciiJiiniin
nllfiiiiiitits (47" 25'). Ei/i'liii In nun-ilia (47 25'), Haloptilus aculi-iitim (46" -lii'i. HeteroVfhabdMA xjiii>/>, (50)
Lucicutiii flaviconiis (17" 2.V). Pli'iii-fiiniiiiiiiii iilxlominale it'i.'i 12'). Cojiilin &tylif,-rn ifii; '20 i, (
Belgica ' report, p. ">.
COPEPODA.
(49), Oithona >/////'//'< (52), Paracalanus //<'////- (52),
nasutw (52), R. giyi* (65 42'), Scoledthrix minor (46 46').
This list contains a .striking number of forms which are usually associated with
more temperate regions, and, as Dr. Giesbrecht remarks, the failure in agreement with
the pelagic species of the ' Belgica ' is very striking, for only two species are common
to all collections. Comparing it with the results of the ' Discovery ' the same
extraordinary differences are manifest, only four species ((.'. yy/-<.y ) /'//y//.-- ) ('. >/'/,/, '//;/////>,
Glausocalanus <'/r///V/v//X <.)itln>nr/'tin/u>i* and C.
xiniillinuix, Clausocalanus , A<>tiu* iinim/u*, Lii*. I'ml, url,/t -/////.-, and in Lat. 49 40' 8., and
Long. 172 18' 30" W., Pleuromamma . \\hich belong undoubtedly to a subtropical or
warm temperate area, and are to lie regarded as accidental.
VOL. IV. K
4 R. NORRIS WOLFENDEN.
While the number of species captured was in each case comparatively small, the
number of individuals in any one haul in the case of the ' Discovery ' was very great.
The rule w y hich appears to hold good for tow-netting in the north part of the North
Atlantic, viz., that the further north we go the smaller the number of species, but the
immensely increased preponderance of individuals of certain species, certainly holds
good as regards the South Polar regions. Immense numbers of the small copepod
Ctenocalanus vanus appear in some of the hauls, to the almost entire exclusion of
any other species, and in other cases the larger copepod Euchceta antarctica appears
in great preponderance. Calanus acutus and, to a lesser extent, Calanus propinquus
also preponderate largely. Similarly, Metridia gerlachei appears in most of the
captures.
The collections of the ' Gauss ' provide information which is not given by those of
the ' Discovery ' or of the ' Belgica,' namely, that several species which appear
in the Southern Polar Sea also occur in the deeper water of the Atlantic Ocean
to the northwards of the Antarctic area. But as this properly belongs to the report
of the ' Gauss ' collections which I have in hand, I forbear its discussion in
this place.
Two questions are suggested by Dr. Giesbrecht in his ' Belgica' report, viz., (l)
Does the Antarctic area possess a peculiar fauna ? (2) Is the small agreement of the
Antarctic copepod fauna with that of the nearest seas due to defective research, or is it
that the area of the pack-ice has its own peculiar fauna ? and the further questions as
to whether the admixture of Polar and Antarctic fauna occurs in the deep ocean, or
whether there are physical and bibgenetic conditions in the Polar regions which differ
from those in the warm seas and prevent such exchange of species, receive some
elucidation from the collections of the ' Gauss.'
With regard to the first question, viz., Does the Antarctic area possess its own
peculiar fauna ? it must be remarked that from the results of the three collections
named the typical copepod fauna (pelagic) of this region consists in the following :
Calanus acutus Oncca curvata, similis, frigida, notopus,
simillimus conifer a,
propinquus Scolecithrix glacialis
Rhincalanus grandis Oithona similis
Euchoeta antarctica ,, frigida
austrina Gaetanus antarcticus
similis Haloptilus ocellatus
Ctenocalanus vanus Paralabidocera hodgsoni
Heterorrhabdiis austrinus Stephus longipes
Eucldrella magna antarcticum
Spinocalanus antarcticus Ectinosoma antarcticum
Metridia gerlachei Microcalanus pusillus
COPEPODA. 5
1. Of the Antarctic Copepoda the following are new species and genera :
Paralabidocera hodgsoni Euchirelli nui/jna
HdloptiluS wi'llut/ix J'"unitl/n/fcckl and 3f. ///>,v//.v).
Oncea curvata ,, 0. subtilis, Giesb.
Rltiiiculnnus grandis ,, /?. gigu*, Brady.
Harpacticus furcifer IT. flexu*, Brady.
3. Of species which occur in the North Polar regions there are only the following
' Discovery ' species, which bear such slight modification as to be practically identical :
Microcalanus pusillus ( = Pscudocalanus pygmseus) ; Oith<>nj(* ; and in
the 'Gauss' collection, Oncea cunlfera, Gaidius tenuispinus and brevispinus, and
Amaltpphora iinii/nn, that is, seven species of a total of 55-60 species occurring in the
South Polar seas, are all that are identical with the species described by Prof. Sars as
collected by Nansen's Norwegian North Polar Expedition. It would not, however, be
safe to take this list of Prof. Sars' as the ultimate result of copepod research of the
North Polar seas, and other species may yet be found to be identical.
The following table shows the comparative relationship of species of the more
frequently occurring genera :
N. Polar. S. Polar.
Citltnttis fitimarr/iicus replaced by <'. /roiiin\.
Eurluetn fintitrctira.
sii/iilix.
Metridia gerlacltei.
princeps.
di Ian us aattits.
tonsil*.
similliunix.
rn mix.
Oithona similix.
Harpa<-ti<-iixfnr,-ifii-.
Microedlanus pusillus.
tSttpli nx lo/iffijies.
1 antarcticus.
Paralabidocfra hodgsoni.
Glausocalanus arcuicornis.
Faroella anliir<-li<-iNc,>rrs.
(Plate I., figs. 1,2,3, 3", 4.)
Calanus propingwa, Brady, Rep. ' Challenger ' XIX., Copepoda (1883), p. 34.
Giesbrecht, Fauna u. Fl. Neap. XIX. (1892), p. 91.
,, Giesbrecht und Schmeil, Das Tierreich, Copi-poda (18!)8), p. 15.
Giesbreclit, ' Belgica ' Report, p. 16.
T. Scott, Trans. Linn. Soc. VI. (1800), p. 25.
The length of this copepod given by Brady is 5 * 5 mm., by Giesbrecht 4 9-5 3 mm.
The majority of the ' Discovery ' and ' Gauss ' specimens measure under 5 mm. The
cephalothorax is rather over three times as long as the abdomen, furcal segments twice
as long as broad. Head evenly rounded, without any trace of crest. Head separate
from first thoracic segment. Last thoracic segment laterally produced a little, and
ending in short points. Anterior antennae not reaching beyond the furca (thus shorter
than described by Giesbrecht, in these specimens). Relative proportions of 24th to 25th
segments as 18 : 21.
Second feet Be 3 divided into proximal portion 24 ; distal 25.
Third 20; 23.
Fourth 36 : 20.
Ri 3 with seven bristles, end saw two-thirds the length of Re 3. Fifth feet, B 1
with inner margin convex below, concave distally, with thirteen or fourteen teeth, and
distally with three teeth, larger than the others. Ri 3 with five bristles, two outer,
two apical, one inner. This animal is much more robust than the next species
(simillimus).
The is about the same size as the 9 , and the chief differences consist in the
structure of the fifth feet and shape of the head and thorax. The latter is very like C.
finmarchicus, the head oval, rounded, and produced, the first thoracic segment with deep
indentation between it and the second, and its posterior margin protruded. The
anterior antenna? with the basal joints much coalesced. Posterior footjaw with a long,
stout, densely-feathered dorsal bristle. Fifth left foot much prolonged, Re 1 and 2
elongated, R<" 3 very short and pyriform shape, with short distal bristle. R! only half
the length of R>', with respectively 1:1:6 short weak bristles. Right foot Ri nearly
as long as Re, with 1:1:6 bristles. Re not much more than half as long as Re of
opposite side. First basal with inner margin armed with seventeen to twenty large
teeth ; in its upper part slightly concave, in its lower part slightly convex, the teeth
all of pretty much the same size.
10 R. NOKRIS WOLFENDEN.
2. CALANUS ACUTUS.
(Plate L, figs. 9, 10.)
Calanus acutus, Giesbrecht, ' Belgica ' Report,* p. 17.
This copepod was first described by Giesbrecht from the Belgian South Polar
Expedition, and in the ' Discovery ' collection it forms the chief constituent of the South
Polar copepod plankton, along with Euchceta antarctica.
The majority of the animals I have met with both in the ' Gauss ' and ' Discovery '
collections are smaller than the size given by Giesbrecht, viz., 4 '7-5* 3 mm. But few
of our specimens reach 5 mm. length, the majority being from 4 '5-4 '7 mm. The
cephalo thorax is not quite four times as long as the abdomen, the head divided from
the first thoracic segment, the last segment of the latter produced laterally, but with
evenly rounded margins and no points. In lateral aspect the head is slightly
produced forwards, and more inclined to be oval than rounded. In the dorsal aspect it
has a distinct triangular appearance, with slight crest in the mid-line. In its broadest
part the thorax is 1 15 mm. broad, that is, three times as long as broad.
The furcal segments are a very little longer than the anal, and nearly twice as long-
as broad. The anterior antennae vary in length in different animals, in some being only
as long as the furca, in others one or two joints longer, and are distinguished by the
comparative length of the last joint, which is about twice as long as the one before it.
In the second feet the Re is divided into two portions, proximal = 24 ; distal = 13.
third ,, ,, ,, = 29 ; ,, =15.
,, fourth ,, ,, = 15 ; = 15.
The jRi'3 has in the second and third feet eight bristles, in the fourth only seven,
and in the fifth only four (with no outer marginal bristle). The end saw of the Re 3 of
the second feet is shorter than the Re 3 ; in the third and fourth pair longer.
The fifth feet are distinguished by the absence of the outer marginal bristle of the
Ri 3, and the total absence of teeth or hairs on the inner margin of the first basal. All
males appeared to be immature.
3. CALANUS SIMILLIMUS.
(Plate L, figs. 5, 6.)
Calanus simillimus, Giesbrecht, ' Belgica' Report, p. 17.
$ 2 5-2 9 mm. ; cephalothorax, 1 9 mm. ; abdomen, G mm. long.
Head separate from first thoracic segment, evenly rounded, without any trace of
crest. Head not quite as long as the rest of the thorax (as 18:21). Last thoracic
* " Belgica Report " is throughout this monograph used to indicate " Eesultats du Voyage du S. Y. ' Belgica '
en 1897-1898-1899." Kapports Scientitiques. 1902.
COPEPODA. 11
segment laterally produced somewhat and ending in short points. Anterior antennas
about as long as the furca, or about one joint longer.
Genital segment as long as the next two. Furca three times as long as broad, and
longer than the anal segment.
Second feet, Re 3 divided by the marginal spine into two about equal parts.
Third feet, Ri 3 with eight bristles (four outer, two apical, two inner), J'< :\
divided into two parts, of which proximal : distal = 17:13.
Fourth feet, Re 3 divided by the marginal spine, proximal : distal = 20 : 11.
Ri 3 with seven Si (two outer, two apical, three inner), terminal saw of Re 3 only
three-quarters as long as Re 3.
Fifth feet B 1 toothed, with fourteen teeth on the rather convex margin, and at
the distal end a slight break in the continuity, with three rather larger teeth somewhat-
hidden, in front view, by the upper teeth of the marginal surface. Ri 3 with five bristles
(two inner, thin and short, two apical, and three outer). In the second pair the Ri
reaches about the end of the He 2, in the third pair to the first inner marginal bristle
of the exopodite, and in the fourth pair as far as the second inner marginal bristle, in
tin 1 fifth pair beyond the origin of the first inner marginal bristle. The endopodites
are therefore proportionately larger than in C. tonsus, and the third segment of the
exopodite is not four times as long as broad.
While this species agrees with C. pn>/>////f//>i\ in many particulars, the proportions
of the third and fourth feet differ, also the toothing and convex margins of the basals
o O
of the fifth feet, and the size. Many of the examples were quite adult females with
sperrnatophore attached, so there can be no question of their being merely undeveloped
examples of C.propinquus,KO.di,&& before mentioned, this species has a considerable area
of distribution in the southern oceans.
4. CALANUS TONSUS.
(Plate I., figs. 7, 8.)
I'lihinmt tinisiis, Brady, Rep. XIX., '('hull.' Report, p. :;i.
Scott, Tr. Linn. Soc. VI. (1803), p. 25.
Giesbrecht, Famui u. Flora Neap. XIX., p. !>:.'.
Diihl. Verb. Dcutsches Zool. (irsdls. IV. (1894), p. 11.
Brady's original description of this species is very incomplete, and he merely states
that it is "like C. finmarchicus and j>r/>ii/tftitix, except that the anterior antennae are
almost devoid of setae, except on the three apical joints; the posterior antenna- are like
those in C. propinquus, the fifth pair without basal teeth, and the first abdominal
segment large and tumid. The anterior antennae are as long as the body ....
9 Size 3' 6 mm." He gives only two figures r!:., of the anterior antennae and the
abdomen.
Giesbrecht includes it under the " I'nbestimmbare species," remarking that the
VOL. IV. I,
1:2 R. NORRIS WOLFENDEN.
first character probably results from the bad preservation of the animals ; the second is
a character of other members of this genus ; and the third point is characteristic of
gracilis and rdbustior.
Scott merely mentions that " the large and tumid first abdominal segment seems
to be a fairly good character " (loc. cit.). However, this is not a characteristic of this
species any more than of C. rdbustior, and the existence of this species up to the present
time therefore must be regarded as extremely doubtful. However, the ' Discovery '
collections contain several examples of a copepod, which, if it is not Brady's species,
answers fairly well to it so far as his description goes.
9 3 '5-3 '6 mm. long (cephalothorax, 2' 75; abdomen, '75. Body broadest at the
end of the first thoracic segment ( 1 1 mm. broad). Abdomen short, genital segment
broad, and one-third broader than the following segment. Furcal segments not quite
twice as long as broad, and nearly twice as long as the anal segment. Head evenly
rounded, without trace of crest, separate from the first thoracic segment, last thoracic
segment only slightly produced, and with rounded margins. Anterior antennae only
reaching the end of the third abdominal segment, the only long bristles on the twenty-
third, twenty-fourth, and twenty-fifth segments, the twenty-fourth joint twice as long as
the twenty -fifth. Mouth parts resembling C. finmarchiws.
Second feet second basal with four large spines on the distal margin at the inner
side. Re 3 divided by the external outer spine into proximal and distal portions
respectively as 23:16. Ri 3 with eight bristles. The whole endopodite does not
reach beyond the distal margin of Re 2. Re 3 as large as Re 1 + Re 2.
Third feet four large spines on B 2 distal inner margin, endopodite reaching a
little beyond the distal margin of Re 2. Re 3 divided into proximal part = 32, distal
portion = 16. Ri 3 with eight bristles.
Fourth feet Re 3 divided into proximal part = 37, distal = 15 ; apical saw only
seven-ninths as long as Re 3. Ri with seven bristles only (three inner, two outer, two
apical). B 2 with one or two spines on distal inner margin.
On the second, third, fourth and fifth feet the outer margin of the second basal is
distally armed with a spine; in second fourth, the lie 3 = lie I + 2, and is about
three times as long as broad.
Fifth feet first basal with straight inner margin without teeth or hairs, B 2 with
five spines on the distal inner surface. .K/ 3 with six bristles (two inner, two outer,
two apical).
The only Calanus with which this shows agreement is, possibly, Brady's C. tonsus ;
but Brady's description is so fragmentary that it may well be another species. It
occurred in some numbers at Station, 22 -11 -01, Lat. 56 31' S., Long. 156 19' 30".
Such males as were observed were all immature.
13
RHINCALANUS (DANA).
(Plate II., fig. G.)
Rhine, gramlis, Giesbrecht, 'Belgica' Rup., p. 18.
? Rh. ffiffas, Brady, ' Challenger ' Rep. XIX., p. 42.
Scott, 19th Rep. Scotch Fishery Board (l'.)"l), p. 237.
Giesbrecht, Fauna u. Fl. Neap. XIX. (1 *'.>:.'), p. 153.
Hli. i/tijas was described l>y Brady as 'distributed over a very wide area between
lmg. 53 32' W. 130 52' E. and lat. 36 44' S 65 42' S. Much doubt has been
expressed by Giesbrecht as to the validity of tin's species, and the figures given by
Brady of abdomen and of the whole animal are those, in Giesbrecht's opinion, of
immature animals, and this author thinks that Brady's figure of the first feet is really
of one of the other pairs of feet.
Scott's specimens (Fair Isle and Firth of Forth) are regarded by Giesbrecht as
Rh. nasutus (Tli. .'! and 4 with dorsal or with a lateral spine, as in nasutus, and a pair of
small dorsal points on the genital segment). Mobius's specimen from the north of
Scotland is also identical with tinsutus. Rh. na-mtus is very common in the Faroe
Channel and seas off the north of Scotland, and occurs abundantly in my collections
made in these regions and along the Atlantic trough, west of Ireland, and also
appears in the ' Gauss ' collections as far south as lat. 20 N., while in the same
collections Rh. grandis (Giesbrecht) appeared. From the remarks of Sars in
"Crustacea of Norway," Vol. IV., p. 15, it might be inferred that Rh. nasutus is of
rare occurrence in the Northern Ocean (" two specimens were taken east of Iceland,
one specimen by Hjort between Scotland and Norway, and it has not yet been found
in the immediate vicinity of the Norwegian coast.") However I have taken it in
abundance on many occasions throughout the Faroe Channel. It is rather important
to establish the identity of Brady's Rh. //'//'-, and of two preserved specimens at the
British Museum, which I have examined, one measured 5'8 mm. and another 6'0 mm.
Both were immature females with four-jointed abdomen, lateral spines on Th. 3
(small), and on Th. 4 (large), with none on the fifth segment, resembling Rli. ijnDul/*,
one dorsal spine on the first abdominal segment (no dorsal spines on the thoracic
segments), and so far as could be seen without dissection, the first feet had an
exopodite of two segments only, and the fifth pair consisted each of only one ramus
of three segments. These two animals were, of course, very much smaller than
described by Brady (8'5-10 mm.) and were undoubtedly immature, and the species
may well be identical with young /t'//. i/nimUs (Giesbrecht).
L 2
14 R. NORMS WOLFENDEN.
RHINCALANUS GRANDIS.
Rhincalanus grandis, Giesbrecht, 'Belgica' Rep., p. IS.
? 7'2-S'O mm. Head produced in front, dorsally roughly triangular in shape,
with large lateral swellings at the base, rostrum not visible from behind. The cephalo-
thorax is over six times as long as the abdomen, which is composed of three segments.
A pair of short spines on the anterior margin of the third thoracic segment, and a pair
of strong and longer spines on the fourth segment, differentiate this species from naxitfu*,
also the absence of any spines on the abdominal segments. The first pair of feet have
the Ri and Re of only two segments, Re three with two marginal spines ; other feet
(except the fifth) have three-jointed rami.
The fifth pair, of one branch only on each side, with three segments, have on the
second segment a long inner marginal bristle, and on the last segment three bristles of
nearly equal length, two apical, of which the outer is the thickest and the middle one a
little the longest, and one on the inner distal margin. A short spine is present on the
outer margin in its upper third. The anterior antennae are about six joints longer
than the furca. Adult males were absent.
These examples are absolutely identical with Giesbrecht's species.
METRIDIA (BOECK).
One of the most remarkable things about Brady's ' Challenger ' Copepoda is the
omission of mention of any example of this genus from his report. Distributed
throughout the Atlantic from the North to the South Pole, and in the Pacific, and
throughout the track followed in the Atlantic and Southern Ocean by the ' Challenger,'
the absence of mention of any species of this genus is certainly extraordinary. In the
northernmost regions Mftridia lon/ja occurs (Sars, Norwegian North Polar Expedition)
throughout the Faroe Channel and the Atlantic trough as far south as Valeutia in
Ireland ; and south of the Wyville Thompson ridge, M. lucens, normani and curticauda
(Wolfenden) ; while south of Lat. 40 and throughout the Atlantic occur M. curticauda,
brevicauda, princeps and veiiuxta ; but south of Kerguelen appears a new and charac-
teristic species, M. gerlachei, which replaces all others. This is the representative
species of the South Polar seas, and it appears abundantly in the ' Gauss,' ' Discovery,'
and ' Belgica ' collections, and it is as characteristic of this area as M. longa and lucens
are of the northern cold area. M. princeps occurs seldom, and M. brevicauda as a
straggler, outside its proper area of distribution.
COPEPODA. 15
METRIDIA GERLACHEI.
Metridia gerlachei, < < iesbreekt, 'Belgiea' Report, p. -2~.
9 3 '5-3 "8 mm., very variable iii size, occasionally a little larger and often smaller.
but the average of size of examples in the 'Discovery' and 'Gauss' collections is
rather less than that given by Giesbrecht for the ' Belgica ' specimens. The cephalo-
thorax is one and a half times as long as the abdomen, head separate from h'rst thoracic
segment, last segment with rounded margins. The abdomen has the proportional
length of its three segment ~ as 9 (genital) : fi : 4 (anal), and tin- i'urca i- one-lift h longer
than the anal, and thivr limes as long as bmad. It is divided into two portions by the
outer marginal bristle, <>f which the proximal is twice the length of the distal.
The shape of the head and thorax is in this species characteristic, the back being
extremely gibbous, and the head with very bold curve, which makes it easilv recov.ni.~-
able at sight from any other species of this genus. The anterior antenna' are com-
paratively short and do not reach beyond the genital openings. The first and second
segments are coalesced ; the eighth, ninth, tenth, eleventh, have only faint indications
of separation ; the thirteenth and fourteenth joints are not so clearly divided as the
others. There are strong teeth on one, two, three, rive, seven (one cadi), those of
the third, fifth, seventh joints the strongest, and directed straight forwards. The
sesthetasks are numerous.
The endopodites of the second pair of feet have the usual excavation and hook
process on the first segment, but in this species the inner hook is exceedingly strong.
In the fourth pair the end saw is only two-fifths of the whole length of the fie 3
(shorter than in Giesbrecht's examples).
The fifth pair consists on each side of three segments, but the distal segment is
more or less completely divided into two, the division however is not complete. The
two basal joints are of about the same length and breadth, each as broad as a little
over half the length. The third joint, however, is not more than four-fifths as long as
the basals, and only half the breadth. The second joint bears one short distal bristle,
the last joint one outer marginal bristle in the proximal half, and three distal bristles,
of which the innermost is the longest and thickest, the outermost the shorte.-t.
MKTKIUIA PRIXCKPS.
(Plate III., figs. 3, 4, 5.)
Sfetridia princeps, Giesbrecht, Atti Line. Ki/ml.. SIT. I. v. ."i. p. i'i.
Fauua 11. !'!. Xrap. XIX.. p. ::in.
ravran, Ann. l!r|>. Fir-h. Iivlaml I'.IOL' OS, PI. II. \[>\<. II. < l'J05).
ilii/ nitii-rtirii, Sal's, Bull, iln Mils. Ocuimog. Monaco, I'.HI.'P, no. In, p. 7.
Though only one example of this species occurred in the 'Discovery' collection, it
was fre<|iient in the 'Gauss' collection, not only at several Atlantic stations, but also
1G K. NOERIS WOLFENDEN.
at the southernmost stations. Northwards it ranges to the west coast of Ireland, and,
as has been mentioned, has an extreme southern distribution. Giesbrecht's description
was given from only one specimen, and compared with Bars', very briefly described,
M. maenu'ii.
M. princeps, Giesbrecht. M. macrura, Sars.
Cephalothorax one and a half times as long as the
abdomen
Anterior antennas extend beyond the furca
Short teeth on 1, 2, 4, 5, and C, the 2nd the
Tail about as long as the anterior division
Longer than the body
Only feeble traces of the, strong teeth of princpps
longest
Genital segment longer than both the following
Anal segment half as long as the preceding one
Furca twice as long as anal, and 5 times as long as About as long as the two preceding segments
broad
5th feet like priitcrjis, but less unequal
Size 8' 5 mm.
Size 10 '50 mm.
Of the many examples which have come under my notice I find that the relative
sizes of the abdominal segments and furca are subject to some variation, thus :
1. G.S. 30. Ab. 2, 16. Anal 6. Furca 28 long, 3 broad. Size 8 '15 mm.
2. ,, 28. 16. 8. 25 ,, 3 ,,8 mm.
3. 27. 16. 7. 23 3 ,,8 mm.
4. 22. 11. 4. 12 2 6mm.
The genital segment is thus always twice as long as the two succeeding segments,
the anal is not more than half the preceding segment, the furca is generally longer
than the two preceding segments, and usually from seven to nine times longer than
broad. The teeth on the antennae are weak, and entirely resemble the figure given by
Giesbrecht in Plate 33, fig. 3 (op. cit.).
It is difficult to resist the conclusion that these are one and the same species, and
not two different species. The male was not described by either Giesbrecht or Sars,
but I have met with several examples.
? . The largest adults were from 8-9 mm. in length.
The body is very transparent, the head evenly rounded, cephalothorax (4'15-
4 '5 mm. long, abdomen 4'0 mm. long), only slightly longer than the whole abdomen,
and a little over one- third as long as broad.
The genital segment is larger than the next two, the anal not more than half
o o o '
as long as the preceding, often much more than the combined length of the two
preceding segments and 6-9 times as long as broad, and divided into two portions
by the marginal bristle, of which the proximal is to the distal as 8 : 5. The right
furcal segment is sometimes a little longer than the left. Anterior antennte at least
three joints longer than the furca, the basal joints broad, with short teeth on the
basal seven joints, of which those on the first two joints are the largest. The
17
distal joints taper and arc very slender. Proportional length of joints of anterior
antennae :
1&2 [ 3 | 4 | 5 | 6 | 7 | 8&9 | 10 | 11 | 12 | 13 | 14 [ 15 | 16 | 17 | 18 | 19 [ 20 [ 21 [ 22 | 23 | 24 | 25
1-2 | 4 | 5 | 5 | 5 | 5 | 14 | G j 9 | 9 | 12 | 12 | 13 | 13 | 14 | 14 | 15 | 10 I 9 | 10 I 10 j 7 j 3
The eighth and ninth joints arc quite coalesced, but in some there is a weak line of
division.
The second pair of feet have each the characteristic hooks on fill, and the
outer one is the longest. The surface of the second basal is beset with short spines,
but not the Re 1. The third feet are normal and with shortened end saw. The fifth
pair each consist of four segments, of which the basal is greater than the second,
this longer than the third, and third longer than the fourth and terminal segment.
The first joint has on its ^surface a considerable bunch of long stiff hairs (as in
princeps), the second joint has a long stiff feathered bristle on the outer distal margin,
and the third joint has a short upright spine on the outer distal margin, in all specimens
(not on the inner side as figured by Giesbrecht), and the end joint has three rather
long fine bristles, of which the innermost is the longest. The spine on the third joint
was in one example replaced by two very short spinules on the right foot, while none
were present on the left side.
$ 5 8-G mm. long (cephalothorax 3 '25 mm. Abdomen 2' 3 mm. long).
Relative lengths of the abdominal segments =14, 10, 10, 4, and the furcal segments
13. The left furcal segment is a little the largest and thickest, and six times as long
as broad, and three times as long as the short anal segment.
The anterior anteniuu extend for about three joints beyond the furca, as in the
female, and the left one is a clasping organ with weak joint between the seventeenth
and eighteenth segments. The segment beyond the elbow is very long and thin, and
as long as the next two distal joints. There are four joints beyond the elbow. The
conjoined first and second (basal) joints have two strong teeth, the distal one the
largest and curved slightly forwards. The fourth joint has a smaller tooth. Fifth feet :
The right foot with very long first joint, the second short, but with very strong, broad-
based curved and long hook, the third joint nearly twice as long as the second, and tin-
fourth and end joint a long simple spoon-process twice as long as the third. In
the left foot the first joint is very small, the second nearly twice as long, the third
a short joint, the fourth a very long simple curved spoon-shaped process. On the
inner margin (proximal) of the fourth, of the third, and the distal foot of the second
arc fine hairs. Both feet are of nearly similar length.
EUCHAETA (PHILIPPI).
Two representatives of this genus appear in the ' Discovery ' collections
E. antarctica, and another which appears to have constant differences, and to which I
have attached the name E. .*///////*. 1 do not in this collection find any example of
18 R. NORRIS WOLFENDEN.
Giesbrecht's species E . amtrina, though I have found it in the ' Gauss ' collections.
E. antarctica appears in many stages, extensive captures consisted wholly of
immature specimens, but there are many adult examples. Many males appear amongst
these, and while the females are very distinctively different, I arn not able to discrimin-
ate between those males, as to which definitely belong to aiiturctica, and others
which might belong to sim'di*. Both species are very closely allied, and differ
very considerably from the large species of the northern cold seas, viz., norireijicn,
glacial! s and l>arl>ata.
EUCHAETA ANTARCTICA.
(Plate IV., figs. 5, fi.)
Ein-litcta antarctica, GiesbrccLt, 'Belgica' Report, p. 21.
This is one of the most abundant copepods in the ' Discovery ' collection, appearing
in all stages of growth, and in some samples almost to the exclusion of other species.
