EXCHANGE

R E P O K T

ON THK

SCIENTIFIC RESULTS

OF THK

..

VOYAGE OF S.Y. 'SCOTIA.

SCOTTISH NATIONAL ANTAECTIC EXPEDITION.

EEPOKT

•

ON THE

SCIENTIFIC RESULTS

OF THE

VOYAGE OF S.Y. "SCOTIA"

DURING THE YEARS 1902, 1903, AND 1904,

UNDER THE LEADERSHIP OF

WILLIAM S. BRUCE,

LL.D., F.R.S.K

Volume IV.— ZOOLOGY.

PARTS II.-XX.— VERTEBRATES, by DAVID HEPBURN, M.D., F.R.S.E. ;
ROBERT B. THOMSON, M.B., Ch.B. ; D. WATERSTON, M.A., M.D.,
F.R.C.S.E. ; A. CAMPBELL GEDDES, M.D. ; W. A. HERDMAN, D.Sc.,
F.R.S. ; R. S. CLARK, M.A., B.Sc. ; HAROLD AXEL HAIG, M.B., B.S.,
M.R.C.S., L.R.C.P. ; WILLIAM S. BRUCE, LL.D., F.R.S.E.; R. N.
RUDMOSE BROWN, D.Sc.; W. EAGLE CLARKE, F.R.S.E., F.L.S.;
L. N. G. RAMSAY. M.A., B.Sc.; C. TATE REGAN, M.A. ; S. F.
HARMER, Sc.D., F.R.S., V.P.Z.S. ; W. T. RIDEWOOD, D.Sc.., F.L.S.;
and THEODORE E. SALVESEN, F.R.S.E.

62 Plates, 31 Text Figures, and 2 Maps

EDINBURGH :

SOLD AT

THE SCOTTISH OCEANOGRAPHICAL LABORATORY;

OLIVER & BO YD, EDINBURGH AND LONDON;
JAMES MACLEHOSE & SONS, 61 ST VINCENT STREET, GLASGOW.

1915.
Price Fifty Shillings.

EDITOEIAL NOTE.

ALTHOUGH absolutely essential in the interests of the Empire that the resources of
the country should be husbanded, it was most unfortunate that, just when the Scottish
Members of Parliament were unanimously resolved to press a vote for a further grant
of money for Scotia publications through the House of Commons, it was found neces-
sary to abandon the intention on account of urgent naval and military expenditure.

The original expedition of the Scotia depended to a very great extent upon the
munificent support afforded by the late Mr JAMES COATS, junior, and Major ANDREW
COATS, D.S.O. Thanks to the generosity of their brother, Sir THOMAS GLEN COATS, it
is now possible to place before the scientific world in particular, and the public in
general, this, the most comprehensive and not the least important volume of Scotia
research yet published, and that too at a period when, owing to the great European
crisis, it was scarcely possible to hope that further progress could, for the time being,
be made.

It is the most complete treatise on Antarctic vertebrate fauna yet published in
any country. Sir THOMAS GLEN COATS has thus helped to perpetuate the work begun
by his family, and, as a Scotsman, has also emphasised the national importance of
Scotland as a potent unit of the British Empire. Material for other six volumes
awaits publication, and it is hoped that this good example at a critical period will
stimulate others to provide the necessary funds to complete publication.

Volume IV. (Vertebrates) of the Scotia Reports is a contribution to Antarctic
Vertebrate Zoology. It also includes the consideration of Atlantic fishes besides those
of Antarctic regions.

I have to thank those who have voluntarily given their time and who have put
such excellent work into its pages.

It happens that this volume is the work entirely of British zoologists —
Wales, England, and Scotland having each taken part in its compilation.

In addition to the generous contribution of Sir THOMAS GLEN COATS,
the cost of publication has been aided by the original Government Grant, and
by a further Grant from the Common Good given by the Corporation of the City
of Glasgow out of the surplus of the Scottish Exhibition of Natural History, Art,
and Industry of 1911, and has also been considerably reduced by the co-operation of

VI EDITORIAL NOTE.

the Royal Society of Edinburgh, the Society having helped by bearing most of the
primary cost of setting up type and illustrations, several of the communications having
been passed through its Transactions. The Royal Society of London has paid the
entire cost for the production of Mr C. T. REGAN'S monograph on "Antarctic
Fishes" out of the Government Publication Grant which it administers. The
Carnegie Trust of the Universities of Scotland has paid the cost of Parts XL, XII.,
and XIII. The text figure in Mr R. S. CLARK'S " Atlantic Fishes " is reproduced by
permission of Mr M. J. NICOL and Messrs WITHERBY & Co. from Three Voyages of
a Naturalist. Special praise is due to Messrs HISLOP & DAY for their excellent
colour work in plates of the Ornithological Report.

Mr W. EAGLE CLARKE'S contributions originally appeared in the Ibis. Mr
R. S. CLARK'S "Fishes of St Helena" appeared in the Proceedings of the Royal
Physical Society. The other ornithological contributions by Mr L. N. G. RAMSAY,
Dr R. N. RUDMOSE BROWN, and myself, as well as the greater portion of Mr R. S.
CLARK'S "Atlantic Fishes," Dr RUDMOSE BROWN'S "Seals of the Weddell Sea,"
and Mr THEODOEE E. SALVESEN'S " Whale Fisheries of the Falkland Islands and
Dependencies," are published directly by the Scottish Oceanographical Laboratory
for the Reports.

Although obvious to many, the relationship between Science and Commerce
must be continually insisted upon, otherwise the sources are apt to get dried up
whence are drawn the funds upon which the scientist so largely depends.

As a vivid example of this relationship, Mr SALVESEN'S article is of special
interest in demonstrating how commercial enterprise follows scientific investigation.
Mr SALVESEN is the head of a large commercial business having its headquarters in
Leith, and it is a direct result of reports brought home by the Scotia naturalists
and others that he now has such large interests in the Antarctic regions. The great
whaling industry at present prosecuted in the neighbourhood of the South Shetlands,
South Orkneys, and South Georgia followed directly in the wake of the scientific
discoveries of the Swedish and Scottish Expeditions in the Weddell Sea. Before
the Scotia sailed there was not a deep sounding taken south of latitude 40° S. in
the Atlantic Ocean. The South Orkneys and South Shetlands were practically
unvisited, and almost entirely unknown. South Georgia was little known. Now over
a thousand people live under the British flag in South Georgia, and the South
Orkneys and South Shetlands are a busy hive of industry during the summer
months. Traders have even turned their attention to the west coast of the mainland
of Graham Land, a direct result of the scientific expeditions sent out by Belgium
and France, and altogether over a million pounds sterling of gross annual revenue
is now taken in these regions previously considered worthless by business men.

Some of the monographs published in these scientific results have previously
appeared in other publications, and as a consequence it has been suggested that

EDITORIAL NOTE. vii

although it is valuable to have memoirs such as are included in these Scotia volumes
collected together, yet it is unfortunate that the species recorded in them are
described in the work as new species. With regard to this, I may be allowed to
say that many of the monographs are avowedly reprints. This is expressly stated
not only in the "Editorial Note" of each volume, but also at the foot of every
page, where the volume and page of the Transactions of the Royal Society of
Edinburgh, or other scientific society, are clearly given. It has been further
suggested that " Much trouble may be thus caused by the annual biological records
again cataloguing these species, or of their being subsequently assigned to wrong
dates." I would .point out, moreover, that much expense would be incurred if
the necessary alterations had to be made out of extremely limited funds already
quite inadequate for the work yet to be done.

WILLIAM S. BRUCE,

Editor.

CONTENTS.

PART II. — SCOTTISH NATIONAL ANTARCTIC EXPEDITION : OBSERVATIONS ON THB ANATOMY OF
THE WBDDBLL SEAL (Leptonychotes Weddelli). By DAVID HEPBURN. M.D., Professor of
Anatomy, University College, Cardiff (University of Wales). (With One Plate). PART I. 1-12

(MS. received February 22, 1909. Read June 7th, 1909. Issued separately May 25, 1909.)

PART III. SCOTTISH NATIONAL ANTARCTIC EXPEDITION : OSTEOLOGY OF ANTARCTIC SEALS. By

ROBERT B. THOMSON, M.B., Ch.B., University of Edinburgh. (With One Plate) . . 13-32

(MS. received April 26, 1909. Read July 4, 1909. Issued separately October 12, 1909.)

PART IV. —REPORT UPON THE ANATOMY AND EMBRYOLOGY OF THE PENGUINS COLLECTED BY
THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION: Comprising — (1) Some Features in the
Anatomy of the Penguin ; (2) The Embryology of the Penguin : A Study in Embryonic
Regression and Progression. By D. WATERSTON, M.A., M.D., F.R.C.S.E., and A. CAMPBELL
GBDDBS, M.D. (From the Laboratory of the Anatomical Department, The University,
Edinburgh.) (With Three Plates) ...... 33-58

(MS. received March 11, 1909. Read February 3, 1908. Issued separately October 21, 1909.)

PART V. — SCOTTISH NATIONAL ANTARCTIC EXPEDITION : OBSERVATIONS ON THE ANATOMY OF
THE WEDDELL SEAL (Leptonychotes Weddelli). By DAVID HKPBUBN, M.D., C.M., F.R.S.E.,
Professor of Anatomy, University College, Cardiff (University of Wales). PART IT. :
GENITO-URINARY ORGANS ... ...... 59-66

(MS. received December 4, 1911. Read January 8, 1912. Issued separately January 19, 1912.)

PART VI. — SCOTTISH NATIONAL ANTARCTIC EXPEDITION : OBSERVATIONS ON THE ANATOMY OF
THE WEDDELL SEAL (Leptonychotes Weddelli). By DAVID HEPBURN, M.D., C.M., Pro-
fessor of Anatomy, University College, Cardiff (University of Wales). PART III. : THE
RESPIRATORY SYSTEM AND THE MECHANISM OF RESPIRATION .... 67-82
(MS. received March 28, 1912. Read June 3, 1912. Issued separately July 18, 1912.)

PART VII. — THE TUNICATA OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. By W. A.
HERDMAN, D.Sc., F.R.S., Professor of Zoology in the University of Liverpool. (With
One Plate) .......... 83-102

(MS. received January 8, 1912. Read February 19, 1912. Issued separately July 3, 1912.)

PART VIII. — SCOTTISH NATIONAL ANTARCTIC EXPEDITION : NOTES ON SOME FRESHWATER
FISHES FROM BUENOS AIRES. By R. S. CLARK, M.A., B.Sc., Scottish Oceanographical
Laboratory, Edinburgh. (With One Plate) . ... 103-110

(Handed in July 2, 1912. Issued separately August 12, 1912.)

X CONTENTS.

PAGES

PART IX. — SCOTTISH NATIONAL ANTARCTIC EXPEDITION : OBSERVATIONS ON THE ANATOMY OP
THE WEDDELL SEAL (Leptonychotes Weddelli). By DAVID HEPBURN, M.D., C.M.,
Professor of Anatomy, University College, Cardiff (University of Wales). (With One
Plate). PART IV. : THE BRAIN ... . 111-136

(MS. received June 18, 1912. Read December 2, 1912. Issued separately February 8, 1913.)

PART X. — SCOTTISH NATIONAL ANTARCTIC EXPEDITION : A CONTRIBUTION TO THE HISTOLOGY
OF THE CENTRAL NERVOUS SYSTEM OF THE WEDDELL SEAL (Leptonychotes Weddelli}.
By HAROLD AXEL HAIG, M.B., B.S. (Lond.), M.R.C.S. (Eng.), L.R.C.P. (Lond.), Lecturer
in Histology and Embryology, University College, Cardiff. Communicated by Dr W. S.
BRUCE. (With Two Plates and Nine Text Figures) ..... 137-158

(MS. received June 18, 1912. Read December 2, 1912. Issued separately February 17, 1913.)

PART XI. — MEASUREMENTS AND WEIGHTS OF ANTARCTIC SEALS TAKEN BT THE SCOTTISH
NATIONAL ANTARCTIC EXPEDITION. By WILLIAM S. BRUCE, LL.D., F.R.S.E., Director of
Scottish Oceanographical Laboratory, Edinburgh. (With Two Plates and One Text Figure) 159-174
(MS. received 17th February 1913. Read March 17, 1913. Issued separately August 4, 1913.)

PART XII. — SCOTTISH NATIONAL ANTARCTIC EXPEDITION : ON THE SKULLS OP ANTARCTIC
SEALS. By WILLIAM S. BRUCE, LL.D., Director of the Scottish Oceanographical
Laboratory. (With Five Plates) ........ 175-180

(MS. received March 13, 1913. Read May 5, 1913. Issued separately June 27, 1913.)

PART XIII. — THE SEALS OF THE WEDDELL SEA : NOTES ON THEIR HABITS AND DISTRIBUTION.
By Dr R. N. RUDMOSE BROWN, D.Sc., University of Sheffield, Naturalist, Scottish National
Antarctic Expedition. (With Nine Plates) . . . . . .181-198

(MS. received February 15, 1913. Issued separately May 20, 1913.)

PART XIV. — ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION : Sections I.
to XIV. By W. EAGLE CLARKE, F.R.S.E., F.L.S. ; L. N. G. RAMSAY, M.A., B.Sc. ;
R. N. RUDMOSE BROWN, D.Sc. ; and WILLIAM S. BRUCE, LL.D., F.R.S.E. (With Four
Coloured Plates, Three Black and White Plates, and Two Maps) .... 199-306
(MS. received July 31, 1913. Issued separately December 31, 1913.)

PART XV. — THE ANTARCTIC FISHES OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. By
C. TATE REGAN, M.A., Assistant in the British Museum (Natural History). Communi-
cated by Dr W. S. BRUCE. (With Eleven Plates and Six Text Figures) . . . 307-374
(MS. received June 18, 1912. Read December 16, 1912. Issued separately May 23, 1913.)

PART XVI. — SCOTTISH NATIONAL ANTARCTIC EXPEDITION : " SCOTIA '' COLLECTION OF ATLANTIC
FISHES. By R. S. CLARK, M.A., B.Sc., Zoological Assistant, Scottish Oceanographical
Laboratory, Edinburgh. (With Five Text Figures) ..... 375-402

(MS. received January 14, 1913. Issued separately June 30, 1913.)

PART XVII. — THE PTEROBRANCHIA OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.
By S. F. HARMBR, Sc.D., F.R.S., Keeper of the Department of Zoology in the
British Museum; and W. G. RIDEWOOD, D.Sc., Lecturer on Biology in the Medical
School of St Mary's Hospital, University of London. Communicated by Dr ASHWORTH.
(With Two Plates and Five Text Figures) ....... 403-442

(MS. received February 15, 1913. Read March 17, 1913. Issued separately July 4, 1913.)

CONTENTS.

XI

PART XVI 1 1. — SCOTTIMI NATIONAL ANTARCTIC EXPEDITION : A DESCRIPTION OF THB SYSTEMATIC
ANATOMY OF A F<KTAI. SEA LEOPARD (Stenorhynchus Itptonyx), WITH REMARKS UPON THE
MICROSCOPICAL ANATOMY OF SOME OF THE ORGANS. By HAROLD AXBL HAIG, M.B., B.S.,
M.K.C.S., late Lecturer in Histology and Embryology, University College, Cardiff; M'Robert
Research Fellow, University of Aberdeen. Communicat»l by Professor ARTHDR ROBINSON,
M.D. (With Four Plates and Three Text Figures) ..... 443-474

(MS. received January 26, 1913. Bead February 16, 1914. Issued separately April 30, 1914.)

PART XIX. — THE WHALE FISHERIES OF THE FALKLAND ISLANDS AND DEPENDENCIES. By

THEODORE K. SALVBSBN, F.B.8.E., Leith. (With Ten Plates and One Text Figure) . 475-460

MS. received December 9, 1913. Issued separately May 12, 1914.)

PART XX. — ANTARCTIC CETACEA. By WILLIAM 8. BRUCE, LL.D., Director of the Scottish

Oceanographical Laboratory. (With Two Plates and One Text Figure) . . . 487-504

MS. received October 21, 1914. Issued separately March 31. 1916.)

PARTS II. TO XX.
VEKTEBEATES.

PART II.

SEALS.

SCOTTISH NATIONAL ANTARCTIC EXPEDITION.
IL-ANATOMY OF THE WEDDELL SEAL

(LEPTONYCHOTES WEDDELLI).

(a) PERITONEAL FOLDS AND REFLECTIONS;
(b) ALIMENTARY ORGANS.

BY DAVID HEPBURN, M.D., F.R.S.E.,

Professor of Anatomy, University College, Cardiff (University of Wales).

Scottish National Antarctic Expedition : Observations on the Anatomy of the
Weddell Seal (Leptonychotes Weddelli). By David Hepburn, M.D., Professor of
Anatomy, University College, Cardiff (University of Wales). (With One Plate.)

(MS. received February 22, 1909. Issued separately May 25, 1909.)

INTRODUCTORY.

The anatomy of all marine mammals presents an interesting field of observation
upon their structural adaptation to a particular environment, and naturally, therefore,
an extensive literature already exists in respect to these mammals, notwithstanding the
many difficulties connected with their detailed examination. Their large size and the
rapidity with which their tissues undergo decomposition have been serious obstacles in
the way of prolonged dissection both of Cetacea and of Seals. Consequently, the
examination of many parts of their anatomy has, of necessity, been more or less hurried.
Accordingly, in the course of the preparations for the Scottish Antarctic Expedition,
led by W. S. BRUCE, LL.D., etc., arrangements were made for the preservation of
mammalian specimens by the same injection methods that are now commonly used for
the practical study of human anatomy, and the medical officer to the Expedition
received practical instruction from myself in the application of these methods.

The animal now under consideration was preserved in the following manner : — One
of the vessels between two of the toes on a hind flipper was opened and a fine canula
inserted. Through this, a quantity of an arsenical preservative, containing glycerine,
was introduced under gravitation, and to-day the tissues are as soft and free from
putrefaction as they were on the day the creature was killed.

The animal is a young male Weddell seal (Leptonychotes Weddelli), one of the
earless seals. It was born on or before the 29th of August 1903, and was killed by
hydrocyanic acid on the 31st of August 1903, and immediately injected with the pre-
servative solution.

Its dimensions are as follows :—

From tip of snout to end of tail ..... 5H ins.

„ „ „ hind nipper . 60i „

;, „ „ fore-limb . 13 „

„ „ genital aperture . . . . 39 „

From genital aperture to anus . . . . . 8 „

„ anus to tip of tail . . . . . . 4J „

Interorbital width . . . . . . 3f „

Axillary and greatest girth ..... 34J „

Fore flipper : —

Length on outer edge . . . . 11£ „

„ „ inner „ . 6£ „

Stretch (expanded) . . . . . 7£ „

(REPRINTED FROM THE TRANSACTIONS OF THE ROYAL SOCIETY OF EDINBURGH, VOL. XLVH., PP. 57-63.)

. •.

.

- . •

6 PROFESSOR DAVID HEPBURN ON

Hind flipper : —

Length on outer edge . . 13f ins.

„ „ inner „ . . . . 12 „

Stretch (expanded) .... 13J ,,

Circumference at base . . . l^i »

Its weight was 83 Ibs. A portion of umbilical cord about 3 inches in length was
still dangling from the umbilicus. Its fur was greyish yellow, mottled by darker spots,
suggesting leopard characters.

Abdominal Viscera. — On opening the abdomen, the general shape of the cavity was
seen to be ovoid, being wider in front and narrowing behind to the well-marked inlet
of the pelvic cavity. Regarded as a whole, the abdominal cavity presented a close
resemblance to that of the porpoise,* of which an account appeared in the Transactions
of the Royal Society of Edinburgh, vol. xl., part ii., p. 315.

Running forwards from the umbilicus, the umbilical vein, already nearly obliterated,
formed a very distinct object, suspended as it was in the free edge of an extensive
ventral mesentery constituting the falciform ligament of the liver. Running back-
wards from the umbilicus, the urinary bladder, with its associated hypogastric arteries,
was likewise suspended in a mesial ventral mesentery whose depth from the abdominal
wall to the bladder measured from an inch to an inch and a half. The bladder was
long and narrow, and at the umbilicus its lumen was distinct. No part of the bladder
had become obliterated to form the urachus, so that the bladder represented the entire
intra-abdominal extent of the allantois.

The liver and stomach occupied the wide anterior end of the abdominal cavity, and
in rear of them only the mass of coiled intestine was visible.

PERITONEAL FOLDS AND REFLECTIONS.

At first glance there was no sign of a great omentum, but on separating the coils of
intestine from contact with the stomach, the great omentum was found crumpled up
against its attachment to the greater curve of the stomach. Its greatest depth from the
stomach to its free margin was 3 inches. It was devoid of visible fat, and was there-
fore an extremely thin and translucent membrane. There was a well-marked gastro-
hepatic or lesser omentum presenting the usual gastric and hepatic connections. The
gastro-splenic omentum was also well marked, and by it the spleen was attached to the
great curve of the stomach. From the dorsal aspect of the spleen the peritoneal
membrane extended to the dorsal wall of the abdominal cavity without coming into
relation with the left kidney, which was situated opposite to its fellow and considerably
further back in the abdomen.

The liver was attached to the sub-diaphragmatic surface by the usual suspensory,
coronary and lateral peritoneal ligaments, to which further reference will be made in
describing the liver.

* HEPBURN and WATERSTON, loc. cit.
(ROY. soc. EDIN. TRANS., VOL. XLVII., 58.)

OBSERVATIONS ON THE ANATOMY OF THE WEDDELL SEAL. 7

The duodenum was suspended in a dorsal mesial mesentery whose base of attachment
measured nearly 5 inches, and this represented the distance between the pylorus
(gastro-duodenal junction) and the duodeno-jejunal junction which was situated close
behind (tailwards of) the superior mesenteric vessels and immediately to the left of the
mesial plane. To the right side of the duodenal mesentery there was a large peritoneal
recess whose right boundary was formed by a tailed lobe of the liver extending back-
wards along the dorsal wall in intimate association with the inferior vena cava.

At the first glance there appeared to be no naked-eye distinction between the small
intestine and the colon. At no point were tsenia coli, sacculations, or appendices
epiploicse visible.

As measured from the duodeno-jejunal junction or flexure, 46 feet of gut were
suspended from the dorsal wall in a mesentery practically corresponding to the entrance
of the superior mesenteric vessels. A closer examination revealed a distinct lateral
diverticulum rather more than 1 inch in length but of the same calibre as the gut,
situated on that part of the gut, which was suspended in the mesentery and at a
point 9 inches from the hinder end of the mesentery. This diverticulum evidently
represented the coecum and the vermiform appendix in their most primitive form, and
in the same condition as I have formerly described in the case of the grey seal
(Hcdichasnts grypus).* We may therefore conclude that this diverticulum marks
the commencement of the colon.

Consequently the first 9 inches of the colon are suspended in the same mesentery
as the small intestine, exclusive of the duodenum. The remainder of the gut, i.e. colon,
was suspended in a dorsal mesial mesentery extending tailwards to the entrance of the
pelvis, while through the greater part of the pelvic cavity a dorsal mesial mesentery
supported the pelvic part of the colon, i.e. the rectum.

The foramen of Winslow was very distinctly defined in relation to the free right
border of the gastro-hepatic omentum. Similarly the lesser sac of the peritoneum was
equally distinct, although the great omentum was not in any sense a gastro-colic structure.

The inlet of the pelvis measured 1 inch in the transverse diameter and 3 inches
in the conjugate diameter. Its lateral boundaries were well defined by the hypogastric
(umbilical) arteries, each of which was supported in a peritoneal sling or ligament attached
to the dorsal wall along the pelvic inlet.

The pelvic peritoneal pouch extended backwards between the gut and the bladder to
a distance of 3^ inches from the pubic crest, and so reached a point posterior to the
bulb of the urethra, i.e. to a point which corresponded with the central point of the
perineum. On the ventral abdominal wall, 1 inch to the outer side of the hypogastric
arteries, there was the opening which led into the inguinal canal (see fig.). Each
opening was circular and half an inch in diameter. It led into a circular peritoneal tube
which extended through the abdominal wall in relation to the hinder free border of the
musculus transversalis abdominis and the musculus obliquus interims abdominis. At

* HEPBCRX, " The Grey Seal (Halichcerus grypus)," Jour. Anat. and Phys., voL xxx.

(HOT. SOC. EDIN. TRANS., VOL. XLVIL, 59.)

8 PROFESSOR DAVID HEPBURN ON

the plane of the musculus obliquus externus abdominis, and close to its pubic attach-
ment, the peritoneal tube passed through a muscular slit corresponding to the external
inguinal ring and thereafter terminated in a blind end situated close to the posterior
end of the testis. The length of this tube of peritoneum was 2^ inches, and its
testicular part formed the tunica vaginalis testis.

Each vas deferens entered the abdomen through the internal inguinal ring, being
suspended from the ventral wall in a short mesentery or peritoneal sling by which each
vas was carried across the abdominal aspect of the hypogastric artery to the pelvic
surface of the urinary bladder about the level of the pubic crest. These two mesenteries
for the vasa deferentia formed a free transverse fold of peritoneum on the pelvic surface
of the urinary bladder at the level indicated. From this level the vasa deferentia passed
backwards on the pelvic surface of the bladder under cover of the peritoneum. On the
pubic aspect of the bladder there was a triangular non-peritoneal surface extending for-
wards from the pubic crest for If inches, at which point the ventral mesentery of the
bladder commenced and continued to the umbilicus.

ALIMENTARY ORGANS.

The stomach (see fig.) presented a single chamber situated with its long diameter
in the axis of the trunk. The oesophagus entered the stomach slightly to the right side
of the most prominent or anterior part of the fundus. The greater convexity or curve
was well defined, and measured 17 inches from the oesophagus to the pylorus. The
distance between the same points along the lesser curve was 1 0 inches. Each of these
curves was associated with the usual omenta. A very sharp bend occurred in the lesser
curve, 5 inches in a straight line from the oesophagus and 3 inches from the pylorus.
The sides of this bend were held in close apposition by the peritoneum, and the general
appearance produced was that of a constriction in the course of the cavity of the
stomach. The greatest width of the stomach on the oesophageal side of this bend was
5^ inches, whereas on the pyloric side of the bend the greatest width was 4 inches, and
opposite the bend the width was 2^ inches. The practical result of the infolding of the
stomach wall was therefore to produce two chambers communicating with each other
by an aperture considerably narrower than either of the chambers. A second slight
constriction was present in the pyloric section, and thus as a whole, from oesophagus to
pylorus, the stomach suggested three imperfectly separated compartments or chambers.

The contents of the stomach consisted of a quantity of a thick pasty substance of a
somewhat light earthy colour. It was uniformly smooth, and contained no evidence of
bones. In all probability it represented partially digested coagulum of milk.

The pylorus was placed in the mesial plane, and was recognisable both to the eye
and to the touch as a constricted ring.

The duodenum extended from the pylorus in the form of a horse-shoe loop 1 2 to
1 3 inches long. It was attached dorsally in the mesial plane by a mesentery. The

(ROY. soc. EDIN. TRANS., VOL. XLVII., 60.)

OBSERVATIONS ON THE ANATOMY OF THE WEDDELL SEAL. 9

proximal and distal ends of the loop were from 4 to 5 inches apart. To the right side
of the duodenal mesentery there was a large peritoneal recess or pouch the mouth of
which was directed anteriorly, and to which reference has already been made.

The duodeno-jejunal flexure lay immediately to the left side of the vertebral column
and marked the beginning of the mesentery proper and of the small intestine suspended
in it. This mesentery had a very short base which practically corresponded to the
entrance of the superior mesenteric vessels. It was twisted to the right and supported
the jejunum and ileum, which together measured almost 46 feet in length.

The coacal diverticulum formerly referred to as representing the combined ccecurn
and vermiform appendix, was taken to indicate the termination of the small and the
beginning of the large intestine. This primitive developmental form of the coecum
and vermiform was rather more than an inch in length, while in calibre it corresponded
with the gut. From this diverticulum to the end of the gut there were no other
external evidences of any distinction between small and large intestine.

From the coacurn the large intestine or colon pursued the first 9 inches of its
course suspended in the same mesentery as the small intestine. Thereafter the colon
assumed a mesial position and, as far as the pelvic inlet, i.e. for a distance of 18 inches,
it was suspended in a dorsal mesial mesentery. The pelvic portion of the colon was
also placed mesially, and the greater part of it was also suspended in a dorsal mesentery.
From the pelvic inlet to the anal aperture the gut measured nearly 10 inches, so that
the entire length of the colon from ccecum to anus was practically 3 feet. Thus the
total length of the gut from pylorus to anus was : —

Duodenum ...... 1 foot.

Small intestine . . . . . .46 feet.

Colon . . . . . 3 „

Total . . . . . 50 „

The liccr (see figs.) was large and deeply fissured, thereby presenting very dis-
tinct lobes. It was intimately associated with the diaphragm, and occupied the anterior
end of the abdominal cavity from side to side. It was provided with the usual peritoneal
ligaments. The falciform and coronary ligaments presented no special features as
regards their arrangement, but the left lateral ligament extended from the sharp left
margin of the liver whereas the right lateral ligament was short and extended from the
smooth surface of the right aspect of the right lobe. The diaphragmatic surface of the
liver was smooth and convex, adapting itself to the abdominal surface of the diaphragm
and presenting right and left lobes in relation to the suspensory or falciform ligament.
The right lobe was divided into mesial and lateral portions by a deep dorso-ventral
fissure, and the left lobe was imperfectly divided by dorsal and ventral notches which,
however, did not meet each other.

On its visceral aspect the liver was much subdivided, particularly in relation to the
right lobe. The right and left lobes were marked off from each other by the ligamentum
teres (obliterated umbilical vein) on the ventral aspect of the hilum, and by the hepatic

(ROY. soc. EDIS. TRANS., VOL. XLVIL, 61.)

10 PROFESSOR DAVID HEPBURN ON

sinus on the dorsal aspect of the hilum. The right lobe presented the same mesial and
lateral portions which were noted on the diaphragmatic aspect. The mesial portion
was divided into dorsal and ventral parts by the hilum. To the ventral portion the
gall-bladder was attached, and this part of the right lobe was connected to the left lobe
by a pons hepaticse. The dorsal portion was to a large extent concealed by the large
hepatic sinus. The lateral part of the right lobe was also divided into dorsal and
ventral segments by the extension of the hilum across its visceral aspect.

A tailed lobe extending from the dorsal segment of the mesial portion of the right
lobe passed backwards on the ventral surface of the inferior vena cava, which was thus
embedded in the liver substance. This tailed lobe expanded so as to conceal a consider-
able length of the inferior vena cava, and into this hidden part of the cava there opened
directly the hepatic veins from this particular lobe, as also the hepatic veins from the
lateral part of the right lobe.

The inferior vena cava itself opened into the large hepatic sinus situated close to the
diaphragm and extending right and left of the suspensory ligament. This sinus
received the hepatic veins from the right and left lobes of the liver, with the exception
of those veins already mentioned as opening directly into the inferior vena cava. The
interior of this sinus was imperfectly divided into right and left parts by a crescentic
partition which was situated to the right side of the line of attachment of the suspensory
ligament, so that the part of the sinus to the left side of the crescentic fold was the
larger. The inferior vena cava opened into the hepatic sinus on the right side of the
imperfect partition, while on its left side it received the mouth of the ductus venosus.

The hepatic sinus narrowed for the purpose of passing through the diaphragm in
order to enter the right auricle of the heart, but the general size of its lumen was so
much greater than that of the inferior vena cava that it is more accurate to say that
the inferior vena cava opened into the sinus and the sinus joined the auricle.