Size of adult examples 7 6 mm.-8 mm. Head evenly rounded, without frontal
prominence and with short rostrum directed forwards. Last thoracic segment with
rounded margins, produced forwards and with a bunch of hairs on each side. The
abdominal segments have the postero-distal margins armed with rather strong bluntly
conical and striated teeth, and the two middle segments have on the ventral side
bunches of long hairs. The furca and bristles are the same as in E. similis. The
genital protuberance occupies the lower half of the segment, its upper margin is not
deeply concave like similis, but the whole swelling is directed downwards, and its upper
margin is slightly convex. Above the genital swelling is a secondary prominence,
which in the ventral aspect is seen to consist of two valve-like chitin thickenings. The
lower part of the protuberance has two lateral lobes, the upper are small, and above this
a prominent horn directed straight forwards and never absent in adult specimens of
E. antarctica, making it quite characteristic of the species.
In the ventral aspect the appearance is quite different from that of similis. The
genital opening is oval, almost round, with lateral cushions, and above the upper edge
of the genital opening guarded by a chitin ridge, is the horn.
The whole swelling is quite symmetrical, rather conical, and occupies quite the
lower part of the segment.
The first pair of feet have the outer margin very concave above and very convex
below, with a bunch of hairs on the Re 1 , and a long seta. The seta of Re 2 is also
very long and thin. The Re 3 is only half the length of the coalesced Re 1 and 2.
In the second pair the Re 1 has a very short Se, that of Re 2 is very long and
more than twice as long as the Se 1 of Re 3.
In Re 3, the Se 1 does not reach the origin of Se2, the Se2 does not quite reach
the end of the segment, and is three times as long as Se 3 and twice as long as Sel.
The Se 1 and 2 are very greatly curved and almost sickle-shaped.
COPEPODA. 19
In the fourth pair the /?c 3 is not three times as long as broad (1G : G). The
anterior antennas are a little longer than the cephalothorax.
The (? is a little smaller than the 9, and presents the same sexual differences as
in other Euehaetae. The bunches of hairs on the last thoracic segment, so prominent in
the 9, are absent in the .
The first feet have a three-jointed exopodite, the outer margin of which is not so
concavo-convex as in the 9 , and its Si' are short,
In the second pair, the Se of the exopodite are also smaller, the /v of Re 2 only
reaching the origin of the Si 1 1 of Re 3 ; the Se2 of Re.% being little more than half
the length of tlie distal part of the segment. The fifth feet are characteristic. The
penultimate segment of the left foot is prolonged on the upper margin into a strongly
toothed process, and has a setose conical uuhaired process on the distal margin, the
last segment into a long process, narrow and with a strong bunch of hairs at the distal
extremity, and with a large conical and strongly haired process. (This process is
sometimes nearly as long as the principal process of the penultimate joint.)
The first basal is short, the second basal long, and with very small and rudimentary
endopodite.
The right foot has short first basal, very broad second basal, long first and second
Tic (which are coalesced), and with the last segment blunt and rounded.
EUCHARTA SIMILIS.
(Plate IV., figs. 1, 2, 3, 4.)
This species occurs plentifully in the same samples in which E. antarctica is
present. For a long time I regarded them as merely different stages in the history of
the same animal, but the careful examination of a great number of individuals from
different tow-nettings proves the constancy of the points of difference between the two.
and as many of the E. antarctica and E. xiuiilix have spermatophorcs or egg sacs
attached, I have come to the conclusion that, though so very similar in most characters,
the two species must be separated on account of the invariably different characters of
the abdomen and genital segment.
E. similis is constantly rather larger than E. anfit/rfii'ii, 8'G mm. -8 '8 mm., and
more robust, the head flat and rostrum small, but strong and directed forwards and
rather upwards. The cephalothorax is two and a half times as long as the abdomen.
The last thoracic segment is produced forwards, slightly triangular shaped, with evenly
rounded margin, no spine, but a bunch of long hairs on each side. The abdominal
segments, of which the genital is twice as long as the next, are covered with fine hairs,
nowhere with large bunches, and the posterior distal margins have only very small
teeth, not large, as in antarctica. Furca with, on each .side, a very short dorsal bristle,
the ventral accessory bristle not geniculated at the base, though bent outwards, its
length not more than about half of the two long tail bristles (next to the innermost).
VOL. IV. M
'
20 R. NORRIS WOLFEXDEN.
Genital segment with very prominent genital tubercle, occupying half the segment,
(the lower half only), directed slightly upwards in lateral aspect, with apparently three
lobes, an upper and lower, each large with small middle lobe, and without any eminence
on the upper part (of the ventral surface) of the segment as in antarctica, and also
invariably without the ventral horn which is so characteristic of antarctica. In the
ventral aspect the genital tubercle appears to be obliquely placed, directed downwards
towards the left, the vulva guarded by two prominent flaps of which the right one is
below the left one. The whole tubercle occupies more of the left than the right side of
the segment, which in the dorsal view is only slightly swollen at each side.
The first and second pairs of feet and the number of bristles on the. maxilla are the
same as in antarctica.
While the female is so distinctly different from that of E. antarctica I am unable
to find similar distinction in the males. Both kinds appear frequently to occur together
in the same sample, and all the males appear to be alike.
ONCEA (GlESBRECHT).
ONCEA CURVATA.
Onrea cvrmta, Giesbrecht, 'Belgica' Report, p. 42.
According to Giesbrecht, this species differs from 0. subtilis in the following points :
Length, O'G-0'8 mm., the body lengthened, the three segments posterior to the genital
are about as long as broad, and comparatively longer than in any other Oncea species
except subtilis (in which they are longer than broad), all three segments together are
shorter than the genital (in subtilis , the genital segment is only a little longer than the
two succeeding segments), the furca is as long or a little longer than the anal segment
(iu subtilis shorter). The posterior antennae in both kinds are similar, the maxillipedes,
however, differ ; the terminal hook which, in subtilis, is thin and unarmed, is strong
and beset with pretty long teeth on the concave side, in curvata. The swimming feet
are similar in both species, except that in curvata the proximal outer marginal bristle
on Ri 3 occurs in all four pairs, while it is absent in subtilis, and the lancet-shaped
apical bristle of the fourth foot is in curvata longer than in subtilis.
The few examples that occur in the ' Discovery ' collections are smaller than
Giesbrecht's examples, none being more than "56 mm. in total length (?s with egg sacs
attached).
The genital segment is a little longer than the next three segments, the furcal and
anal segments of the same length, the second abdominal segment as long as broad, and
rather larger than the third segment, the relative lengths of Ab. 2:3:4 and furca
being as 4 : 3 : 4 : 4, the latter nearly three times as long as broad.
In the posterior antennae, the first basal joint is the largest, and the inner margin
of the second basal has a few fine teeth ; the distal segment has three proximal bristles
COPEPODA. 21
of unequal length, the middle uiie very short, and distal to it, a comparatively
and slightly armed bristle, terminally four long and two shorter bristles.
The maxillipede is armed with a strong claw bristle, denticulated on the inner
margin, and the second basal has two comparatively stout bristles, the proximal of
which is armed with wide-apart bristles.
In the fourth pair of feet the apical bristle of the exopodite is longer than the third
exopodite segment by one-third of its length.
The agreement, therefore, between this species and Giesbrecht's examples is very
close, the only difference being one of size, and there is no doubt that the two species
are identical.
STEPHUS.*
Mobianus, Giesbrecht, Fauna u. Fl. Xcap. XIX. (18H2), p. 205.
Helios, Th. Scott, 10th Rep. Scotch Fishery Board, Vol. X. (1892), p. 2-15.
Stephus=Stepho8 i Giesbrecht, ' Belgica' Rep., p. 20.
Stej)hos, Bars, "Crustacea of Norway," Vol. IV. (1903), p. 61.
Since Giesbrecht described the genus Mi)l>innus, which was subsequently identified
with Stcphos (Scott), several other examples of the same genus have been described.
Scott himself recorded three specimens, viz., S. minor, S. fultoni, and 8. ///>.
Sars has described two new species from Norway, rlz., S. lamellatus and
Scott), which latter is again identical with Stephos gyrans Scott (not Giesbrecht).
Giesbrecht's species '(gyranx) is said by Sars to differ in the asymmetrical last thoracic
segment and genital segment, the latter with " a number of irregularly arranged
spiniform processes not found in any of the northern species," the last feet of the
male also differing from S. scotti.
The 'Belgica' report contains yet another species described by Gic-lnvcht as
Antarctic, viz., S. longipes. This species recurs also in both the 'Gauss' and
'Discovery' collections, and in the latter I find a further and considerably lar-cr
example, to which I have given the name antareticum.
As these descriptions are scattered over six different volumes, I think it may be
of service to bring them together here.
1. S. GYRANS.
S. gyrini*. Hi'-sbrecht, Fauna u. Fl. Xcap. XIX. (I*'.':.'), p. 205; Giesbrecht, u. Schrneil, Das Ticrn ich,
Copep., p. i".i.
Furca longer than broad, anterior antennas reaching to end of genital segment,
genital segment with a curved hook on ventral side, shorter hook on dorsal, fifth feet
* The author, in agreement with Sars. pri-fcn's the name originally given to the genus by Scott, but it is
perhaps better to observe the ordinary rule. En.
M 2
22 R. NORRIS WOLFENUEN.
in female, end segment rather curved and broad basally ; iu male, left foot with several
long appendages, right foot, thin appendages at end and rest foliate ; size =0'8-1 mm.
(Naples.)
2. S. SCOTTI.
8. gy rans, Scott, Nineteenth Rep. Scotch Fishery Board (1901), p. 237.
S. scotti, Sars, " Crustacea of Norway," Vol. IV., p. 63.
Slender ; cephalothorax symmetrical ; genital segment without spines, furca longer
than broad, anterior antennse reach end of abdomen 2, Re of posterior antennse twice
as long as Ri ? , fifth, with denticles on last segment, which is elongated and
pointed ; $ , fifth, penultimate joint of left foot tumefied, last segment with about half
a dozen short processes ; last joint of right foot with long, sickle-shaped process.
Size= "85-'95 mm. (Loch Fyne ; Norway.)
3. S. MINUS.
S. minor, Scott, Tenth Rep. Scotch Fishery Board, 1892, p. 245.
Robust, cephalothorax symmetrical ; genital segment without spines, anterior
antennae about as long as the thorax, furca as long as broad, fifth feet in ? with
elongated last segment with two little lateral spinules ; in $ , right foot a long foliate
joint at end, left foot with two digitiform processes at end, penultimate joint only
slightly tumefied. Size = 074 mm. (Firth of Forth.)
4. S. LAMELLATUM.
and spine, but no processes. Size '8-' c J in.
(Antarctic). ('Discovery' and 'Gauss' collections. Wolfenden.)
7. S. ANTARCTICUM.
Robust, cephalothorax a little unsymmetrical, right side prolonged; genital segment
swollen laterally, with bunch of spine-like bristles each side ; fiuva us broad as lon^;
anterior antenme reach to Ab. 2 ; Re of posterior antennae longer than Ri. Fifth feet
in 9 right side longest, each with three end spines, innermost hook-like ; in <$ ,
right with third joint elongated and club-shaped distally, with a large, roughly
triangular plate, and last joint a strong, curved hook ; left foot, no tumefied segment,
and last joint with distally a short-stalked haired knob, no appendages. Size = 175-2
mm. ('Discovery' collection.)
STKI-HUX LONGIPES.
(Plate V., iigs. 1, 2, 3.)
sir/iinis lui/i/ijH's, Giesbrecht. ' Bdgicu ' llq>., p. L'H.
9 '75-' 80 mm. <$ '65-70 mm. Cephalothorax rather more than twice as long as
the abdomen ; head separated from first thoracic segment, but all segments very
difficult to determine, owing to the indistinctness of the lines. Last thoracic segment
with rounded margins and symmetrical. Abdomen in the female four, in the male five
segments, f'urcal segments only as long as the anal, as broad as long, and with rounded
margins, each with four tail bristles and a short fifth inner marginal bristle, (ieiiital
segment in the female as long as the next two, laterally with roughly triangular
swellings, and in its greatest breadth, broader than long. Anterior antenna) not as
long as the thorax, and of twenty-three joints, the first, second, eighth and ninth
coalesced, with few bristles, the longest on the seventh and eighteenth joints, but well
supplied with long ;esthetasks.
Posterior antemuu with exopcdite about one-third longer than the endopodite.
Posterior foot-jaw with rather thick first basal and />' I : /> -2 : /i'/as S : 7 : <> ; mandibles
with broad-ended masticatory plate, one pointed outer tooth, with considerable space
between il and the middle stout, com para lively broad teefh, and three pointed inner
teeth.
24 R. NOERIS WOLFENDEN.
First feet Ri=. 1, Re= 3 ; no Se on Re 1, and very short Se on Re 2.
Second feet Ri = 2, .Re = 3.
Third and fourth feet Ri and Re ^.
Fifth, each of three segments, two basal, each short and comparatively thick,
terminal segment longer and thinner, prolonged, with a stout curved hook with short
bristles on the upper margin and an external marginal thin and short bristle.
The male is rather smaller than the female, the abdomen has five segments, the
mouth organs are as in the female, but the fifth feet are transformed into clasping
organs, that, of the right side of four segments, the left of five. The second and
third segments of the left are elongated, the distal segment short and broadened
out, the distal extremity ending in a spine, and at the opposite side a short knob
process, apparently without marginal hairs.
The right foot with short broad second basal, and two distal very elongated and
thin segments, with a long thin sickle-shaped process at the end, which appears to be
a continuation only of the joint above it, and though forming an elbow, does not
articulate.
In the general structure this small Copepod bears great resemblance to the
species Stephus antarcticum, which, however, is twice as large ; the fifth feet in the
female as well as the male are different, and the two species are therefore quite distinct.
STEPHUS ANTARCTICUM.
(Plate V., figs. 4, 5, 6, 7, 8.)
9 1'85-2'0 mm. long, cephalothorax about three times as long as abdomen, and
in its broadest part distal of the middle line one-third as broad as long. Head evenly
rounded, a little produced in front, but without trace of rostrum, a weak line indicating
its division from the first thoracic segment. Last two segments of the thorax im-
perfectly divided, and last segment a little unsymmetrical ; on the right side a little
longer than on the left, produced into a round-ended margin, on the left side more
acutely pointed, which is most marked in lateral view. Abdomen of four segments
respectively proportioned: genital segment, 2:3 and anal as 20:13:8:8; furcal
segments as long as the anal, and as broad as long. Genital segment laterally swollen
in the upper part (genital protuberances) and again slightly swollen laterally in its lower
part, with on each side a bunch of rather long spines, none dorsally. Furca with four
tail setae on each side, outer margins haired and with, on each side, a short lateral
spine instead of the usual bristle, and on the ventral side a very short accessory
bristle. Of the tail bristles, the two middle ones are much the longest and thickest,
and those of the right side more so than those of the left.
Anterior antennae reach in both sexes to about the end of the second abdominal
segment, having twenty-four segments, the eighth and ninth coalesced. In the posterior
antennae the exopodite is longer than the endopodite. The mandibles with branches
COPEPODA. 25
nearly of same length, masticatory plate with strong teeth. The posterior foot jaws
with first and second basal and Ri in proportion of 24 : 11 : 20 ; maxillae, 132 with 5 ;
Re with 10; Pi I with 4, Ri2 with 3, Ri '3 with six bristles, Li 2 and 3 present, and
Lc 1 with eight bristles.
The first feet have one jointed Hi and three jointed Re, without Se on Re 1.
The second feet have two jointed Ri and three jointed Re.
The third and fourth feet have both rami three-jointed.
Fifth feet comparatively large, that of the right side a little longer than the left.
Each of three segments, of which the two basals are equal in size, but the last segment
on the right foot is a little longer than in the opposite foot. Each has terminally
three spiny processes, the innermost comparatively thick, curved, and hook-like, and
with hairs only on the outer margin. The two outer spines are neither much more
than half the length of the inner one, and only half as thick.
The 3.
First pair of feet, B 1 and B 2 with only slightly convex inner margin and no Si.
Ri 1 longer than Re 2 + Re 3, outside margin haired and long marginal bristle. Re 2
and Re 3 with similarly long marginal bristles. Se = 1 : 1 : 2 and Si 1 : 1 : 5. Ri 2
nearly twice as long as Ri 1 and with 5 Si.
In the second pair B 2 is broader than long, the inner margin convex and without
hairs or bristles ; B I is also without Si. Ri 1 prolonged, Ri 2 shorter (as 7:10). Ri 1
with 2 Si, Ri 2 with 7 Si.
Re 1 : 2 : 3 as 1 1 : G : I), with respectively 1:1:1: Se, that of YiV I the largest and
of Re 3 the smallest; the end saw about two-thirds as long as the whole /c'< ; a small
curved spine distal margin of Re 3 just external to the saw. 5 Si on Re 3.
3rd feet. B I and Ji 2 without bristles or hairs, Ri as in the preceding pair, Re
as in the preceding pair and with 1 : 1 : 1 Se only.
4th feet. B I and B 2 as before. Ri 1 with three Si, Hi '1 with only six Si. The
three joints uf R<> rather more equal in length, Se 1 : 1 : 1 as in the other feet. Ri I is
more elongated than in the other feet and twice as long as L'i _'.
In the second to fourth pairs the Ri is mure than half as long as the Re and
extends beyond the distal margin of Re 2. In all feet the number of external marginal
spines is certainly peculiar, the usual rule being three marginal spines on the Re 3, so
that the animal differs from any true Ltibiflocem.
The 5th feet ('(insist of a common basal and one ramus on each side of two
B 2 and Re. l\i represented onlv by a spine. The first and second basal. s are
L. iv. N
28 R. NORPJS WOLFENDEN.
nearly equal in length and each about as broad as long. The second basal segment has
at its distal inner margin a very stout articulating spine, JKi four-fifths as broad as this
joint is long. On its outer surface, near the distal and outer end, is a delicate bristle.
The third segment is nearly twice as long as the second basal, tapers to a point, and just
below the distal end is a delicate little bristle. Near the end of the joint and on the
inner side is a very stout broad-based spine, not articulating, and nearly half as long as
the whole segment. Frequently the foot of one side is a little longer than of the
other.
The whole animal is very unsymmetrical, especially in the shape of the last
thoracic segment and the genital segment of the abdomen. The characters of the
swimming feet, as to proportions, and especially as to tlie absence of spines on tin' lu*t
joint of tin' f.rnpodite, and the absence of anything like the usual ocelli of Labidocera,
are points which appear to remove it from that genus. The abdomen of most females
is more or less enveloped in a mass of colourless, structureless membrane. The $ is
distinctly five-jointed in the abdomen, whereas in Labidocera this is four-jointed. The
right anterior antenna especially also differs from other species. From the characters
of the female feet and the abdomen of the males I have thought it justifiable to create
a new genus for this animal. I name the species after Mr. Hodgson, the naturalist of
the ' Discovery ' Expedition.
$ smaller than the 9 (1'6 mm). Cephalothorax with head separate from next
segment, two dark lateral spots, but not ocelli, in front of the head. Abdomen little
more than half as long as the thorax. -Head evenly rounded, without side hooks, last
thoracic segment rounded and not produced. Abdomen of undoubtedly five segments,
of which the second is about as long as the third and fourth together, the first is very
short, the fourth is twice as long as the anal, which is a short segment ; the furcal
segments, of which the right is a little larger than the left, are twice as long as broad
and three times the length of the anal segment.
Right anterior antenna a clasping organ, the middle joints swollen, the joint
before the elbow with a marginal row of small teeth and with only two distinct
segments beyond the elbow, of which the distal is very long and thin (over three times
as long as broad), and in its distal part carrying on the inner margin a very long spine
tapering to a fine whip-like extremity, but broad in its basal portion. This appendage
is half as long again as the whole joint. The joint immediately distal to the elbow has
on its margin proximally a short but thick spine. I find it very difficult in any of the
specimens, of which there are several, to agree with any degree of accuracy upon the
exact number of segments in this antenna owing to the fact of its being curled up
in every case. First to fourth feet and mouth organs as in the female.
5th feet, powerful clasping organs, the right one of four segments, the left of
three, with a common basal. The first segment of the right foot has on its inner
margin a small knob projection, the second has two short, thick spines, the third a fine
spiiii-, mid the last joint is curved into a strong hook, without any appendages.
COPEPODA. 2!>
The first joint of the left foot has a short spine 011 the external margin, arid the
last joint is broad proxiinally and foliaceous, and on its inner surface is a row of fine
bristles, with three or four stiff and longer than the rest.
EUCALANUS (DANA).
That any species of Eucalanus should be found at extreme southern latitudes is
certainly curious, E. elongatus certainly occurs south of lat. 40, and about 40 W.
long., ' Gauss' collection ; and in the 'Discovery' collection I have found about half a
dozen examples of a Eucabinu* which 1 regard as a variety of K. subtenuis or >niti:n>-
natm. This occurred at station marked 21. x. 01, lat. 57 25^' S., long. 151 f E.,
and station lat. 56 31' S., long. 156 19' 30", 22. xi. 01 ; in both cases a long
distance outside the Antarctic Circle.
The 9 (no males were found) is 4 mm. long. The head is very triangular,
elongated, and produced in front into a blunt point slightly bent downwards; there are
lateral swellings as in attenuatus, the part behind is not, however, indented. The last
thoracic segment is rounded. The abdomen has three segments, and one tail bristle on
the left side is a little thicker and longer than the rest. The genital segment is
laterally swollen and broader than long. Posterior antenna: with first and second
joints of the exopodite coalesced, the first joint of the endopodite about three times as
long as broad, and about the same length as R(2. The mandibles with very short Hi,
the proximal part about three times as long as the distal, the whole Ri very much
shorter than the distal part of the basal, and with four bristles and 'two short marginal
bristles. Maxilla, B 2 with five, Ri 1 with four, Ri 2 with four, Ri 3 with five bristles.
With some resemblance to E. subtenuis, pileatus, and mueronatii*, it is larger than anv
of them. The five bristles on the B 2 of the maxilla cause it to differ from either
inuaronatus or mhtcnuis, and it has considerable differences from pileatus in size,
posterior antenna and mandible. The shape of the head is certainly not that of
subtenitix, nor is it so triangular and pointed as
CTENOCALANUS (GIESBRECHT).
CTENOCALANUS VANI -.
Ctenocalitnua rinmx. Gksbrucht, Atti Ace. Lincci Rend., Ser. 4, 1888, p. 335.
Fuuua u. Fl. Neap. XIX. (1MH'), p. I'.M.
This is extremely abundant in the ' Discovery ' collections, but does not differ in
any material particular from the species well known in the Atlantic. Its range of
distribution is very great, extending from the Faroe Channel (Wolfeudeu) throughout
the Atlantic to the southernmost parts of the Antarctic area. /.<. to the ice region.
x 2
30 R. NORRI8 WOLFENDEN.
CLAUSOCALANUS (GIESBRECHT).
CLATJSOCALANUS ARCUICORNIS.
Clausocalanus amdcornis. Giesbrecht, Atti. Ace. Lincei Rend., Ser. 4., vol. 4, p. 334.
Giesbrecht, Fauna u. Fl. Neap. XIX. (181)2), p. 50.
Giesbrecht u. Schmeil, Das Tierreich, p. 27.
That this species should occur so far south is rather peculiar. It was found in the
collections made at :
Lat. 49 40' S. Long. 172 18' 30" W.
Lat. 55 44' S. Long. 95 43' 30" W.
Lat, 5G 12' 45" S. Long. 136 18' 30" W.
Lat. 57 25' S. Long. 151 43' E.
Lat. 58 49' 45" S. Long. 154 48' W.
Lat. 59 19' S. Long. 120 24' 30" E.
Lat. 63 5' S. Long. 175 43' E.
Lat. 84 01' S. Long. 170 49' E.
and does not differ essentially from the species common in the Atlantic. It has a
considerably greater range than was thought, since I can record it from the Irish coast
to nearly the Antarctic Circle.
GAETANUS (GIESBRECHT).
GAETANUS ANTARCTICUS.
(Plate III., fig. 6.)
G (tetanus antarcficits, Wolfenden, Plankton Studies, Part I. (1905), p. 7.
Size 8 mm. The body is very robust and dorsally very gibbous. The head and
first thoracic segment are coalesced, and together much longer than all the rest. The
last thoracic segment carries two short stout curved spines, directed backwards. The
head is in its upper part quite square, and with short stout curved spine, directed a
little forwards. The abdomen is not a quarter the length of the cephalothorax.
Anterior antennae not as long as the body, of twenty-three segments, with the
eighteenth, nineteenth and twenty-first segments longer than the twentieth, and all
joints with very few setse. Ei of the posterior antennae more than half the length of
Rt j . Posterior foot jaws with lamellar process on the first basal.
Maxillse ; Li 1 and Li 3, each with four bristles ; B 2 with five, Ri small and two-
jointed. Re small, and less than half the length of B 2.
First feet, Re of three segments with three marginal spines, the segmentation being
complete ; Ri of only one segment.
COPEPODA. 31
Second feet, Rl distinctly two-jointed.
Third and fourth feet, Ri and Re of three joints each. B 2 of the fourth feet with
tubal bristles as in Gaid'm*.
The extraordinary size of this animal makes it the largest known species of
Gaetann*. It occurred once only in the 'Discovery' collection, and also appeared in
the ' Gauss ' collection, and is probably Antarctic in its habitat Several new species of
Gaiiiiniix have been described recently, and it may serve some useful purpose to
recapitulate here the different species discovered since Giesbrecht and Schmcil's last
work (Tierreich, 1898).
1. (TARTAN us MAJOR.
G. major, Wolfe-mien, Proc. Zool. Soc., London, Feb. :!, I'.titt, in Dr. Fowler's paper.
Farran, Ann. Rep. Fish. Ireland, l!)n^-ii:;. Part II., App. II., I'.Mi.V
Size 5 mm. and over. Anterior antenme larger than the body by about one joint ;
lamellar appendage of posterior foot jaws absent ; Re of first feet of three segments,
and with three Se. Cephalic spine short, and as in G. ariii!;/> r.
2. GAETANUS CAUDANI.
i-i/mluiii, Canu, Ann. Univ. Lyon, V. :.'4, p. 24.
,, (?) pi'lci/ti/s, Favran, ihiil.
Like G. mil/'*, ljut anterior antennae only one-and-half times as long as the body :
lamella of posterior foot jaw like that of G. milt's. Rn of first feet, three segmented
basals of fourth feet with tubal bristles, 5 mm. and over. Cunu's original description
was of one immature cJ . Farran's were also immature specimens.
3. GAETANUS HOLTI.
Gftftanus holti, Farran, ibiil.
litifrons, Sars, Bull. Mus. Monaco, No. -'(I, March, 11)05.
s, Wolfenden, Plankton Studies, Part I. (1!)05), p. 7.
Cephalic spine strong- and directed backwards with long interval between the
frontal part and base of the horn. Anterior antennae not as long as the body-spines of
the last thoracic segment, strong, long, and directed backwards. Small lamella on
posterior foot jaw. First feet with three segments and three 6V. Fourth feet with
basal tubal bristles. Sixe 4 '74 mm.
4. GAETANUS ANTARCTIC! s.
Gaetunii* untaretints, Wolfenden, Plankton Studies, Part I. (1'Jn")), p. 7.
Thorax gibbous, very stout short curved cephalic spine directed forwards, head
square, not like G. armiger. Abdomen short and thick, not a quarter of whole length.
U. NORRIS WOLFENDEN.
Anterior antenna not as long as the body. Posterior foot jaw with lamella. Re of
first feet with three segments and three Se. Fourth feet with tubal bristles. Size
8 mm.
5. GAETANUS MINOR.
Gnetanus minor, Farran, Ann. Rep. Fish. Ireland, 1902-03, P. II., App. II. (1905), p. 34.
Cephalic spine as in G. armiger, long slender spines of last thoracic segment.
Anterior antennae reaching only to genital segment. First feet with Re of two
segments and only two Se. Second feet with one-jointed Re. Size 2'4 mm.
6. GAETANUS ROBUSTUS.
G (tetanus rolustus, Sars, Bull. Mus. Oceanographique Monaco, no. 26 (1905), p. 11.
Robust, cephalothorax a little swollen in middle. Cephalic spine small, curved,
or sometimes absent. Spines of last thoracic segment very strong and divergent.
Abdomen thick and about one-third of whole length. Anterior antennae scarcely as
long as the body. Size 8 mm. (? possibly the same as G. antarcticus, but the
description of G. robustus is insufficient.)
7. GAETANUS INERMIS.
Gaetanus inermis, Sars, op. cit., p. 12.
Body very thick, anterior division swollen. No trace of cephalic spine, last
segment of thorax rounded and without spines. Abdomen very short, not a quarter of
whole length. Anterior antennas not longer than body. Structure of other parts not
different from other species of this genus (?) Size 6 30 mm.
(In the absence of cephalic and thoracic spines, which are constant in this genus,
this is probably not a Gaetanus.)
8. GAETANUS CURVICORNIS.
Gaetanus curvicornis, Sars, op. cit., p. 11.
Body like G. miles (Giesbrecht), short curved cephalic spine. Spines of last thoracic
segment moderately large and divergent. Very short abdomen only a quarter the
whole length. Anterior antennae scarcely longer than the body. Size 4 '35 mm.
9. GAETANUS KRU^PPI.
Gaetanus Jcrwppi, Giesbreclit, Mitt. Zool. St. zu Neapel, XVI. (1903), p. 202.