The conditions presented by the hepatic veins afford some interesting light upon
the question of their development. Clearly the large hepatic sinus has resulted from
the fusion of the two embryonic vense reveherites, although, from the position of the
imperfect crescentic partition, it is evident that the left vena revehens was the larger of
the two and therefore received the smaller or right vena revehens. This arrangement
would therefore appear to provide a variation upon the current statement that " the
left vena revehens loses its connexion with the sinus venosus and opens into the right
vena revehens" (Cunningham's Text-Book of Anatomy, 2nd ed., p. 935). Again, the
ductus venosus is described as passing directly " from the left vena advehensto the right
hepatic vein" (vide ibid.), whereas, in the liver under consideration, the mouth of the
ductus venosus opens to the left side of the crescentic fold, which appears to represent
the remains of a fusion between the right and left hepatic veins. Further, to quote
again from the same authority, " The upper part of the inferior vena cava is developed
as an outgrowth from the common trunk formed by the fusion of the ductus venosus
with the right hepatic vein." From the present dissection the inferior vena cava would

(BOY. soc. EDIN. TRANS., VOL. XLVII., 62.)

OBSERVATIONS ON THE ANATOMY OF THE WEDDELL SEAL. 11

appear to arise from the right hepatic vein independently of the ductus venosus, more
especially in view of the fact that the inferior vena cava receives direct tributaries from
the caudate lobe adherent to its ventral aspect.

The gall-bladder, which was empty, occupied a fossa on the visceral aspect of the
mesial portion of the right lobe of the liver. It thus lay to the right side of the
ligamentum teres, from which it was separated by a projecting portion of liver
substance, connected to the left lobe by a pons hepaticae. This portion of the liver
corresponded in general position with the quadrate lobe of the human liver.

The cystic duct passed towards the hilum of the liver and, having been joined by
hepatic ducts corresponding in number with the three chief lobes of the liver, the
common bile-duct was formed. This duct pursued its course on the ventral aspect of
the foramen of Winslow and, immediately after passing this foramen, i.e. just before
reaching the duodenum, it was joined by the pancreatic duct. The duct thus resulting
came in contact with the duodenum a little more than half an inch from the pylorus.
It perforated the duodenal wall very obliquely, and opened on the summit of a papilla at
a distance of 2 inches from the pylorus.

The pancreas presented a characteristic appearance. A small part of this gland
was found within the duodenal mesentery, but the greater part of the organ extended
headwards behind the lesser peritoneal sac. The pancreatic duct emerged from the
substance of the organ on the cephalic (anterior) side of the foramen of Winslow (not
from that part within the duodenal mesentery), and, extending to the right side, it
formed a junction with the common bile-duct on the cephalic side of the foramen of
Winslow and fully 2 inches from the common orifice by which both ducts pour their
contents into the duodenum.

The spleen measured from 9 to 10 inches in length, from 1 to 6 inches in width,
being narrow anteriorly and wide posteriorly. It was extremely thin, being not
more than a quarter of an inch in thickness. It was situated between the stomach and
the dorsal wall, being attached to the former by the gastro-splenic omentum and to the
latter by a dorsal mesial mesentery. There was no intimate relation between the spleen
and either of the kidneys, because these organs were removed from all immediate
association by reason of their situation considerably nearer to the pelvic region. The
tail end of the pancreas extended into the posterior splenic mesentery but did not come
in contact with the spleen.

(ROY. soc. EDIX. TRANS., VOL. XLVII., 63.)

Scot. Nat. Ant. Erp., Vol IV.

Prof. DAVID HEPBURN on "Observations on the Anatomy of the Weddell Seal."'

CESOPH.

Stomach.

LEFT.

DORSAL.

LIG. TERES.

VENTRAL.

Liver of seal. View from abdominal aspect.

-LOBE ON VENTRAL
ASPECT OF INF. V.
CAVA (TURNED TO-
WARDS DORSUM).

RIGHT.

>

i

§

CO

\^

LIG. TERES.

Liver of seal (seen from ventral aspect).

URINARY BLADDER.

HYPOGASTRIC A.

INT. ING. RING.

VAS DEF.

Pelvic cavity of seal (seen from pelvic inlet).

PART III.

SEALS.

SCOTTISH NATIONAL ANTARCTIC EXPEDITION.
III.-OSTEOLOGY OF ANTARCTIC SEALS.

BY ROBERT B. THOMSON, M.B., Ch.B., University of Edinburgh.

(WITH ONE PLATE.)

Scottish National Antarctic Expedition : Osteology of Antarctic Seals. By
Robert B. Thomson, M.B., Ch.B., University of Edinburgh. (With One Plate.)

(MS. received April 26, 1909. Read July 4, 1909. Issued separately October 12, 1909.)

Through the courtesy of Dr W. S. BRUCE, the leader of the Scottish National
Antarctic Expedition, and of the late Professor D. J. CUNNINGHAM, the skeletons of the
seals collected in the Antarctic during the voyage of the Scotia have been placed in my
hands for study, and I wish to express my gratitude to these gentlemen for their great
kindness in placing all the necessary requirements at my disposal. I also wish to state
that Dr BRUCE has given to the Museum of the University of Edinburgh a choice of
the various skeletons, which, together with the specimens already obtained, will form
a comprehensive and almost unique collection of the Antarctic seals.

The seals obtained during the stay of the Scotia in the Antarctic seas numbered
forty-four, and embrace all the Phocidae found there, with the exception of the
elephant seal. Of the forty - four, thirty-four are Weddell seals (Leptonychotes
Weddelli); four are saw-toothed seals (Ogmorrhinus or Lobodon carcinophaga) ;
three are leopard seals (Stenorhynciis leptonyx) ; one is a sea-lion (Otaria Juhata) ;
while two are Ross seals (Ommatophoca Rossi).

A most careful account of date of capture, sex, probable age, full measurements, and
weight was recorded in a special log-book. In some cases the skeletons were cleaned
by immersing the carcases beneath the ice, when amphipods and cushion-starfish did
the work ; in other cases the carcases were conveyed in special tanks and cleaned after
their arrival home.

The osteology of the more common species of the Phocidse represented in this collec-
tion, such as the Weddell seal, saw-tooth seal, sea-lion, and leopard seal, appears to have
been pretty fully worked up, and especially so in a minute and careful description by
Principal Sir WILLIAM TURNER ; for the present, therefore, the endeavour of this work is
to present a detailed description of the Ross seal, as I have not been able to find such
of the skeleton of this species, with the exception of the skull. In the second place, I
purpose making a summary of the large collection of other seals with regard to such
points as dentition and vertebral formula, since the collection is large enough to obtain
a satisfactory average.

Ross SEAL (Ommatophoca Rossi).

The first specimen of Ommatophoca was obtained by Sir JAMES Ross in 1840, in
the pack-ice in the Ross Sea, and is now preserved in the British Museum as the
type-specimen. I am indebted to the authorities of the British Museum and to
Dr 0. THOMAS for the privilege of examining this specimen for purposes of com-

FROM THE TRANSACTIONS OF THE ROYAL SOCIETY OF EDINBURGH, VOL. XLVII., PART I., PP. 187-201.)

18 MR ROBERT B. THOMSON: SCOTTISH NATIONAL ANTARCTIC EXPEDITION:

parison. Unfortunately, it is not a complete skeleton : both fore-limbs with the
exception of one scapula, both hind-feet, one dorsal vertebra, and two pieces of sternum
are awanting. A description of the skull was made by Dr J. E. GREY in 1844. For
over fifty years it remained the sole representative of its race in natural history
collections, as not till 1899 was the Ross seal again captured by the Belgian Expedi-
tion (1898-9). Since then numerous specimens have been obtained and the skins
and crania brought home by the Southern Cross (1898-1900), Discovery (1901-4),
Morning (1902-4) ; while the Scotia captured two specimens — a male and a female, of
which the male skeleton is complete with the exception of two segments of the sternum
and one patella. The female skull has been unfortunately damaged, the occipital
region being wanting ; but a perfect set of teeth, of so much importance in Om/matophoca,
is preserved. Further, the whole respiratory apparatus of the male Ross seal has been
brought home, and is of great interest — as the voice of this animal, which has been
described by Dr RACOVITZA, with the peculiar inflation of the pharynx, has attracted
the notice of, and been recorded by, most Antarctic explorers.

The method of describing the skeletons is that adopted by Sir WILLIAM TURNER
in Challenger Reports, and I wish to express my appreciation of its arrangement and

carefulness.

Surface Measuremenls (from Log-Book).

No. 2. No. 43.

Dale of Capture— 6ih February 1903. 28th February 1904.

Sex — Female. Male.

Age — Adult. Adult
Length —

Nose to tip of tail, 90". 89 -5".
Girth—

Anterior or neck, 41". 38'5".

Axillary, 52". 55'5".

Greatest (9 ins. behind axillary), 54". (12 ins. behind axillary), 56'5".
Tail, 5".
Fore flippers —

Outer edge, 16". 17-2''.

Base, 8|".
Hind flipper —

Outer edge, 16". 18'4".

Inner edge, 14|". 18'7".

Stretch, 25". 24".

Base, 10-7". 11".

Eye, l|"xl". l-4"x8".

Interorbital space, 6". 6-5".

Total weight, 450-500 Ibs. 400 Ibs.

Remarks.

Intestines of No. 43 measured from pyloric end of stomach to anus 268". Cuttlefish
beaks, pieces of cuttlefish, and what appeared to be scales of fish were found in the
stomach. A great quantity of tape-worms were present in the gut.

(HOY. soc. EDIN. TRANS., VOL. XLVII., 188.)

OSTEOLOGY OF ANTARCTIC SEALS. 19

No. 43 OF COLLECTION (MALE).

The skull is characteristically short and broad. The greatest length was from
premaxilla to the occipital condyles, whilst the greatest breadth was between the two
zygomatic arches, just at the articulation between its two component parts — the zygo-
matic process of the temporal and the malar bones.

A comparison of this skull was made with those of an adult Weddell and sea-leopard
in regard to length and breadth — an index being framed. Thus, the length -breadth
index of the Ross skull was 7 2 '4 ; of the Weddell skull, 62' 13 ; of the leopard seal, 5 5 '07.
The nasals measured 79 mm. long, and were completely ankylosed. They articulated
with the frontal and superior maxillary bones. The part between the two frontals
amounted to about three-quarters of the entire length of the bones (57 mm.), and was
triangular in form, while the remaining quarter between the two superior maxillary bones
was quadrilateral.

The distance between anterior edge of the outer border of the nasals and the tip of
premaxilla was 18 mm., the superior maxilla thus forming to this extent part of the
outer boundaries of the anterior nares. " The length of the section of the boundary
formed by one of these bones varies in the different specimens from 9 to 1 7 millimetres "
(BARRETT HAMILTON, Resultats du Voyage du S.Y. " Belgica," 1897-1899, p. 5).
The anterior nares sloped downwards and forwards at an angle of 59° with the hard
palate. In Weddell seal this angle was 48°, in leopard seal 35°.

The anterior nares were bounded from above downwards by the anterior border of
the nasal bones, part of nasal borders of superior maxilla, and by the premaxillary bones.
Looking into the anterior nares, one was struck with two points — the thickness of the
meso-ethmoid, and the extremely convoluted arrangement of the turbinate bones. The
anterior edge of the vomer was received between the meso-ethmoid and the premaxillary
bones. The premaxillary bones supported the two incisor teeth ; their palatal parts were
triangular in form, and, as before mentioned, their nasal parts did not extend so high as to
meet the nasal bones. The ante-orbital process of the superior maxilla was well marked,
and lay in the same transverse plane as the infra-orbital foramen, below which is a
definite depression from which a groove leads to the orbital floor. The widest part of
the hard palate was situated well behind the last molar, and was 7 mm. in front of outer
end of articulation of palatal processes of the superior maxilla and palate bone. It was
almost flat, showing only a trace of a concavity. The posterior border of the hard
palate was concave, and possessed a faint post-nasal spine. On the norma verticalis of
the skull, at the junction of the frontal bones, i.e. at the seat of the anterior fontanelle
in the young, is situated an opening between the bones. Its margins are irregular and
spiculated (see fig. 1). As there is no history of an injury, the animal not being
clubbed, but shot in the thorax, it appears to me to be a persistent anterior fontanelle.
In the female Ross seal this condition is only faintly represented, but in the plate of
the Ross seal of the Belgian Expedition a similar well-marked deficiency is to be

(ROY. soc. EDIN. TRANS., VOL. XLVIL, 189.)

20 MR ROBERT B. THOMSON: SCOTTISH NATIONAL ANTARCTIC EXPEDITION:

observed. The occipital bone presented occipital crests which were well marked, as also
were the par-occipital processes. Two well-marked supra-occipital venous foramina
were to be noticed in the middle line of the occipital squama. The condylar articular
surfaces were highly convex and approached close to each other on the inferior aspect
(7 mm.) ; above, they were distant 41 mm.

The basi-occipital was quadrilateral in form, Hat and thick. The tympanic bulla
was smooth and rounded. The carotid canal was separated from the jugular foramen
by a thick bar of bone. In the interior of the skull the tentorium cerebelli and falx
cerebri were partly ossified.

In the female skull the nasal bones were not ankylosed, and measured 55 mm. long.
The distance between the anterior edge of the nasals and the premaxilla was 20 mm. on
right side, 18 mm. on left.

The Lower Jaw.

The body was long and somewhat slender, due to small size of teeth, with lower
border slightly incurved. The ramus formed with the body a moderate angle and
possessed a well-marked sub-condyloid process. The condyle, which was convex and
elongated transversely, was separated from the coronoid process by a shallow sigmoid
notch. The coronoid process was short and pointed.

Dentition.

The dentition of the Ross seal is remarkably feeble, and varies more especially with
regard to the number of post-canines. Dr E. A. WILSON, in vol. ii. of Natural History
of National Antarctic Expedition, 1907, states : "The food of the Ross seal consists
mainly of soft-bodied cephalopods, and to this end has developed the incisors and canines
into needle-pointed recurved hooks of great delicacy and has allowed its post-canines to
degenerate. The gums presumably can manipulate such food as well as could molar
teeth, and so we find in some cases the post-canines are small and insignificant, whereas
in others they are loose and useless, and occasionally absent altogether." He also gives a
complete table of the dentition of all the Ross crania which have been brought home,
and which shows the great variations one meets in the dental formula. To this list I
would add the formulae of the two specimens of the Scotia :—

No. 2. Female, 2- L 5

No. 43. Male,

2. 1. 5

2. 1. 5

2? 1. 5?-

No. 2 skull presents for examination a perfect set of teeth, both upper and lower.
The upper incisors are more powerful than the lower, but both present, like the canines,
the well-marked recurved, needle-pointed character. The post-canines are recurved

(ROY. BOC. EDIN. TRANS., VOL. XLVII., 190.)

OSTEOLOGY OF ANTARCTIC SEALS.

21

backwards and three-cusped, the central one being longest. In all but the first the
post-canines were double-rooted.

In No. 43 the teeth are not so well developed, and in the upper jaw the left outer
incisor and caniue are broken. The dentition is not quite easily made out. There is
no fifth post-canine, but small sockets are to be observed ; hence we may presume that
they were present, having been partly fixed in the natural . state by the fleshy gum, a
condition pointed out by WEBER, BENNET, and WILSON (National Antarctic Expedition :
Natural History, vol. ii. p. 46).

Again, in the lower jaw no central incisor is present, but the two halves of the jaw
have been disarticulated. On placing the two halves together, it would seem that no
room at all is left for central incisors. A comparison of the measurements of the two
lower and upper jaws between outer side of canines shows :—

No. 2. Upper diameter, 44 mm.

Lower „ 30 „
No. 43. Upper diameter, 48 ,,

Lower „ 26 „

In No. 2, which has not been disarticulated, there is a fair amount of fibrous tissue,
so it is possible to explain the difference in diameters by the absence of this. Hence,
with the fibrous tissue in place in No. 43, room would be made for central incisors.

Against this may be stated the fact that the upper central incisors are very feeble,
and, as the whole of the upper teeth are stronger than the lower, it is possible that the
lower central incisors may not have developed.

Measurements of Skulls.

Ross Seal,
No. 43.

Ross Seal,
No. 2.

Weddell
Seal.

Leopard
Seal

Extreme condylo-premaxillary length .

mm.
250

mm.

mm.
272

mm.
39*

„ inter-zygomatic width ....