Like G. armiger, but larger, viz., 3 '6-4 mm. long, thorax shorter. Anterior
antennae reach three joints beyond furca, twenty-second segment longer than in G.
armiger. Feet like G. milt'*. <$ 3'7 mm. long, thoracic spines shorter than ? and
antennae shorter than body. Se of Re 2 of first foot rudimentary ; fourth feet without
tubal bristles. Fifth feet Ri of one segment, Re of right foot of two ; of left, of three
segments. Mediterranean.
CUFEi'OUA. 33
X ANTHOCAL AN US (GIESBREUHT).
There are only two examples of this genus in the 'Discovery' collection. Since
the publication of Giesbrecht and Schmeil's " Copepoda " (in " Tierrcii-h ") the genus has
received many additional species. To the originally described species, viz., X. ag'dix
and AT. minor (Giesbrecht), are now added X. /j>iiKjn/i. (Sars),
A", niiitii-iix (Sars), AT crixtatus (Wolfenden), X. xnli,-rixiiitux (Wolfenden), X. tfim/ili-.r
(Wolfenden), AT. ninijinix (Wolfenden), AT: calaminus (Wolfenden), X. ntl), p. 40.
Size 2 "4 mm. Furcal rami little longer than broad, anterior antennae reach genital
segment, feet very spinulose. Fifth feet, three segmented, spinulose, and with two
terminal and two lateral spines on last segment ; second joint longest, and spiuulose on
both margins. (Atlantic. Ireland.)
12. X. PINGOIS.
X.pinguis, Farran, Ann. Rep. Fish., Ireland, 1002-03, pt. ii., App. II. (1905), p. 40.
Size 4'5 mm. Head imperfectly separated from thorax ; lateral processes of
last segment blunt ; furcal segments slightly longer than broad ; anterior antenna;
short, not as long as thorax ; rami of posterior antennae short and broad ; feet spinulose.
Fifth feet of three segments, and two lateral and two apical end spines ; surface of
third and margins of first (inner) and second (outer) spiuulose. (Atlantic. Ireland.)
13. X. GREENI. X. MUTICUS.
X.ijreeni, Farran, Ann. Rep. Fish., Ireland, 1902-03, pt. ii., App. II. (190;>), p. 40. X. muticus, Sars,
Bull. Mus. Monaco.
Size 5'30-G mm. ; last two segments of thorax united, last segment with obtusely
triangular margins ; furcal segments short, broader than long; anterior antenna; little
longer than budv. Fifth feet small, two-jointed (Sars), or three-jointed (Farran), without
spines on inner margin ; last joint with three small apical spines. (Atlantic. Ireland.)
14. X. XTPICUS.
Amalli>linr< >me time when
Professor Sars' supplement appeared with the description of u:t til !<>/>*!*. Consequently
I do not kuow to which name priority should be given, nor do I feel yet certain that
the genus described briefly by Sars is identical with the Faroella described by me in the
J. M. B. Ass. of 1904. Certainly the Faroella of the Antarctic Sea has some differences,
and I therefore retain the name for the genus which I originally gave, more especially
as Professor Sars, who has examined some Irish specimens, states, as I am informed,
that they are not identical with his.
FAROELLA ANTARCTICA.
(Plate II., figs. 1, 2, 3, 4.)
? $ size 4'3 mm. (cephalothorax 3' 3 mm., abdomen I'O mm.). The fore-bod v is
therefore over three times as long as the abdomen. The head and first thoracic segment
are united, the two last segments of the thorax separate, the anterior segment over twice
as long as the four last segments ; the most posterior of these is well defined from the
one in front, small, and laterally prolonged into stout spines which are about three-
quarters as long as the genital segment. In dorsal aspect the head is rather triangular-
shaped, and on each side below the level of the posterior antenna', laterally expanded.
In the lateral aspect the head is evenly rounded, oval, and with stout two-pointed
40 R. NOERIS WOLFENDEN.
rostrum directed forwards, with slrght curve downwards, the rostral spines not at all
divergent, as in Sars' picture of ^Etidiopsis. The whole cephalothorax is studded with
fine and closely-set prickles. Abdomen of four segments, slender, the genital only a
little larger than the next, with strong ventral protuberance, > Alt 2 > Alt 3 > Al 4.
Furcal segments as long as the anal, and twice as long as broad. Tail bristles four on
each side, with very short and delicate veutrally placed accessory bristles. Anterior
antennas reaching just beyond the end of the genital segment, the first two joints
comparatively large and as long as the next five joints, the combined eighth and ninth
joint as long as the two joints either proximal or distal to it, the eighteenth and nine-
teenth joints longer than those proximal or distal, and the twenty-fourth separate from
the twenty -fifth. All joints only sparingly setiferous.
Posterior antennae with exopodite a little longer only than the eudopodite.
Anterior foot-jaws with the outer margin of the basal only a little convex, the
fifth lobe longer than the four proximal and nearly equally-sized lobes, the Ri small
but distinctly segmented. Each lobe with three bristles, two each on the first, second,
third, and one on the fourth, being stout, long, and with wide apart stiff marginal
bristles. Bristles of Ri long, not feathered, but slightly serrated marginally.
Posterior foot jaws having proportionately B 1 : B 2 : Ri = 10 : 12 : 5. 132
therefore a little longer than B 1, and over twice as long as Ri; B 1 two and a half
times as long as broad, with two small lobes with respectively two and three short
bristles ; B 2 four times as long as broad, its marginal bristles very small and distal of
the middle. Ri short and distinctly five-segmented.
Maxillse, Le 1 with nine bristles and its outer margin nearly straight ; B 2 with
five,, and not segmented from Ri with thirteen bristles ; Le2 a small lobe, but without
bristles ; Re small, longer than broad, and with ten bristles ; Li 1 with nine large hooks
and four bristles ; Li 2 and 3 well-formed lobes.
First feet. Ri one-jointed ; Re three-jointed, with three long thin marginal
spines.
Second feet. Ri two-jointed, Ri 1 short, Ri 2 very elongated, and nearly four
times as long as Ri 1. The whole Ri only extends to the distal end of Re 2. On the
distal part of the surface of Ri 2 is a bunch of fine hairs ; Re 3 is as long as Re 1+2,
and its end saw is longer than the Re 3 and beset with a great number (exceeding
fifty) of closely-set teeth, of which those in the middle are the largest.
Fourth feet. Ri and Re of three segments each. Ri proportionately longer than
in the other feet, and the third segment as long as Ri 1 + 2, and a little over three
times as long as broad, with fine hairs on the surface distally. Re 3 much longer than
Re 1 + 2 and over three times as long as broad. Its end saw three-quarters as long
as the Re 3, and with over fifty closely-set teeth. No fifth feet.
The chief points in which this Antarctic species differs from that of the northern
seas are in its greater size, the greater strength of the rostrum, the rather different
body proportions, and the more equal size of the rami of the posterior antennae. The
COPEPODA. 41
chitiu everywhere in the ecphalothorax is almost covered with prickles, and the
whole animal is more robust.
MICROCALANUS (SARS).
MlCROCALANUS PUSILLUS.
(Plate II., fig. 5.)
Microcalanus pusillus, G. 0. Sars, Crnstac. of Norway, IT. (1903), p. ir>i;.
Pseudocalanus pygmteus, Sars, Norwegn. N. Polar Expdn., Vol. V. Crustacea, I'.HMI.
Giesbreeht, ' Belgiea' Report, p. L'I.
Sars originally described a small Calanoid, Ps&udocalanus pypmseus, from Nansen's
Polar Expedition, which he subsequently re-named Microcalanus, and included in the
new genus a second and still smaller form, of M. puxillu*. Griesbrecht described a small
(.'alanoid from the Antarctic seas, which agreed generally with Sars' Polar species, except
for very small differences, './/. the length of the anterior antennae, and the length of the
outer marginal spines of the exopodites of the feet. The size varied, mostly from
0" 7-0 "75 mm. ; some were even smaller. P. pijijmseu*, Sars (= .17! }>>/fjmceus, Sars), is
a little longer, viz. 86 mm. (Sars).
The 'Discovery' collection contains many examples of a very small Calanoid
which agrees so closely with Sars' M. ]>u*i///<), occurs very sparingly in the ' Discovery ' collection.
HARPACTICUS (DANA).
HARPACTICUS FURCIFER.
Jfiir/itn-tintsfurcifer, Giesbrechf,, ' Belgica ' Report, p. 37.
The 9 of this species was first described by Giesbrecht in the ' Belgica ' report ;
and in the 'Discovery' collection, marked 4. i. 02. W.Q., occurred three specimens of
the male, though female examples were conspicuously absent.
The 9, according to Giesbrecht, is 1'55 mm. long; the rostrum small, the
series of points on the abdominal segments not numerous, the furca as long as both last
abdominal segments, narrowing distallv and about three times as long as broad ; the
VOL. IV. P
44 R. NORRIS WOLFENDEN.
anterior antennae nine-jointed, the exopodite of the posterior antennae like H. chelifer,
but smaller, the second basal of the mandible like H. brevicornis ( If. fulvus), and the
exopodite scarcely half as long as the endopodite ; both rami of the maxillae are about
equal ; the first lobe of the anterior foot jaws has three bristles, the fourth lobe is long,
and its hook short, the posterior foot jaw is much thinner and weaker than in H. chelifer
and IT. brevicornis and more like PL flexus. The first feet have thin and weak terminal
claws, both rami of only two segments, and the endopodite is short, the joints of both
branches being broader than in flexus ; the endopodites of the second and fourth feet
are larger in proportion to the exopodites than in chelifer and brevicornis, and in the
fourth pair reach to the middle of the last joint of the exopodite ; the bristles on
the second endopodite joint are, however, two, instead of one as in chelifer ; the last
joint of the fifth feet is comparatively small, and is scarcely half so broad as this ;
its last joint has five, the process of the basal joint, four bristles.
The striking feature of H. furcifer is the length of the furcal segments, which are
usually very short in this genus, and though related to PL flexus, it differs in the size,
which, in the latter species, is only '64 mm. in length, compared with 1'5 in
PL furcifer.
. \UUirllr
|-\|,.
Mai iun
up. ill. l jil. I.
f'al.mus propiri'i
simillim
] itis i
tU i '. 1O)
Antarctic (Discovery) E\p.
Marlon
jiuila pi. II.
Faroella amaivtioa (1-4). Microcalanus pusillus (5).
Khinocalamis yramlis (6).
COPEPODA.
45
EXPLANATION OF THE PLATES.
PLATE I.
C itl'in us jiriipim/Kim, h'g. 1. Whole animal, 9- Oc. 3, obj. 2 in.
fig. 2. Whole animal, $ . Oc. 3, obj. 2 in.
fig. 3 and 3a, $ . 5th pair of feet. Oc. 3, obj. i in.
fig. 4. 9- Basal joint of 5th foot. Oc. 3, obj. i in.
fig. 5. Whole animal, 9 Oc. 3, obj. 2 in.
fig. (1. Basal joint of 5th foot. Oc. 3, obj. \ in.
fig. 7. Whole animal. 9 Oc. 3, obj. 1 in.
fig. 8. Basal joints of 5th foot, 9 Oc. 3, obj. \ in.
fig. 9. Whole animal, 9 Oc. 3, obj. 2 in.
fig. 10. Basal joints of 5th foot, 9 . Oc. 3, obj. \ in.
tonsus,
acutus,
PLATE II.
initiiriiirii, fig. 1 ,
fig- 2.
% 3.
fig- -t.
, fig. 5.
Rhincalanus grandis, fig. 6.
9 , whole animal. Oc. 3, obj. 2 in.
9 , Posterior foot-ja\v. Oc. 3, obj. 1 in.
9 , 1st foot. Oc. 3, obj. 1 in.
9 , 2nd foot. Oc. 3, obj. 1 in.
9 , whole animal. Oc. 3, obj. i in.
9 , whole animal. Oc. 3, obj. 2 in.
Mftri/lin /'//// ''r/is,
PLATE III.
fig. 1. 9 , whole animal. Oc. S, obj. 2 in.
lig. 2. 5th foot, 9. Oc. 3, obj. 1 in.
fig. 3. Whole animal, 9 . Oc. 3, obj. -1 in.
lig. I. 5th pair of feet, 9 . Oc. 3, obj. \ in.
tig. 5. 2nd pair of feet, 9 Oc. 3, obj. 1 in.
anlarcticiis, fig. C>. 9 > whole animal. Oc. 3, obj. 2 in.
si mil i*.
PLATE IV.
fig. 1. 9 , whole animal. Oc. 3, obj. 2 in.
fig. 2. 9 , 1st foot. Oc. 3, obj. 1 in.
fig. 3. 9 , 2nd foot. Oc. 3, obj. 1 in.
liir. I. 9- abdomen ami genital segment. Oe. :',. obj. 2 in.
lig. ">. 9 . abdomen and ircnital segment. Oc. 3, obj. 2 in.
fig. i;. 9 , abdomen, lateral view. Oc. 3, obj. 2 in.
P 2
46
R. NORRIS WOLFENDEN.
Stffhus lonyipes,
PLATE V.
fig. 1. Whole animal, $.
fig. 2. Whole animal, 9 .
fig. 3. Whole animal, 9 .
Oc. 3, obj. in.
Oc. 3, obj. | in.
Oc. 3, obj. \ in.
antard'icum, fig. 4. Whole animal, 9 Oc. 3, obj. 1 in.
fig. 5. Whole animal, 9 , dorsal. Oc. 3, obj. 1 in.
fig. 6. 5th feet, 9 . Oc. 3, obj. i in.
figs. 7, 8. 5th feet, <$ . Oc. 3, obj. i in.
Paralabidocera hodgsoni, fig. 1.
fig. 2.
fig- 3.
.) fig- 4.
fig. 5.
fig- 6.
fig- 7.
fig- 8.
fig- 9-
PLATE VI.
9 , last thoracic segment and abdomen. Oc. 3, obj. -i in.
Whole animal, 9 Oc. 3, obj. -i in.
5th foot, 9 . Oc. 3, obj. J in.
4th foot, 9 i exopodite. Oc. 3, obj. i in.
1st foot, 9 . Oc. 3, obj. \ in.
2nd foot, 9 . Oc. 3, obj. in.
Posterior antennas, 9 Oc. 3, obj. \ in.
Maxilla, 9 . Oc. 3, obj. \ in.
Mandible, 9 . Oc. 3, obj. \ in.
fig. 10. Anterior foot-jaw, 9 . Oc. 3, obj. \ in.
fig. 11. Terminal lobes of post-footjaw, 9 Oc. 3, obj. \ in.
fig. 12. Whole animal, $ . Oc. 3, obj. ^ in.
fig. 13. 5th feet, 9 . Oc. 3, obj. K in.
PLATE VII.
Xantlioi'iiliiinix magnits, fig. 1. Whole animal. Oc. 3, obj. 2 in.
fig. 2. Rostrum. Oc. 3, obj. 1 in.
,. fig. 3. Posterior foot-jaw. Oc. 3, obj. I in.
fig. 4. Anterior foot-jaw. Oc. 3, obj. 5 in.
fig. 5. Posterior antenna. Oc. 3, obj. 1 in.
fig. 6. Maxilla. Oc. 3, obj. i in.
fig. 7. 1st foot. Oc. 3, obj. 1 in.
fig. 8. 2nd foot. Oc. 3, obj. 1 in.
fig. 9. 5th foot. Oc. 3, obj. i in.
antarcticus, fig. 10. 5th foot. Oc. 3, obj. in.
,, fig. 11. Posterior foot-jaw. Oc. 3, obj. i in.
Antarctic (Discovery i Exp.
Marion I y
)1. III.
Haloptilus ocellatus
Metndia princeps (3 5).
Gaetanus antarcticus
;
. \iit.n Disci '\viy) K\p.
poda pi. IV.
Kuchaeta similis (i
,, antarctica 15, '>)
Antarctic (Discovery) K\]>.
.i pi. V.
lon^ipes (i -j).
antarcticum (.( s )
Antarctic (Discovery) Kxp.
.l [ll. VI.
1'aralabidocera.
Antarctic ( Discovery) K\p.
-.1111.1
pi. \ 11.
Xanthocalanus magnus (19).
antarcticus (10, 1 1)
ECHINODERMA.
I.
By F. JEFFREY BELL, M.A.
Emeritus Professor and Fellow of King's College (University of London}.
(5 Plates.)
HAD it not been for the earlier return of the 'Gauss,' this report would probably
have had the distinction of signalising the re-discovery of the interesting Criuoid
genus, Promaekocrinus, which, as its name denotes, was one of the prizes of the voyage
of the ' Challenger.'
As in the collection of the ' Southern Cross,' 1 am again able to call attention to
.some remarkable variations within what are obviously the limits of single species.
The specimens which exhibit these variations could not have been obtained but
by very careful collecting, and in the case of CtjCethm it is of importance to note
that they were all taken in a comparatively small area; the examples of this genus
received by two naturalists, who made a number of species with them, were all taken
from stations comparatively close to one another,* but it is to be hoped that such a
course will never be taken again ; evidence as to the variability of species of
Echinoderms is now beyond question.
A.-ANACTINOGONIDIATA.
I. HOLOTIIURI01DEA.
The collection of Holothurians is small, and the points of greatest interest are
revealed by Prof. MacBridc and Mr. Simpson in their valuable report on the larvae,
which follows this memoir.
CHHUDOTA.
I submitted an example of a form taken at 100 fins., <>!!' Coulman lsl;md, to
Prof. Lud wig, who has made Antarctic Synaptids one of his special domains. Owing,
apparently, to the unfortunate use of formol the spiculrs are so disintegrated that ;i
definite judgment is impossible, but it seems probable that, the species is (_'. j>ixtti/ii or
allied thereto.
* It is true that C. simplrjc was fnmnl ;it Triniilail Channel, and the 'Challenger' specimens on the eastern
side of Patagonia ; but he who will look at a map of South America will smile at a " Chorological Synopsis of
the species " which gives three to the Atlantic and one to the Pacific.
F. JEFFREY BELL.
CUCDMARIA CROCEA.
Holothvriit- (t'lii'umctriii) crncpa, Lesson, Cent. Zool. (18:i2), p. 153, pi. lii. fig. 1.
Cucumariii rruirn.,* Ludwig, Hamb. Magalh. Sammelreise, Holothurien (1S98), p. 15 iliiiji" 1 ntnln.
In the succeeding memoir by Prof. MacBride and Mr. Simpson there will be
found an interesting account of the brood-pouches and young of this form. It was
taken at Coulman Island, 100 fins., and on various dates at Winter Quarters down
to 41 fms.
. CUCUMARIA LAEVIGATA.
PentacMla laorigata, Yen-ill, Bull. U.S. Nat. Mns. i. 3 (1876), p. 76.
Cucummia latvigata, Ludwig, op. eit., p. 32 ibique citata.
The characters of these two species have been fully discussed by Prof. Ludwig,
and it is quite unnecessary for me to enter again on the subject.
Taken in AViuter Quarters to 41 fms. and in McMurdo Bay.
Several distorted Cucumariae and some young were taken at various dates while
at Winter Quarters, and at 78 25' 40" S., 185 39' 06" E.
PsEUDOPSOLUS FERRARI.
(Plate V., fig. 3.)
It is unfortunate that this new species should be represented by a single specimen
only, but there is no doubt that it is unlike any form that has as yet been described.
Tentacles ten, two much smaller than the rest ; trivial pedicels in three complete
longitudinal series ; dorsal perisome devoid of large scales or grains, flexible and soft
to the touch, and having only a few trellis-like spicules. There are some distinct
pedicels scattered on the dorsal surface. 73 mm. long, 32 '5 mm. greatest width.
The important differences between this species and the only form belonging to the
genus which has yet been described are of varying value ; the distinctness of the dorsal
pedicels is of great significance, for their presence in P. macquariensis is so inconspicuous
that Prof. Deudy denied their existence, on which Prof. Ludwig has remarked
" Zu eiuer Entschuldiguug will ich aber gleich hinzufiigen, dass die dorsalen Fiisschen
sehr gering an Zahl und sehr retraktil sind, sodass sie sich der Beobachtung leicht
entziehen konnen." f
The spicules are more delicate and trellis-like in the new species, and there is a
* In my ' Southern Cross ' Report I cite Wyv. Thomson as the first author of the combination ' Cucumaria
crocea" ; he wrote Cladoilncfi/Jn crocca in 1876 (not 1878, and not on p. 55) ; as all my blunders were made by
Dr. Lamport in his " Seewalzen," I stand convicted of neglecting the legendary advice of the venerable gentleman
who was once President of my College at Oxford.
t Hamburger Magalh. Sammelreise Holothurien (1898). p. 49.
ECIIINODERMA.
very considerable difference in size. As there is but a single specimen, I thought
the question whether or no it is hermaphrodite might be left over for the present.
The only suggestion that I have to make is that Prof. Ludwig's definition of the
genus (<>p. cit., p. 50) should be altered by the omission of the words " Sehr wenigen,"
as applied to the dorsal " Fiisschen."
As the type of the genus came from Macquarie Island, the two are not widely
separated ; no information is given as to the depth of the more northern species ; the
Antarctic form was taken at 20 fins., while at Winter Quarters.
THYONE sp.
A young specimen was taken off Cape Adare.
MESOTHURIA MAGELLAN:.
Iliilitliiii-iii HKii/fUitiii, Ludwig, Ber. Oberbess. (k-s. xxii. (1883), p. 17">.
Mi'fiithnria niui/clltmi, Oustenrren, Festskrift fOr Lilljeborg (18%), p. 350; Ludwig, Ergebn. Hamburg.
Summelreise Holotbur. (1898), p. 8.
Two much-injured specimens are assigned to this species ; I was unable to find
spicules, but I do not always expect to find specimens taken out of ice-cold water and
from 100 fms. equal to the best products of the laboratories of Naples or Plymouth.
Coulman Island, 100 fms.
B -ACTINOGONIDIATA .
II. CRINOIDEA.
('OMATULIDAE.
PROMACHOCRINUS KERGUELENKNSIS.
(Plate I.)
frergueletisis, P. H. Carpenter, Proc. Roy. Soc. xxviii. (1870), p. 385.
This is certainly one of the most interesting finds of the ' Discovery,' for the genus
was, till the German South Polar Expedition brought back specimens, only known from
the collections of the ' Challenger,' which found two species in the Southern Seas, and
one a few degrees north of the Equator. The only addition to our knowledge since
Dr. Carpenter's report has been the note by Dr. Minckert of Greifswald,* while Dr.
Bather lias suggested that it may be a " permanent meristic variation.''! I give a figure
* Zool. Anzeig. xxviii. (1905), p. 490.
t Treatise on Zoology iii. (1900), p. 195.
4 P. JEFFREY BELL.
of a self-dissected specimen which is, I think, clearer and more illuminating than the
originals of the ' Challenger ' report. It would be of particular interest to discover
fossil examples of this ten-rayed form ; at present, all we can say as to the
" permanency " of the form is that it has endured for thirty years ; as to the
polyphyletic origin of the genus which Dr. Bather suggests,* we can only say that
we have only such data as are given by the distribution of the genus. Though
Dr. Minckert had made two genera, he has not contributed to the elucidation of
Dr. Bather's suggestion.
Taken at East end of Barrier, 100 fms. ; off Coulman Island, 100 fms. ; and at
Winter Quarters.
ANTEDON ADRIAN:.
(Plate II.)
This new species is represented by a number of specimens ; as I have already
named a member of the genus A. wilsoni, I take the specific name from the second
Christian name of Mr. E. A. Wilson, M.B., surgeon and naturalist to the Expedition.
It was taken at various depths, down to 130 fms., at Winter Quarters, and at
500 fms. off Mts. Erebus and Terror.
At first sight this species does not recall the northern A. eschrichti, but it shows,
on analysis, some striking points of resemblance the shape of the ceutrodorsal, the
length (but not the number of the cirri), and the shapes of their joints, as well as
the form of the first brachial ; but what distinguishes the creature from all Antedons
known to me is the saw -like appearance of the arms when viewed from the side,
and well shown in the figure ; this is due to the presence of a well-marked square
protuberance on the dorsal face of most of the arm -joints.
Arms ten, ceutrodorsal semicircular, with three rows of cirri ; these are from
50-60 in number, may be 60 mm. long, and may have as many as 60 joints ; the
axillary has a prominent knob, and the outer edge of the first brachial is twice as
long as the inner ; the arm-joints are stout, laterally compressed, and nearly all have
one or two dorsal spiuous processes ; syzygies are extraordinarily rare in the
proximal part of the arm, where, too, the pinnules are rather short ; further out
they are better developed. No specimen is anything like perfect.
ANTEDON ANTARCTICA.
Antedon antarctica, P. H. Carpenter, Chall. Rep. Comatulae (1888), p. 144.
I have compared a single specimen taken at Winter Quarters with those
collected by the ' Challenger ' at Heard Island, and have no doubt of their identity.
Some young specimens, which appear to belong to this species, were also taken.
* Treatise on Zoology iii. (1900), p. 195.
BCHINODBEMA. 5
ITT. ECHINOIDEA.
ClDARIDAE.
ClDARIS CANALICULATA.
Temnui'/il/ir/s cfi/uilici/h/t/r, A. A^assiz, Bull. M.C.Z. i. (1863), p. 1*.
tociiliiri* canaticulata, <<>. IVv. Krhin. i 1*7.';, p. 131 ; Wyv. Thomson, Journ. Linn. Soc. xiii. (1876),
p. r.. r > ; Lovt'ii, Bib. Svrask. Akad. Hillg. xiii. iv., 1. p. it ; Agass., Mom. M.C.Z. xxxi. (1904- ), p. 4.
x ( Duroritlarix) i-iiiin/ii'////i/ii, Doderlein, Jap. Secigel, i. (18x7), p. 10.
i* canaliculate, Mcissnur, Ergebn. Hainb. Miigalh. SainnielreisL-, v. (190u), ]., p. :; // y w ,-ltfita.
nniiilii-nliitii, Mortensen, lugolf Echinoid. (1903), p. 29.
I have given the name of C. canaliculata to a number of specimens of an
Echiuoid, which were mainly collected at a depth of 100 fms., but I must own to
grave doubts as to the correctness of the name. It seems to have escaped notice
that this consensu omnium (with the exception of Dr. Mortensen*) circumpolar
Antarctic form was first described from the " Caroline Islands," which Caroline Islands
we surmise to be those in the Pacific, as in the Revision of the Echini we find
" Caroline and Sandwich Islands " ; elsewhere, Zanzibar and the Navigator Islands are
given as habitats, "if the localities are to be trusted"; that is to say, the species
was founded on specimens said to be found within the tropics.
I am not going to join those who claim that forms must differ specifically, either
because they are found at different spots or different depths ; but, as I showed many
years ago, there is a distinct intertropical fauna in the Great Ocean ,f and the fact, if
fact it be, that a member of that fauna is also a circumpolar Antarctic form ought to
rest on the firmest possible basis. Unfortunately, the National Collection has no
examples from any station further north than Tasmania.
Again, the original diagnosis, based on the Caroline specimen or specimens, is too
short for a form which every student who has examined it, except Dr. Mortensen, allows to
be eminently variable ; with the exception of Prof. Doderlein, none of these students has
given a serious diagnosis of the species, and even his is not altogether what one wants.
Yet another difficulty remains to be mentioned ; it is generally agreed that
C. iiutri.r^. and C. ricijiura are synonyms of C. canaliculata; in other words, the form
has a marsupial habit, but I cannot detect signs of it in the specimens before me ;
it may be, of course, as it curiously is in the case of Jfemiaster cavemosus, that this
collection consists of males only ; but Mr. Hodgson tells me that he observed no signs
of viviparous habit; our experience, in fact, is the same as that of the hte Felix
Bernard : " Quoique G. canaliculata soit signale coiume vivipare, je n'ai pas constate
aucun jeune sur le corps des adultes."
* Op. cit., p. 27.
t For some modification of this doctrine, see Prof. Koehler in " Die Fauna Sudwest-Australiens " I. iv.
U907), p. 242.
J Prof. Lyman Clark doubts this (see f.-n. next page) ; of C. vivipura of Studcr he makes no mention.
Bull. Mus. P;m<. i. iis'.).-,i. p. 272.
VOL. IV. '.'
6 F. JEFFREY BELL.
Specimens were taken at Coulman Island, 100 fms. ; Winter Quarters, 100 fms.;
East end of Barrier, 100 fms.
This species is one that has lately been the subject of some dispute ; Prof. Agassiz,*
who originally described it, suggests that it be placed in a new sub-genus Centrocidaris,
while Dr. Mortensen f finds it to consist of several species, but he does not appear to
be sure of the exact number. J
ECHINIDAE.
ECHINUS MARGARITACEUS.
Echinus marffaritaceus, Lamk. An. s. V. iii. (1816), p. 47; Bull, Coll. ' Southern Cross' (11102), p. 219
i/iiijuf ntiitu ; Mortensen, Ingolf Echinoid. (1!)03), p. 101.
Echinus dhHlcina. Studer, MB. Akad. Berlin (1876), p. 456 ; Al. Ag. Chall. Rep. Echin. (1*81), p. 117.
Echinux liorriilus, Al. Ag. op. fit. p. 117 ; Mortensen, op. rit. p. 102.
Sterechinm antarefitiis, Koebler, Echin. Toy. 'Belgica' (1901), p. 8.
This appears to be a circumpolar species, as I remarked when I reported on the
collections of the ' Southern Cross.' It was taken by the ' Discovery,' not only at
several dates in Winter Quarters, but at the East end of the Barrier Reef, 100 fms. ;
at Cape Wadsworth, 8-10 fms. ; off Coulmau Island, 100 fms. ; and South of
Antarctic Circle, 254 fms.
HEMIASTER CAVERNOSUS.
(Ti-ipyhis) carernosus, Philippi, Arcb. f. Nat. si. (1845), p. 345.
ntrernosits, A. Ag. Rep. Ech. (1872), p. 132 ; Meissner, Ergebn. Hamburg Magalh. Sauimelreise,
v. (1900), p. 13 iblqm citata ; Bell, Coll. 'South. Cross' (1902), p. 219.
Abattis cavernosits, Loven, Bih. Svenska Vet. Akad. Hdlgr. xviii. 4, no. 1, p. 3.
This species is not very well represented in this collection, and, curiously enough,
all the examples are males.
If the late M. Bernard was right in regarding Tripylus excavatus as a synonym,
the name of the species ought to be excavatus, as that was the first of the three species
described by Philippi ; for myself I am inclined to abide by Prof. Agassiz's view.
Taken at Coulman Island, 100 fms. ; Winter Quarters, 20 fms. ; and East end of
Barrier Reef, 100 frns.
IV. ASTEROIDEA.
The collection of starfishes was rather large, and contained some very fine
specimens ; but, as will be shown later, there is very great difficulty in coming to
definite conclusions regarding them, and I have preferred to be vague rather than
dogmatic in treating of them.
* Mem. Mus. Comp. Zool., xxxi. (1904), p. 32. t Ingolf Echinoid. (1903), pp. 25-7.
t As the final proof was passing through the press I received Prof. Lyman Clark's latest memoir on the
Cidariilaf (Bull. Mus. C. Z. li. (1907), no. 7) ; he has instituted a new genus, Austrocidaris.
Bull. Mus. Paris, i. (1895), p. 247.
ECHIXODERMA. 7
AsTERIAS BRANDTI.
brandti, Bell, Proc. Zool. Soc. (l**i;, p. :H.
Asterias ;m//.
1 >r. Meissner lias suggested that J. negleeta, which I described at tlie same lime
as A. ftninilfl, is the same species. I think he is right; A belli of Prof. Studer and
^.4. glomerata of Sluden appear to be also synonymous ; with regard to the first three
Prof. Perrier suggested the same view in 1891.
My species were described on pp. 91 and 94 of the Proc. Zool. Soc. for 1881 ;
Dr. Meissner's synonymy will be found on p. 7 of tlie Ergeb. Hamburg Magalluui.
Sammelreise, Lief. vii. (1904). I suggested in the report on the 'Southern Cross'
collections (p. 215) that the species was circumpolar in its distribution, and so it
certainly is.
A large number of specimens were collected; the largest, from 4-10 fins.
McMurdo Bay, was the " mother of a number of young after capture" * ; and some other
large specimens were taken at No. 3 hole, and some smaller at No. 11. Flagon Point
(10-20 fms.), Cape Wadsworth, and Hut Point also yielded examples ; the lar-je
examples marked 4H and 49 are only said to be from Winter Quarters.
From 100 fms. (East end of Barrier) and 130 fms. (X<>. 2) specimens were
collected which, but for the formidable synonymy of the species. 1 should be inclined (o
regard as distinct.
ASTERIAS L< >.\<;STAFFI.
This line >pecies is represented bv a single specimen, which presents well-marked
features. 1 name it after the generous friend of science whose contribution to tin-
funds of the Royal Geographical Society made the vovage of the 'Discover.-
possible.
A large stout form with A' about equal to I! / ; the whole of the dorsal surface i-
uuiformly and densely covered with paxilliform spines of some size. The madreporite is
of moderate size ; the podia are enormous, the adambulacral spines are loni;.
cylindrical, blunt at the tip, so irregularly set that it is ditlicult to say whether then-
are two or three rows : beyond them there are shorter and more delicate, but otherwise
similar cylindrical spines verv doselv set; there are two tjuitc well-marked fets of
marginal plates; the inferomarginals are very narrow in the an^le of the arm, while
the superomarginals in the same region are verv .-hort ; both sets are covered with
coarse granules, among which no pediccllariae are to be detected. The podia within
the oral circlet are of enormous si/e. If = 210 ; / = 70.
\V.( V >. : 10-20 fms.
As there is only a single specimen, no comparisons are possible; but it muv be
' These luivc IHTII in:uli- tin- -uliji-ct ..1 ;ui iiitcivstin^ separate iv[>it \<\ I'mf. M;i.-]Hiili' and Mr. '. ('. Simpson.
F. JEFFKBY BELL.
pointed out that the closely packed arrangement of the podia, and the distinctness of
the two rows of marginal plates are, on Mr. Sladen's basis of classification of Starfishes,
mutually destructive ; but the well-known observations of Prof. Ludwig on Echinaster
sepositus justify us in supposing that we have here a case of retarded disappearance of
the marginals ; the crowding of the podia appears to lie a much more important
morphological character ; but the union of these two strikes, I think, a final blow at
the current classification, against which Prof. Ludwig has already raised his voice, and
the adoption of which by MM. Delage and Herouard in their " Zoologie Concrete "
came as a great surprise to me.
HEURESASTER* HODGSON i.
(Plate III.)
Two specimens of a very fine starfish were taken at about 25 fathoms while in
Winter Quarters ; they appear to me to form the type of a new genus, as to the
general position of which there can be little doubt ; a still larger specimen was taken
from McMurdo Bay at 2 fms.
It has somewhat the appearance of Porania, but has, in the larger examples, spines
in the interambulacral actinal areas. Prof. Perrier defines the Poraniidae as follows :
" Squelette masque par les teguments ; margiuales apparentes, mais formant au corps
uu bord tranchant ; squelette ventral forme de se'ries de plaques allant de chaque
adambulacraire a une marginale, squelette dorsal reticule."
I have invented for it a name which will remind the student both of the name of
the ship, and of its indefatigable biologist.
The larger specimens may be thus described : Arms long, tapering to a rather fine
point, R is about = 3. The upper surface is smooth and soft to the touch, and has
papulae, in ill-defined areas, spread over the whole of it ; the edge is quite sharp,
forming almost a ledge, and made up by a large number of small supero- and infero-
marginals. The lower interambulacra covered with some eight rows of small regularly
set plates covered with rather coarse granules, and, in the angle, with short spines
which give a hairy appearance to these areas. The ambulacral groove is bounded by
rows of four or five spines, of which the outermost is small and the innermost spatulate
and fluted at its free end. The podia are stout. At each oral angle there is a huge
spine, the distal third of which is glossy. These specimens are flat, but the smallest
example has the disc arched, and this is probably more natural
R = 200 r = 70.
R = 130 r = 50.
R = 90 r = 30.
Accepting Prof. Perrier's family diagnosis of the Poraniidae, we may make the
generic diagnosis of Heuresaster: Abactinal surface only invested by integument,
discovery.
BCHINODBBMA. '.'
margin very .sharp, the inner actinal ambulacralfl with spines, the oral angle provided
with a large spine. While the specific characters would appear to be that the longer
radius is more than twice as long as the shorter radius, the marginal plates are MTV
numerous, and the innermost ambulacra] spine is spatulate and fluted at its free end.
The smallest specimen has not quite acquired the generic characters of the larger;
the abactinal integument is not so thick as to altogether hide the superficial granules,
and the granules on the actinal interambulacra have not developed into spines, so that
there is no marked difference between the outer and the inner parts of these areas.
PEXTAGONASTER INCERITS.
The single small specimen, is, I think, an ally of the Australian species of Penta-
gonaster (sens, lat.}, but the arms are proportionately longer than they generally are
in this genus ; it is, possiblv, an immature specimen in which R would gradually
increase in proportion to r. If it should prove to be an adult, its proportions mav be
compared to those of P. duebeni and P. ) p. ->\->.
(Hitarctii'iix, Sladen, n/>. fit. p. 190.
I must own to some temerity in associating a specimen in which R = 212 and
r = 58 with a species whose type had Li = 38 and r = 12 '5, and a representative
of which, hardly much larger, was found to be bearing young; but even the most
recent writers on Echinoderms have not yet promulgated the. doctrine that difference
in size is a specific character, though I am not quite sure that in practice they do
not sometimes act as though they had. However, one has only to get a clear idea
of the essential characters of this genus to feel sure that one lias it here ; as to specific
characters, it is first to be said that most of the L. kerguelpnensis material is badly
preserved, while the condition of L. niifdrrticus is particularly good. Though the
differences between the two species appear, from Mr. Sladen's lengthy description, to
be considerable, it will, I think, be found on examination of the specimens preserved
in the Museum, that L. jiliio:ona inermis, Bell, Rep. 'Southern Cross' (1902), p. 217.
This species, which was not well represented in the earlier collections, was found
on numerous dates during the stay of the ' Discovery ' at Winter Quarters.
It exhibits a very wide range of variation, so great indeed as to have been at first
quite bewildering ; and it may be noted that Mr. Hodgson did not recognise any of
them as belonging to an already known Antarctic form.
In the ' Southern Cross ' report I figured some of the remarkable variations in the
arrangement of the plates of the dorsal surface of the disc of Ophiosteira antarctica ;
a similar, but less striking, variation obtains in this Ophiozona. In 0. antarctica I
described the serrated and keeled appearance of the upper surface of the arm ; this is to
be seen also in some examples of 0. inermis. The most striking differences are to be
seen in the height of the disc, and the size of the so-called radial shields ; so that it
is by no means always true that the shields are inconspicuous.
Winter Quarters, 10-178 fins, (various dates) ; MacMurdo Bay, 96-120 fms. ;
East end of Barrier, 100 fms. ; off Barrier, 300 fms. ; Coalman Island, 100 fms.
In some specimens the dorsal plates of the disc are deeply incised ; this appears to
be due to want of sufficient calcareous matter, but as others are infested by a sponge it
is possible that it is the cause.
Ol'HIOSTEIRA ANTARCTICA.
(Plate V., fig. 2.)
(>l'liinsteira antarctica, Bell, Rep. ' Southern Cross' (1002), p. 218.
It appears that the ' Southern Cross ' did not exhaust the variations in the disc
plates of this wonderful species ; the upstanding plates shown in Plate V., fig. 2, are
worthy of record, and it may be noted that the single arm left to this disc has the
dorsal serration extremely well marked.
ECHIXODERMA. 13
In the original diagnosis of the genus I spoke of " a large keel-like plate on the
disc, which completely overshadows the radial shields," although I was aware of and
figured (Plate XXVI., fig. 3) an angle of the disc in which there appeared to lie two large
radial shields ; in this case, however, three of the angles of the disc conform exactly to
the diagnosis.
I have now before me a specimen in which all five angles have a pair of plates.
Were it not for the ' Southern Cross ' specimens it would not be possible to put this
specimen with the genus Ophiosteira at all.
When we are asked whether " specific characters are useful,' 7 we may retort that
generic characters even are not always constant.
AVinter Quarters, 13 fms. Coulman Island, 100 fins.
OPHIONOTUS VICTORIAE.
Oji/iiniiotiix rii'/nridf, Bell, Rep. 'Southern Cross' (1902) p. 210.
This species was not so abundant; it was taken off the Barrier at 300 fms., near
Franklin Island, and from 254 fms. at an unrecorded locality.
OPHIACANTHA IMAGO.
'i/i/tiacantha imago, Lyman, ' Chall.' Rep. Ophiur. (1882), p. 18i; i/>iq//i' ,-ilahnn.
I hope I am right in referring to this species specimens from Winter Quarters,
30 fms. ; Hut Point, 77 12' 12" S., 167 27' W., 77 50' 30" S., 165 40' K, 100 fms. ;
but, in sooth, some of the ' Challenger ' types of Ophiuroids are hardly mature.
OPHIACANTHA VIVIPARA.
Ophii'i>l. lid. vii. il'.HMi. |>. 684.
VOL. iv. K
14 F. JEFFBEY BELL.
The single* six-armed specimen taken by the 'Discovery' at 8-15 fms. off Cape
Wadsworth is a good deal stouter than Ljungman's types, but I can see no reason for
making it a new species ; the arms of the young may be seen projecting from some of
the bursae.
OPHIACANTHA COSMICA.
Ophiacantha cosmica, Lyman, Cball. Rep. Oplriur. (1882), p. 194 ibi^ue citatum.
This widely spread species was taken on various occasions, but many of the
specimens have been marked by me " immature " ; in connection with this I should
like to quote some words of Mr. Lyman : " The specimen just described is unusually
large for this division of the genus, which leads me to think that the bulk of those
now known are immature, and therefore to be treated with all the more caution."
Winter Quarters. Off Coulman Island, 100 fms. Off Barrier, 300 fms.
OPHIOCONIS ANTARCTICA.
Ophioconis antan'tita, Lyman, Chall. Eep. Ophiur. (1882), p. 107 ibiqite citatum.
If I have correctly determined some specimens from Winter Quarters, the types
of Mr. Lyman 's are very far from reaching the size to which this form attains.
AMPHIURA BELGICAE.
i
Amphiura belyicae, Koehlcr, Res. Voy. 'Belgica' (1901), p. 27.
The Antarctic area appears to be the home of large specimens of Amphiura ; not
only did the ' Discovery ' bring home several examples of what appear to be the
species obtained by the ' Belgica,' but there is a single specimen off the Barrier at
301 fms., which has a disc diameter of 18 mm.,f and which can hardly, I think, be
the fully adult example of the somewhat smaller specimens ; but, as it is solitary, I
hesitate to regard it as an undescribed species.
Several specimens of A. belc/icae were taken at Cape Wadsworth, 8 to 15 fms., and
off the Barrier at 300 fms. ; the latter is, as already said, the locality of the single
larger form. Like Solaster octoradiatus, A. belgicae is probably a circumpolar species.
YOUNG OPHIURID.
(Plate IV.)
I give some figures of a remarkable Ophiuroid, to which it is very difficult to
assign a systematic place ; it appears to be a young form in which some of the arm-
plates are still not consolidated. The exceedingly large size of the bursal slits is
perhaps only due to the mode of preservation.
Two specimens, both broken, were taken in Winter Quarters on 2. 1. 04, and two
off Barrier.
* As there is but a single specimen this collection throws no critical light on Prof. Koehler's just-made
suggestion that this species exhibits sexual limorphism ; cf. Bull. Sci. France xli. (1907), p. 322.
t That is exactly the same as the disk diameter of A. magnified; of which Prof. Koehler has lately published
a brief diagnosis. See Xool. Anz. xxxii., p. 146 (17 Sept. 1907), but the species is not the same.
ECHIXODERMA. 15
AsTIHiToMA A6ASSIZI.
Astrotoma agassizii, Lyman, 111. Catal. Mus. C. Z. No. viii. II. (1*75), p. 24.
Mr. Lyman calls attention to the coarse granulation on his single specimen,
which came from the Straits of Magellan ; the series now collected shows that the
granulation may be more or less obscured by thick skin. It is remarkable that the
species was not collected by the ' Gauss.'
McMurdo Bay, 9G-120 fms.
Specimens from off Coulman Island, and from 77 12' S., 1G7 27' E., 2 fms. seem
to be sufficiently different to lie regarded as varieties.
R 2
16 F. JEFFREY BELL.
DESCRIPTION OF PLATES.
ECHINODERMA.
PLATE I.
Promacorhrinus Jcerguehnensis.
FIG. 1. Disc and base of arms from the side to show the insertion of the cirri, and the form of the
cirrus pit, and the shape of the disc, x 2.
FIG. 2. Disc and base of arms from below ; the central boss is more distinctly seen than in fig. 1.
x n.
FIG. 3. An arm from the side, x 3 ; the roughened edges of the joints are well seen in this figure ; the
proximal pinnules only are completely shown.
PLATE II.
Antedon adriani.
General view of this Crinoid. x 2.
PLATE III.
Heuresasfer hodgsoni.
X not quite f, seen from below, to show the breadth of the ambulacra! grooves, their size and
arrangement.
PLATE IV.
The young Ophiurid (see p. 14).
FIG. 1. Aboral view of disc and arm. x f.
FIG. 2. Oral view of same. X f .
FIG. 3. View of mouth to show arrangement of the oral and adjacent plates, x 8.
FIG. 4. 1st, 2nd, and 3rd upper arm-plates, x 10.
FIG. 5. View of arm from above, at about its middle, x 10.
FIG. 6. 1st, 2nd, and 3rd lower arm-plates, x 10.
FIG. 7. View of arm from below at about its middle, x 10.
PLATE V.
FIG. la. Variety of Cycethra (see p. 10) from above ; fig. Ib from below ; both slightly reduced.
FIG. 2a. Disc of an Ophiosteira (see p. 12), magnified to show keel-like plates ; 2a, seen from above ;
2b, seen from the side.
FIG. 3. Pseudopsolus f&rrari. x ?,.
Antarctic (Discovery) Exp.
Echinoderma pi. I.
l>.TJ>'au del. Ituttcrworth. ST.
--
Antarctic ( Discover)') I'.xp.
Echinoderma pi. II.
lierjeau. del. Batlerworth, so.
A
Antarctic (Discovery) Exp.
Echinoderma pi. III.
' del I Illtll \\orth. BC.
Antarctic (Discovery) K\p.
Echinodenna pi. \\ .
HtterrtOl th BC
\a
-*
'-
>
*
Antarctic (Discovery) Exp.
n iderma pi. V.
Tcrzl, del. Huth. del. Buttervvorlh.se.
ECHINODEEMA.
IL-ECHINODEBM LAEV^E.
By E. W. MACBRIDE, M.A., D.Sc., F.R.S., Professor of Zoology, McGill University; and
J. C. SIMPSON, B.Sc., Demonstrator of Zoology, McGill University.
(1 Plate.)
THE collection of larval Echinodenns brought back by the National Antarctic Expedition,
when received by us from the authorities of the British Museum, was found to consist
of forty tubes of material collected during the years 1902-1904. Of these forty tubes,
however, by far the greater number contain nothing but early segmenting eggs showing
no distinctive features ; the present paper is, therefore, in reality based upon the
contents of eight tubes only.
A preliminary examination showed that, though the collection contained but
four species of Echinoderms, these differ widely from one another, there being one
representative of each of the four great classes of the sub-phylum Eleutherozoa. These
four species are: Asterias brandti Bell, Cucurnaria crown Lesson, a new Echim>-
pluh'ii*, and an Ophiopluteus which may also be new.
At first sight the Asterias seemed to promise much, for there were two tubes
containing a couple of hundred specimens which had been born after the capture of the
mother. More careful examination, however, disclosed the fact that they were all
of approximately the same age, only two closely related developmental stages being
represented. In the case of the Cucumarin, besides a number of new-born young, the
collection contained an adult female which was the mother of some of them. To this
specimen considerable interest attaches, from the fact that in it an unsuspected brood-
pouch containing a great number of embryos was found. Of the two plutei it need only
be said in this place that, since they are probably the first free-swimming Echinodenn
larvae found within the Antarctic Circle, they are not without considerable importance.
In the matter of preservation the material leaves much to be desired ; but when
one considers the difficulties of collecting under Antarctic conditions, it could hardly be
expected that this could be otherwise. Of necessity the only preservatives used were
formalin and spirit, and as neither of these is at all suitable to Echinoderm larv;v, the
specimens show very little histological detail.
In examining the material it was found that the preservation of the plutei was
such as to make sectioning useless, so they were studied /// fafn, first unstained, and
then stained in eosin and mounted in balsam. This method was found to give very
satisfactory results. The Asterias and Cuciinmria, however, were treated quite
differently. Selected specimens of these were first carefully drawn at a magnification
2 E. w. MACBRIDE AND j. c. SIMPSON.
of 12| diameters l>y the aid of the camera lucida, and then, after double imbedding in
celloidin and paraffin, were cut into serial sections and stained with Delafield's
hsematoxylin.
In the succeeding pages the four species will be taken up and described in detail.
1. ASTERIAS BRANDTI.
Asterins Irandii, Bell, P.Z.S., 1881, pp. 94 and 50C.
The young of this species contained in the collection consist in a brood " born
after the capture of the mother," which was taken in McMurdo Bay, February 13th,
1902.
The parent was identified by Professor F. Jeffrey Bell as A. brandti, but as it was
not sent with the embryos, nothing definite can be said as to its " brood-care," except
that this species must now be added to the list of viviparous Asteroidea. It is
probable, however, that according to the analogy of its family, the young are attached
to the lips of the mother during the earlier stages of development.
All the embryos of this brood are of approximately the same stage ; the difference
between the youngest and oldest being simply a matter of size. The youngest is
shown in Fig. 1, which gives a ventral view. It measures 2' 4 mm. from the centre of
the disc to the tip of the arms, while the radius of the disc is 1 5 mm. (R. = 2 4 mm.,
r. = 1*5 mm.). The only external trace of the embryonic condition is in the closed
mouth and the small pre-oral lobe 'or larval organ. The oldest embryo is much larger
than the preceding (R. = 3 2 mm., r. = 1 6 mm.), but apart from the smaller size of the
pre-oral lobe there is little difference. Internally, too, there is little difference in the
development of the organs, so that they may very well be described together.
As may be seen from the figure, metamorphosis is complete in these young starfish,
and an examination of serial sections shows that the internal organs have nearly all
attained the adult relations. As these relations are, moreover, practically the same as
those in a young Asterina gibbosa of the same stage, it will only be necessary to mention
here the few points which show peculiarity.
In the first place it will be noticed that the great size attained before the mouth
and anus are formed must be looked upon as an expression of the favourable conditions
under which development of this species proceeds. Another indication of the same
thing may be seen in the amount of yolk which still remains in the closed gut, and
which may even be seen outside the gut in the space between its dorsal wall and the
inner wall of the coelom.
The pre-oral lobe (larval organ) in the oldest stage is little more than a vestige,
but in the youngest its connection with the inner perihtemal ring can still be traced.
The water-vascular system is well developed, and one can follow it quite easily in
vertical sections. The madreporic pore, pore-canal, and stone-canal may be said to
have assumed the adult relations. The axial sinus can be seen adjacent to the stone-
HcnixoDERM LAKV.I:. 3
canal, and can be traced from its opening into the latter to its junction with the inner
perihaemal ring. In none of the sections, however, can one be sure of the aboral sinus
or of the madreporic vesicle, though this is doubtless due to the preservation.
Several points in the structure of these young Asterids suggest a relation to an
embryonic starfish studied in this laboratory some time ago by Dr. E. II. Henderson (4)
These were large yolky eggs of an unknown species, belonging, probal.lv. to the "enus
Anuteruu, collected off the Franklin Islands. In the remarkably close resemblance to
the development of Aster ina gibbosa of the same age, and in the similarity in the
appearance and distribution of the yolk, we have indications of a near relationship
between this species and Aortas bmndti, but the great difference in age between the
two lots ,,f embryos makes it difficult to institute a closer comparison.
2. OPHIOPLUTEUS sp.n. (?).
(Fig. 2.)
Two specimens of this presumably new Op/iinpluteus were taken at the Winter
Quarters of the Expedition ; the first on December 15th, 1903, from No. 12 hole in
3 fin*. O f water; the other on January 4th, 1904, from No. 13 hole, in G fras. The
first is in a comparatively good state of preservation : the second is badly disintegrated
and from it alone little could be made out. It is, however, of approximated the same'
stage of development, though a little larger, than the first. The following description
is, therefore, based upon the better preserved specimen.
The body, exclusive of the arms, measures 2' 3 mm. in length, and is marked by
a strongly developed conical papilla at its posterior end.
Three pairs of arms are developed ; of these the antero-laterals and post-orals
are only slightly developed, but the postero-lateral pair are already equal to the body
in length and are widely divergent. This development of the postero-lateral arms
reminds us of the great size of the corresponding arms in the larva of Ophiothrix
fragilis, which is such an early and marked characteristic of this species. In fact, tl.r
whole appearance of our specimen is strongly reminiscent of the larva of Ophiothrix
The ciliated band can be traced quite distinctly bordering all the arms.
In our better specimen the coelom can just be made out, On the left side it has
already divided into anterior and posterior cceloms, but on the ri-ht side it is still
undivided.
The gut is well developed and shows the characteristic division into oesophagus,
large spherical stomach, and intestine opening on the ventral surface.
No trace of the skeleton is preserved in these specimens, and this, together with
their young stage, makes classification impossible. Judging from their habitat
however, and from their general appearance, they are quite possibly examples of a new
species.
4 E. w. MACBEIDE AND j. c. SIMPSON.
3. ECHINOPLUTEDS ANTAECTICUS.
(Fig. 3.)
The material contained only two specimens of this larva : one taken in
lat. 78 29' 10" S., long. 103 38' 18" W., on January 25th, 1902, and labelled "30,"
the other marked " 42," taken in the D-net at Hut Point, near the Winter Quarters
of the Expedition, on February 19th of the same year. Both specimens are of
approximately the same age.
In the stained and mounted condition the specimens measure 0'72 and 0'84 mm.
in length, including the arms ; but in each case the body, which is sub-spherical in
form, is 0'43 mm. in diameter.
Four pairs of arms can be made out ; of these, the post-oral pair are the longest,
being a little more than two-thirds as long as the body. Of the others, the autero-
laterals are about half and the postero-dorsals about one-quarter the body-length.
The pre-oral arms are very rudimentary, appearing only as small outgrowths at the
bases of the antero-laterals. All the arms are close together and are approximately
parallel.
The ciliated band, which borders all the arms, extends back between the post-oral
and postero-dorsal arms of either side, so as to form lateral bays running about half
way along the body.
Three pairs of epaulettes are present, and are arranged so as to form two
incomplete ciliated bands encircling the body. The anterior circle consists of a
ventral pair situated at the bases of the post-oral arms, and a dorsal pair in a
somewhat corresponding position on the dorsal surface of the body at the bases of the
postero-dorsal arms. The posterior circle is made up of a single pair only, and these
are smaller and much narrower than either of the other pairs, and are laterally placed
near the posterior end of the body.
We find that three Echinoplutei have been described in which the presence of
six ciliated epaulettes is noted. These are : E. coronatus, E. theeli, and the pluteus
of Echinus csculentus. From each of these, however, the present larva differs
considerably. In Echinus esculent us the presence of the posterior pair of epaulettes
was first pointed out by MacBride (8), and a great deal of material of the larvae of this
species has been at our disposal for comparison with our new type. Selecting an
individual which, judging by the development of the so-called " Echinus-rudiment,"
is of the same age as our specimens of E. antarcticus, we find that the size of the
characteristic larval arms is quite different. The post-oral and postero-dorsal pairs
are of about the same length, and are already about twice as long as the body, instead
of two-thirds and one-quarter the body-length, as they are in E. antnrcti<-ux. The
antero-laterals are also of great length, being more than twice as long in comparison as
are the corresponding arms in our new species. The most marked difference, however, is
ECHIXODERM LARY.K. 5
in the pro-oral pair ; for while these are mere rudiments in K antarcticus, they are
almost equal to the length of the body in E. esculentm.
The other two larvae, E. coronatm and E. theeli, were described by Mortensen (9)
The first of these is a larva of about the same or a slightly older stage than our
specimens. In it, however, the post-oral and postero-dorsal arms are about one and a
times as long as the body, while the antcro-laterals and pre-orals are equal to it
m length. Besides this great difference in the arms there is also a difference in the
posterior pair of epaulettes ; for Mortensen describes these as " very broad/' while in
E. antarcticus they are much the smallest of the six.
In E. t/u-i-li we are dealing with a larva which is much younger than the others
mentioned, hut even here we see certain features in the arms by which to
ifferentiate the two; for while three pairs of the arms are of about the same
proportionate length as in our specimens of E. antarcticus, the fourth, the pre-orals,
are long and well-developed, not at all resembling the mere rudiments by which they
are represented in the latter species.
As in the case of the Ophiopluteus, so here nothing can be made out of the
skeleton, the result, probably, of the method of preservation.
On the left side of the larva a well-developed " Echinus-rudiment " can be seen,
but of this no details can be made out in either of the mounts.
4. CUCUMARIA CROCEA.
Holothuria (Cucumariif) crocea, Lesson, Cent. Zool. (1832), p. 153, pi. Hi., fig. i.
Cucumaria crocea, Lampert; Seewalzen (1885), p. 214.
Our collection contains some two hundred specimens of the young of this
Holothuriau, "born after the capture of the mothers," and one adult female, the
parent of some of these young ; all taken at Hut Point, near tl..- Winter Quarters of
the Expedition, on the 13th and 28th of February, 1904.
The young of this species were first dredged by the ' Challenger ' at the Falklands
in January, 1876. Since that time nothing, so far as we can learn, has been added to
our knowledge of their development, and as the specimens then found were of a
considerably older stage than those brought back by the ' Discovery,' it may be well
to recall certain of the observations made by Sir Wyville Thomson at that time, which
have a special bearing upon the developmental history.
In his " Voyage of the ' Challenger,' " Sir Wyville says : " In a very large
proportion of the females which I examined, young were closely parked in two
continuous fringes adhering to the water-vascular feet of the dorsal ambulacrum.