181

169

217

„ width behind external meatus

167

177

198

Greatest width of palate ..... 76

71

58 90

Width between outer side of base of upper canines 48

44

52

75

„ ,, „ lower canines 26

30

30

50

Length of palate in line of suture to central incisor

81

82

105

157

Height of skull — basion to mid-occipital crest

83

...

84

110

Smallest interfrontal diameter ....

45

49

30

45

Length of nasals .......

79

55

61

103

Greatest width, anterior nares

43

34

36

44

Length of mandible ......

168

170

171

301

Width between outer end of condyle .

182

168

57

205

The vertebral formula was :—

D

Cd,

'15 ^5 "3 ^"10'

(BOY. soc. EDIS. TRANS., VOL. XLVIL, 191.)

22 MR ROBERT B. THOMSON : SCOTTISH NATIONAL ANTARCTIC EXPEDITION :

Cervical Vertebrae.

The bodies of the posterior six cervical vertebrse were keeled on their ventral aspect,
the keel being more pronounced in the anterior than in the posterior series. On con-
trasting the bodies of the Ross seal with those of the Weddell and leopard seal, a
great difference was noticed with regard to their length and breadth. If the 4th
cervical be taken as a typical vertebra, we find the length index of the body to be :—

In the Ross seal— Length 33 ^J^° = Index 145'4
Breadth 48 33

In leopard seal — Length 51 47 x 100^ jn^ex 92-16
Breadth 47 51

In Weddell seal — Length 38 _ jndex 1QO

Breadth 38

If these indices be compared with the length-breadth index of the skull, one is
justified in saying that the short and broad character of the Ross skull can also be
applied to the bodies of the vertebrse. The neural ring was triangular. A depth-
breadth index shows much the same difference as the bodies in the three species
contrasted. In the Ross seal, the index was 35 'I ; in Weddell seal, 54'5 ; and in the
leopard seal, 50.

The transverse processes in all except the 7th were perforated at their roots by a
vertebraterial foramen, and possessed at their extremities well-marked tubercles in all
except the 2nd and 7th. The spinous processes exhibited a marked difference in the
various vertebrse. Thus the axis possessed a massive spine with four well-marked
tubercles — two anteriorly and two posteriorly. The 7th had a well-marked spine, not
bifid ; the 3rd and 4th had only a trace of a spine, while the 1st possessed a well-marked
tubercle representing a spine. The 5th and 6th presented fairly prominent spines, bifid
—a character not shown by either the Weddell or leopard seals. The type-specimen
in the British Museum also exhibits this character. The anterior articular processes
looked upwards and inwards, the posterior downwards and slightly forwards.

The atlas has powerful lateral masses supporting anteriorly deeply concave articular
surfaces for occipital condyles, while the posterior articular facets for the axis were flat.
The transverse processes were plate-like, and projected outwards and downwards. The
lamina was perforated on each side by a foramen for the vertebral artery, which foramen
was situated internal to the upper ends of the articular facets for the occipital condyles.

The axis did not present so well-marked an odontoid process as either the Weddell
or leopard seal. On the inferior aspect of the odontoid process was a triangular facet
for articulation with the atlas, whilst superiorly it had a small oval facet for articulation
with the transverse ligament. The transverse processes were feeble and pointed, whilst
their superior roots were mere spicules of bone. The ventral aspect of the body was
markedly keeled.

(ROY. soc. EDIN. TRANS., VOL. XLVIL, 192.)

OSTEOLOGY OF ANTARCTIC SEALS. 23

Dorsal Vertebrae.

The bodies of the anterior four were keeled, the keel becoming less pronounced as
we proceeded from 1 to 4. At the posterior end of the dorsal region, 14 and 15 pre-
sented a slight keel, more accentuated in 15 than in 14. The side of the 1st body had
one and a half facets ; 2 to 10 inclusive (two half-facets, 11 to 14 inclusive) a whole facet
for the corresponding ribs. The 1 5th dorsal vertebra showed a peculiar arrangement ; thus,
on the right side of its body a well-marked facet was present for articulation with the 15th
rib of that side, while on the left a well-marked plate-like costal process, representing
the last rib, projected from the junction of the body and pedicle outwards, forwards, and
slightly downwards. It measures 42 mm. in length and 25 mm. in breadth. (See
fig. 5.)

The spines of the dorsal vertebrae were relatively feeble compared with those of the
Weddell and leopard seals. They were low, and projected upwards and backwards as
far as No. 11 ; while 12, 13, 14, 15 projected upwards.

The transverse processes of 1 to 10 were prominent and presented rounded facets for
articulation with the tubercles of the ribs. In 11, 12, and 13 the transverse processes
were feeble but possessed facets. In 14 and right side of 15 they were rudimentary
and non-articular, while the character of the transverse process of left side of 15 has
already been noticed. From 9 to 15 the mammillary processes were well defined, while
accessory processes were faintly marked in 9 to 14.

Lumbar Vertebrx and Sacrum.

Each body of the lumbar vertebrae was elongated antero-posteriorly and was keeled
on its ventral aspect. The spines were broad and more pronounced than in either the
Weddell or leopard seals. The mammillary processes were rounded and directed
forwards and outwards. The transverse processes were thin and flat, and curved out-
wards, forwards, and downwards. The sacrum was composed of three segments fused
together. The ventral surface in its anterior part was broad and flat, with a faint keel,
while its posterior part, corresponding to the 2nd and 3rd segments, was protuberant
in the centre, concave laterally, where it presented the openings of the two anterior
sacral foramina. The posterior surface showed mesially three spinous processes, while
the laminae of the three segments were only fused laterally. The fused articular
processes were prominent at the junction of S: and S2, less so at S2 and S3. The
direction of the 1st posterior sacral foramen was horizontally outwards, in the 2nd and
3rd upwards and only slightly outwards. Laterally the sacrum presented a definite
U-shaped articular surface for ilium, while the interval between the two limbs of the
U and the area posterior to it were hollow for ligamentous attachment. The articular
surface for the ilium was confined entirely to the first sacral segment.

(ROY. soc. EDIN. TRANS., VOL. XLVII., 193.)

24 MR ROBERT B. THOMSON : SCOTTISH NATIONAL ANTARCTIC EXPEDITION :

Sacral Index.

Ross seal . . . . . 108

Weddell seal ... 102 -9

Leopard seal . . . . . 114'8

The caudal vertebrae were ten in number. The posterior six were represented by
elongated bodies alone — the last being cartilaginous. The 1st possessed, in addition to
the body, a definite neural arch terminating above in a rudimentary spinous process.
The laminae in the 2nd did not unite superiorly, so that the neural groove was not con-
verted into an arch. The third possessed on each side of its body rudimentary laminse.
The bodies of the first four caudal vertebrae were keeled on their ventral aspect. The'
sternum was incomplete, two segments being awanting. Their articulations could not
be made out definitely, so I refrain from describing them.

Ribs.

The ribs were fifteen in number on the right side and fourteen on the left, this anomaly
being explained in connection with the thoracic vertebrae. The 1st, 12th, 13th, 14th,
and 15th ribs possessed a single facet each on their heads for articulation with the
corresponding vertebrae. The 2nd to llth inclusive possessed two facets separated by
a distinct ridge. The 1st to 12th ribs inclusive possessed well-defined necks, while the
1st to 13th showed an oval concavo-convex facet for articulation with the corresponding
transverse processes. In the anterior series of ribs was a slight depression, in some a
roughness, on each side of the articular surface of the head, for attachment of ligamentum
conjugate costarum described by Sir WILLIAM TURNER (Challenger Reports, vol. xxvi.,
Report on Seals, p. 14). The shafts of the ribs were slightly curved, the most pro-
nounced curvatures being observed in the 2nd, 3rd, and 4th, while the greatest in
length were the 8th and 9th. The last rib on the right side possessed a head with an
articular facet, no neck, and a feeble shaft. Its length was 127 mm.

The Fore- Limb.

The scapula presented the well-marked sickle shape so characteristic in the Phocidse.
The anterior and superior or vertebral borders were convex and thin, the inferior or axil-
lary was concave and rounded. On the axillary border at junction of lower and middle
thirds in the Weddell and leopard seals was a prominent ridge for muscular attachment.
No such ridge was present in the Ross scapulae. The dorsum scapulae was divided into
a pre-spinous and post-spinous area by a feeble spine. This spine was relatively longer
than in the Weddell and leopard seals, but not so well developed. The pre-spinous area
was concavo-convex, the post-spinous was flat. There was no acromion process, but in
the Weddell and leopard seals this was present although feeble. The coracoid process
was rudimentary and projected forwards, measuring from the bottom of the shallow

(ROY. soc. EDIN. TRANS., VOL. XLVII., 194.)

OSTEOLOGY OF ANTARCTIC SEALS. 25

notch at its base 18 ram. The ventral surface showed a slight subscapular angle cor-
responding in position to the attachment of the spinous process on the dorsal surface.
Two faint muscular ridges were present, directed towards the glenoid fossa. The
glenoid fossa was pear-shaped, deeply concave, with a prominent margin.

Index.

Ross seal . . . . .85

Weddellseal ..... 82'14

In the leopard seal the epiphysis of the posterior angle was wanting, hence the
index could not be ascertained.

The humerus measured 115 mm. long. The head was convex, and its articular
surface was continued forwards on to the base of the lesser tuberosity, which was
equally as prominent as the greater one. The shaft was slightly twisted, and pre-
sented a strongly marked deltoid ridge. The bicipital groove was in the form of a
shallow groove. The lower end presented a well-marked trochlea for the ulna, a slight
coronoid fossa, but no radial or olecranon fossae. The capitellum was circular and
convex. The internal condyle was more pronounced that the external, the external
supra-condyloid ridge more so than the internal, which did not possess a supra-
condyloid foramen or even a process.

Bones of Forewarn.

In neither were the distal epiphyses fused with the shaft. The ulna measured
152 mm. and was expanded above from before backwards so as to form a long but
narrow olecranon process. A coronoid process was not present. The articular surface
for humerus was concavo-convex, while the facet for radius above was flat and con-
tinuous with that for the humerus. The shaft was flat, with a rounded anterior border
and a sharp posterior border. The lower end was conical and articulated with radius
slightly, and with cuneiform and pisiform. The radius was 135 mm. long. The head
possessed a deep cup-shaped cavity for articulation with the capitellum of humerus,
while the margin of the head was non-articular except on its inner side, where there
was a well-marked facet for articulation with the ulna. On the neck a faint tubercle
was noticed, while the shaft was paddle -shaped, with a definite roughness on its outer
aspect for pronator radii teres. The lower end was grooved on its posterior aspect, and
articulated with the ulna and scapholunar.

Humero-Radial Index.

Length of radius x 100
Length of humerus

Ross Seal. Sea Leopard. Weddell Seal.

117-4 107-1 98-1

(ROY. soc. EDIN. TRANS., VOL. XLVII., 195.)

26 MR ROBERT B. THOMSON : SCOTTISH NATIONAL ANTARCTIC EXPEDITION :

Carpal Bones.

These were seven in number, the scaphoid and semilunar being fused together to
form a single bone. The carpal bones were devoid of ridges or processes. The articu-
lations were difficult to make out, but I have referred these as follows :—

The scapholunar articulated with radius, trapezium, trapezoid, os magnum, and
unciform.

The cuneiform articulated with the ulna, pisiform, and unciform.

The pisiform was a mere nodule, and articulated with the cuneiform and ulna.

The trapezium articulated with scapholunar, trapezoid, and 1st and 2nd metacarpals.

The trapezoid articulated with scapholunar, trapezium, os magnum, and 2nd meta-
carpals.

The os magnum was small, and articulated with the 2nd, 3rd, and 4th metacarpals,
and with the trapezoid, scapholunar, and unciform.

The unciform articulated with the scapholunar, cuneiform, and os magnum, and with
the 4th and 5th metacarpals.

Metacarpal Bones.

The 1st metacarpal was the longest, and possessed both a proximal and a distal
epiphysis not united with the shaft. The 2nd was longer than the 3rd, 4th, or 5th,
which were of about equal length, and the epiphysis for the head of each was separate
from the shaft.

The Hind-Limb.

The innominate bone, measuring in length 215 mm., consisted of three parts — ilium,
ischium, and pubis. The ilium was short and stunted. The ischium and pubis were
long, somewhat narrow, and enclosed between them a very large obturator foramen-
larger indeed than the ilium itself.

The acetabulum was circular in form, entirely articular except an elongated, hollow
area at the bottom with a definite margin. The brim of the acetabulum was deficient
in its lower aspect, thus forming a cotyloid notch. The ilium measured from centre
of acetabulum to the highest part of the iliac crest 74 mm. The length of its crest was
102 mm,. Its dorsal and ventral aspects were flattened and did not possess any definite
muscular ridges. At the posterior part of the ventral surface was a U -shaped articular
surface for that part of sacrum corresponding to its first segment, while there was a
roughened area for the attachment of the posterior sacro-iliac ligament.

The os pubis projected backwards and slightly downwards from the acetabulum,
and its junction with the ilium was marked by a prominent pectineal eminence. From
this tubercle to the symphysis, which was small, the pubis measured 130 mm.

The ischium passed backwards and slightly upwards from the acetabulum, and
possessed a pointed spine.

The femur measured 114 mm. long. The head was entirely articular except in its
postero-superior quadrant, where there was a non-articular area continuous with the

(ROY. soc. EDIN. TRANS., VOL. XLVII., 196.)

OSTEOLOGY OF ANTARCTIC SEALS. 27

non-articular neck for attachment of the ligamentum teres. Dr KEITH, in Human
Embryology and Morphology, p. 385, states: "The ligamentum teres is isolated
from the capsule by the development of the head of the femur, which expands as a wing
on each side of the ligamentum teres, and by fusion of the wings Isolates it from the
capsule." In the Ross seal femur the wings have not yet met; hence, according to
KEITH'S view, the ligamentum teres would still be in connection with the capsule. (See

%- 6.)

The trochanter major was a broad, rough prominence projecting from the outer side of
the junction of the neck of the bone with the shaft. There was no trace of a trochanter
minor, as is to be found in Otaria Jubata ; neither was there a third trochanter.

The shaft, flat and expanded, possessed an anterior and a posterior surface, both
being very slightly rounded and devoid of muscular ridges. The outer border was
short and concave, the inner convex and very rough for muscular attachment.

At the posterior end of the bone, the trochlear surface for the patella was flattened
and ascended as high on its outer as on the inner side. It was quite distinct from the
condylar surface, but closely approximated to the external one, being separated by a
very narrow groove.

The internal condylar surface was flat and circular, the outer one oval and slightly
convex.

The intercondyloid fossa was shallow.

The patella was 34 mm. long and 24'5 mm. broad. It was oval in form, and its
articular surface was slightly concave, and did not exhibit any subdivision into special
articular areas. Its anterior end was much thicker than the posterior.

The tibia and fibula were fused together at their upper ends. The tibia measured
247*5 mm. Its upper end was elongated transversely and possessed two articular areas
for the condyles of the femur, separated by a rough intermediate area. The shaft was
slightly curved, flat and smooth on its ventral aspect, deeply concave on its dorsal
aspect for origin of the tibialis posticus muscle. The nutrient foramen was a large one.
The inner border was broad and rough, the outer one thin and rounded. The lower end
was not united with the shaft, and from it projected a slight malleolus, which was non-
articular. The ventral and dorsal aspects of the lower end were deeply grooved, while
the articular area for astragalus was reniform and concave.

The fibula was 233 mm. in length. The upper end was fused with the tibia, whilst
the lower end was grooved on the outer side by the peroneal tendons, and had on its
inner aspect an oval facet for outer surface of astragalus. The shaft was straight,
and its surface broad for muscular origins. The lower epiphysis was not united to

the shaft

Tibia- Femoral Index.

Length of tibia x 100
Length of femur

Ross Seal. Weddell Seal. Leopard Seal.

217-1 203-9 213-5

(BOY. SOC. EDIN. TRANS., VOL. XLVIL, 197.)

28 MR ROBERT B. THOMSON : SCOTTISH NATIONAL ANTARCTIC EXPEDITION :

Tar sal Bones.