The young were in all the later stages of growth, and ,,f all sizes, from 5 mm. up to
40 mm. in length; but all the young attached to one female appeared to I..- of the
same age and size All the young I examined were miniatures of their parents; the
only marked difference was that in the young the ambulacra of the bivium were quite
VOL. IV.
c E. w. MACBRIDE AXD .1. c. SIMPSON.
rudimentary." They " attached themselves by the tentacular feet of the trivial
ambulacra, which are early and fully developed." He says, however, that they " were
too late at the Falklands (January 23rd) to see the process of attachment of the
young in their nursery .... There can be little doubt that, according to the analogy
of the class, the eggs are impregnated either in the ovarial tube or immediately after
their extrusion, that the first developmental stages are run through rapidly, and the
young are passed back from the ovarial opening, which is at the side of the mouth,
along the dorsal ambulacra, and arranged in their places by the automatic action of
the ambulacral tentacles themselves." (Vol. II., p. 215 et seq.}
Now the young brought back by the ; Discovery,' with two exceptions, all
measured from 2.? mm. to 3 mm. in length, and had been born in the interval
between capture and preservation ; it seems, therefore, that we had here a key to the
solution of the manner in which the early developmental stages are passed over, and
that we might thus get a more complete history of the " brood-care " of this species.
A careful examination of the adult female specimen, made with this object in view,
showed two prominent openings, each about 3 mm. in diameter, situated in the dorsal
interambulacrum, a short distance behind the peristome (Fig. 4). Looking into these
openings, one could see a number of embryos of about the same size as the new-born
specimens of the collection. At first sight these openings seemed to lead into the body-
cavity, but a careful dissection revealed the fact that they were in reality the openings
of two large brood-pouches situated in the dorsal body-wall (Fig. 5). Owing to the
extreme contraction of the specimen, the extent of these pouches can be made out only
approximately. The right, which is by far the larger of the two, extends from the
mid-dorsal line downwards to the right ventral interambulacrum, and from a point just
behind the peristome backwards for about one-third of the body length. The left sac,
however, is only about half as large. Both pouches are divided up by a number of
pillars representing the podia, which thus seem to traverse the sacs. In reality,
however, the podia are probably only surrounded by the pouches as they develop.
Some idea of the extent of these spaces may Vie gained from the figure, which is drawn
of the natural size. No communication was found between the two sacs, nor was any
opening into the body-cavity or genital ducts made out.
In these two pouches 140 embryos were found 110 in the right and 30 in the
left. These, however, are all of the same size, and of a comparatively late stage of
development, with the exception of two specimens which are much less developed than
the rest.
Through the kindness of Professor F. Jeffrey Bell we have also been able to examine
three other specimens of C. crocea belonging to the British Museum. It was hoped
that they might show some trace of these brood-pouches, but the most careful search
revealed nothing.
If we now try, from the facts before us, to get a connected idea of the " maternal
care " of this species, we shall probably find that its course is somewhat as follows :
ECHINODERM LAKV.K. 7
The eggs, either before or after fertilization, are shed into the water and are attached
to the mother's back, possibly by means of a mucoid secretion. They are soon,
however, surrounded by the brood-pouches, which probably develop by a sinking-in of
the dorsal ectoderm, and here they are retained until they have readied a condition
where the tentacles are able to serve as organs of attachment to the mother. This we
know must be when the little animals are between 3 mm. and 5 mm. in length. They
then pass out of the brood-pouches and are arranged along the dorsal ambulacra of the
mother as Thomson has described.
The presence in the sacs, among a large number of older embryos, of two which
are comparatively young, naturally brings up the question of the number of broods in
this species. Are all of the young brought forth in a single brood, or is there a
succession of broods such as we find in Si/ixiptn r/r/jinni, Clark (3) ? If these two
embryos represented the beginning of a second and younger brood, one would expect
to find the gonad of the mother showing some sign of activity ; but, as this is not the
case, we must conclude that they are members of the same brood as the larger embryos,
and that their development has been, for some reason or other, retarded. What
evidence we have, therefore, seems to be in accord with Thomson's observation that all
the young attached to one mother were of the same age and size.
Coming now to the embryos themselves, it is somewhat disappointing to find that
among the large number brought back only two stages are represented. Turning our
attention to the younger of these, we see that of the two specimens in the collection
one was somewhat distorted through the pressure of the surrounding embryos in the
brood-pouch ; but this does not at all affect the disposition of the internal organs as
seen in sections. The other specimen, apparently quite normal, is oval in form, and
measured before sectioning 1 8 by 1*2 mm. No evidence of any external opening
could be made out.
When sections of this stage are examined it is seen to be a typical pentactula, and
the only points which need be specially mentioned are in connection with the water-
vascular system. The ring canal is well developed, and the radial canals coming off
from it can easily be followed as they arch back and run towards the aboral pole. As
yet there is no sign of the development of podia, but the tentacular canals can be seen
to originate one on either side of each radial canal, close to where it comes off from the
ring canal, and to run out into the tentacles which do not reach the surface of the
embryo, but still project into the "atrium.' Even at this stage valves may be seen
guarding the entrance to these tentacular canals. The stone-canal is well formed, and
runs out in the dorsal inter-radius to open into a large, clearly-defined axial sinus.
From the latter the pore-canal can be traced outwards for a short distance, but we were
not able to follow it to its external opening in this stage, though in the older embryos
it is very easily followed. No Polian vesicle has yet been developed.
Of the later stage there were a great number of specimens, for it was found that
all the embryos in the brood-pouches (with the exception of the two ahvadv mentioned),
s -.:
E. W. MAcBRIDE AND J. C. SIMPSON.
and a second brood dredged a fortnight later, are of approximately identical
development. These embryos all measure from 2' 5 to 3 mm. in length, and have
assumed the typical ten-teutacled form. The tentacles are still simple, unbranched,
peg-like outgrowths (Fig. G), but there is some slight indication of a differentiation in
size, the two ventral tentacles being just a shade smaller than the rest.
In sections the digestive tract is seen to be well developed, though the oesophagus
is still solid, and the anus has not yet been formed. The stomach walls are thrown
into heavy folds and the intestine shows the characteristic double twist.
The water-vascular system in this stage shows many advances. The pore-canal
can be distinctly seen to open to the outside, and can be traced into the axial sinus,
which runs for a short distance in the body-wall before it communicates with the stone-
canal. The presence in Holothuriaus of an axial sinus opening, on the one hand, into
the stone-canal, and on the other to the outside by means of the pore-canal, has been
described in two cases only once by Bury (2) and once by Ludwig (6). As Bury's
observation, however, has been very generally questioned, and as Ludwig, though
noting the presence of these structures, does not attach any very great weight to them,
the present discovery becomes of extreme interest and importance, as a confirmation
which places the matter beyond doubt. As has been noted above, in the younger
embryos the pore-canal cannot be traced throughout its whole length, but this is in all
probability due to bad preservation.
A large Polian vesicle is now seen in the left dorsal interradius, a position which
Ludwig (5) says it occupies in some Cucumariae, though he does not mention C. crocea.
Tube feet have made their appearance, and we now find four arising from each radial
canal, though they have not as yet reached the surface.
The state of the preservation prevents any very minute study of the nervous
system, but one can readily see the well-developed nerve-ring and the five radial
nerves, as well as the branches to the tentacles.
The epi-neural ring and canals are also clearly seen, and we find that the ring also
gives off branches which accompany the tentacular nerves.
Of the condition of the mesenteries little can be made out. A bunch of cells in
the dorsal mesentery close to the stone-canal may very possibly be the rudimentary
gonad, but of these points it is impossible to be sure.
In conclusion, it may be said that the collection, though small, has yielded many
interesting points. Chief among these may be mentioned the discovery of two
pelagic Echinoderm larvse within the Antarctic Circle, and of the brood-pouch in
Cucumaria crocea. The first are important because they disprove the generally
accepted theory that none of the Antarctic Echinoderms have free-swimming larvoe,
the second because it fills a gap in our knowledge of the life-history of a well-known
form. The well-marked axial sinus and pore-canal in the Cucumaria embryo is also
ECMiXODEKM LAKV.H.
a point of importance, since it gives positive evidence of the existence of these
structures among the Cucunmm.Uu, a fact which has been asserted, hut never
conclusively proven.
Zoological Laboratory, MeGill University, April 1st, I!>o7.
LIST OF PAPERS QUOTED.
1. DELI..^ J.--" E rhinoderms " in K,port on the collections of the ' Southern Cross," London, 1902,
2. IJniY, II.-" Studies in the Embryology of the ly:hinodmns." Q.J. Micros. Sri V ol 29 fi 8 891
It- 09-44'J.
3. CI.AKK, II. L-Synapta vivipara : a contribution to the morphology of Erhiuodermata," Boston
Memoirs Soc. Nat. Hist., 5 (mis), pp. ;,;;_*, 5 p l s .
4. HKXI.ERSON E. H --"Some observations on the development of an Asterid with large yolky eg**
from the Franklin Islands," Ann. Ma- Xat. Hist, vol. 10 (I'll,;,)
s, > M., * Abth.,
6. Lu.-"Zur Entwicklnngsgeschichte dcr Holothurien." Sitzuugsber. JVenss. Akad. d. \Viss.
Jt>erliD, 1^91^ p. do;j.
-m W -" Devel P ment of Astt ' rina t'il'bosa." Q.J. Micros. St-i., vol. 38 (1896),
-
8. MAC_BHI,, E E W- -The Development of Echinoids," pt. l. -The larva, of Echinus miliuris and
E. esculentus." Q.J. Micros. Sci., vol. 4i> (1801)), pp. 835-S8M
9. MOETENSE-N, Tii " Die Echinodermenlarven der Plankton-Expedition d. Ilnmboldt-Stiftung "
iW. II. J., Kiel and Leipzig, 18'J8.
10. THOMSON, C. WYVILLK.-" The Voyage of the Challenger-the Atlantic," London, 1x77.
TABLE OF PLATES.
FIG. 1. Astern bramHi liell. x U>i. Youngest einl^ryo of brood.
ilG. 2. Oplnuplufrtis sp. n. ? x 100.
FIG. 3. Eehmopluteus mitim-tii-iia sp. n. x 100.
' 'M crocea Lesson, x 1. Adult female, showing openings of the two brood-pouches.
MTb MiM Lesson. < 1. Another view of the same specimen, with the body-wait cut
thiough so as to expose the interior of the brood-pouches
Fl %L n l:t:r; ;:;;" Lcsso "' x 15 '- YM emb ^ *- "- * <"< *-. -^b
F ''i I""""""" '""" LeSSO ' T 'e < through the a,,tc.,i,,r region of eml.rjo Ao.vn in
Preoral Lobe - -f
FlG.I. ASTERIAS BRANOTI.
Youngest efnbr^yo of brood
Fie 2 OPHIOPLUTEUS N SP 7
K/00
Openings of
' Brood pouches
Position of openings
*> shown in> Fiy 4.
.-Left Broad-pouch
Riaht Brand-pouch
FlG 3. ECMINOPLUTtUS ANTARC1I
X 1(10
Ventral
Tentacles.
FIG 5. CUCUMARIA CROCEA
Adult female to show
interior of brood -pouches.
x/
' . }', ',l~ru 7
Fic 4 CUCUMARIA CROCEA
Adult female to show
openings of brood-pouches,
x 7.
Part Can n I
, Axial Sin it *
. ^<"
Polian. Vesicle -
FIG 6. CUCUMARIA CROCEA.
Young embryo born after
capture of mother
x IZi.
' '
Ring Canal
" IE soph a y
FIG.?. CUCUMARIA CROCEA
Transverse section through
anterior region of embryo
shown in Fig 6
To be bound with plalr. [..irval l-icliiiioderius.
Antarctic (Discovery) Exp
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MYZOSTOMIDAE.
BY DR. RUDOLF RITTEI; v. STUMMER-TRAUNFKLS (Graz, Austria).
(1 Plate.)
ElNLEITDNG.
DAS von der ' Discovery ' heimgebrachte uud mir von Prof. F. Jcft'rey Bell in
liebenswiirdigster Weise zur Bearbeituug iibergebeue Myzostoma-MateTia] umfasst
nur eine relativ geringe Anzahl von Exemplaren, die sich auf zwei Species, eine
freilebende und eine encystirte, verteilen. Die erstere ist eine neue Art : M.
(iiitiirctii-nni, die k-tztere halte ich fur ideutiscli mit dem schon von Prof. Graff
beschriebenen M. cysticolum.
AVenngleich also die Ausbeute nur eine geriugfiigige Erweiteruug unserer
beziiglichen systematischen Kenutnisse zu bringen vermockte, so erscheint sie doch
in tiergeographischer Beziehung reclit benierkenswert : einerseits, weil die beiden
Arten aus dem siidlichsten bekannten Gebiete des antarctischeu Uzeans stammen,
aus welch' letzterem bislier noch nie Myzostomen heimgebracht wordeu sind ; *
und andererseits, weil dadurch ein neuer Beweis fiir den Kosniopolitisnius
des M. cysticolum geliefert wird. Diese Art kennt man nun von drei geographisch
ausscrordentlich weit von eiuander entfernten Fundorteu (tropiseher West-Atlantic,
Ostkiiste v. Japan, Ross-Meer iu der Autarctis), ein Befund, welcher durch das
hohe geologische Alter der Gruppe sowie durch die bekanntlich sehr uniformen
Lebensbediugungen des Stillwassers und der Ticfsee seine Erklarung findet.
Da, wie schon gesagt, das mir zur Bcarbeituiig iibcrgebene Material nur eine
geriuge systematische Ausbeutung zuliess, so beschriiukt sich die vorliegeudc Arbeit
hauptsiichlich auf die anatomische Beschreibung der beiden Arteu. In dieser Hinsiclit
vermochte ich insbesondere das M. cyxtic.olum genauer zu untersucheu, als dies bisher
* Das siidlichste Gebiet, aus welchera man bis jetzt Myzostomen kannte, niimlich die See um die Prince
Edward- und die Crozets-Inscln, liegt nocli in der Snbantarctia u. zwar nur wenig siidlich von der Nordgrenze
der treibenden Eisberge und der Sttdgrenze des Wcltverkehrs. Von liicr wurden durch die ' Challenger '-
Expedition folgende Arten gesammelt :
)[. i-niii/ireiiniim GmtT . . -3, pag. 42.
M. coronatinii dr.- ill' . . . 3, ., 60.
8tf!i'i-li >
I
1
6-0
7-5
Die Untersuchung erstreckte sich auf je ein in Xylol aufgehelltes Exemplar der
Classe a, d, e, i)
zuerst in lateraler Richtung an der Dorsalseite des Magens, um dann an dessen
seitlicher Circumferenz ziemlich vertikal nach abwarts zu ziehen. Nacb Umgreifung
des hintersten Hauptstammes der Darmverzweigung sehlangeln sie sick dann langs
der Ventralseite des Magens zur Kloake hin, in welche sie schliesslich von beiden
Seitcn her mit je einem spaltformigen Nephroporus ausniiindeu.
VERWANDTSCHAFTLICHE BEZIEHUNGEN zu ANDEREN MYZOSTOMA-ARTEN.
Die Griiude, welche Graff seinerzeit (3, pag. 22, Abs. 2) veranlasst haben, von
einer Untertheilung des provisorischeu Sarumel-Genus Myzostoma abzuseheu, bestehen
auch noch heute, da fiir die Hauptmasse der von ihni beschriebenen Arten noch
immer eingehende anatomische Untersuchungen fehlen, ohne die eine systematische
Die einzelnen Hodenfollikel sind von einander relativ weit entfernt und duroh mehr oder minder breite
Bindegewebspartien oder durch andere Organteile getrennt. Die Speruriducte sind im Allgeuieinen verlangert
und in einzelne Abschnitte (Vasa efferentia und deferentia, Samenblase, Ductus ejaculatorius), differenziert.
Der rnaunliche Geschlechtsporus ist ventralseitig, zumeist ziemlich weit nach innen vom Kb'rperrande und fast
regelmassig an der Spitze einer kegelformigen von der Basis des mittleren Parapodiurns nach aussen hin
vorspringenden Genitalpapille (Penis) gelegen.
(6) Compacter Hoden (" compact roundish glands occupying definite areas in the lateral part of the body " :
Graff, 3, pag. 11, abs. 1)
Typus : M. cysticolum (vorliegende Arbeit, Pag. 13-14).
Die Hodenfollikel sind auf eineui deutlich lungrenzten rundlichen Bezirk in den Seitenteilen des Korpers
concentrirt. Sie liegen enge neben einauder und werden nur durch diinne Bindegewebsbalkchen oder -lamellen,
nie durch andere Organteile getrennt. In der Mitte des Hodens bildet sich durch Platzen der dortselbst
gelegenen bindegewebigen Follikelwandungen ein centraler Sammelraum, in welchem sich die reifen Geschleehts-
producte anhiiufen. Dieser steht entweder vermittels eiues ganz kurzen unbedeutendeu Ductus ejaculatorius
oder auch direct mit der miirmlieheu Geschlechtsomiung in Verbindung. Letztere ist am Seitenrande oder in
unmittelbarer Nachbarschaft desselben gelegen. Eine Genitalpapille scheint nie ausgebildet zu werden.
(c) Den dritten von den zwei vorigen Modificationen principiell verschiedenen Typus der Hodenanordnung
hat Wheeler (8, pag. 247, 248) bei dem M. belli u. bei deni M. cryptopodium beobachtet.
Bei diesen zwei Arten soil die Gesammtmasse der Hodeufollikel dorsal von den Darmiisten in der mittleren
Korperregion gelegen und durch zwei dorsoventrale Septa in drei Lappen (einen medianen u. zwei laterale)
geteilt seiii. Die Hodenfollikel springen in den dorsalen Theil der Leibeshiihle (" Uterus ") vor, m welchen
auch die fertiggebildeten Spermien entleert werden, die dann offenbar (Wheeler spricht sich dariiber nicht aus)
durch die weibliche Geschlechtsoffuung (oder Kloakalofmung ?) ihren Weg nach aussen finden.
Bei der Isoliertheit dieser Beobachtung und bei der Wichtigkeit, welche dieselbe hinsichtlich der Leibes-
hohleiifrage sowie der Genese der miinnlichen Keiinzellen bei den Myzostomen besitzt, erscheint eine
Nachuntersuchung dieser beiden vou AVheeler nur oberfliichlich beschriebenen Arten dringend geboten.
Anhangsweise mochte ich hier noch die sogenannten subectodennalen Hoden erwiihnen, welche Nansen
(5, pag. 78 u. 79) bei M. gigas, M. gigantcum, M. graffi, u. AT. carpenteri beschrieben hat. Diese liegen als
vollkommen von dem normalen verzweigten Hoden getrennte Follikel dicht unterhalb des Integumentes. Sie
schemen dies liisst auch ihre eigentiimliche Structur vennuten (Nansen, 5, pag. 79, zeile 3-5) abgetrennte
und in Riickbildung begriffene Terminalfollikel des normalen Hodens zu sein.
MTZOSTOMTOAB. 7
Bearbeitung der Myzostoniiden undurchfiihrbar ist. Diescr Mangel lasst duller im
AUgemeinen vorderhand nur eine additionelle Einreihung einer ncucn Species in
diese Gattung geboten erscheinen. Im Speciellen wird man sich darauf beschranken
miissen die ueubeschriebene Form mit den wenigeu Arten zu vergleichen, deren
Organisation genauer bekannt ist und ihre Beziehungen zu ihnen festzustellen.
Von diesem Standpunkte aus betrachtet steht das M. antarcticum jencr Gruppe
von Arteu am niichsten, zu welcher das M. cirriferum (Gratf', 2, div. pag., Nansen,
5, div. pag., Wlu-el..r, 8, pag. 229-236 ; pag. 276, abs. 3), ferner das .]/. 'gigas, das
M. giganteum, das .]/. graft und das M. carpenten gehoren. (Die anatomische
BeschreilMin- fiir .lie vier letztgeuannten Arten hat Nansen 5, pa". G)-70 u. \\ mr
T f i \
geliefert.)
2. MYZOSTOMA CYSTICOLUM.
(Plate, fig. 2-10.)
Myzostoma cynticolum, Graff, Cliall. Rep. (1884), p. KG.
" Mts. Erebus u. Terror; 22. i. 1902 ; 500 fms."
Unter dem von der 'Discovery' heimgebrachten Crinoiden-Material fanden
sich an den Armen eines Exemplares von Anted on adriani Bell zwei Cysteu vor,
\\-elche eine Mi/;,>.stoma-Ait beherbergten, die ich fiir identisch mit dem M. cysticulum
Graff (3, pag. 66-68) halte.
HlSTORISCHES UND KRITISCHES.
Diese cystenbewohnende Species wurde von dem genannteu Autor naeh einer
Serie von Exemplaren aufgestellt, die aus dem westlichen Gebiete des tropischen
Atlantic's (" Cabo Frio," Brasilien ; " Insel Grenada," kleine Antillen) stammten. Als
\\ irt derselben fuugierte durchwegs Actinomttra meridwruili* var. cm-inata P.H.C.
Graff hat aus Griinden, welche er in der Eiuleitung zu seinem ' Challenger '-
Report (3, pag. 22, abs. 2) auseinandersetzt, bei der Beschreibung der lietreffenden
Art fast ausschliesslicli uur ihre aiisserlich erkennltaren Merkmale sowie ihre
biologischen Eigentiimlichkeitcn beriicksichtigt. Seine Angaben iiber die innere
Organisation clieser Form beschranken sich auf eiuige die Darmverzweigung sowic den
Genitalapparat betrcffende Befunde. 01)\vohl sich dieselben durchwegs als richti^
beobachtet erwiesen habeu, so hat sich inzwischen doch herausgestellt, dass jene
iiuter ihnen, \velrhe sich auf die Geschlechtsorgane des sogenannten " Weibchcns "
beziehcn, von Gruff in Ubereinstimmung mit den damals noch herrschcuden
Ansichten iiber die Morphologic des weiblirhen Sexualapparutes missverstandlich
gedeutet \vi >rden sind. Dies gilt insbesondere von den angeblichen " Hodenrudimeuten,"
welche, vie spater (Pag. 27) dargeleot wei-deu wird, als die functionierenden Ovarien
aufgefasst werden miissen.
Seit dem Erscheinen der ' Challenger '-Publication ist das J/. cy*t'm>htw, suwcit
mir bekannt, uur uoch eiumal und zwar an der pacifischen Kiiste von Japan in einer
DR. RUDOLF RITTER v. STUMMER-TRAUNFELS.
Armcyste von Antedon discoidea P.H.C. angetroffen worden. McClendon, welcher
die beiden in der Cyste vorhandenen Individuen untersucht hatte, beschrieb dieselben
als eine neue Varietat : M. cysticolum var. orientale (4, pag. 120-121), und zwar mit
der Begriindung, dass :
(a) die Wirtsformen der Graff schen und jene der japanischen Exemplare
eine verschiedene systematische Stellung einnahmen ;
(b) die betreffenden Fundorte in tiergeographischer Hinsicht differierten, und
(c) die japanischen Exemplare gegeniiber den Graff'schen Typen einige
unwesentliche morphologische Unterschiede zeigten.
Abgesehen da von, dass das McClendon vorgelegene Material (eine einzige
Cyste !) an Zabl weitaus zu gering gewesen 1st, um die Aufstellung einer besonderen
Varietat zu reclitfertigen, so halte ich die hierfiir angezogenen Argumente des
genannten Autors nicht fiir eiuwandfrei.
(ad a) Beziiglich des ersten derselben erinnere ich an die durch Graff (3,
pag. 21) bekannt gewordene Tatsache, dass ein und dieselbe Myzostoma-Axt auf
1-4 Criuoidenspecies vorkommen kann (in einigen Fallen sogar auf Vertretern
zweier Genera). Graff, gewiss der erfahrenste Kenner der Myzostoma-Systematik,
hat sich jedoch nicht veraulasst gefiihlt, auf Grund dieses Umstandes verschiedene
Varietaten der betreffenden Species zu unterscheiden.
(ad l>) Was die von McClendon hervorgehobeue zoogeographische Differenz der
Fundorte anbelangt, so diirfte dieses Argument gerade bei den Myzostomeu nur
mit grosster Vorsicht anzuwenden sein, da das Verbreitungsgebiet dieser Parasiten
noch sehr wenig bekaunt ist, aller Voraussicht nach aber mit jenem ihrer Wirte
zusammenfallt, von welchen weitaus die meisten sich als charakteristische Vertreter
der einen cosmopolitischen Charakter besitzenden " Stillwasserfauna " erwiesen habeu
(Doflein, 1, pag. 251 u. pag. 272-273). I in iibrigen kann man der tiergeographischen
Differenz der Fundorte als solcher allein keine Bedeutung fiir die Aufstellung von
Varietaten beimessen, wenn sich nicht die verglichenen Localformen auch in
morphologischer oder in biologischer Hinsicht als solche documentieren. Diese
Bedingung erscheint meines Erachtens nach bei den Exemplaren McClendon's nicht
erfiillt,
(ad c) Die Besonderheiten, durch welche sich die letzteren vor den Graff'schen
Typeu auszeichnen solleu, beschranken sich auf die bedeutendere Grosse sowie auf
den etwas differenten Bau der Cyste, ferner auf die hervorragenderen Dimensionen der
Parasiten selbst, schliesslich auf eine geringfiigige formale Differenz der sogenannten
" Mannchen."
Die Grosse der durch das M. cysticolum hervorgerufenen Cyste hiingt in erster
Linie von der Grosse des ihr Lumen fast vollstandig ausfiillenden " Weibchens " ab.
Da dieses jedenfalls von kleineren Dimensionen zu grosseren heranwachst und auch
sonst in dieser Beziehung individuelle Verschiedenheiten zeigen kann, so werden
MTZOSTOMIDAE.
demnach auch verschieden grosse Cysten vorkommen.* Da wir nun derzeit iiber die
maximale Grosse, welche das M. cysticolum erreichen kanu, noch vollkommen im
Unklaren sind, so erscheint es vorderhancl untunlich eine iiber die bekaunten Masse
hinausreichende Cystengrb'sse als Kriterium zur Aufstellung einer neuen Varietat zu
verwenden.
Ebensowenig geeignet sind zu diesem Zwecke auch kleinere, den formalen Typus
der Cyste nicht beeintrachtigende Differenzen im Baue derselben. Deu physiologischen
Anlass zur Bildung einer Cyste gibt allerdings das Myzostoma, da es entweder durch
seine Bisse oder warscheinlicher durch Absonderung toxischer Substanzen, wie es z. B.
die Excretstoffe sein kb'nnen einen localeu Reiz auf den Wirtskorper ausiibt, auf
welchen dieser dann durch Wucherung des geschiidigten Gewebes reagirt. Da wir aber
den hauptsiichlichsteu, das ist den formativeu Anteil an der Cystenbildung der
Wirtsform zuschreiben mussen, so werden wir kleiue, unwesentliche Veranderuugen
in der Morphologic der Cyste iu erster Lime als den Effect einer iudividuellen
Verschiedenheit des Wirtes zu betrachteu haben und erst in zweiter Linie, wcun sich
eine solche ausschliessen lasst, an eine Verauderung des reizenden Substrates denken.
Voraussetzung fiir diese Erwaguug ist uatiirlich, dass sich die Cysten an verschiedenen
Individuen ein und derselben Wirtsspecies vorfinden.f In weit hoherem Grade
niusseu wir jedoch die Wirtsform fiir etvvaige Veriinderungen im Bau der Cyste
verantwortlich machen, wenu ein und dieselbe Myzostoma-Att verschiedene Wirts-
species befallt. Es kommt denn eben die artliche Difi'erenz der letztereu auch im Baue
der auf ihnen entstehenden Cysten zum Ausdrucke. Von diesem Gesichtspunkte aus
muss auch die von McClendon beschriebene Cyste beurteilt werden. Sic fallt
luiuptsachlich dadurch auf, dass sie im Gegeusatze zu den mit einem einzigen Porus
versehenen Exemplaren Graff's, zwei derartigc Offnuugen und zwar eine an jedem ihrer
Schmalenden besitzt.
Als morphologischeu Unterschied zwischen den von Graff und deu von McClendou
untersuchteu Parasiten erwahnt der letztgenannte Autor vor allem die bedeutendere
Grosse seiner Individueu. Diese kaiiii jedoch ebensogut durch individuelles AVachstum
oder durch besondere Eruahrungsverhaltnisse bediugt sein. Weiters hebt dersclbe
Autor die Beobachtung hervor, dass bei dem mannlich fuuctiouierendeii (" small
individual") seiner Exemplare die zwischen der Korperperipherie und der Eeihe der
* Dies ist auch bei den von Graff untci-suchten Cysten der Fall gewesen. Vergleioht man die auf die
Cystenliingc beziiglichen Angaben des genannten Autors mit dem von McClendon gegebenem Masse, so ergiebt
sich, dass die Grossendifferenz zwischen dem kleinsten und dem grossten Exemplare Graffs genau dieselbe ist,
wie zwischen dem letzteren und dem Exemplare McClendon's :
/ kleinstes Exemplar . 1 mm.
Graffscher Typ : ( ^..^ . 3 mm. < Differcnz = 2 mlu -
McClendon's Exemplar ... .5 rum. "^ ^ mm-
t Die Cysten von M. cysticolum, welche Graff untersucht hatte, fanden sich samrntlich an Actinomrlr.i
mn-idionalis var. carinata P.H.C. Obgleich einige von ihnen kleinc Verschiedenheiten in ihrer Gestalt und
ihrer Lage am Crinoidenarmc aufwiesen, so ist cs dem genannten Autor doch nicht beigcfallen, die Insasscn .In-,
sulbon uls dififerente Varietaten dor typisL-hcn Form v.\\ beschrcibcn.
10 DK. RUDOLF HITTER v. STUMMER-TRAUNFELS.
Parapodien gelegene Randpartie (" edge of the disc ") sich welter gegen das Centrum
der Bauchseite bin ausdehnte, als dies bei der von Graft' gelieferten Abbildung (3, taf.
xni. fig. 5) der Fall ware. Diesem Umstande vermag ich durchaus keiue Bedeutung
zuzumesscu. Jeder Beobachter, welchem zaMreiche Exemplars eiuer Myzostoma-Sipecies
zu Gesicht gekommen sind, weiss wie modulationsfakig hinsichtlich seiner centripetalen
Ausdehnung gerade dieser Korperabschnitt 1st, dessen Areale durcli starkere Con-
tractionen der " bauchstandigen Muskelmasse" (Graff, 2, pag. 41-42) vergrossert, durch
Entspannung derselben verkleinert werden kann.
Von den eben dargelegten Erwagungen ausgehend glaube ich also dass eiu
zureichender Grund zur Aufstellung einer neuen Varietart fiir die von McClendon
beschriebenen Exemplare des M. cysticolum uicht vorliegt. Ich hoffe durch die
Beschreibung der von der ' Discovery ' heimgebrachten Individuen, fiir welche alle die
von dem obgenannten Autor zu Gunsten seiner Auffassung ins Treffen gefiihrteu
Argumeute in gleicher Weise, ja vielleicht in noch hoherem Ausmasse passen, wie fiir
die von ihm untersuchteu Exemplare, einen weiteren Beweis fiir meine Ansicht
erbringen zu konnen.
Von der inneren Anatomie des M. cysticolum beriicksichtigt McClendon im
wesentlichen nur den Geschlechtsapparat. Er berichtigt und erganzt die beziiglichen
Angaben Graffs und bestatigt auch fiir die genannte Species das Vorhandensein von
weiblichen Gonaden bei einem jeden der beiden, friiher als " Weibchen " und als
" Zwergmannchen " unterschiedenen Individuen des die Cyste bewohnenden
Parasiten-Parchens.
BESCHREIBCNG DER ' DISCOVERY '-EXEMPLARE.
CYSTEN.
Die beiden mir zur Untersuchuug iibergebenen Cysten waren als eiformige
Auftreibungen des Wirtsintegumentes ihrer Lange nach dem Crinoidenarme etwas
seitlich von dessen Ambulacralrinne angeschlossen (Fig. 3).
Sie massen 6 '2 mm. beziiglich 5 mm. in der Lange uud 3'2 mm. beziiglich 2'5
in der Breite. Die Wandungsdicke der grossereu Cyste betrug durchschnittlich
0' 15 mm.,- war also verbal tnismassig viel geringer als bei den Cysteu, welche Graff
beschrieben hatte (3, pag. 67, zeile 9-10 ; pag. 68, abs. 2 ; taf. xni., fig. 4).*
An jedem Cystenende zeigte sich in gleicher Weise wie bei den Exemplaren
McClendon's eine ins Innere fiihrende Offnung, von welchen die von der Muudscheibe
des Wirtes abgewendete in beiden Fallen viel grosser als die andersseitige war und
sich vor dieser ausserdem noch durch eine schwache Aufwulstung ihres Randes
* Dies hiingt mit der ansehnlichen Grosse der antarctischen Myzostoma -Individuen zusarnmen ; denn eine
Cyste, welche ein grosseres Myzostoma beherbergt, wird auch eine ausgedehntere, daher relativ dtinnere Wandung
besitzen als eine solche, welche einen kleineren Parasiten umschliesst. Auch unter den von Graff beschriebenen
Cysten besass die kleinste (3, ' Blake '-Exemplar, pag. 68, abs. 2) eine derbere "Wandung als die iibrigeu.
MYZOSTOMIDAE. 1 1
auszeichuete. Die kleinere der Cystenoffnuugen wurde erst daim sichtbar, wt-nn man
die in der Cyste enthaltenen Parasiten entfernt liatte, so dass Licht durch den
punktforinigen Porus fallen konnte.*
A.USSERE MORPHOLOGIE DER PARASITEN.
Jede Cyste umschloss, wie bei den Exemplaren Graft's u. McClendon's, je ein
grosseres weiblich- und eiu kleineres niauulich-functionicrendes Individuum. In
folgendem will ich nach dem Beispiele der friiheren Autoren der bequemeren
Ausdrucksweise halber das erste als " Weibcheu," das letztere als " Mannchen "
bezeichnen, trotzdem beide, wie spater nachgewiesen werden soil, als morphologische
Hermapliroditen organisiert sind.
A. Das Weibchen (" female " : Graff, 3, pag. 67 ; " large individual " : McClendon,
4, pag. 121), (Fig. 2 u. 4), war der Lange nach in der Cyste gelagert, deren Innenraum
es dabei so vollstaudig ausfiillte, dass seiu Hinterende in ihm keinen Platz mehr fand
und aus der grosseren der beiden Cystenoffnungen etwas herauszurageu gezwungen war.
Auch der Breite nach entsprach das Cystenlumen nicht den Dimen.siouen des Tieres,
weshalb dessen laterale Korperpartieu, wie dies schon die beiden vorgenannten Autoren
geschildert hatten, dorsalwarts aufgerollt waren, so dass die Seiteuriinder in der
Medianebene nicht bloss zur Beriihrung sondern auch stellenweise (am Hinterende und
in der Mitte des Korpers) zur Uberlagcrung gelangten (Fig. 7 u. 8).
Wie ein Blick auf das in Fig. 4 von der Veutralseite her abgebildete Weibchen
zeigt, ist der (in der Fig. nach oben gerichtete) Hinterrand desselben medial
ziemlich tief eiugezogen. An dieser Stelle springen die benachbarten hintersten
Abschnitte der aufgerollten Scitenrander nach riickwarts vor und legen sich zugleich,
wie dies aus Fig. 6 zu erkennen ist, nach Art von zwei einander tiberdeckenden
Fliigelklappeu von seitwarts und hiuteu her, iiber den trichterformig rnvcilrrten
After (aiJ), sowie iiber die unmittelbar olterhalb von diesem gelegcne wcibliche
Geschlechtsb'ffuung (6).
Oberhalb von diesem miindet der dem sogenanuten Uterus des Weibchens
entsprechende mediale Dorsalabschuitt des Coeloms durch eine kleine Offuung aus, die
wir demgemass als ein Homologou der weiblichcn Geschleclitsiill'nuug betraclitm
miisscu (Fig. 9, >H). Die beiden maunlicheu Genitalporen (Fig. 5, d<">), welclic in der
die mittlcreu Parapodien vcrbiudendeu Transvcrsanebeue gclegen sind, offuen sich
nicht, wie Graff uud McCleiidou fiir ihrc Exemplare augebeu, marginal, sondern auf
der Dorsalseite des Tieres, siud indess dem Korperrande sehr uahe geriickt. Von
diesem aus zieht zu eiuem jeden von ilmeu eine kurze rinuenformige Vertiefung des
Integuments, welche moglicherweise von den beiden obgenannten Autoren als Ductus
ejaculatoriua aufgefasst wordeu ist, woraus sich ihre von ineinem Befuude differierende
Angaben erklaren liesseu.
Eiii Excretionsporus ist auch beim Manncheu nachzuweisen. Er liegt wie beim
Weibchen median auf der Ventralseite dicht vor dem After (Fig. 10, nj>).*
* Wio aus dem Vorstehenden zu entnelimen ist, findcn sich simitlirli.- ;un Kiirpov des Weibchens
autretende ()ffnungcn auch iri gleichcr Lage brim Mannchcn vor, ein Vcrbaltcn, welches mit d.-m lliinveis auf die
spiiter zu bcsprechcnden uiorphologischcn uud biologischen Bcziehungen der beiden Qesohlechtsformen sclion
hier hervorgehobeu sei.
U 2
14 DR. RUDOLF RITTER v. STUMMER-TRAUNFELS.
INNERE ORGANISATION.
AA 7 ie in der Lage der Korperoffuungeu so besteht auch hiusichtlich der iuneren
Organisation beim Mannchen uud beim Weibchen trotz deren aiisserlichen Grossen- und
Formdifferenz eine principielle Ubereiustimmung. Diese aussert sich vor allem am
Sexualsysteme, das bei jeder der zwei Geschlechtsformen durch zwei Apparate, einen
niannlichen und eiueu weiblichen, vertreten ist, von welchen aber beim Mannchen nur
der erstere, beim AA r eibchen uur der letztere reife Sexualproducte liefert. Die
Differenzen zwischeu den Geschlechtsorganen des Mannchens und jenen des Weibcheus
sind also hauptsachlich functionelle uud dies gilt irn Grunde genomnien auch fiir alle
iibrigen Organe. Diese erscheinen bei den zwei Geschlechtsformen nach demselben
Plane angelegt uud unterscheiden sich nur durch ihren Ausbildungsgrad, der seiner-
seits von ihrer Functionsintensitat abhangt. Diese wird wieder durch die physiologische
Sexualitat des betreffenden Individuums bestimnit*
Aus diesem Grunde halte ich es fiir zweckmiissig in Nachfolgendem die einzelnen
Organe gemeinsam fiir beide Geschlechtsformen zu beschreiben und zwar in folgender
durch obigen Gedankengang begriindeten Eeiheufolge :
(1) Weiblicher Geschlecht*apparat.
(2) Mdnnlicher Geschlec.htsapparat.
(3) Ernahrungsapparat.
(4) Excretionsapparat;
Auf eine Schilderung des Nervensystems beabsichtige ich aus dem Grunde nicht
einzugehen, weil sich dasselbe in seiner groberen Morphologic nicht von jenem der
anderen Myzostomen unterscheidet, und weil andererseits eine Untersuchung seines
feineren Aufbaues, der bei dem Maimchen und dem AVeibchen moglicherweise
interessaute Differenzeu vergeben hatte, wegen der quantitativen und qualitativeu
Unzulauglichkeit des Materials aussichtslos schien.
AVEIBLICHER GESCHLECHTSAPPARAT.
Derselbe wird bei den Myzostomen, wie bekannt, von einem Teile der Leibes-
hohle (dem Uterus mit seinen Diver tikelu), und weiters von bestimnit localisirten
Wucherungen seiner peritonealen Auskleidung (den Ovarien) gebildet ; (vergl.
diesbeziiglich : Stummer, 6, pag. 574 u. ft'.).
Mannchen : Bei diesem erscheint der zu den weiblichen Geschlechtswegen
umgebildete Leibeshohlenanteil, ini Gegensatze zu den ausschliesslicB mauulich
* Durch die physiologische Unisexualitat werden abgesehen vom entsprechenden Geschlechtsapparate
hauptsachlich die Stoffwechselorgane (Darni- u. Excretionssystein), in ihrein Ausbildungsgrade beeinflusst, und
zwar in der Weise, dass die Functionsintensitat dieser Organe beim Weibchen gegeniiber jener beim Mannchen
erheblich gesteigert erscheint. Der miitterliche Korper, welcheni die Ausbildung der grossen und dotterreichen
Eier obliegt, besitzt eben ein weit grosseres Stoffwechselbediirfnis als der vaterliche, der durch die Production
der winzigen Spermien diesbeziiglich nicht so sehr beansprucht wird (vergl. Pag. 20 und 21).
MYZOSTOMIDAE. 15
functiouiercndeu Individuen anderer Myzostoma-Alten, bei welchcu sich bisher nur
sparliche Andeutungcn dieser Coelompartie nachweisen liesseu, wohl entwickelt ; er
1st zwar raumlich nicht so ausgedehnt, wie beim Weibchen, zeigt aber dicsclbe Unter-
gliederung wie bei dicsem. Er setzt sich aus einem dorsalcn, zwei seitlichen und
eiueni veutralen Abschnitte zusammen.
Der erste entspricht der bei anderen hermaphroditisch functionierenden My:ostnin-
Arten gemeiniglich als Uterus bezeichneten Leibeshohlenpartie. Er stellt ein medianes,
dorsoventral abgeplattetes Rohr dar, welches ungefahr oberhalb der Mageninitte
beginnt und von hier aus unter allmahliger Verschmiilerung nach riickwarts bis zur
weiblichen Geschlechtsoft'uung verlaiift, durch welche es ausmiiudet (Fig. 9, ua + ul>]
Bald liinter seinem verbreitcrten Vorderende erscheint seine Seitenwauduug rechts und
links in einen kurzen Zipfel ausgezogen, an dessen Lateralende sich je ein Nephrostom
(Fig. 9, ns) offnet.
Vorne steht der Uterus jederseits durch einen kurzen und breiteu Querast mit den
beiden Seiteuabschnitten der Leibeshohle in Verbindung. Diese verlaufeu zunachst in
dorsoventraler Richtung langs der rechteu und liuken Seitenwand der vorderen
Magenhalfte nach abwarts, um sodann beiderseits nach Umfassuug der Wurzeln des n.
und in. Hauptdarmastes in den Ventralabschnitt der Leibeshohle iiberzugeheu.
Letzterer besteht aus zwei symmetrischen, durch cine schmale suboesophageale
Quer-Commissur verbundeue Halfteu (Fig. 10, vl). Eiue jede von dieseu beginnt als
ein langlicher, unter und seitwarts von den lateralen Magenpartien gelegener
Hauptraum, von dem weiterhin drei Uivertikel gegen die Hauptdarmaste ausgehen,
welche die letzteren unter entsprechender Teilung langs ihrer ganzen Verzweigung
ventralseitig begleiten (Fig. 10, 1,2,3).
An der Wurzel des ersteii uud dritten dieser Divertikel liegt nun je ein aus einer
localeu Wucherung des Peritonealepithels hervorgehender Zellhaut'cn, welcher von
seitwarts und von unten her in den Hauptraum einspiiugt. Im Ganzen sind daher
vier derartigc Gebilde vorhanden (Fig. 10, o). Hirer Lage und ihrem histologischen
Aufbaue nach miissen wir sie in Ubereinstimmung mit McClcndon (4, pag. 121, abs. 3)
mit den von Nansen (5, pag. 78, abs. 4) bei anderen Myzostomen entdeckten
" problematischeu Organen " homologisieren, die spatcr von Wheclor (7, pag. 178,
abs. 2) als die echten Ovarien erkannt worden sind. Dass sic bcim ^liiunchen von
M. cysticolum tatsachlich als solche friiher odor spater functiouieren lasst sich jedoch
nicht uachweiseu. Sie machen im Gegensatze zu den productiven Ovarien )
augehorenden Hoden liegt eiu deutlicher centraler Sammelraum, der direct mit der
Geschlechtsoffnung communiciert und sowohl Spermien als auch cine Meuge von
sich nicht weiter eutwickelndeu, viellcicht auch degenerierten Spermatocytcn
(Stummer, 6, pag. 583) enthalt.
Beim Weibchen fand ich die beiden Hodeu genau in derselbcn Lage und in der
formell gleichen Ausbildung, wie beim Mannchen. Sie liegen jederseits in der ]\Litte
des aufgerollten Seiteurandes, dicht nel)en dem mannlicheu Geschlechtsporus, und
stellen eine deutlich vom iibrigen Korperparenchym abgegrenzte Masse von reticu-
larcm Bindegewebe dar, dcssen Intercellularraume zu grosseren Blasen ausgedelnit
sind (Fig. 7, t). Die letzteren flicssen in der Mittc dieses Complexes zu einem dicht an
die mannliche Geschlechtsoffnuiig anschliessenden centralen Sammelraum zusammen.
Wahreud aber die Intercellularraume beim Mannchen dicht mit Keimzellen bezie-
hungsweise mit deren Teilungsproducten crfiillt sind, cuthalteu sie l)eim Weibchen
relativ uur wenigc Zellen. Von dicseu lasscn sich zweierlei Formen uuterscheiden :
Die eincu gleichen auftallend den von Graff (2, pag. 64, Abs. 1) als " Ruudzelleu "
* Benierkenswort ist, dass schon Graff die Ovarien bei M. cysticolum gesehen hat. Nur wunlcn sic von
ihm zufolge der damals noch herrschenden Anschauung, dass die Vcrzweigungcn des Uterus (Leibeshohle uucl ilire
Divertikel) das Ovarium bei den Myzostoinen ilarstellten, als rudinicntare Hoden gedcutet (Gruff, 3, pag. 67,
abs. 2). Nach der Abbildung, welche der genannte Autor von diesen Gebilden gelicfert hat (3, taf. xm., fig. 4, t),
erscheint es jedoch zweifellos auch McClendon ist dieser Meinuug (4, pag. 121, abs. 3) dass sic mit den oben
gcschilderten echten Ovarien idcntificicrt \verden miissen. Die Prioritiit, die wahren .1A/^ns/oia-Ovarien zuerst
gesehen zu haben gebiihrt daher Graff, und nicht Nansen, dern sie bisher zugeschrieben worden ist.
18 DR. RUDOLF HITTER v. STUMMER-TRAUNFELS.
beschriebeuen freien Bindegewebselementen (Stammer, 6, pag. 589, letzt. Abs.). Die
auderen machen jedoch vollstandig den Eindruck von degenerierten Zellen. Sie
besitzen einen relativ kle.ineu uuregelmassig gestalteteu Kern, der sich mit Haema-
toxylin intensiv farbt und dabei neben einem homogenen Aussehen auch ein
eigentiimliches Lichtbrechungsvermogen entwickelt. Ihr Cytoplasma 1st nur in sehr
geringer Meuge vorhanden und im giinstigsten Falle als ein kaum tingierbarer,
sehmaler, den Kern umhullender Saum zu erkenneu. Von Spermien fand sich
innerhalb des ganzen Bindegewebscomplexes keine Spur. Der centrale Sanimelraum
desselben entliielt dagegen bei jedem der mir vorgelegeuen Exemplare eine feinkornige
oft fadig angeordnete und auch aus der Geschlechtsoffhung dringende Masse, in
welche zahlreiche der oberwahnten anscheinend degenerirten Zellen eingebettet
waren.
Ich stehe nicht an, die eben geschilderten Organe auf Grund ihrer Lage uud ihrer
Ausbildung als functionslose Hoden zu betrachten, die jenen des Mannchens vollstandig
homolog sind. Es ist infolge des ungeniigenden Materials selbstverstandlich sehr
schwer zu entscheiden, ob sie als Riickbildungen von mannlichen Gonaden aufzufassen
sind, die in einem friiheren Lebensalter des Individuums functioniert haben, oder ob
sie auf einern friihen Entwickelungsstadium stehengebliebene, niemals zur sexuellen
Betatigung gelaugte Apparate darstellen. Meiner Meinung nach dtirfte die erstere
Annahme die zutreffende sein, insbesondere deshalb, weil sich ahnliche Verhiiltnisse
(Proterandrie mit spaterer Riickbildung des Hodens) bei zahlreichen frei lebenden
und auch bei einigen entoparasitischen Myzostoma-Axten, mit Sicherheit haben nach-
weisen lasseu (Wheeler, 8, pag. 288-289). Schliesst man sich dieser Ansicht an, so
konute man von den oberwahnten, in den Intercellularraumen vorhandenen Zellen,
die von mir als auscheinend degenerirt bezeichneten Elemente als riickgebildete
Keimzellen, jene aber, welche den " Rundzellen " Graff's ahneln, als secundar in die
Follikel eiugewanderte freie Biudegewebszellen auffassen. Die feinkoruige, im
Sanimelraum enthaltene Masse wiirde danu auf Reste von abgestorbeneu Spermien
oder von zugrundegegangeueu Sperm atocy ten etc. zuriickzufiihren sein.
Derartige fuuetionslose Hoden sind weder von Graff noch von McClendon bei den
von ihnen untersuchten " Weibchen " beschrieben worden. Ich glaube aber, dass diesbe-
ziiglich ein Uberseheu von Seiten der genannten Autoren vorliegt, da diese Gebilde bei
dem vollstandigen Fehlen von tatigen Keinizelleii und von Spermien nicht sofort als
Hoden zu erkennen sind. Graff diirfte ihnen schon deswegen keine weitere Beachtung
gescheukt haben, weil er iiberzeugt war, in den von ihrn beim Weibchen beschriebenen
"collections of small cells closely resembling the immature testicular follicles . . . ."
(3, pag. 67, abs. 2) Hodenrudimente vor sich zu haben. Dass diese seine Auffassung
eine irrtiimliche war, uud dass diese Gebilde die Ovarien darstellen, habe ich schou
friiher (Pag. 17) erwahnt.
MYZOSTOMIDAE. 19
ERXAHRUNGS-APPARAT.
Derselbe ist nach dem schon von den iibrigen Myzostomen her bekannten Typus
gebaut.
Wegen der viel einfacheren Verhaltnisse, welclie er beim Mdnnchen aufweist, sei
er zuerst an diesem geschildert : Beziiglich YOU Muud und After vcrweise ich auf die
schon friihcr (Pag. 13) gelieferten Angaben. Der lange, eylindrische, an seinem
Vordereude nicbt mit Tastpapillen ausgeriistete Pharynx (Fig. 9, bin), steht
vermittela eines kurzeu " Oesophagus " mit dem Magen in Verbiudung. Dieser
(Fig. 9, mi/), zerfallt durch eine seichte, querc Einschniiruug in eineu vorderen
\veiten uud einen hiuteren eugeren Abschnitt. Vom crsteren gehen ventral jederseits
drei Hauptdarmaste (Fig. 9, I, II, ill) ab, welche sich nach kurzein Yerlaufe teilen
und mit ilm-n wcuig zahlreicheu, stumpf eudigenden Divertikeln bis zur Randzone des
Korpers reicheu. Von dieaen drei Hauptdarmasten verzweigen sich jederseits die zwei
ersten in der vor dem Hoden gelegenen, der dritte in der liiuter diesem befindlichen
Korperpartie. Der hintere Magenabschmtt ist kleiner als der vordere uud geht
vermittels eiues nur am oontrahirtcn Tiere (Fig. 5) sichtbaivu Absatzes in das
rohrenformige dorsoventral compresse Eectum (Fig. 9, r) iiber. Die Afteroffuung
kann durch einen sich aus dem Hautmuskelschlauche ditiereuzierenden schwachen
Spliinktcr gescliLisscn \vcrdrn.
Der gesamte Verdauungsapparat ist bcim .Mauuchen entsprechend dessen
geriugem Ernahrungsbedurfnisse recht einfach gebaut. Seiue cinzeliii-u Abschnitte
setzen sich aus einer weit geringcreu Auzalil von Zellelcmcuten zusammeii \vic jene des
Weibchens. Auch seine Muscularis ist nur schwach entwickell und an manchen
Stellen iiberhaupt uicht nachzuweisen.
Beim Weibchen hingegeu dessen Stoffwechselbediirfnis infnl-i' der reichlichen
Eierproduction uugeniein gesteigert ist, zeigt der Eruiihrungsapparal eine vicl
machtigere Ausgestaltuug : vor allcm ist der Magm (Fig. (>, 7, 8, mi/) relativ
bedeutend voluminoser wie beim Mannchen. Er wird seiner ganzeu Ausdehuung nacli
sanit dem Oesophagus (Fig. 6, c) von einem sehr auftalleudeu Wimpercpithd
ausgekleidet, desseu zahlreiche uud dicht stehende Cilien wegen ihrer uugewohnlichen
Lange bemerkenswert siud (Lange der Epithelzellen : 0'OG8 m., jeue der C'ilien :
O'l m.). Die Hauptdarmaste sind distahvarts ausserordentlich reich verzweigt.
Ihre Divertikel liegen vollkommen in die latcraleu Leibeshohlensacke eiugebettet
(Fig. 7 u. 8, (In), uud verbreiten sich in grosser Xahl iiislpfsoinlrrc iimcihalli <\>T
aufgerollten Seiteupartien des Korpers, wo sie sich mit ihreu Terminalenden bis
kuapp unter deu Hautmuakelschlauch erstn-rken. Das Hiutrrcndc des Magens
erscheint scharf von Eectum (Fig. 6, r) abgesetzt. Das letzterc verliiuft in zicinlii-h
gerader Richtuug bis zuni After. Es wird von einem Wimperepithel ausgekleidet,
das in radiiir in sein Lumen einspriugcude Langsfalten gelegt ist. Eim-
durch aiissere Ring- uud iiinere Langsfaserii hergestellte Muscularis fmdet sich
20 DR. RUDOLF EITTER v. STUMMER-TRAUNFELS.
sowohl am Magen als auch am Rectum. Sie erscheint insbesondere am crsteren
kraftig ausgebildet ; an letzterem ist sie jedoch schwach entwickelt, ihre beiden
Schichten werden aber hier noch durch eine periphere dritte Lage vervollstandigt,
die aus locker vcrfilzten Fasern besteht. Ein besonders differenzierter Sphinkter
findet sich weder an der Grenze zwischen Magen und Rectum, noch um die
Afteroffnung. Fiir die letztere wird die Function eines solchen wohl von den benach-
barten Partien des Hautmuskelschlauches iibernommen. Ausserdem erscheint aber
der After fur gewohnlich auch noch durch die friiher (Pag. 11) beschriebenen
Analklappen verschlossen.
NEPHRIDIEN.
Dieselben beginnen jederseits mit einem sich in dem vorderen Uterusabschnitt
ofihenden cylindrischen Nephrostom (Wimpertrichter) (Fig. 8, 9, ns). Dieses setzt
sich in einen engen, beim Mannchen kiirzeren, beim Weibchen, wegen dessen
bedeutender Dicke, laugereu Nephridialcanal fort, welcher in ziemlich geradem
Verlaufe langs der Seitenwand der hinteren Magenhalfte nach abwarts zieht (Fig. 8),
um sich etwa in halber Magenhohe plotzlich (diese Stelle ist in Fig. 8 und 10 mit *
bezeichnet) zu eiuer grossen, vielfache Windungen und Einschuiirungen zeigenden
Endblase zu erweitern. Die letztere (Fig. 6, 8, 10, nb) besitzt in ihrem proximalen
an den Nephridialcanal anschliessenden Abschnitt eine bedeutende Ausdehnung,
verschmalert sich aber distalwarts . allmahlig zu einem relativ langen, ziemlich gerade
nach hinten verlaufenden Canal, welcher mit jenem der auderen Kb'rperseite gegen die
Medianebeue convergiert und sich schliesslich mit ihm zu einem kurzen unpaaren
Endstiick (Fig. G, 10, nu) vereiiiigt, das durch den median, knapp unter der
Afteroffnung gelegeuen Excretionsporus (Fig. 6, 10, np) ausmlindet.
Wahrend der Nephridialcanal von eiuem verhaltnismassig niedrigeu Wimper-
epithel ausgekleidet ist, findet sich in der Endblase bis zuni Excretionsporus hiu ein
deutliches Driisenepithel, dessen hohe keulenformig gestaltete Elemente (aus Fig. 8 ist
die Lagerung und die relative Hohe derselben ersichtlich) der Wimpern entbehreu.
Eine besoudere Muscularis liess sich nur an der Endblase nachweisen, aber auch
hier ist sie sehr schwach ausgebildet uud besteht lediglich aus einer Ringfaser-
schichte.
So wie der Ernahruugsapparat, zeigeu auch die Nephridieu des Weibchens
gegeniiber jenen des Mannehens eine viel machtigere Entfaltuug, die sich insbesondere
in der relativen Lauge des Nephridialcanals sowie in der Grosse der Endblase aussert.
SEXTTALVERHALTNISSE TJND DIMORPHISMUS DER GESCHLECHTSFORMEN.
Graff (3, pag. 11-12) unterscheidet unter den von ihm beschriebenen cysticoleu
Myzostomen zweieiiei in Bezug auf ihre Sexualverhaltnisse divergente Formen und
zwar rein dioecische und rein hermaphroditische.
MYZOSTOMIDAE. 21
Zur ersteren Gruppe gehoren : M. tenuispinum, M. w[ll<-n\ag. <>7, Abs. 2), da er vermutete, dass bei ihr das " Weibchen " Hodenrudimente
besasse.
Auf Grund unserer heutigen Kenntnisse iiber die Anatomie dieser Art diirfen wir
jedoch behaupten, dass ihre beiden dimorphen Greschlechtsindividuen morphologisch
als Hermaphroditen organisirt sind, in physiologischer Beziehung jedoch nur
eingeschlechtlich und zwar entweder nur als Mannchen oder uur als Weibchen
functionieren.J Von einem Sexualdimorphismua in der gebrauchlichen Bedeutung
kann daher bei ^f. ci/sticolum nicht die Rede sein. Man wird vielmehr die bei
dieser Species herrscheudeu Geschlechtsverhaltnisse als das Endergebnis einer
physiologischen Sexualdifferenzierung betrachten miissen, die sich im Laufe der
Ontogenese auf (morphologisch-) hermaphroditischer Grundlage vollzieht und dabei niit
einer Dimorphosierung der sich entwickelnden Geschlechtsindividm-n vcrbuudeu ist.