The astragalus was the largest of the tarsal bones. It consisted of a body and two
large processes. The posterior process projected backwards beyond the hinder end of
os calcis, and presented a well-marked groove on its under aspect. Its anterior process
or head passed as far forward as did the os calcis, and articulated with the scaphoid,
cuboid, and fore-part of os calcis. The superior surface of the body presented a quadri-
lateral convex surface for articulation with the lower end of tibia, continuous externally
with a triangular facet for fibula. Its internal surface did not articulate with the
internal malleolus. The inferior surface articulated with os calcis and presented a deep
groove for an interosseous ligament.

The os calcis possessed an attenuated posterior or peroneal process, and a feeble
internal or sustentacular one. It articulated with the astragalus and fibula above, with
the cuboid in front, and was grooved on its outer aspect by the peroneal tendons.

The cuboid presented on its inferior aspect a well-marked plantar tubercle and a
deep peroneal groove. It articulated with the os calcis, astragalus, scaphoid, external
cuneiform, and 4th and 5th metatarsal bones.

The scaphoid articulated with the astragalus, cuboid, and the three cuneiforms. Its
posterior surface was not entirely concave, being convex in its outer part.

The internal cuneiform articulated with the scaphoid, middle cuneiform, and 1st and
2nd metatarsals.

The middle cuneiform was the smallest of the three, and was completely shut out of
the plantar aspect of the foot. It articulated with the cuneiform on either side of it,
with scaphoid behind and 2nd metatarsal in front.

The external cuneiform articulated with the scaphoid, cuboid, 2nd and 3rd
metatarsals.

Metatarsal Bones.

The 1st was the longest, the 5th next in order, then 4th, 2nd, and 3rd, which was
the shortest.

None of the epiphyses were fused with the shafts. The base of the 2nd showed the
characteristic hook-shaped base.

The articulations of their bases were as follows : —

1st with internal cuneiform and 2nd metatarsal,
2nd with the three cuneiforms 1st and 3rd metatarsal,
3rd with external cuneiform 2nd and 4th „
4th with cuboid 3rd and 5th ,,

5th with cuboid 4th „

while its base projected outwards as an elongated process.

The terminal phalanges of both fore and hind limbs were retained in connection with
the skins.

(BOY. SOC. EDIN. TRANS., VOL. XL VII., 198.)

OSTEOLOGY OF ANTARCTIC SEALS. 29

Summary.

The skulls correspond in their distinguishing characters from other Antarctic seals
to those tabulated by Sir WILLIAM TURNER in the Challenger Reports ( Voyage of
" Challenger" Reports on Seals, p. 66).

With regard to the rest of the skeleton, one specimen alone does not provide
altogether sufficient evidence ; still, I note the following points which may be of interest
on further skeletons being brought home and examined : —

1. Much greater length-breadth index of vertebrae.

2. Bifid character of 5th and 6th cervical spines, also present Ln type-specimen in

British Museum.

3. Sacral index.

4. Scapular index.

5. Humero-radial index.

6. Non -articular area for ligamentum teres on head of femur continuous with non-

articular neck.

7. Ti bio-femoral index.

NUMBER OF VERTEBRAE IN PHOCID^E.

In the Resultats du Voyage du S.Y. " Belgica" : Zoologie, p. 7, Mr BARRETT
HAMILTON states that in most seals the numbers of the thoracic and lumbar vertebrae
appear to be usually 15 and 3 (5?), more rarely 14 and 6. In the Antarctic seals
without exception I have found the vertebral formula in the adult to be C7 D]5 L5
S3 Cd10 or u.

With a view to obtaining as large a number of specimens as possible, in order to
form an average, I have made a careful examination of the seal skeletons in the
University Museum here, the Museum of Science and Art of Edinburgh, British
Museum of London, and give a table of the vertebral formula in 47 specimens : —

Weddell seal : 27 with formula C7 D15 L5 S3 Cd10 or n.
Crab-eating seal : 2 with formula C7 D15 L5 S3 Cd]0.
Sea-leopard : 2 with formula C7 D15 L5 S3 Cd10.
1 ,, „ C7 Du L6 S3 Cdn.
Ross' seal : 1 with formula C7 D15 L5 S3 Cd10.
Sea-lion : 2 with formula C7 D15 L5 S3 Cd10.
Fur seal : 1 at C7 DJ5 L5 S3 Cdn.
Crested seal : 1 at C7 D15 L5 S3 Cd10.
Common seal : 2 at C7 D15 L5 S3 Cdn.
Elephant seal : 1 at C7 D15 L5 S3 Cd10.
New Zealand fur seal : 1 at C7 D15 L5 S3 Cd10.
West Indian seal : 1 at C7 D15 L5 S3 Cd12.

(ROY. soc. EDIN. TRANS., VOL. XLVTL, 199.)

30 MR ROBERT B. THOMSON : SCOTTISH NATIONAL ANTARCTIC EXPEDITION :

Grey seal : 1 at C7 D15 L5 S3 Cd12.

Bearded seal : 1 at C7 D15 L5 S3 Cdn.

Northern fur seal : 1 at C7 D15 L5 S3 Cd10.

Altogether 46 specimens with formula of C7 D15 L5 S3 Cd10 to 12.

One specimen with formula of C7 D14 L6 S3 Cdn.

The number of caudal vertebrae seems to vary from 10 to 12, while in some
cases one could not be positive, as frequently the terminal vertebrae are preserved
in the skins.

SUPRA-CONDYLOID FORAMEN.

With a view to the formation of a percentage of such foramina found in the humeri
of the Phocidae, I have examined altogether 100 humeri, made up as follows :—

60 Weddell seals.
4 crab-eating seals.
6 leopard seals.
4 sea-lions.
2 Ross seals.
4 elephant seals.
4 South American fur seals.
2 New Zealand fur seals.
2 West Indian seals.
2 common seals.
2 North American fur seals.

Total, 92

In these 92 no foramen was present, but in 4 Weddell humeri there were small
supra-condyloid processes.

In the following 8 humeri, supra-condyloid foramina were present :—

2 grey seals.
2 bearded seals.
2 crested seals.
2 common seals.

Total, 8
thus giving a percentage of 8.

SUMMARY OP DENTITION.

As the dentition of the Phocidae is of some importance, especially with regard to
variation, I have formed a table of the dentition of the seals under my observation. I

(HOY. soc. EDIN. TRANS., VOL. XLVIL, 200.)

OSTEOLOGY OF ANTARCTIC SEALS. 31

have also examined the crania of the seals in the University Museum of Science and
Arts, Edinburgh, and include such in the following list :—

2. 1. 5

Weddell seal : 2 1 specimens with formula .

1 specimen

2. 1. 5
2. 1. 6

2. 1.5

The fourth post-canine in the upper jaw was obviously split into two, thus account-
ing for the variation.

2. 1. 5

Sea-leopard : 5 specimens with dental formula
Crab-eating seal : 7 specimens with formula
Sea-lion (Otaria Jubata) : 2 specimens with formula

1 specimen „ „

2. 1. 5

2. 1. 5

2. 1. 5

3. 1. 5
3. 1.5

3. 1. 6
3. 1. 5

LITERATURE.

The Zoology of fhe Voyage of the " Erebus " and " Terror," pp. 7-8, plates vii. and viii.

J. E. GRAY, Ommatophoca Rossi, 1844.

Challenger Reports, vol. xxvi., Sir WM. TURNER, 1888.

RtmUats du Voyage du S.7. " Belgica," G. E. H. BARRETT HAMILTON, 1901.

National Antarctic Expedition : Natural History, vol. ii., E. A. WILSON.

DESCRIPTION OF PLATE.

FIG. 1. — Ross skull, No. 43. Superior aspect.
FIG. 2. — Ross skull, No. 43. Lateral aspect.
FIG. 3. — Ross skull, No. 43. Anterior aspect.

non-articular area on head.
FIG. 4. — Ross skull, No. 43. Inferior aspect.

FIG. 5. — Ross seal, No. 43. 15th dorsal vertebra.
Showing costal process on right side, and
facet for rib on left side.

FIG. 6. — Ross seal, No. 43. Femur showing

(ROY. soc. KDIN. TRANS., VOL. XLVTL, 201.)

Scot. Nat. Ant. Erp. VOL. IV.

ROBERT B. THOMSON—" Scottish National Antarctic Expedition : Osteology of Antarctic Seals."

^4'V

FIG. 1.

FIG. 2.

FIG. 3.

FIG. 4.

FIG. 5.

PART IV.
PENGUINS.

SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

IV.-ANATOMY AND EMBRYOLOGY OF ANTARCTIC

PENGUINS.

BY D. WATERSTON, M.A., M.D., F.R.C.S.E., AND
A. CAMPBELL GEDDES, M.D.,

Anatomical Department of the University of Edinburgh.
(WITH THREE PLATES.)

Report upon the Anatomy and Embryology of the Penguins collected by
the Scottish National Antarctic Expedition, comprising : (1) Some Features
in the Anatomy of the Penguin ; (2) The Embryology of the Penguin : A
Study in Embryonic Regression and Progression. By D. Waterston, M.A.,
M.D., F.R.C.S.E., and A. CampbeU Geddes, M.D. (From the Laboratory of
the Anatomical Department, The University, Edinburgh.) (With Three Plates.)

(MS. received March 11, 1909. Read February 3, 1908. Issued separately October 21, 1909.)

Among the specimens collected by the Scottish National Antarctic Expedition and
brought back to this country was a series of the embryos of penguins of various species,
collected at different stages of development by Mr R. N. RUDMOSE BROWN and Dr J. H.
HARVEY PIRIE. These embryos were, for the greater part, of the species Gentoo
(Pygoscelis papua, Forst.), the remainder of the species Adelia (PygosceUs adelicB,
Hombr. and Jacq. ). Two Emperor penguins, which had been preserved for examination
by the injection of the blood-vessels, were also brought home. Through the kind-
ness of Dr W. S. BRUCE these specimens were entrusted to one of us (D. W.) for
examination and report, but as the work involved in this proved to be very
extensive, Dr A. CAMPBELL GEDDES was asked to undertake a share in it, and this
he agreed to do.

This report contains the results of the combined investigations.

PART I.

ON THE ANATOMY OF THE ADULT PENGUIN.

INTRODUCTORY.

Upon the return of the Challenger Expedition to this country, the late Professor
MORRISON WATSON had handed over to him the valuable collection of adult penguins
which had been made. Upon these he based his classical memoir, " A Report on the
Anatomy of the Spheniscidse collected during the Voyage of H.M.S. Challenger"
(Challenger Reports, vol. vii. p. l). To that report little, if anything, can be added.
There are, however, two points which a detailed dissection of the adult penguin has
brought to light, which seem to us to be worthy of being placed on record. These are,
first, the peculiarities of the cervical portion of the vertebral column ; second, some points
in connection with the arrangement of the fascial layers in relation to the pectoral
muscles. In all the other points illustrated by our dissections we can merely corrobo-
rate the description given by Professor WATSON.
(REPRINTED FROM THE TRANSACTIONS OF THE BOTAL SOCIETY OF EDINBURGH, VOL. xi/vn., PART IL, PP. 2-23-244.)

38 DR D. WATERSTON AND DR A. CAMPBELL GEDDES ON EMBRYO PENGUINS

SECTION I.— SOME PECULIARITIES OF THE CERVICAL PORTION OF THE

VERTEBRAL COLUMN.

The cervical vertebrae are thirteen in number : the portion of the vertebral column
which they make up is remarkable for the development of its antero-posterior curves.
In the lower half there is an extraordinary antero-posterior curve the convexity of
which is directed forward ; in the upper half there is a second antero-posterior curve the
convexity of which is directed backward.

These curves are present in all birds ; in none, however, do they approach in intensity
the curvatures found in the penguin. The lower curvature is so pronounced that the
bodies of the vertebrae come in contact with and fill up the angle formed by the limbs
of the clavicle, and, in the living bird, actually cause a projection on the front of the
neck. As a result the trachea and oesophagus, instead of passing into the thorax in
front of the vertebral column, are pushed away to the right side and actually lie on a
plane posterior to the vertebrae of the convexity. The knowledge of this extraordinary
condition is not new ; it is fully described by Professor WATSON, but the point seems to
have been entirely overlooked, for in all the specimens in museums that we have had
the opportunity of seeing the cervical vertebrae are incorrectly mounted.

Professor MORRISON WATSON was of opinion that these curvatures were associated
with the maintenance by these birds when on land of the erect attitude, and that they
served to bring the centre of gravity of the head and neck over the base of support
formed by the feet. To us it appears more probable that the curvature is useful to the
bird when in the water, giving it a greatly increased displacement forward and reducing to
a minimum the fatigue of carrying the head and heavy beak in the long ocean voyages
which it undertakes. These may apparently be very prolonged, for the birds have been
seen no less than 600 miles from land.

SECTION II. — SOME PECULIARITIES IN THE ARRANGEMENT OF THE FASCTAL LAYERS
IN RELATION TO THE PECTORAL MUSCLES.

The most striking feature with regard to the pectoral muscles is their enormous
size and power. There are four muscles belonging to this group : first, the pectoralis
major ; second, the dermo-humeralis (muscle des parures) ; third, the pectoralis medius ;
fourth, the pectoralis minor. The origins and insertions of these muscles are given in
detail and with great accuracy by Professor MORRISON WATSON. Their action is not
without interest. The pectoralis major is divided into two distinct parts : an anterior
which arises from the clavicle and from the outer surface of a strong aponeurosis
which separates it from the medius and is inserted through a special tendon into the
whole length of the anterior or radial margin of the bones of the wing, and by an
aponeurotic extension from the tendon which covers both surfaces of the wing and
conceals the blood-vessels and nerves, and a posterior which terminates in a V-shaped

(ROY. soc. EDIN. TRANS., VOL. XLVII., 224.)

OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 39

tendon inserted into the humerus. These two portions have different actions ; the
anterior carries the wing forward to a position at right angles with the trunk, while the
posterior draws the limb back in the effective part of the swimming stroke, and by its
insertion into the anterior margin of the humerus it " brings about that rotation of the
wing round its long axis which, combined with the backward stroke, gives rise to the screw-
like motion of the organ observable when the bird is progressing through the water." *

The nerve supply is provided for by a special branch from the large cord of the
brachial plexus.

The dermo-humeralis arises from a strong fascia which covers the external oblique
muscle ; at the posterior margin of the sternum it is reinforced by a number of fibres
arising from the subcutaneous tissue covering the knee joint. It passes forward parallel
to the outer border of the pectoralis major to be inserted along with the posterior fibres
of that muscle into the anterior margin of the humerus. Its action apparently is to
co-operate with the external part of the pectoralis major in depressing the wing. It is
supplied by a twig from the nerve to the pectoralis major.

The pectoralis medius passes over a pulley formed by the bones of the shoulder and
acts as the great elevator of the wing ; by some authors it has been called levator
humeri. It also is supplied by a twig from the nerve to the pectoralis major.

The pectoralis minor is inserted into the outer margin of the tricipital fossa ; its
chief action appears to be to rotate the humerus in a direction opposite to that of the
rotation produced by the action of pectoralis major. In addition, however, it must help
in producing the effective, propulsive stroke of the wing. Its nerve of supply is a twig
from the nerve to pectoralis major.

The arrangement of the fascia in regard to these muscles is striking and peculiar.
Tracing the deep fascia of the neck downwards, it passes deep to the clavicle, to which
it is lightly attached ; and, passing to the thoracic region, it lies superficial to the
pectoralis medius and minor and becomes continuous with the strong fascia which
covers the external oblique. Superficial to this fascial layer lie the pectoralis major and
dermo-humeralis, and from it they both obtain a portion of their origin. The super-
ficial aspects of pectoralis major and dermo-humeralis are in close contact with skin — so
close, indeed, that they are marked by regular rows of pits which accommodate the
rounded ends of the feather quills.

From this arrangement it appears to us possible that the dermo-humeralis, pectoralis
major, and clavicle are in reality cutaneous structures. That the dermo-humeralis is in
series with the dorsal cutaneous muscle and panniculus carnosus is certain ; that it is
closely associated with the pectoralis major in position, in action, and nerve supply is
also certain ; but whether the pectoralis major can justly be associated with the same
group is, on the present evidence, not certain, although it appears to us probable. Un-
fortunately the evidence from nerve supply is so weakened by the association of
pectoralis medius and minor with the same nerve as to be valueless.

* Quoted from Professor MORRISON WATSON'S memoir.
(ROY. soc. KDIN. TRANS., VOL. XLVIL, 225.)

40 DR D. WATERSTON AND DR A. CAMPBELL GEDDES ON EMHRYO PENGUINS

The suggestion that the pectoralis major is in reality a cutaneous muscle seems to
us to be not without interest in view of the occasional occurrence in man of the
abnormal muscle, sternalis. Sir WILLIAM TURNER has suggested that it is to be
regarded as a vestige of the panniculus carnosus. Professor D. J. CUNNINGHAM has
pointed out that there is considerable evidence to show that it is formed by a devia-
tion or dislocation of a portion of the pectoralis major ; Mr F. G. PARSONS has shown
that in rodents the panniculus carnosus possesses two strata, and that there is
evidence to show that in man the deeper stratum of the pauniculus forms the fascial
sheath over the external oblique and that possibly a portion of the sterno-mastoid is
derived from the same stratum. He has also shown that there are good reasons for
regarding the pectoralis major as derived from the panniculus. To these observations
we now add the facts of the anatomy of the penguin, which appear to us to suggest that
the apparently antagonistic views of Sir WILLIAM TURNER and Professor CUNNINGHAM
are in reality not opposed, but complementary.

PART II.

ON THE EMBRYOLOGY OF THE PENGUIN.

INTRODUCTORY.

As has been shown, the anatomy of the adult penguin has been previously very
completely described, and little remains to be added to complete our knowledge of it.

Of the embryology, on the other hand, the existing knowledge is incomplete, owing
presumably to the great difficulty in bringing back the necessary material in a con-
dition which allows of a detailed examination being made. A small number of
embryos was collected by the Challenger Expedition, but the condition in which they
were received by Professor MORRISON WATSON rendered them useless for description.

The material which was put into our hands consisted of a number of specimens each
one of which had been removed from its egg upon a different day of incubation. It
would, therefore, appear that we had a specimen for each 24 hours of development. In
one sense this was so, in another not. The method of collection which was of necessity
adopted was as follows : — The nest was watched, and the eggs were marked with the
date of their first appearance and were subsequently collected upon the desired day.
The difficulties of so doing and the fallacies necessarily attendant upon it are obvious.
First, it was quite impossible for the observer to know in which of the 24 hours preced-
ing the marking the egg had been laid ; secondly, it is known to be no uncommon thing
for the males and females to fight for possession of their egg and for the privilege of
incubating it. In the course of these struggles the eggs are apt to be dropped and to
lie for some time directly upon the ice. The result of this must be to retard for a time
the processes of development, and therefore, although an egg may have been laid for 3,

(ROY. soc. EDIN. TRANS., VOL. XLVII., 226.)

OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 41

6, 9, or 12 days, it does not follow that the development has been proceeding for the
same number.

Be this as it may, we found it necessary, especially in dealing with the early
specimens, to revise the sequence of specimens so as to avoid the absurdity of embryos
of an obviously earlier stage of development being classified as older than specimens
of a later stage. The number of days of incubation is, according to the series of
specimens, thirty. This is surprisingly short, and as there are some obvious gaps
in the series, we are not prepared to express an opinion as to whether it is correct
or not.

The embryos were preserved in 2^ per cent, formalin, and we desire to place on
record our sense of real obligation to the collectors for the care and trouble which they
took, and for their great skill in packing the specimens, which alone is responsible for
the excellent state of preservation in which many of these exquisitely delicate specimens
were when put into our hands.

As a result of the long time which elapsed between the date of collection and
examination and of the prolonged immersion of the specimens in formalin, it has not
been possible to carry out, satisfactorily, any very fine histological examination nor to
determine with absolute accuracy the exact date of the first appearance of the primary
ossific centres, the formalin, apparently, having exercised a slight decalcifying action.
Our investigations have, therefore, of necessity been somewhat restricted, and many
important problems in connection with the development of the penguin remain unsolved.

In dealing with the embryology of the penguin the following points seemed to us
important : —

1. The enormous length of time through which the penguin has persisted without
undergoing important modification. In the Quart. Journ. Geol. Soc., vol. xv. p. 670,
Professor HUXLEY described a penguin of large size whose remains were obtained from
the pliocene strata of New Zealand. For the reception of this bird he established the
genus Palseeudyptes.

It is exceedingly doubtful whether the establishment of a new genus was necessary,
and Professor MORRISON WATSON in his report on the anatomy of the Spheniscidse
(Challenger Report, vol. vii. p. 46), expresses the opinion that this pliocene bird
might very well be regarded as belonging to a species of the existing genus Eudyptes.

From the outset, therefore, we were prepared to meet with some peculiarities of
development of a more primitive avian type than that obtaining in modern birds.

2. The enormously high specialisation undergone by the forelimb of the penguin in
the course of which it has been transformed from an organ of serial to an organ of sub-
aqueous flight.

3. The modification undergone by the cervical spine to secure sufficient water dis-
placement anteriorly to carry the head and heavy beak without fatigue during the long
ocean voyages which the penguins undertake.

4. The skeletal arrangements necessitated by the quasi-erect attitude.

(ROY. soc. EDIN. TRANS., VOL. XLTIL, . 227.)

42 DR D. WATERSTON AND DR A. CAMPBELL GEDDES ON EMBRYO PENGUINS

METHODS OF EXAMINATION.

The earlier specimens were prepared for examination by being stained in borax
carmine or alum carmine and mounted entire. Duplicate specimens for examination
by section were not available. The later specimens were cleared so far as possible,
but owing to the long immersion in formalin the clearing did not yield very satisfactory
results. A figure illustrating the development of the limb bones is included among the
illustrations (fig. 26). The decalcifying action of the formalin has, unfortunately,
made any definite statements as to date of appearance and order of appearance of the
ossific centres impossible, and we have had to content ourselves with the negative
observation that no unusual or abnormal processes or sequences were to be observed,
although such were looked for as carefully as the material available would permit.

In order to carry out a comparison regarding the dates of acquisition of the external
adult features and the progressive development of the limbs and trunk during the
later stages of development, we found it necessary to provide a control by carrying out
observations on the same lines upon the embryo of another genus of birds.

Data for this purpose were not available in the literature, and we therefore prepared
and examined, by methods identical with those used for the penguins, a series of duck
embryos at every 24 hours of incubation. The period of incubation in the duck (28-
30 days) approximates sufficiently closely to that of the penguin (30-32 days ?) to
enable comparisons to be drawn with accuracy between embryos of corresponding age
taken at each 24 hours.

The data which we obtained appear to be new, and we have, therefore, included an
account of the external form and the measurements of the embryo-duck with those of
the penguins.

SECTION I. — EARLY STAGES OP DEVELOPMENT.

EMBRYOS SHOWING EARLY STAGES, COMPRISING SPECIMENS UP TO THE TWELFTH DAY.

Specimen I. — In the first of these, which was the youngest specimen examined,
there was an oval area pellucida measuring 3 mm. in length on the surface of the
blastoderm. This specimen illustrates the earliest stage after the close of segmentation
before the formation of the primitive streak, and it closely resembles in size and shape
the corresponding area in the ovum of the chick and the duck. No area opaca could be
made out, nor was there any sign of the embryonic shield. The shape of the area
pellucida is comparable to that seen in the blastoderm of the chick at about 18 hours,
but in the absence of primitive streak and embryonic shield it resembles the chick
blastoderm at the commencement of segmentation.

Specimen II. (fig. 1). — The second specimen showed a more advanced stage. The
germinal area was somewhat pear-shaped. At its wider end was a narrow, semi-

(KOY. soc. EDIN. TRANS., VOL. XLVII., 228.)

OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 43

circular, deeply marked line running round the .periphery of the clear area.' This line
appeared to be the commencement of the amniotic fold. Behind this line was a
narrow area in which there were a few dark spots, which marked apparently an early
stage of the formation of a vascular area. This area, in turn, merged into a crescentic
clear area which was limited on the opposite side by a short crescentic dark line
parallel to the first line. This line occupied the central part of the wider end of the
clear area, and lay across it, and it appeared to be the rudiment of a commencing head
fold. From the concavity of this fold a darker streak passed for a short distance
backwards towards the narrow part of the clear area, and this structure appeared to us
to be an indefinite stage in the formation of the primitive streak.

Specimen III. (fig. 2). — This specimen resembled that of the chick of 22 hours.
The head fold was well formed, and behind it lay the medullary folds. These diverged
from one another at their posterior extremity, and behind them lay the remains of the
primitive streak stretching to the posterior end of the clear germinal area. The
primitive streak had the form of two narrow dark lines, enclosing between them a
narrow clear streak.

Specimen IV. (fig. 3). — Labelled as 3 days. — The embryo was now 3 mm. in length
and presented well-formed medullary folds, a notochord, seven pairs of somites, and also
the remains of the primitive streak which was separated by a narrow clear area from the
posterior end of the notochord and extended for a distance of I mm. behind it. The
appearances were very similar to those of the chick at 25 hours. The medullary folds
were as yet ununited, but showed at the cephalic end evidence of distinct thickening,
while at the posterior ends they diverged from one another in a V-shaped manner. The
notochord could be traced as far forward as the cephalic enlargements (but this part of
the embryo was rather damaged), while posteriorly it terminated in a club-shaped
enlargement.

Specimen V. (fig. 4). — The succeeding specimen, labelled as being 5 days,
showed a slightly more advanced stage of the same condition, closely similar to the
chick at 26 hours.

The embryo measured 3 '5 mm. in length, and nine pairs of somites were present.
The posterior end of the notochord was enlarged and shaped like an Indian club, and
extended beyond the termination of the medullary folds, which diverged outwards on
either side of it. Behind this extremity was a clear area, and behind it again lay two
short parallel dark lines, the representatives of the primitive streak.

The cephalic ends of the medullary folds were considerably enlarged, to form the
cerebral vesicles. These vesicles were not distinctly marked off from one another, but
the anterior part was bent forwards, to form the earliest cephalic flexure. In front of
this again was a narrow clear crescentic area — the pro-amniotic region.

Specimen VI. (fig. 5). — Labelled 6 days. — -The next specimen was somewhat
larger, measuring 6 mm. in length. Twelve pairs of somites were present, and the
neural folds were larger, and had apparently united with one another at the fourth

(ROY. HOC. EDIH. TRANS., VOL. XLVH., 229.)

44 DR D. WATERSTON AND DR A. CAMPBELL GEDDES ON EMBRYO PENGUINS

somite. The folds approached one another closely at the posterior end, and in that
region they enclosed a clear area, shaped like an arrow head, in which lay the expanded
posterior end of the notochord. The end of the notochord projected backwards beyond
the medullary folds in the form of a bulbous extremity. Behind it there was no
distinct evidence of a primitive streak. The whole of the posterior portion was
enclosed in a clear oval area. The somites were well formed, and their internal
structure showed them to consist of a clear central portion and a periphery or
cortex arranged in columns radiating outwards.

At the outer margin of the somites was a clear area in which lay a narrow darker
line, extending along the side from the third anterior somite backwards to beyond the
hinder somites.

In position this structure corresponds to the primitive cardinal vein.

The neural folds showed evidence of segmentation, being constricted opposite the
centre of the somites, and they were united about the middle. The specimen described
was one said to be of the sixth day, and the subsequent specimens, which showed stages
of gradual advance, were in harmony with this statement.

If these dates be accepted, it is obvious that the early changes in the penguin
embryo are very much slower than the corresponding changes in the chick, for the
same degree of development is shown in the chick of 40-44 hours.

Specimens VIL, VIII., IX., and X. (fig. 6).— Labelled 7-1.1 days.— These
specimens showed that the next stages of development are very similar to
corresponding stages in the development of the chick, and do not require detailed
description, except in so far as they show differences from it. The head and trunk
show gradual increase in length and in thickness.

At the hinder end of the notochord a dark area shaped like a spear-head persists
for some time, but eventually becomes merged in the hinder end of the trunk.

Figs. 7, 8, and 9, which illustrate the development of the duck at 5, 6, and 7 days,
are introduced for comparison with fig. 6, which is a photograph of a penguin embryo
labelled as 8 days.

SECTION II. — KATE OF GROWTH OF THE TRUNK AND LIMBS FROM THE

TWELFTH DAY ONWARDS.

Our observations were directed first to an examination and measurement of the
developing trunk and limbs.

The observations begin with the twelfth day of incubation, which is the date of
appearance of the limb buds upon the surface of the body.

The details of the measurements are recorded in the appended table (Table
No. I.).

(ROY. soc. EDIN. TRANS., VOL. XLVII., 230.)

OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 45

NOTES UPON AND COMPARISON OF THE MEASUREMENTS.
The measurements were analysed along several different lines.

1. General Growth of the Trunk.

A. Penguin. — The vertex to coccyx length of the specimen of 12 days (fig. 10), taken
as it lay in the egg, i.e. in a flexed position, was 15 '5 mm., while that of the duck of
corresponding age was practically twice as great, namely, 30 '6 mm. In the penguin
the rate of growth from this date onwards was fairly uniform and progressive, and the
largest embryo, that of 30 days, measured 61 '2 mm. in the same position. The
specimen labelled as being 24 days was larger than those of 25 and 26 days, and
among the other specimens similar anomalies were present. Taking the maximum
measurements of specimens which seemed to be typical in their rate of growth, we
found that the length attained at 12 days was doubled at 19 days and doubled again at
27 days, the progression being thus almost in geometric ratio.

The same ratio is found to exist when measurements of the breadth of the specimen
are taken. The maximum breadth attained at 12 days was rather more than doubled
at 19 days, and rather less than doubled again at 27 days.

B. Duck. — The maximum length of the embryos of this species was uniformly
greater than that of the penguin of same age. Even the largest fully developed
penguin embryo was smaller than the largest duck embryo. At 12 days (fig. 20) the
duck embryo was almost exactly twice as long as that of the penguin of corresponding
age. The rate of growth from this date onwards, as ascertained by the same criteria
as in the case of the penguin, was, however, much slower than in that creature.

At 19 days the length had increased only by rather less than one-half, i.e., was
4'36 cms. as against 3-06 cms., and not until the 23rd day of incubation was the length
doubled, and from this time until the end of incubation the increase again was rather
more than half, from 43'6 mm. to 76'8 mm. The breadth figures showed a rate of
growth very similar to that found in the penguin. The breadth at 12 days was 10 mm.,
and this figure was practically doubled at the 19th day and almost doubled again at
the end of incubation.

2. Rate of Growth of Different Segments of the Trunk.

Penguin. — Analysing the rate of growth of the different segments which make up
the vertex to tail measurement, the head segment is found to increase slowly, at first,
but grows rapidly in the last few days, while the neck increases rapidly through all
stages.

The growth of the trunk and tail follows the general rate of the whole body, the
length attained at 12 days being doubled at 19 days, and doubled again at 27-28 days.

The figures for the head are modified by the fact that measurements were taken

(ROY. 8OC. EDIN. TRANS., VOL. XLVII., 231.)

46 DR D. WATERSTON AND 1>R A. CAMPBELL GEDDES ON EMBRYO PENGUINS

TABLE

Adelia.

Days 12

13

14

15

16

17

18

18

P.

D.

P.

D.

P.

D.

P.

D.

P.

D.

P.

D.

P.

D.

P.

D.

Vertex to coccyx length, flexed

1-55

S-OG

1-94

S-36

1-03

3-32

1-94

4-07

2-42

S'95

2-58

4

3 35

2-86

4-45

Maximum breadth ....

•64

1-28

*l-5

1-41

*-89

1-6

"•91

1-5

1-09

1-8

1-53

1-8

1-46

1-35

2-42

Length of head .....

•76

1-47

•34

1-24

1-01

1-3

•94

1-34

1-05

1-SS

1-49

1-6

1-25

...

1-21

4 1

,, neck .....