Der typische Myzostoma-TLabitua uud die relativ geringc Grosse de.s Manncheus,
ferner die verhaltnismassig sehr eiufache Ausbildung seiner iunereu Organe, sowie
der primitive Aufbau seiner Korpcrgewebe lassen dasselbe viel urspriiuglicher und
unverauderter erscheinen als das Weibchen, weshalb wir annehmcu miisseu, dass die
Dimorphosierung hauptsachlich an dem letzteren wahrend seiner Entwickelung Vdll-
zieht. Es wird hier die allmiihlig sich eiustellende physiologische Praeponderanz
der weiblichen Gonaden nicht nur den hciheren Ausbildungsgrad dos weiblicheu
Sexualapparates (Leibeshohle), sondern auch die functionellc Ausgc-staltung jeuer
* Nur eininal wurde von Graff (3a, pag. 8) in einer durch If. willemocsi hervorgerufcnen Cyste bloss ein
einziges und zwar uiiinnlich functionierendes Individuum vorgefunden.
t In diese Gruppe gehoren noch : M. bcardi und M. platypus (Graff 3a, pag. 13 u. 14 ; fiir M. platypus siehe
auch : Wheeler 8, pag. 214), ferner M. Irlli, M. cryptopodium und AT. rremUn (Wheeler 8, pag. 246-251) sowie
warscheinlich auch das M. clm-ki (McClendon 4, pag. 121-122), obwohl es dem letztgcnanntjn Autor nicht
gegliiekt ist, bei den von ihm untersuchtcn Individuen Hoden oder Reste von solchen aufx.ufmden.
i Ich vermute, dass ahnliche Vcrhiiltnisse auch bei den ubrigen von GralT als " dioecisch " bezciclmeten
Arten herrschen. Eine diesbeziigliche Entscheiduug ist allerdings nur init Hilfe der Schuittmethode zu
erbringen, die der genannte Autor, \vie aus dem betreffenden Texte und aus den von ihm gelieferten Abbildungen
hcrvorgeht, bei dieseu Species nich angewandt hat.
x 2
22 DR. RUDOLF EITTER v. STUMMER-TRAUNFELS.
Organe bedingen, welche fiir die Eierproduction eine nur indirecte Bedeutung haben :
Darm und Excretionsapparat (Stoffwechselorgane). In zweiter Liuie wird der
correlative Einfluss der auftretenden physiologischen Unisexualitat bei dem sich zum
Weibchen entwickelnden Individuuin auch aiisserlich und zwar an der relativen
Dimensionierung, der Consistenz und der Form des Kb'rpers, ferner am Bewegungs-
apparate zum Ausdrucke kommen : Die reichliche Auzahl der producierten Eier,
welche innerhalb der Leibeshohle infolge der Aufnahme von Dotter zu einem
Vielfachen ihrer ursprunglichen Grosse heranwacbsen, bediugt einen verbal tnismassig
umfangreichen miitterlichen Korper mit derber dem bedeutenden Innendrucke
gewachsener Wanduug. So kornmt es deun schliesslich dazu, dass die Dimensionen
des Weibcliens jene des Manuchens weit iibertreffeu. Die Breitenausdehuung des
erstereu wird sogar eiue so grosse, class ihr das Cystenwachstum nicht zu folgen
vermag, weswegeu die Seitenrander des Tieres gezwungen sind, sich medianwarts
in der friiher beschriebenen Weise einzurollen (Graff, 3, pag. 12, abs. l). Da ferner fiir
das Weibchen die Bewegungsmoglichkeit innerhalb der dem Korper enge anliegenden
Cyste verloren gegangen 1st, so werden die Parapodien als fiir die Eiproduction
iiberfliissig gewordene Korperteile eingeschmolzen. Es findet sich von ihnen nur
mehr der Hackenapparat, wahrend die hautigen Teile des Fussstummels fast
vollstandig in die Leibeswand aufgenommen erscheinen. Dementsprechend ist auch
die Stummel- und Hackeumuskulatur rudimentiert.
Bei dem sich zurn Mauuchen entwickelnden ludividuum braucht sich die
Sexualdifferenzierung fast nur auf die functionelle Ausgestaltung des mannlichen
Apparates zu beschranken ; denn der Darm und der Excretionsapparat werden
hier jeneu Ausbildungsgrad kaum iiberschreiten, welchen sie bereits bei dem der
Geschlechtsform unmittelbar vorangehenden Jugendstadium besassen. Beim
Mannchen erfahrt eben das Stoffwechselbediirfnis durch die Erzeugung von Sperma
keineswegs eine so wesentliche Steigeruug, wie beim Weibchen durch die
Versorguug der Eier mit Dotter. Die Production der kleiuen, wegen ihrer Form in
compendio'sester Weise zusarnmenpackbareii Spermien benotigt nur relativ geringe
KOTperraume, wogegen fiir die Absatzmoglichkeit des Sameus eine unbehiuderte
Bewegungsfahigkeit des Tieres vorteilhaft erscheint. So besitzt denn das
Mannchen einen kleinen, zarten und abgeflachten Korper mit wohlentwickeltem
Parapodialapparate. Es spricht vieles dafitr, dass sich dasselbe auch im Habitus
nicht weseutlich von jener Jugendform unterscheidet, die ihm in der Ontogenese
uumittelbar vorausgegangen ist.
Die Entwickelungsstadien, auf welchen sich die Sexualdifferenzieruug bei den zwei
Geschlechtsformen herausbildet, sind uns leider noch vollig unbekannt und daher bleibt
die Frage noch offen : Gilt auch fiir das M. cysticolum die Teorie Wheeler's von den
successiven Sexualphasen (8, pag. 288-289), oder herrschen diesbeziiglich bei dieser Art
andere Verhaltnisse ?
Nach den Beobachtungen des ebeugcnannten Autors ist die Sexualdifferenzierung
MYZOSTOMIDAE. 23
der Myzostomen eine successive, das heisst, es tritt im Leben eines jeden Individuums
nach Ablauf eines gescklecktlick indifferenteu Jugeudstadiums zuerst eine mannliche,
spiiter hingegen eine weibliche Gescklecktsperiode auf. Bei den moisten Arten fand
Wheeler, dass diese beiden " Sexualphasen " durch eineu iuterrnediareu functionell
hermaphroditiscken Zustand verbuuden sind ; er beobachtete jedock auck Species (z. B.
3[. pulvinar, 8, pag. 289), bei welchen diese Zwischenperiode ausfallt, so dass die
mannliche und die weiblicke Gescklecktspkase direct an einander ansckliessen.
Ick halte es nun durckaus nickt fiir unwarsckeinlick, dass diese Teorie "Wheelers
auck fiir das M. cyxticolum Gcltung besitzt. Das Yorkandcnsein von Ovarien l)eim
Manucken und von Hodenresten beini Weibcken lassen diese Annakme zu, insbesondere,
wenn man geueigt ist, die Ovarieu des Mannckens als sick erst entwickelnde, aber nock
nickt functionsfakige Organe aufzufassen. Weuu \vir demgemass das Manucken als
ein frillies Entwickelungsstadium des Weibckens betrackten, so ware aber auck der
auffallende Umstand zu erklareii, dass in alien bisker beobackteten Fallen (1G)* in
jeder Cyste uur die beideu extremdimorpken Individual, aber keine ontogenetiscken
Zwiscken- oder Ubergangsstadien, deren Vorkommeu man ja in Consequeuz der obigeu
Teorie voraussetzen muss, augetroffen wurden. Wheeler, \velcher das Bcdiirfhis
diesbeziiglich zu einer plausiblen Vorstellung zu gelangen wohl gefiiklt hatte, spricht
sick dariiber folgeudermassen aus : " v. Graff's supposition tkat tke young Myzostomes
associate in pairs aud together take part in forming a gall seems to me hardly
plausible. Judging from my observations on M. ijlabrum and pulvinar, both of which
show a distinct tendency to occur in pairs, each consisting of a senior and junior
individual, I believe tkat in tke case of tke cysticolous species tke gall must be formed
by a single individual, and tkat later a young Myzostome, wkeii it abandons its
pelagic trockopkore stage, must enter through tke aperture of tke gall and settle down
to a quiet life witk tke senior individual. Tke latter probably dies at tke end of its
female stage, and, undergoing decomposition, may perhaps serve as food for its still
vigorous junior partner. This one in turn may thereupon become the senior partner
of another young Myzostome. and so on. According to this view, all the r >/*//< Wc//x
Myzostomes of a given species would not bo /*/-///-,/////.
6. R. RITTER v. Sxi'.M.MEU-TuAT'Ni'HLS. Beitrilge zur Anatomic und Histologic der Myzostomeu :
I. Myzostoma asteriae Marcnz. Zeitschr. f. wissensch. Zoolog. LXXV. (l'J) ; von der Yentralseite gesehen. Man erkenut die dellenformig einge-
buchtete Stelle, an welcher das Mannchen gesessen liatte. Yergr. 10-facb.
FIG. 5. Mannchen (ans Cyste ) ; von der Yentralseite her abgebildet. Das Exemplar ist rostro-caudal
etwas contrahiert. Yergr. 27i-fach.
FIG. 6. Weibcben (ans Cyste a). Medianer Liingsschnitt. Yergr. 11-fach.
FIG. 7. Weibchen (au3 Cyste &). Etwas sthief zur Liingsaxe ausgefallener Querschnitt in der die mittleren
(dritten) Parapodien verbindenden Trausversauebtne. Von den beiden Ovarien der linken
Korperseite ist nnr das riickwiirtige, von den beiden Hoden nur der linke getroffeu. Vergr. 32-fach.
FIG. 8. Weibchen (aus Cyste i). Hinter dern vorigen, in der Gegend der Nephridialcaniile gefiihrter
Querschuitt. Nur der reuhte Nephridialcanal ist in demselben getroffen. Vergr. 32-fach.
FIG. 9 und 10. Manuchen (aus Cyste b). Aus Flachenschnitten combinirte und etwas schematisirte
Teilbilder. In Fig. sind die mehr dorsal gelegenen Organe, wie der Eruiihrungsapparat, der
milnnlicbe Geschlechtsappurat uud'der Dorsalabschnitt des weiblicbeu Geschlechtsapparates, ferner
die Xephrostomata dargestellt. Fig. 10 zeigt den A'entralabschnitt des weiblichen Geschlechts-
apparates mit den Ovarien, ferncr die Eudblasen mil dern uupaaren Eudabsclmitte der Xephridien.
Zwischen den Nephrostornatis (tis in Fig. 9), und den in Fig. 10 mit * bezeichneten Durch-
scbnittsstellen durch die Xephridien verlaufen in dorsoventraler Eichtung die kurzcn Xephridial-
caniile. Yergr. fiir beidu Fig. 79-fach.
6o' ao
.ns
Aut del
MYZOSTOMA.
SIPUNCULOIDEA.
By W. F. LANCHESTEI;, M.A.,
King's College, Cambridge.
I 'MA si i H.OSOMA SOC'IUM.
THLS collection consists of some thirty specimens, all of small size, which were
mostly taken )>y means of a net from holes in the ire. while the 'Discovery' was
in Winter Quarters. One specimen, however, was captured with the dredge, the
depth being given as 100 fathoms. That the bulk of them belong to a single
species of Phascolosoma is quite certain, both from their general appearance and
from a study of the relations of the various organs, though it is to be noted that
these relations have been found to be far from invariable. Some half-a-dozen,
however, have been the cause of considerable difficulty in the exact determination of
their specific identity ; yet it has seemed best to include them under the above
species. In the first place, they agree in the main details of their structure with
the typical forms; and I have not been able to place them without hesitation
under any of the other closely allied species (sub-species or varieties) which centre
round the Arctic Ph. margaritaceum, namelv. Ph. antarcticum, c//.v//' ( . ////-. fuxcum,
<'<'//. group to render any detailed comparison with this species
unnecessary. On the whole then, it seems reasonable enough to consider this form as
a type differing from Ph. margaritaceum and Ph. capsiforme ; and for much the same
reasons that we can so separate P/t. antarcticum and Ph. fuscum. But the comparison
with the two latter forms is a much more difficult matter. In the first place, much the
most striking feature in the majority of these specimens, as viewed with the naked eye
or the lens, is (a) the thinness and semi-transparency of the skin, and (b) its extreme
smoothness, the papillae only being barely visible with the aid of the lens (they are all
small specimens) on the hind end, or sometimes also at the base of the introvert.
In these features of their general appearance, in their light, somewhat straw-like
colouration, coupled with the abruptly pointed termination of the body, and the
localisation of the papillae, they recall the general facies of Ph. vulffare more than that
of any other species with which I am familiar. And it is in just these features that
they seem to differ so markedly from Ph. untarctieum and Ph. fuscum, in which the
body-wall is relatively thick, the colour is dark, and a system of cross-striation is
present at the hind and front ends of the body, which appears to be at best but weakly
represented in this form. "NVo may then proceed on the conceivable presumption that
these are a single (i.e. from the specific point of view mouovalent) group of points ; in
other words, that we are dealing with one specific point only, and that that point may
be either variable according to size and age, or variable within the species. Passing
on then to other points, we find a clear resemblance to Ph. fu*cum in the close
approximation of the openings of the segmental organs to the line of the anus.
Y 2
4 AV. F. LANCHESTER.
Further, on opening up the body it is found that the ventral retractors arise
from one-third to nearly one-half of the distance between the segmental openings
and the end of the body, which again suggests Ph. fuscum ; but that the segmental
organs are very short and do not reach as far as the origin of the ventral
retractors, which accords with the arrangement in Ph. antareticum. The latter
point is complicated by the fact that in three of the specimens the segmental
organs do over-reach the ventral retractors by as far again, yet are not so much
as half the length of the body in Ph. fitxcttin. And the number of coils of
the gut points to neither the one nor the other, as they vary from about fifteen to
about twenty-five (about eighteen in Ph. fuscum, about twenty in Ph. antareticum).
Turning lastly to the papillae, we find that the description of them in the other two
species accords fully with their appearance in this, except that here they are variable in
one point. Dr. Michaelsen tells us that in Ph. antarcticum .they are '027 mm. wide
and '08 mm. high (/.<>. three times as high as wide) ; in Ph. fuscum, he only says that
they are ''bis '07 mm. laug, also nicht gauz so laug wie die entsprechendeu von
Ph. antareticum" but the comparison of absolute lengths where the difference is so
small is valueless, assuming the proportions to be the same ; so that we can only
conclude that they are practically identical in the two species of Michaelsen in regard
to this point. Now in our species the papillae in some individuals show this ratio of
3 : 1, but in others the ratio of the height is less (2*5 : 1, 2*25 : 1, 2 : 1).
In Pit. georgianum the introvert is only half the length of the body, and the
papillae arc much longer than in the other species quoted ; and these points, taken
along with other smaller differences, seem to clearly separate our form from it.
From Ph. Id gen*? Fischer, these specimens are very little distinct ; but this form is
itself hardly to be distinguished from Ph. antareticum. The thinness and clear
colouration seem to be the main distinction in most of the specimens ; and in all
of them the relative unimportance of the system of striation on the hind end. In
Pit. lagense, moreover, the segmental organs are as long as half the body.
I subjoin a list of measurements taken from a few of the specimens ; the
sign " S.O." is meant to indicate the " opening of the segmental organ," and the
measurements are in millimetres and taken as accurately as the conditions would
permit.
The letters A, B, etc., correspond to different localities, while the figures
1, 2, 3, etc., merely indicate different specimens. At the same time it may be
pointed out that the A and B specimens are all obviously of one type, viz.,
thin-skinned and straw-coloured, while those marked E and F are those that I
have already mentioned as differing from the rest in general fades, viz., dark in
colour and apparently thicker-skinned.
The most remarkable difference is to be found in the specimen marked E 2.
Here the extended introvert is much longer than the body, 26 3 mm. and 8 5 mm.
On the other hand, in the one marked El, which presents all the general facies
SIPUNCULOIDEA.
From and to S.O.
Distance of retractor origins
from S.O.
Number of
Distance of
Length of
Body. Introvert.
Dorsal. Ventral.
tary canal.
auus from S.O.
Al
19
22
8 19
1
A 2
1 1; :,
17-:.
2 7 21
j:.
A3
18
15
2-25 7-7:> I'M
1
A4
1 1 :<
13-5
2 l'7.'i 17
2:.
A:.
l :, :,
17-:.
2-:> 1-7:. is
l
A 6
in
18
2 5 2:.
25
AT
10-5
s 1-25 4-25 17
25
Bl
21
22
:! 9-5 Hi
"5
B2
12
15
i :, in
- 5
E l
9
in
2 6-25 ?
EL'
,x ;>
26-5
l-7.'i 3-5 21
F 1
14
11-5
1-25 4 11-12
and the same internal structure as in E 2, together with closely similar papillae,
the retracted introvert is shorter than the body, 14 mm. : 11-5 mm. In El,
moreover, the dorsal retractors originate from a point half-way between the opening
of the segmental organ and the ventral retractor, while in E 2 they originate at a
position only one-third approximately (as in the other specimens) of the distance
between these two points. It is a little difficult to see how to deal with those
anomalies satisfactorily ; and I have preferred to consider these two provisionally
as within the limits of the present species. In two of the specimens I find the
segmental organs reaching about as far again beyond the insertion of the retractors.
A spindle muscle, not attached to the hind end, is generally present, though in one
or two cases I have not been quite able to decide as to its presence or absence.
In two cases I have found two distinct muscles attaching the front portion of the
gut to the body-wall; in the other cases I have seen no muscles at all, though it
seems quite possible that they may easily be torn, and so escape notice, in these small
specimens.
The following list of localities and notes may be added :
A. in specimens .... Winter Quarters. I. is. 03. No. 12 Hole. I) net.
S. of Hut Point. 2.")-30 fins. 100 yds.
B. 2 specimens ... 28. ii. 02. Up to In fins.
C. 3 specimens .... ,. ll.xi. 02. I) net. Hut Point.
6 W. F. LANCHESTER.
D. 1 specimen .... Winter Quarters. 17. i. 03. Flagon Point. 102
fins. Dredge.
E. 4 specimens .... 2!). viii. 03. Same as A.
F. 4 specimens .... ,, 3. ii. 04. D net. Hut Point.
G. 1 specimen . . . . ,, ,, 19. iii. 02. D net. Hut Point.
H. 3 specimens .... 19. xi. 02. D net. Hut Point.
K. 1 specimen .... 19. iii. 02. Up to 10 fms.
Finally, reference should be made to a very brief preliminary description of
Phascolosoma charcoti, obtained by the Charcot Expedition, and described by
M. Marcel Herubel in the "Bulletin du Mus. d'Hist. Nat," 1906, p. 127, together
with Ph. fuseum and Ph. antarcticum. Since Ph. charcoti is there said to resemble
Ph. georgianum closely, I do not think our species can be synonymous with it. On
the other hand, we are told, in regard to the skin of Ph. charcoti, that it is
"a peu pres lisse," with " quelques papilles espacees et petites"; in which it both
resembles our species and differs from Ph. georgianum, of which Michaelsen says,
" Der ganze Korper ist mit grossen dunklen Papillen besetzt." This discrepancy
in one point may not invalidate its general resemblance to Ph. georgianum, but,
combined with the brevity of the diagnosis, it makes it impossible to assert
similarity or the reverse as between our form and Ph. Charcoti.
COELENTEBA.
IV.-ACTINI/E.
BY JOSEPH A. CLUBB, M.Sc.
(3 Plates.)
THE Actiuiau collection of tlie ' Discovery ' Expedition consisted of upwards of fifty
specimens, which have been allocated to eight species, representing six genera and four
families. With two exceptions, all were taken at or near to the Winter Quarters,
McMuido Bay. The two exceptions * are Port Harris, Falkland Islands, containing
two species, and Enderby Island, Auckland Islands, containing one species, and from
both places a considerable number of specimens, more than half the total, was obtained.
The specimens were for the most part in a fair state of preservation, but I must express
regret that no memoranda were made of the colours of the living animals. In the
preserved condition absolutely no trace of colour remains, and while admitting that
anatomical characters are the only sure guide in Actiuological studies, the colours of
living Actiniaus, when known, serve at least as a valuable clue to identification with the
species of the older authorities, who give very little but external characters as their
diagnoses.
The work has been carried on in the laboratories of the Liverpool Museums,
and I have again to express my thanks to the Museums Committee for permission to
use apparatus, etc. , and to Dr. Forbes, the Director of Museums, for other facilities.
FAMILY ACTINIID.dE, Gosse, 1858.
ACTINIARIA with an adherent base ; column wall smooth or provided with verrucas,
but never with hollow vesicular outgrowths ; sphincter muscle endodermal, generally
diffuse and weak ; tentacles simple ; margin smooth or provided with simple acroragi ;
mesenteries in several cycles, of which usually more than one is perfect : longitudinal
muscles of tentacles usually diffuse ; no acoiitia.
Genus PARANTHEOPSIS, McMurrich.
Actinudse without acroragi, but with conspicuous verrucas on flic distal portion of
the column ; no collar, but the margin a distinct parapet, within which is a well-marked
fosse; tentacles of moderate length and rather slender, capable nf being concealed in
contraction ; sphincter diffuse.
* These forms arc so interesting that, after some consideration. I hiive allowed tln-ir niti-mrr into this
report. ED.
JOSEPH A. CLUBB.
PARANTHEOPSIS CRUENTATA.
Actinia, rruentata Conthouy in Dana., U.S. Exploring Expedition, Zoophytes* (1S4R), p. 138, PI. 3,
Figs. 23 and 23A.
t'l-ri'iix n-tifittritiis Milne Edwards, Hist. Nat. Corall. (1857), 1, p. 2G8.
Bit nodes cruentata Gosse, Actinologia Britannica (1860), p. 194. Verrill, Trans. Connect. Acad. (186!)),
1, p. 467. Andres, Fauna und Flora des Golfes von Neapel (1884), p. 215.
Comlylactis cruentata McMurrich, Scientific Results of Explorations U.S. Fish Commission Steamer
'Albatross' (1893) p. 15<). Carlgren, Zoantharien, Hamb. Maghal. Samrael. (1898), p. lo, Figs. 13
and 14.
Parantheopsis cruentata McMurrich, Zoolog. Jabrb. Suppl. VI. (1905), p. 233.
I have adopted the course taken by McMurrich (10, p. 233), and place this species,
previously known as Condylactis cruentata, under this genus.
Six specimens were taken by the ' Discovery ' at Port Harris, Falkland Islands,
along with a number of specimens of Bunodes octoradiatus. In my description of the
latter species (p. 7) I refer to the extreme similarity in external appearance of these
two species.
The largest specimen measures 2 5 cm. in height of column and 1 5 cm. in
diameter. Proximally the body-wall is longitudinally grooved, corresponding to the
insertion of the mesenteries ; distally the corrugations are replaced by distinct rows of
verrucaj. The papillfe or pseudoacroragi terminating the rows have no trace of
nematocysts, and are simply outgrowths of the body-wall, being essentially identical in
structure (PI. 1, Fig. 3) with it. They vary considerably in size in different individuals,
evidently depending on their state of contraction. In the specimen from which the section
figured (PI. 1, Fig. 1) is taken the pseudoacroragi are much inflated. Considerable
variation exists in the number of tentacles. The largest specimen has 48 tentacles
(6 + G + 12 + 24), the smallest has only 26, and an intermediate size 32. This
certainly suggests that the adult condition is hexamerous, and that any octamerous
condition is simply a stage in development and transitory, as Carlgren supposes. No
sphincter muscle is developed, there being no indication in transverse sections through
the parapet and adjoining body-wall of any modifications more than the ordinary
muscle bands found generally (PI 1, Fig. 1). The. circular muscles of the body-wall
are moderately developed, and the mesogloeal layer is relatively thick (PI. 1, Fig. 1).
The longitudinal muscles of the tentacles and oral disc are ectodermal, and the
longitudinal and parieto-basilar muscles of the mesenteries are well marked.
FAMILY PARACTID.E, E. Hertwig, 1882.
ACTINIARIA with a mesodermal .sphincter muscle ; no acontia ; mesenteries arranged
in several cycles, of which usually more than one is perfect ; longitudinal muscles of
the mesenteries usually diffuse ; column wall usually smooth, sometimes with sub-
marginal plication and solid ridges : no acroragi.
* Dana (Amer. J. Sci. (2| i. (1S46|. p. 178| cites his Zoophyte work as of this year ; the copy in the Zool. 1'ept.
ol the Museum bears date 1848. ED.
A< TI.M.K.
Genus PAKACTIS.
Paractidae with thin and smooth column wall ; tentacles of moderate length and
of uniform thickness throughout : margin not lobed ; individual mesenteries of cadi
paii' cojiiallv developed.
PAKACTIS I-OI.AKIS.
One specimen labelled " Winter Quarters, 24. 8. 03," and taken in 25-30 fathoms
(PI. 1. Fig. 2).
Height of column 2'3 cru. ; diameter of foot-disc 35 cm., diameter of oral disc
'2'5 cm. Tentacles of uniform size, about 0'-'! cm. long, in four cycles
12+12 + 24 + 48 = fM>. The base is adhesive and thin, the insertions of the
mesenteries being visible through it. It is produced all round wider than the column,
which narrows considerably. The bodv-wall is firm, and near the parapet somewhat
thick, where it is slightly puckered by contraction. The mouth opening is visible in
the centre of the tentacles, crowded together bv contraction. The oesophagus is
plicated and the two siphonoglyphs are well marked.
The sphincter is fairlv strong, tuesogloeal, and produced to a tine termination.
A transverse section (PI. 1 , Fig. 3) shows that it lies nearer to the endoderni. and is
reticulate in appearance, giving indistinct traces of layering. (ireatcr details of
structure are given in Fig. 4.
The mesenteries are membranous, and the longitudinal muscles are somewhat
diffuse, but well marked (PI. 1, Fig. 5). The meso1),
in his comments on the genus /'. fit.
/'in-ai-tix /Hi/inrrf Milne Kilwank-s, p. :M'.i, Imn. tit.
A single specimen labelled '' McMurdo Bay, Winter Quarters, 20 fathoms," only
slightly contracted, with tentacles, oral disc and resophageal opening plainly visible,
the last-named being 1*5 cm. in diameter.
-1 JOSEPH A. CLUBB.
The height of body-wall is 4' 5 cm., diameter of column 5'5 cm., and of oral
disc very slightly less. The body-wall is quite smooth and extremely membranous,
giving the animal quite an inflated appearance. The upper margin is slightly
crenulated by contraction of the sphincter ; the pedal disc is adherent and a little
firmer and thicker than the body-wall ; the oral disc has slight radiating folds running
outwards, up to and between the tentacles ; the oesophagus is strongly corrugated,
and the two siphonoglyphs are visible, but not markedly distinct. The tentacles are
stout, in four cycles (12 + 12-1-24-1-48 = 96); in the innermost cycle about 1 cm.
long in the contracted condition, and diminishing slightly in length in the outer cycles.
The sphincter is mesogloeal, occupying almost the whole of the mesogloeal layer,
and projecting as a distinct collar, with the ectoderinal epithelium, and forming a deep
fosse between it and the tentacles. Fig. 7 on PI. 2 shows a radial section through
the sphincter. The muscles of the oral disc (PI. 2, Fig. 7) and tentacles (PI. 2, Fig. 8)
are mesogloeal. The longitudinal muscles of the mesenteries are in the form of slender
strands, which may be seen running more or less parallel to each other over the extremely
delicate and membranous mesenteries. In transverse section (PI. 2, Fig. 11) it is seen
that the mesogloea is extremely thin throughout, but is slightly thickened and bears
short slightly branched lamellas for the muscle strands. The parieto-basilar muscle
is small and insignificant. Two cycles of mesenteries are complete, and the pairs of
directive mesenteries bear the usual relation to the siphouoglyphs. All the complete
mesenteries are fertile, including . the directives, and ova in various stages are seen
(PI. 2, Fig. 9), and a figure is given of a section of the somewhat characteristic
mesenterial filaments (PI. 2, Fig. 10).
I have identified this specimen with Paractis papaver, the Actinia paparer of
Drayton, briefly described and figured by Dana (4, p. 143, PI. 4).
Genus ACTINOSTOLA, Verrill.
Paractidaj usually of large size, with a firm leathery wall, which may be
corrugated or folded, but with no verrucas Margin tentaculate. Tentacles relatively
thick, and not enlarged at the base. The pairs of mesenteries in certain cycles are
unequally developed, having one of each pair smaller and narrower than the other.
ACTINOSTOLA CHILENSIS.
Actinostola chilensis McMnrrich, Zoolog. Jahrb., Supp. VI. (Fauna Chilensis Hi.), (1905), p. 247, Pis. 15
and 16, Figs. 30-33.
Five specimens of large size contained in one bottle labelled " 4.3.04, 254 fathoms,
mud and stones." The locality is not stated on the label, but I am informed that the
position was 67 21' 46" S. and 155 21' 10" E. The specimens vary greatly in form
and size. One specimen measures 5 '5 cm. in height and 4 cm. in diameter of
column ; another is 4 cm. only in height, but 6 cm. in diameter.
The wall is firm and leathery, smooth to the touch, and no verrucas or tubercles
ACTINLE. 5
are present ; hut irregular shallow f'urmws are formed by contraction in these preserved
specimens. In all cases the tentacles are exposed, and they agree generally with the
description given by McMurrich, in his account of the species (10, p. 247) from a
specimen from Calbuco, both as regards number and arrangement and relative size.
The oesophagus is generally widely expanded, and in the majority of the
specimens is almost indistinguishable from the portion of the oral disc inside the
tentacles. But in one of the more contracted specimens the lip of the oesophagus is
easily distinguished, but the corrugations are continued radially over the oral disc,
running between and a short distance up the bases of the tentacles. The siphonoglyphs
are iii no case well marked. The foot-disc is well-marked and strongly adherent.