•48

1-4

•56

1-65

•68

1-6

•63

1-24

•67

1-32

•82

1-35

1-06

1-28

1-74

,, trunk, including tail

1-32

2-22

1-44

2-65

1-41

3-09

1-35

3-65

2-01

3-74

2'13

S-82

2-71

2-42

4-45

Cloaca to furcal angle ....

•78

1-5

1-08

1-82

1-14

1-92

•98

2- 12

1-36

2-6

1'44

2-45

1-92

1-G5

3-29

Tip of beak to occiput

•78

2-05

1-21

2-38

1-29

2-6

•98

2-63

1-68

2-86

1-93

3'9>:

2-28

2-06

3-23

,, to ear .

•52

1-59

•7

1-82

•82

2-04

•7

2-1

111

2-34

1-43

2-3

1-76

1 39

2-7

,, ,, to post angle of eye .

•63

1-44

•86

1-64

•84

1-7

•74

1-87

•97

1-97

1-33

2

1-53

11

2-11

Head-j „ ,, to mouth

•25

•7

•43

•92

•29

1-05

•29

1-1

•61

1-2

1-78

1-17

•99

•78

1-23

,, ,, to nostril

•6

No

N.

•7

No

N.

•64

No

N.

•74

•8

No

N.

No

N.

•87

Breadth between eyes

•64

1'2S

1-5

1-34

•89

1-35

•91

1-35

1-09

1-35

1-33

1-45

1-46

1-35

1-6 i

„ ears .

•35

•84

•55

•74

•46

•9

•54

1-13

•08

1-07

•87

1-1

•97

1-04

1-5

[ Length .
Neck \

•48

1-4

•56

1-65

•08

1-5

•63

114

•67

1-32

•82

1-35

1-06

1-28

1-74

[Width .

•31

•33

•43

•45

•38

'44

•31

•57

•37

•73

•54

•78

•56

...

•53

•9

Upper arm, anterior border

•11

•44

•19

•44

•23

•46

•18

•5

•37

•56

•35

•55

•54

•35

•64

„ „ posterior ,, .

•39

•11

•61

•15

•57

•12

•64

•21

•7

•26

•7

•33

•28

•75

Forearm, anterior border .

•29
incl.

•37

•41

•38

•45

•47

Hi

•6

f79

•6

t'91

•73

tl'21

tl'04

•75

,, posterior „

Hand

•57

•58

t'6

f52

•62

f48

•82

•84

•78

1-1

•8

1-25

tl'19

•37

Fore-

incl.
Hand

limb "

Elbow, width ....

•13

•31

•16

•44

•17

•53

•16

•53

•24

•57

•26

•55

•23

•31

•63

„ thickness

•08

•19

•09

•25

•07

*22

•08

•26

•09

•12

•09

•a

0-9

•09

'28

Length of hand

•72

•72

•86

•9

1'21

1'3

1 23

Width

•19

•si

•18

•19

•2

'32

•35

. Length of pollex

•18

•18

...

•34

•24

•35

•37

•23

Leg, anterior border ....

J-54

1-04

t-72

•98

J-73

1-05

t-61

i-os

•57

1-23

•55

1-35

•77

•86

1-55

» po

iterior „

•63

...

•42

.it

•52

•74

•21

•82

•18

•86

•28

•22

1-12

Plantar, heel to end of third
phalanx

...

1-16

1-45

1-62

1-66

•54

1-97 ,

•69

2-08

•91

•78

2-57

Plantar, length of phalanges

...

•56

•65

•73

i i •

•75

•28

•96

•28

1-15

•45

•37

1-23

Foot •

Plantar, web between outer and
middle

...

•41

•42

...

•47

•18

•5

•17

•7

•19

•17

•75

Plantar, web between inner and
middle

•45

•4

•12

•64

•14

•65

•14

•12

•63

Dorsal, ankle to tip of nail of
middle toe

1-04

...

1-3

1-35

It

•64

i .Off)
J. O%

•54

1-94

•82

•78

2-25

. Dorsal, length of nail . .

•1

•14

•13

•1

•09

•28

* Over eyes.

t Elbow to tip.
(ROT. soc. EDIN. TRANS., VOL. XLVIL, 232.)

+ Knee to tip.

OP THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

I.

1

1

1

0

2

1

•2

2

2

3

2

4

2

6

2

•

a

7

21

3!

3

)?

p.

D.

P.

D.

P.

D.

P.

D.

P.

D.

P.

D.

P.

D.

P.

D.

p.

D.

P.

D.

P.

D.

2-99

4 -36

3'37

6-25

3-62

5-93

3'49

5-05

4-09

6-39

4-64

7-04

4-13

6-06

3-99

7 68

6-07

...

571

...

6-12

...

1-45

3-35

1-52

2-73

175

2-95

1-68

S-8

1-89

3-gg

2-24

3-35

2-21

S-8

2-18

4-46

2-35

3-05

...

3-03

...

11

1-3

1-18

1-2

1-35

1-43

1-24

1-56

1-48

1-97

1-47

g-43

1-54

g-35

1-55

1-95

1-91

...

2-55

2-89

...

1-09

2-05

•99

1-94

1-23

1-95

1-4

S-15

1-66

S-34

1-63

g-68

1-58

2-93

1-39

2-88

1-92

...

1-85

1-92

27
1-84

4
26

3'2
2-24

5-85
4'16

3-47
3'66

5-01
3'46

3-38
2-52

5-4
3-13

4-06
3-21

6-Og
4-66

4-16
3-41

6
4-41

3-55
2-81

5-83
4'61

3-91
3-38

6-32
4' 4

4-91
4-15

5-51

...

5-35

...

2-34

s-i

2-43

*f •*•**

3-45

2-77

*rf

5-53

2-69

3-64

2-81

39

3-2

4'OS

3-12

4-01

3-19

T f

4-4

3-43

3-98

4-21

...

1-7

g-34

1-89

g-78

2-18

2-73

2-08

2-75

2-26

2-9

2-57

2-95

2-48

3

2-48

3-27

272

3-07

3-28

...

1-44

:

1-59

s-s

1-79

2-43

1-71

ft5

1-85

ft

1-93

g-47

1-97

g-66

1-95

g-87

2-07

2-39

2-56

•92
No

N.

1-25
•83

1-1
No
N.

1-43
•91

1-23
•43

1-41
•97

1 28
•52

!•#
•9

1 29

•57

1-58
•94

1-5
•58

1-6
1

1-44

•58

1-66
1-02

1-38
•9

1-SS
1-18

1-66
•59

...

1-91

7

...

2-01
•88

...

1-45

1-45

1-52

1-54

1-66

1-57

172

1-53

17

1-6

1-88

1-77

1-8

1-58

1-82

1-64

1-87

...

2-02

...

1-96

...

•98

1-2S

1-15

t-4

1-15

1-45

1-13

1-43

1-27

1-5S

1-54

1-66

1-42

1-54

1-32

1-73

1-69

1-83

...

1-89

...

1-09

g-05

•99

1-94

1-23

V95

1-4

S-15

1-66

g-34

1-63

TO

1-58

g-93

1-39

2-88

1-93

...

1-85

••

1-92

...

•7

•7

•77

1

•83

1-1

•63

1

•8

1-08

1-09

1-05

•88

1

•91

1-8

1-23

...

1-26

...

1-31

...

•46

•69

•51

1

•64

•53

•65

•5

77

•78

I'll

•56

•58

•64

•72

•74

79

...

77

...

•93

•32

•82

•45

•96

•46

•64

•41

•83

•47

1

•57

•8

•58

1-S7

•72

1-43

•58

...

•81

...

•96

tl'23

•76

tl'34

•97

ti'52

•93

tl'45

•64

tl'65

1-14

tl'87

•9

t'73

1-08

tl'73

1-18

t'2'17

...

•68

...

•68

1-34

1-07

1-44

1-23

1-59

1-07

tl'51

•96

tl'72

i-vr

t2'07

1-1

tl'84

1-S3

tl'93

138

t2'13

...

1-01

•91

...

•32

•f3

•35

•6

•4

•S$

•44

•51

•48

•56

•56

•78

•58

•55

•57

•74

64

76

...

•83

•1

•a

•11

•:•:

•12

• u

•12

•a

•16

•M

•14

*4

•27

•29

•15

•39

•22

...

•23

•36

...

...

1-5
'26

...

1-5

1-55
•A

...

1-3
•37

...

1-46
•a

1-75
•51

...

1-76
•Si

1-83
•62

...

...

1-38

1-5

...

•38

jj*t
•JtS

•T
•5

"SI

*r*

'56

"73

vjt
•fit

'6A

•89

1-18

•91

•T"

g-3

1-01

t-15

1-07

.-•-;

1-21

g-53

1-6

S-8

1-42

v^
8-96

1-57

v+
3-18

1-61

2-18

2-12

...

•3

1

•4

1-54

•37

1-4

•32

1-68

•48

1-55

•46

g-8g

•46

1-88

•78

2-14

•68

...

•73

•66

...

•89

.-•o

1-12

3-56

1-31

3-24

1-15

3-4S

1-25

f»

1-73

3-93

1-5

4-23

1-56

4'61

1-85

2-32

2-43

...

•43

1-14

•54

1-9

•63

1-47

•59

1-55

•61

1-93

•8

113

•66

2-03

•2

8-03

•93

1-01

...

1-07

••-

•29

•85

•34

1-17

•41

VIS

•35

1-S5

•34

1-46

•55

1-54

•47

1-BS

•54

1-7

•65

•81

...

•83

•21

•8

•22

1-16

•29

1-18

•23

1-SS

•23

1-46

•31

1-54

•28

1-52

•42

1-78

•41

...

•53

...

•52

...

•82

:•:•

•93

M

1-15

3-07

1-08

S-S3

1-34

»•&

1-53

3-64

1-47

3-62

1-32

4-1

1-79

...

2-02

2'3

...

•11

•;••

•14

•33

•15

•34

•19

•35

•25

•38

•29

•44

•26

•43

•22

•47

•39

...

•46

...

•41

* Over eyes.

t Elbow to tip.
(BOY. soc. EDIS. TRANS., VOL. XLVIL, 233.)

: Knee to tip.

48 DR D. WATERSTON AND DR A. CAMPBELL GEDDES ON EMBRYO PENGUINS

from the tip of the beak, and consequently the increase in total length is found to be
greatest in the last few days, when the beak elongates rapidly, while the figures
obtained for the growth of the head in width show a more gradual and regular increase,
comparable to the rate of growth of the trunk.

3. Growth of the Limbs.

A. Penguin. — Fore Limbs. — The first appearance of the fore limb had the form
of a bud-like projection from the lateral aspect of the trunk (fig. 10). This process at
first was somewhat spatulate, and then rather bulbous at its free end (figs. 11 and 13).
This shape was in turn soon lost, and the limb rudiment assumed the characteristic
sickle-shaped form found in the adult bird (figs. 15 and 17, 19 and 23). The two
segments of the arm were soon clearly to be recognised and were separately measured.

The measurements taken were those of the anterior border of the upper segment,
together with the anterior border of the forearm from the elbow to the tip. At 1 2 days
the total length was 4 mm., made up of I'l mm. of upper segment and 2 '9 mm. of the
lower.

At 19 days the length was 13'9 mm. and at 27 days 297 mm. The rate of growth,
therefore, had been such that the length of the limb was more than trebled in the first
period, and was more than doubled in the second. Both segments shared the growth,
and did so in fairly equal proportions throughout.

Lower Limb. — The portion of the lower limb which was free from the coverings of
the trunk was measured in one or in two pieces, as the size permitted. At 12 days
it measured 5'4 mm., at 19 days 18'1 mm., and at 27 days 36'2 mm. In the first
period the length was more than trebled, and in the second it was again doubled,
showing a rate of growth almost exactly similar to that found in the upper limb.

B. Duck. — In the duck corresponding measurements showed that at 12 days (fig. 20)
the fore limb measured 15 '3 mm. and was nearly four times as long as in the penguin.
At 19 days the length was almost doubled (28'5 mm.), and at the end of incubation it
had only added one-third to its length (37 '5 mm.). The lower limb, measured in a
similar way at 12 days, was 22 mm. in length, again four times as long as in the
penguin, and in the specimen of 20 days it was more than doubled, and at the end of
incubation it was almost doubled again.

If the proportion which the length of the limbs bears to the total length of the
body be examined, it is found that in the penguin the fore limb at 12 days is almost
one-fourth of the body length, that it increases rapidly for a few days until it measures
nearly one-half of the length of the body, and maintains approximately this ratio till
near the end of the incubation period, when it progresses rapidly and assumes a ratio
of rather more than two-thirds of the body length.

In the duck the ratio at 22 days was one-half, and this proportion became rather
larger, but eventually again became nearly one-half.

(BOY. soc. EDIN. TRANS., VOL. XLVII., 234.)

OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

49

TABLE II. — VERTEX TO COCCYX LENGTH, THE LENGTH OF LIMBS AND THE RATIO BETWEEN THEM,

THE FORMER BEING TAKEN AS 1.

Days.

Vertex to Coccyx
Length.

Length of Upper
Limb.

Index.

Length of Lower
Limb.

Index.

Penguin.

Duck.

Penguin.

Duck.

Penguin.

Duck.

Penguin. Duck.

Penguin.

Duck.

12

1-55

S-06

•40

1-5S

•25

•50

•54

2-20

•34

•71

13

1-94

3-36

•60

1-54

•30

•45

•72

2-43

•37

•72

14

1-93

8-38

•68

1-79

•35

•53

•73

2-57

•37

•77

15

1-94

4-07

•59

2

•30

•49

•61

2-68

•31

•65

16

2-42

3-95

1-16

2-37

•47

•60

Ml

3-20

•45

•80

17

2-58

4

1-26

2-58

•48

•64

1-26

3-43

•48

•85

18

2-85

4-45

1-75

2-62

•61

•58

1-71

4-12

•60

•92

19

2-99

4-86

1-39

2-95

•46

•67

1-81

3-63

•60

•83

20

3-37

6-25

1-85

•54

1-84

5-36

•54

•85

21

3-62

5-93

2-16

3-01

•59

•50

2-42

6-SS

•66

90

•2-2

3-49

5-05

2-10

2-24

•60

•44

2-38

5-72

•68

l-!3

23

4-09

6-39

2-42

3-38

•59

•52

2-62

6-73

•63

1-05

24

4-64

7-04

2-98

3-21

•64

•45

3-52 7-73

•76

1-09

25

4-13

6-06

...

3-47

...

•57

7-19

...

1-18

26

3-99

7-68

2-45

3-75

:6l

•48

3-21

7-79

•80

1-01

27

6-07

2-97

•48

...

3-62

...

•59

...

23

5-71

3-53

...

•61

4-71

• • •

•82

...

30

6-12

...

4-24

...

•69

• • •

4-79

•78

...

SYNOPSIS OF RESULTS FROM COMPARISON OF THE MEASUREMENTS OF THE PENGUIN

AND DUCK.

1. Length of Trunk.

At early stages the length of the trunk of the penguin is much less than that of
the duck of corresponding age — the proportion being that the penguin at the 12th
day is almost exactly one-half of the length of the duck at the same age.

As development proceeds the penguin grows more rapidly, until at the end of
incubation the penguin is only fractionally shorter than the duck.

2. Length of Fore Limb.

At corresponding dates after the limbs first appear the fore limb of the penguin is
only about one-fourth of the length of that of the duck.

Its growth subsequently is much more rapid, and at the three weeks it is almost
the same size, and at the end of incubation it exceeds in length the duck's fore limb,
in some specimens by as much as one-quarter.

3. The Ratio of Length of Upper Limb to Trunk.

At the 12th day of incubation the fore limb of the penguin measures approxi-
mately in length one-fourth (index, "2 5) of the length of the trunk, while in the duck

(ROY. SOC. EDIN. TRANS., VOL. XLVH., 235.)

50 DR D. WATERSTON AND DR A. CAMPBELL GEDDES ON EMBRYO PENGUINS

the limb is equal to one-half of the length of the trunk (index, '50). The ratio of
growth of the fore limb in the penguin is continuous and progressive in relation to the
ratio of growth of the trunk.