The more minute anatomy agrees generally with McMurrich's description. The
mesogloea of the body-wall is variable in thickness, and in one specimen, just
below the sphincter, is nearly 2 mm. thick, from which in an upward direction it thins
rapidly to less than 1 mm. The mesogloeal sphincter exhibits no trace of layering,
having a simple reticulate strueture iu transverse section. Transverse sections of the
tentacles exhibit the same appearance of portions of the ectodermal epithelium being
cut off and apparently enclosed in the mesogloea, as described by McMurrich, due to
contraction and the unusual thickness of the mesogloeal layer.
The mesenteries in number and arrangement agree with McMurrich's descrip-
tion, but I cannot confirm the hermaphroditic condition described for the specie.-.
In the specimen selected for histological work no spermatozoa were found, and the
ova were large and well-developed. Otherwise the ' Discovery ' specimens agree
well with McMurrich's description, and I have no hesitation in placing them under
this species.
FAMILY ALICIID/E. DUERDEN (1895).
ACTINIARIA with a large, flat, contractile base. Tentacles simple, cylindrical and
entacmseous. Column wall with simple or compound hollow tubereles or vesicles,
covering the greater part of the column, arranged mostly in vertical rows. Sphincter
muscle endodermal and diffuse, variable in amount of development. No ciuclides or
aeontia, margin with or without acroragi. Mesenteries arranged in several cycles of
which usually more than one is perfect.
Genus CYSTIMTIS, M. Edw.
Aliciidas having the column covered with simple vesicles. Tentacles of variable
length, in one, two, or three cycles. Numerous perfect mesenteries ; sphincter muscle
moderately well developed.
( 'YSTIAGTIS ANTARCTICA.
A single specimen, bearing the label " McMurdo Bay, "Winter Quarters, 28.2.02."
Depth, 20 fathoms.
z 2
f, JOSEPH A. CLUBB.
External characters Height of body-wall 5 '75 cm., diameter of column 4 cm.,
diameter of oral disc 3 cm., diameter of pedal disc (contracted condition) 2 '75 cm.
There is no trace of distinctive coloration in the preserved specimen, and no
record on the collector's label. The single specimen is poorly preserved, and owing to
the thin and delicate body-wall is quite collapsed. The column is studded with
thin-walled simple vesicles (PL 2, Fig. 12) communicating with the coeleuteron and
arranged in twenty-four vertical and parallel rows, corresponding to the niesenteric
spaces. The attachment of the mesenteries ma}' be seen in places, between the rows
of vesicles. The vesicles of adjacent rows are arranged alternately, and they increase
in size from the proximal to the distal portion of the column (PL 2, Fig. 12), where
the largest are not much inferior in size to and do not differ much in appearance from
the tentacles in the preserved and contracted specimen.
The tentacles are twenty-four in number, arranged in two cycles of equal size,
short and somewhat club-shaped. The largest measures O 1 5 cm. in length. The
oral disc is slightly furrowed radially and the mouth is large and bears a well-marked
hypostome. The oesophagus is short and strongly plicated, and two deep and
well-marked siphouoglyphs are present, and are continued some distance below the
rest of the cesophagus as distinct lappets.
The coelenterou is large, and twelve pairs of mesenteries, all complete, are present.
Two pairs of directive mesenteries bear the usual relation with the siphonoglyphs.
The single specimen taken is poorly preserved and much of the histological detail is
lost. The sphincter is diffuse and endodermal in character, with slightly projecting
mesogloeal lamellae shown in transverse section (PL 2, Fig. 13). The muscles of the
tentacles are ectoderrnal. The muscles of the body-wall (PL 2, Fig. 14) are feeble,
and sections through the vesicles show a similar structure. The ectoderm and
endoderm layers consist of relatively high columnar cells (PL 2, Figs. 13 and 14);
but the mesogloea is thin. The mesenteries are very thin and delicate,' but transverse
sections show a weak but well-marked longitudinal muscle (PL 2, Fig. 14), the
mesogloea in relation with it being thickened and possessing short branching lamellae.
The parieto-basilar muscle is small (PI. 2, Fig. 14).
The specimen is female, and ovaries are found on all mesenteries, including
directives, forming broad bands occupying in their widest portions about a third of
the width of the mesenteries. Ova in all stages of development (PL 2, Figs. 14
and IS) are present.
FAMILY CRIBRINID^E, McMurrich, (1901.)
ACTINIARIA with a well-developed circumscribed endodermal sphincter ; simple
tentacles ; without cinclides and acontia ; verrucas usually present ; adherent base ; no
true acroragi, but frequently pseudoacroragi present.
ACTixi.r..
CKIBRINA.
Cribrinida: with strong endodermal sphincter; frequently with pseudoacroragi ;
ectodermal musculature of discs and tentacles not imbedded in the mesogloea ; column
wall provided with verruca? arranged more or less distinctly in vertical rows :
tentacles simple.
CRIBRIXA OLTORADIATA.
liiniin/fK t>i-ti>ni'tii'tiis Ciirlgren, Hamburg. MagalhaenB. Sammelreise, Zoanth. MSIINI. ],. i>n. i-'j.jx. n ; uid 12,
1898.
This species was formed by ( 'arlgren (1, p. 20) for a number of specimens taken
from the Straits of Magellan and other localities in the South Seas. In one of the
'Discovery' bottles. Iain-lied "Falkland Islands, Port Harris/' were fourteen specimens
of very similar-looking Actinians. Upon dissection and microscopical examination.
however, I find two distinct species Bnnrriini ,,//,//,/ agree with Carlgren's
specimens (1, p. 20). The sixteen rows of verruca; on the distal portion of the column
and the sixteen pseudoacroragi are well seen in all the specimens. Usually gravel
and broken shell are attached to the verruca?. The number of the tentacles
(8 + 8 + 10 = 32) appears quite constant. The two siphonoglyphs are well marked
and quite regular. The sphincter muscle is strongly circumscribed, and may be
described in McMurrich's terms (9, p. 20) as of the pedunculate palmate varietv
(Pl. 3, Fig. 10). The section figured is through the space between the pseudoacroragi,
but sections passing through the latter show the sphincter projecting from the inner
wall, near its base, so that the cavities of the pseudoacroragi project beyond and
above the sphincter. The mesenteries number sixteen pairs (8 + 8 = 10), all
complete ami fertile, including the directives (PL 3, fig. 18). The longitudinal
muscles of the tentacles are ectodermal and transverse sections (l'|. :>, Fig. 17) show
strong mesogloeal lamella;. The longitudinal muscles of the mesenteries and the
paricto-basilar muscles are exceedingly large and strong, and in the ci,nir ;l Hed
specimens are closely packed into a solid mass, so that the various organs are quite
dovetailed together (PL 3, Fig. 18). The mesogloeal lamella? of the muscles are long
and branched. The circular muscles of the body-wall too are very strong. The
strength of the musculature generally is quite a feature of the species.
JOSEPH A. CLUBB.
But the most interesting character of these specimens from the Falkland Islands
is the presence of " brood pouches." Immediately below the pseudoacroragi is a well-
marked constriction of the body-wall, also present, it is interesting to note, in Punui-
theopsix cruentata. In this constriction, in the species under consideration, specimens
were found possessing sixteen pores, one pore to each line of verrucae, leading into
distinct invagiuations of the body-wall, forming characteristic " brood pouches " in the
coelenteron. These " brood pouches " agree in all essential points with the arrangement
shown to be present in the ' Southern Cross ' specimens (3, p. 299). The drawing
(PI. 3, Fig. 16) shows a section of the body-wall, passing through one of these pores,
and the " brood chamber " into which it opens. The three layers of the body-wall may
be distinctly traced as shown, each of them much thinner the mesogloea markedly so.
Usually each "brood chamber" contains two embryos only, and in the section figured
they are well advanced, showing several of the mesenteries complete and others well
developed. Dissections were made so that, looked at from below, the sixteen " brood
pouches " were seen like so many miniature grapes, lying quite regularly in the body
cavity, between the mesenteries. In diameter they average about 3 mm.
In the short description of the species by Carlgreu (1, p. 20) no mention of
them is made. It is somewhat remarkable if none of Carlgren's specimens possess
these "brood pouches," and possibly re-examination may show their presence in some.
CRIBRINA HERMAPHRODITICA.
llanodfs hermaphroditicus Carlgren, Hamburg. Magalhacns. Sammelreise, Zoantli. (1898), p. 23, Fig. 18,
1898.
About forty specimens from " Enderby Island, Auckland Islands, 19. 3. 04" of
sizes ranging from 3 cm. to 2 cm. in height of column. In some cases the oral disc
and tentacles are fully exposed, in others the animals are contracted so that the
tentacles are entirely hidden, and there are many intermediate stages of contraction.
The foot-disc is well marked and strongly adherent. The rows of principal
verrucas, usually twenty-four in number, are well seen, especially on the distal portion
of the column, where also secondary alternating series are visible. The pseudo-
acroragi, terminating each principal row of verrucas, are distinct, emphasizing greatly
the parapet. The tentacles are in four (6 + 6 + 12 + 24 = 48), or five (6 + 6 + 12
+ 24 + 48 = 96) cycles.
The sphincter is endodermal and circumscribed. The oesophagus is longitudinally
folded, and two well-marked siphonoglyphs are present. The mesenteries in the
specimens examined are all hexamerously arranged, and in the larger specimens in four
cycles, of which three are complete. The longitudinal and parieto-basilar muscles
(PI. 3, Fig. 19) are well developed. Two pairs of directive mesenteries bear the usual
relation with the siphonoglyphs. The longitudinal muscles of the oral disc and
tentacles (PL 3, Fig. 21) are ectodermal. Zoanthellse occur throughout the endoderm,
being especially abundant in the tentacles (PL 3, Fig. 21). Reproductive elements
ACTIXLK. 9
arc present in relation with the mesenteries of the first cycle in small specimens, including
direct ivcs, and iu large specimens, with the mesenteries of two cycles ; liut I was not
able to determine to my satisfaction if present in relation with the third complete cycle.
Both ova and spermatozoa are present, often associated together in relation with the
same mesentery (PI. 3, Fig. 19). Occasionally fertile mesenteries are found showing
ova or spermatozoa alone, but in all the specimens examined microscopically the
hermaphroditic condition is constant. Embryos in various stages of development are
present in the body-cavities of nianv.
(icnus Until (ACTINIA, Agassiz.
Cribrinidu3 (Tealiidae, Bunodidre, Bunodactidse), with well-developed foot disc;
distinct verrucfe present on body-wall ; radial muscles of oral disc and longi-
tudinal muscles of the tentacles meso-ectodermal to mesogloeal ; tentacles simple ;
strongly circumscribed endodermal sphincter.
RHODACTINIA rKAssicoiiNis.
Aiii /i in i-nixxii-nniix MiilkT, Prod. Zool. Dan. (177f>), p. 231.
Aiiiiiiii. I'li'iimitixxiiiiii. ]5rauilt, Prod. duscriptionis animnliuni ab H. Mertcnsio oWrvarormn ( ix:?5), p. i:;.
niiiiiliii-tiiiin ilnrixii Agassi/,, I,., Complus Reiidus, XXV. (1847), p. (i77.
Uliniliiiiiiiiii ilnrixii Yen-ill, A. E., Mem. Hoc. Nat. Hist. Boston (1Ki/.:/irr//i-/ixis Kwk'tniewski, Zoo]. Jahrb. (Syst.) (1SI1K), p. 1, p. 121.
Carlgren in 1902, in his report on "Die Actiniarien der Ulga Expedition"
(2, p. 39), thought it necessary to revive the old genus of Agassiz, Illi<>-///.v Carlgren places a number of
.synonyms, including Aiiinin elegantissima, Brandt, and Leiotealia spitzbergensis,
Kwietniewski. 1 have previously been struck with the variation iu the degree in
which the muscles of the tentacles are imbedded in the mesogloea, in specimens of
<:ri/*x/<-i>riiix examined by me, from a well-known local habitat. Ililbre Island, at the
mouth of the Dee. The text-figures (2, p. 41) given by Carlgren are most
instructive, and from llillire Island specimens I can produce examples in which the
longitudinal muscles are quite as much meso-ectodermal as Carlgren's figure 4, if not
as much as his Fig. 3. In my examination of the 'Southern Cross' Antarctic
Actinians (3, p. 294), I was convinced of the many close resemblances to the type
species crassicornis, seen in the specimens then under observation ; and notwithstanding
that the longitudinal muscles of the tentacles were not imbedded in the mesogloea,
I decided to include these species under the genus /"///>///ai-li>/rrlieries, so as to form an incomplete arch over
a sub-dermal space or channel.
A small "silica pearl" was found in one section.
Numerous large ova occur, with oval body and thick pseudopodia-like extensions,
the body being 130 X '65^, and the total diameter over 300/n. In some instances.
the ova are more rounded and compact, with the cytoplasm more drawn together.
I had described the specimens as representatives of a new species, but I now find
that they belong to < 'nmiJIn viyitta (Lendenfeld).
Dr. v. Lendenfeld very kindly sent me the revised proofs of his Report
''Tetraxonia der deutschen Siidpolar-Expedition, 1901-1903,'' thus enabling me to
make the necessarv alterations before it was too late.
4 R. KIRKPATRTCK.
The 'Gauss' Expedition obtained six specimens, which are all young and small,
being from 2-10 mm. in diameter.
The difference in the size of the megascleres of the 'Discovery' and 'Gauss'
specimens may be attributed to differences in the age and size of the specimens :
but the difference in the size of the sigmata of the two sets of specimens is more
notable those of the ' Gauss' being 14-20/x, and of the 'Discovery ' being only 12 '3^
to 13 p. in length as viewed in the C aspect. In my opinion difference in size of
sigmata has not the importance that is sometimes attached to it.
All the specimens were dredged in the neighbourhood of Winter Quarters in depths
of 10-130 fms. Also ' Gauss' Expedition, near its Winter Quarters, 350-385 m.
CRANIELLA SAGITTA var. pachyrrhabdus.
(PL VIII., figs. 14, 15 ; PI. IX., figs. 17, 18, 19.)
1907. Ti'tli/n sriflit/it. Lendenfeld (lla. p. Sue).
This variety is represented by several oblong pieces about 6 cm. in length, deeply
blackened by osmic acid, and forming part of a large sponge, which had been cut up
for preservation in that reagent.
The fragments are sufficient to show that the surface of the sponge was covered
with a spicular pile about 4 mm. in height.
Mr. Hodgson informs me that the bulk of the specimen was lost. He was under
the impression that this sponge, which was obtained from the same locality as the
var. microsigma, and which resembled the latter in general appearance, was identical
with it in all its characters. An examination of a section, however, shows that the
radiating fibres of the skeleton contain numerous large thick strongyles and styles,
about 7 x O'llG mm. in dimensions (VIII. figs. 14, 15); the sigmata are 18-20 /x
long, and slightly thicker and rougher than in the var. microsigma.
Plate IX. figs. 18, 19 show collar cells, stained in gentian violet. The cells are
here seen to be separate, and the collars apparently not coalescent ; but the tissues are
not very well preserved ; and, though much care was taken in the drawing, a renewed
inspection of the preparation from which the illustrations were drawn has shown that
the figures of the collars are not satisfactory. The flagellated chambers usually contain
only a few collar cells in the normal position (IX. 17), the rest forming detached
masses in the centre of the chambers. The flagellum originates from the end of the
large oval nucleus, which is situated a little below the surface of the body of the cell.
Locality. Winter Quarters, Hut Point, 25 fnis.
CRANIELLA LEPTODERMA.
(Plate XI., figs. 4-14.)
isss. Tetillaleptoderma8oliaa(2l. p. 3).
Sponge, club-shaped or pyriform ; with verruculate surface ; with one large oscule
near the summit ; with root-tuft. Surface pile of spicules absent in the adult state,
and only slightly developed in early stages, large protriaenes being very rare or absent.
TKTKAA'oXIDA. 5
SpiculeS. .Megascleres. 1. Large oxeas, !) 7 X 0' 081 mm.
2. Smaller oxeas, scattered in the choanosome, 540 to 1350/x in length.
3. Cortical oxeas (XI. 8), 770 x 30 n, straight, fusiform.
4. Anatriaenes of three kinds: a. (XI. 9) Cladi 170 n long x 20 p thick at
base, slender, sharp-pointed, chorda llOju; rhabdome, about 12mm. long, thick at
junction with cladome, then tapering, again thickening, and finally terminating in a
filiform extremity.
!>. (XI. 11) Cladome with thick conical dadi, 140 X 35 p., chorda 60 //. ; rhabdome
with a thick neck, and then nearly uniform till it tapers to a filiform extremity.
c. (XI. 10) Cladome with very long, straight, slender cladi, 150 x 10 p, nearlv
parallel with, or making an angle of, about 10 with the rhabdome, chorda only 31 p. :
rhabdome slender (of uncertain length).
5. Protriaenes. Rare, usually with three slender equal cladi, 90 /A long ; rhabdome
of nearly uniform diameter : but slightly diminishing near the cladome.
G. Triehodal protriaeues, usually (especially in the case of those forming the inner
oscular fringe) with one ray greatly prolonged to 140 p. or more, the other two having
almost disappeared ; rhabdome slightly thickened at cladal origin.
7. Sigmata, 14 - 25 to 15 /j. long, 0'75/x thick, and 12'3/n broad in the c aspect,
with slightly and finely granulated surface.
There are three specimens of this sponge, the largest (XI. 4) being 12 cm.
in total length and 5 '2 cm. in breadth, with a root-tuft 3' 5 cm. in length:
the second specimen (XL 5) is oval, 2'9 cm. long and 1'7 cm. broad, and the smallest
is U! x 12 mm.
The colour of the first two is dirty gray, and that of the last pale but}'.
The scale-like, occasionally over-lapping, verrucae, attain a height o|' 2 nun., and
a breadth at the base of 4 mm.
The oscule is oval-shaped, edged with a barely visible fringe of tridiodal
protriaenes ; it measures, in the large specimen, 7'5 x 5 mm.
The skeleton. The radiating fibres only vcrv rarely project beyond the surface :
near the oscule are a few tufts of oxeas. Uver the rest of the surface of the sponge,
the dermal membrane covers over the summits of the verrucae, the distal points of
oxeas and the cladomes of anatriaeues showing through ; medium-sized in large
protriaenes, so abundant in < 'r<n section, the ectosome is scarcely differentiate,! from the
choanosome, excepting that sub-dermal spaces are visible just beneath the dermal
membrane.
The cortical oxeas are more densely packed than in the preceding species.
A tangential section shows the pores in longitudinal areas between the densely
serried rows of cortical oxeas, and fan-like tufts of trichodal protriaeues supporting the
6 R. KIKKPATKICK.
sieve-like pore-areas ; the pore-areas are distributed over the surface up to the
neighbourhood of the oscule.
Caual System. The pores lead into sub-dermal channels whence canals descend
into the choanosome. The flagellated chambers are eurypylous.
Small ova, 60 x 30 ^u, with pseudopodic-like processes, occur in the largest
specimen.
A young Specimen is pear-shaped with a granular surface, and without a
root-tuft. Each " granule " is a sharp-pointed conule, with a protriaene projecting
from it.
The young specimen of Craniella sagitta has a very different appearance, being
quite fluffy on the surface owing to the numerous tufts of protriaenes.
Specimens were obtained from : (1) West of Balleney Island in 254 fins., Type
specimen (XL 4) ; (2) W.Q., Flagon Point, 10-20 fms. ; (3) W.Q., Hut Point, 25 fins.
The ' Challenger ' dredged specimens off Rio de la Plata, 600 fms.
ClNACHYRA BARBATA.
(Plate IX., figs. 1-14.)
1888. Cinachym larbata Sollas (21. p. 23. Pis. III., XXXIX.).
190G. CinacJujra barliata Kirkpatrick (10. p. 602. PL XIV.).
Twenty-seven specimens of this sponge, varying in diameter from a few
millimetres up to 14 centimetres, were obtained. In addition to these, the collection
includes a massive root-tuft no less than 32 cm. across and 12 cm. in thickness,
belonging to what must have been a very large specimen, for the concavity into which
the latter fitted occupies nearly the whole area of the upper surface of the root-tuft.
The largest specimen obtained by the ' Challenger ' from Kerguelen was 10 cm. in
its longest diameter.
As in the case of the ' Challenger ' specimens there is a considerable variation in
the shape, which is usually spheroidal ; some specimens are spherical, the nucleus of
the radiating bundles of the skeleton being exactly central, others again resemble an
inverted cone, and very young specimens are oval.
An interesting feature shown in adult specimens is the division of the sponge into
three zones, viz., (1) a basal root-tuft zone; (2) an equatorial zone of porocalyces ;
and (3) a polar zone of oscules.
These zones are clearly perceptible in the fine series of well-preserved Antarctic
specimens, but are not so well marked in those obtained by the ' Challenger.' The
surface pile of spicules in the porocalycal zone is longer and looser than in the oscular
zone, and directed obliquely downwards ; but the pile in the oscular area is shorter,
and more vertical and stubble-like.
The root-tuft. A remarkable feature about the root-tuft is its great size in
some specimens, in which it may be much more bulky than the sponge body ;
TETRAXOXIDA. 7
generally, however, it is of smaller size than the body, especially in large and
old specimens.
The porocalyces. In adult specimens the porocalyces form a broad belt round
the sides of the body. A few, in large specimens, were nearly 2 '5 em. (one inch) in
depth. Ae I have already stated (10. p. 662) the porocalyces are, in all species of
Cinachyra, inhalant areas perforated by true dermal pores, and are never exhalant or
oscular.
The OSCllleS. In Cinachyra iinrlmta these structures are arranged in an obscurely
spiral manner on the upper or " polar " surface of the sponge in adult specimens. In
almost every instance, the oscules are tightly contracted, the spicular fringe being
closed over them in form of a conical stack. It is this strong contractibility that has
led to their being overlooked in most of the species of this genus, the porocalyces
having been regarded as partly inhalant and partly exhalant.
The oscules are very rarely found open. Where this happens, the oscular orifice
is at the summit, of a slight crater-like elevation. Within the circle of long protriaenes
forming the oscular fringe is a circle of fine protriaenes on the edge itself and also
on the gentle slope leading inwards. At a short distance from the edge, the slope
leads to a vertical barrel-shaped tube ending in a well-marked sphinctrate ring, which
is ab'out on a level with the innermost plane of the cortex ; the oscular tube slightly
expands below the sphincter into a sub-cortical cavity in the lateral walls of which are
the orifices of exhalant canals proceeding from just beneath the cortex. At its lower
end the sub-cortical space is continued on into a large main vertical canal, which passes
up radially from the central region, receiving in its turn many affluents. The wall of
the oscular tube is mainly composed of a thick sheath of concentrically arranged
uiyocytes.
The vertical section of the oscular tube, when contracted, shows the almost closed
lumen of the tube surrounded by a dark zone of sigmata, and with spiral bands of
myocytes passing upwards around the lumen.
Two of the eight known Cinachyra species, viz., C. voelt:koici Lendenfeld (11. p.
101) and C. malaccensis I. J. B. Sollas (19. p. 219), are described as having scattered
pores in addition to the pores of the porocalyces. Some of these " pores" will certain ly
be found to be oscules situated, in young specimens, more or less antipodally to the
porocalyces, others possibly belonging to developing porocalyces ; for as W. J. Sollas
observes (21. p. 29), " in their inception, the porocalyces are simply poriferous areas of
the cortex."
Young specimens. Very young specimens (IX. 3) are oval, broader at the
superior end, and without a root-tuft, though the surface spicules are longer at the
narrow end of the sponge. At this stage the sponge lies with its long axis oblique or
horizontal. The smallest specimen in this collection is 3 x 4 mm. in diameter. It
has one porocalyx situated in the centre of one side, and one oscule a little to one side
of the centre of the summit of the broad end. This nearly bilateral symmetry calls to
2 r.
R. KIEKPATRICK.
mind the species of Fangophilina Schmidt, viz. F. submersa 0. Sch. (17. p. 73), and
F. gilchristi Kirkp. (10. p. 667), in which the ovoidal body of the sponge has one
large porocalyx and one large oscule situated on the upper aspect. As the sponge
grows, the porocalyces increase in number and extend on each side of the original one
till they form a complete equatorial belt ; in the meantime, the main axis of the sponge
gradually rotates from horizontal to vertical, so that the crown of oscules comes in
many specimens to lie in a horizontal plane at right angles to the vertical axis.
The cortex. In young specimens the cortical oxeas are arranged taugentially
in a single layer. In the largest examples, the cortex reaches a thickness of 3 '25 mm.,
the densely packed oxeas being arranged vertically and obliquely to the surface.
The skeleton. The only additional observations to be made here are on certain
bodies which Sollas (21. p. 24) refers to as " Globules ; accessory or accidental forms,
0'0535 mm. in diameter." These bodies, which Schulze terms "silica pearls," are now
known to be not uncommon in sponges (Schulze, 18. p. 6, and Weltner, 33. p. 190).
Spheres may be normal spicules of the sponge, as in Caminus sphaeroconia Sollas. In
many instances, however, spheres or globules result from malformation or incomplete
development, as in cases where a tylote spicule is reduced to a knob, or a sphere may
result from the reduction of an oxea, as in Epallax callocyathus Sollas. In some of
the specimens of C. barbata there are a considerable number of pearls and some of them
are double (IX. 6, 7). One example 114 x, 94 /JL is oval, with two nuclei and with
concentric layers of deposition round each, up to the point where the spheres come in
contact ; later there is a single oval layer common to the two centres, but an annular
depression or kink is always visible in the plane midway between the nuclei of the two
original spheres. Sollas (21. p. 214, PI. XXVIL, figs. 8-9) figures composite spheres
occurring in Caminus sphaeroconia, but here the composite sphere does not possess
separate central points round which are deposited concentric laminae, but (apparently)
one centre and an axial line or rod round the end of which the layers are deposited.
Whether these spheres or silica pearls are always spicules or the result of incom-
plete development of spicules, or whether they are sometimes due to deposition of
layers of silex round some foreign organic or inorganic body, has not been determined.
The nucleus is generally a refringeut point, but is sometimes irregularly shaped and
of a faint yellow colour ; attempts to investigate it under high powers generally result
in the pearl being crushed.
Several of the smallest pearls, only 6 or 7 \i in diameter, are associated with fan-
shaped crystalline bodies (IX. 8-12) ; these latter proved to be simply crystals
deposited from the sea-water. Sometimes the pearl is in the centre of a spheroidal
mass of crystals. Under polarised light, the crystals are doubly refracting and are
brilliantly defined on the dark field, while the pearls are isotropic, though usually very
faintly visible ; accordingly a spherical mass of crystals surrounding a pearl has a dark
central space.
In a vertical section of the sponge stained in borax-carmine the spheroidal
TETRAXONIBA. 9
conglomerations of crystals, 30-45 /u in diameter, are clearly defined owing to their being
much more deeply stained than the surrounding tissues. The appearance of capsules
in PI. IX. figs. 13, 14, is due to an optical effect. The deep staining and the well-
defined spheroidal shape might cause these bodies to be mistaken for organic structures ;
they are barely visible when unstained, and it is not easy to separate them from the
dehydrated tissues in which they lie, in order to test the action of reagents ; when
separated they are found to be soluble in water.
Localities. The specimens come from six different localities in the neighbour-
hood of Winter Quarters, from depths ranging from 10 to 25 fms. Also Kergueleu
Island, 25-60 fathoms, ' Voy. Challenger.'
ClNACHYRA VERTEX.
(Plate X., figs. 1-14.)
1907. Cinaclnjra vertex Lendenfeld (lla. p. 310).
Sponge, generally oval, ovoidal, or conical, with long diameter horizontal. Surface
with conules, usually oblique and more or less appressed to the surface, arranged in
spiral rows, and terminating in tufts of spicules, the conules and tufts wrapping round
the body. With well-developed root-tuft.
Porocalyces, varying greatly in size and appearance, 1 icing hemispherical pits with
circular orifice, or obliquely directed pockets with slit-like opening ; surface hispidated
with trichodal protriaenes.
With several oscules situated in a more or less circumscribed area on the side
opposite the area of greatest development of porocalyces, or at or near the summit of
conico-ovate specimens. The oscules were open only in one instance ; they were
3 25 x 2 mm. in diameter, with a marginal fringe, and a platform-like spicule below
the rim.
Ectosome, an outer layer of collenclyrnae, and an inner of fibrous tissue, with
cortical spicules.
Flagellated chambers eurypylous.
Spicules. Megascleres. (1) Somal oxeas, 5'5 x '057 mm., fusiform, tapering
gradually to sharp points.
(2) Choanosomal oxeas 520 x 18 p, slightly curved.
(3) Auatriaenes ; cladome, more or less hemispherical, often with apical umbo, cladi
at first at right angles, then making a sharp bend ; length 210 /u, thickness at base 70 /j.,
chorda 300 n, sagetta 200 /u : rhabdome 6'5/x broad at junction with cladome, then
narrowing slightly, again thickening slightly, and terminating gradually in a fine
sharp point.
(4) Protriaenes, cladome usually with one long clad us 1 70 ^ twice the length of
the other two; rhabdome 7'5 x '019mm. nearly uniform in thickness in distal
half, tapering at proximal end to fine sharp termination.
2 B 2
10 R. KIRKPATRICK.
(5) Trichodal protriaenes with rhabdome 550 /x in length, and with longest
cladus 50 fj,.
Microscleres. (6) Sigmata, 18'36/x in length, 1'8/x in thickness, and 10'5/xin
breadth in the C-aspect ; surface finely granular.
There are twenty-five specimens of this sponge, the largest being 10 '5 x 8 cm.
and the smallest 8x6 mm. The salient external features of the larger specimen