Towards the end of the third week it measures one-half of the length of the trunk
(index, 21st day, -50), and at the end of incubation it measures two-thirds (index, '69) of
the length of the trunk.

In the duck, on the other hand, the length at first increases relatively to the trunk,
attaining in one specimen, at the end of the third week, a maximum ratio of over two-
thirds (index, 19th day, '67) ; but thereafter it slowly loses ground, and at the close of
incubation the fore limb is less than one-half the length of the trunk.

4. Hind Limb.

A. Absolute Length. — In the penguin at the 12th day of incubation the hind
limb is also one-fourth of the length of the hind limb of the duck, the ratio being very
similar to that between the upper limbs of the two animals. In the penguin's limb,
however, growth is more rapid, with the result that at the end of the third week the
hind limb has reached a length equal to nearly one-half of the length of the hind
limb of the duck of corresponding age, and at the end of incubation it is rather more
than half as long as the corresponding part in the duck.

5. Ratio of Length of Lower Limb to that of the Trunk.

At the 12th day in the penguin the length of the lower limb is approximately
one-third of the length of the trunk (index, '34), but the rate of growth is more rapid, so
that at the end of incubation it is nearly as long as the trunk (index, 28th day, -82).

In the duck the limb at 12 days is rather more than two-thirds of the length of
the trunk (index, 71), at the 18th day it is nearly as long as the trunk (index, '92),
and thereafter it continues to increase at a rate relatively rather greater than the trunk,
so that at the end of the third week it becomes actually longer (index, 22nd day, T13),
and retains this superiority to the end of incubation.

In this respect, i.e. in the rate of growth from the 18th day, it is the exact reverse
of the fore limb — it continues to increase in length relatively to the trunk, whereas the
fore limb actually shows a relative decrease.

ANALYSIS OF RESULTS.

Among the four limbs examined there are, therefore, two types of development.

First, there is a progressive type, and to this the development of the fore and hind
limbs of the penguin and of the hind limb of the duck belongs.

Second, there is a partially regressive type, and to this the development of the fore
limb of the duck belongs.

(BOY. soc. EDIN. TRANS., VOL. XLVII., 236.)

OP THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 51

Now, if it be true that ontogeny is an abbreviated recapitulation of phylogeny we
are forced to conclude that the fore limb of the penguin is a progressive structure, the
fore limb of the duck, partially at least, a regressive. The reason for this, doubtless,
is that the mesoblastic portion of the duck's wing has, with the advancing specialisation
and adaptation of the epiblastic structures, become less valuable. To all flying birds
feathers, not bone and muscle, are of prime importance in securing wing spread for
aerial flight : for the penguin, on the other hand, a rigid bony paddle has been evolved,
adapted to resist the pressure of the water to which it is subjected when the bird's
great pectoral muscles are in strenuous action.

And we thus are led to the interesting conclusions : —

1. That the mesoblastic structures in the penguin's "wing" are progressive, the
epiblastic regressive.

2. That the mesoblastic structures in the duck's wing are regressive, the epiblastic
progressive : for in the development of the duck we find that the developmental energy
is suddenly on the 1 7th day switched off, as it were, from the mesoblastic structures on
to the epiblastic ; for from that day onward the down rapidly develops and the meso-
blastic framework loses ground.

With regard to these developmental facts the question arises : —
Is the duck's or the penguin's wing the more direct descendant of the common
ancestor ; or have they both diverged from the common stock approximately equally,
but in opposite directions ?

Embryology alone cannot answer this question, but the evidence is clear in this,
that the fore limb of the penguin in its development goes through a progressive and
continuous series of stages along one unbroken line. The mesoblastic portion of the
fore limb elongates, but its characters do not alter. It elongates, however, with a
relatively greater rapidity towards the end of development, whereas the duck's fore limb,
after being relatively longer than the penguin's ever is, regresses rapidly. So that the
answer to our question, so far as the embryological evidence is concerned, must be that
the wings of both these birds are different from the ancestral wing, and that the
differentiation has been in opposite directions and that the common ancestor was a
flying bird of a somewhat primitive type depending in large measure for the spread of
its wing upon bone and muscle.

SECTION III. — EXTERNAL CONFIGURATION OF THE EMBRYOS FROM THE

TWELFTH DAY ONWARDS.

1. Gentoo Penguin Embryo, 12 Days. (Fig. 10.)

The embryo was not fully formed, the brain was covered by the thinnest of mem-
branes, the beak was short and soft, the eyes were very prominent, with dense white
opaque pupils. The fore limb well developed ; the elbow flexure completely marked.

(BOY. soc. EDIN. TRANS., VOL. XLVII., 237.)

52 DR D. WATERSTON AND DR A. CAMPBELL GEDDES ON EMBRYO PENGUINS

It ended in a somewhat spatula-shaped extremity. The hind limb formed a spatula-
shaped protrusion.

Duck Embryo, 12 Days. (Figs. 16 and 20.)

The embryo was fully developed, the integument was strongly marked, and there
were down papillae on two short lines at either side of the tail. The eye prominences
were very marked ; the head and beak were soft and pliable except at the point of the
beak, where there was a small white nodule of keratin. The fore limb showed distinct
division into upper arm, forearm and hand. The pollex was prominent.

2. Gentoo Penguin Embryo, 1 3 Days. (Fig. 11.)

The embryo was not fully formed. The head was soft and the brain showed
through it. The beak was short, the eyes were prominent, the fore limbs were short,
but showed plainly the elbow joint, and had a flattened, spade-like tip : to the
naked eye there was no sign of digitation. The hind limbs were short, the ankle was
unformed, the end was spatular and showed signs of three digits, the tail was relatively
long and was much curved ventrally.

Duck Embryo, 13 Days. (Fig. 18.)

The embryo was fully formed. Down was appearing in lines on the back and sides
of the embryo, and laterally on the neck ; otherwise as for 12 days.

3. Gentoo Penguin Embryo, 14 Days.

The embryo was not fully formed, the head was soft, the brain showing through it.
The beak was short, the eyes prominent ; the fore limbs short, but well formed ; the
hind limbs were short, spatular protrusions. The heart showed distinctly, shining
through the thin anterior wall of the thorax. The tail was relatively long and much
curved ventrally.

Duck Embryo, 14 Days. (Fig. 21.)

Down papillae were seen all over the embryo, especially along the dorsal tracts and
on the tail. The eyes were widely open.

4. Gentoo Penguin Embryo, 15 Days. (Fig. 13.)

The embryo was small, the beak was beginning to grow forward. The fore limbs
were small and soft, but fully formed. The hind limb showed no division into the
toes, but ended in flat, spatular extremities. There was no sign of the fourth toe
independent of the flattened extremity. The tail was relatively long and curved
ventrally.

Duck Embryo, 15 Days. (Fig. 22.)

Down was appearing upon the head along a well-defined tract which looped round
the eye from behind.

(ROY. soc, KDIN, TRANS,, VOL. XLVII., 238.)

OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 53

5. Gentoo Penguin Embryo, 16 Days. (Fig. 14.)

The embryo was small, the nails were not developed, so far as could be seen on
naked-eye examination. There was a well-marked ridge down the centre of the
abdomen and thorax. The toes did not appear beyond the web, but looked more as if
they were embedded in it. The fourth toe appeared as if the web were continued
round it.

Duck Embryo, 10 Days.
As for 15 days. Down more marked.

6. Gentoo Penguin Embryo, 17 Days. (Fig. 15.)

The embryo was in good condition and the fully formed integument was complete.
The skin was marked by papillae regularly arranged. On either side of the tail there
were several down fibres varying from 17 mm. in length to a length just visible to the
naked eye.

Duck Embryo, 17 Days.

The down was well developed.

7. Gentoo Penguin Embryo, 18 Days. (Fig. 19.)
This specimen was somewhat macerated.

Duck Embryo, 18 Days.
Embryo completely down-covered.

8. Adelia Penguin Embryo, 19 Days.

The embryo was fully formed, the integument was complete. There was a well-
marked median groove over the abdomen and thorax, and from it on either side
extended twelve transverse grooves. The body was completely naked except for eight
fibres on the tail (four on either side of the median line).

9. Gentoo Penguin Embryo, 19 Days.

The embryo was fully formed, and the integumentary covering was complete.
There were no transverse grooves on the abdomen.

10. Gentoo Penguin Embryo, 20 Days.

The embryo was fully flexed and completely devoid of down. The integument over
the abdomen was not quite complete. A bare area in length '28 cm. and in breadth
at base '07 cm. extended forward in front of the umbilicus. There were no transverse
grooves on the abdomen.

(ROT. 8OC. EDIN. TRANS., VOL XIA1L, 239.)

54 OR D. WATERSTON AND DR A. CAMPBELL GEDDES ON EMBRYO PENGUINS

11. Gentoo Penguin Embryo, 21 Days.

The embryo was completely flexed; the limbs were completely formed. The
integument over the abdomen was incomplete; a triangular area 1*23 cms. in length,
•07 cm. in breadth, extended forward in front of the umbilicus and was continued over
the front of the thorax as a well-marked groove from which on either side extended
twelve transverse grooves.

12. Gentoo Penguin Embryo, 22 Days. (Fig. 17.)

The limbs of the embryo were complete ; the integument over the abdominal
regions was incomplete. An area 1 '04 cms. in length and '09 cm. in breadth at the
umbilicus, gradually tapering to a point, was undeveloped. The head and body were
entirely naked. The eye prominences were very marked ; the skull and beak were soft
and pliable. There were five transverse grooves on the abdomen. Flexion was
incomplete.

13. Gentoo Penguin Embryo, 23 Days.

The embryo was not quite fully developed, the integument being incomplete. On
the anterior abdominal region — anterior to the umbilicus — a very acute isosceles
triangle persisted uncovered, the sides of which measured 1'19 cms., the base '12cm.
The eye prominences were very marked, the skull and beak were soft and pliable. A
few short pieces of dark down had developed on the head ; the rest of the body was
absolutely naked. There were no transverse grooves on the abdomen ; flexion was
incomplete.

14. Gentoo Penguin Embryo, 24 Days. (Fig. 23.)

The embryo was fully developed. The prominence of the eye-balls was well marked
and the eye-slit was open, 1 8 cms. long ; its width at its broadest part when stretched
was '08 cm. The head was covered with dark down, and on the back and body there
was a sparse covering of light-coloured down considerably more than on the 2 5-day s'
embryo. The middle line of the front of the abdomen showed a deep groove from which
extended across the abdomen, on either side, six well-marked transverse grooves. The
feet and tail had now assumed the adult relations, and formed the peculiar flat base
which is characteristic of the adult bird and permits of its adopting the quasi-erect
attitude habitual to it.

15. Gentoo Penguin Embryo, 25 Days. (Fig. 24.)

The embryo was fully formed. The prominence of the eye-balls was well marked.
The eye-slit was small, the beak was soft, and the head was sparsely covered with short
black down on the back. The body was fully flexed ; the spine formed a semicircle from
tail to head. There was a well-marked median groove on the front of the abdomen
and thorax, from which extended transversely on either side eight well-marked grooves.

(ROY. soc. EDIN. TRANS., VOL. XLTII., 240.)

OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 55

16. Gentoo Penguin Embryo, 26 Days.

The embryo was fully formed ; the eyes looked large and prominent ; the beak was soft.
The head was covered with short black down ; the rest of the body was practically naked.
Along the middle line of the abdomen, from the umbilicus to the root of the neck, was a
well-marked groove.

17. Gentoo Penguin Embryo, 27 Days. (Fig. 25.)

The limbs of the embryo appeared fully formed. The feet were flexed over the front
of the abdomen, plantar surface outwards. The upper limbs lay close to the side and
over the abdomen under cover of the feet. The neck was flexed so that the beak lay along
the right side of the abdomen immediately internal to the right superior extremity.
Down was plentifully present on the head and back, shading from almost black over the
head to a light slate grey over the sacral region. The down on the abdomen and anterior
aspect of the thorax was sparse and light in colour. The middle line of the body in
front was marked by a very evident ridge in the lower part, flanked on either side by a
groove. These grooves coalesced in the upper part of the abdomen and were continued
over the thorax as a median depression. This could be traced as high as the root of the
neck, where it was lost.

18. Gentoo Penguin Embryo, 28 Days.

The embryo appeared fully formed and was in a fully flexed position, so that the
under surface of the mandible was pressed firmly against the thorax and abdomen ; the
beak lay in a groove formed by the right fore limb and left hind limb. The head was
covered with a dark grey, almost black, down ; the back with slate coloured, the abdomen
with white. The back of the head had been flattened by pressure.

Duck Embryos, 19-28 Days.

The advance was purely one of body bulk. The amount and directions of growth
are shown by the measurements of the specimens. (See Table I.)

19. Gentoo Penguin Embryo, 30 Days. (The specimen in fig. 25 closely
resembled this older specimen, but was rather smaller.)

The embryo appeared fully formed and was in a fully flexed position, so that the under
surface of the left side of the head was pressed against the thorax, the beak lying to the
right side of the abdomen between the right fore limb and the foot of the left hind limb
anteriorly and the right hind limb posteriorly. The head and back were covered with
slate-coloured down, the belly with white down. In the flexed position the greatest
length was 571 cms., the greatest breadth 3'05 cms. In the extended position the
length from the crown of the head to the tip of the tail was 8 '41 cms. When measured
separately the head was 2'55 cms. in length, the neck T85 cms. ; the body and tail

(ROY. soc. EDIN. TRAXS., VOL XLVII., 241.)

56 DR D. WATERSTON AND UR A. CAMPBELL GEDDES ON EMBRYO PKNGUINS

together 5'51 cms. — giving a total length of 9*91 cms. The 1'5 cms. of difference (9'91
cms. — 8 '41 cms.) is accounted for by the marked spinal curves which immediately asserted
themselves on extension, a lumbo-thoracic curve ventrally concave, a cervical curve
dorsally concave.

SUMMARY AND CONCLUSION.

Part 1. — Anatomy of the Adult Penguin.

1. The cervical portion of the vertebral column is possessed of a curve of extra-
ordinary convexity which causes the vertebral bodies to form a visible bulging on the
front of the neck. This convexity we regard as useful to the birds by increasing water
displacement anteriorly and permitting of the head and beak being carried without
undue fatigue on the long ocean voyages which the birds undertake. The furcal angle
being occupied by the vertebral bodies, the trachea and oesophagus enter the thorax on
a plane posterior to the cervical spines. This curvature of the column and arrangement
of the viscera is permanent and characteristic of the adult bird.

2. The arrangement of the fascial layers in relation to the pectoral muscles suggests
that the pectoralis major and clavicle are in reality cutaneous structures. It was found
impossible from the embryological material at our disposal to prove or to disprove this
suggestion.

Part II. — Embryology of the Penguin.

1. The number of days of incubation is, according to the series of specimens, thirty.
This is surprisingly short, and as there are some obvious gaps in the series, we are not
prepared to express an opinion as to whether it is correct or not.

2. The early stages of development are exactly comparable to the corresponding
stages of the chick or duck, though, if the dates of our specimens be correct, they take
place more slowly.

3. The later stages of development are exactly comparable to the corresponding
stages of the duck except with regard to the down, the fore limb, and the foot.

4. The foot of the penguin is, throughout its development, exceedingly clumsy and
primitive, as is the foot of the adult ; the foot of the duck, from its earliest appearance,
assumes a lighter and neater appearance, and is adapted as a paddle, which the foot of
the penguin is not.

5. The fore limb of the penguin is, in its mesoblastic structures, definitely progressive,
whereas the mesoblastic portion of the duck's wing is relatively regressive. Both,
however, have varied from the ancestral type, but their variation is in opposite direc-
tions : the wing of the penguin has required rigidity to subserve its function as a paddle,
and it has acquired it, throughout its length. The wing of the duck, on the other hand,
has required lightness, and it has acquired it — the reduced spread of bone being
compensated for by an increased development of feathers.

(ROY. soc. EDIN. TRANS., VOL. XLVII., 242.)

OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 57

6. The evidence from embryology is in favour of the early divergence of the
Aptenodytidse from the main stem of the flying birds, and of the common ancestor of
both having been a flying bird, though of a clumsy type.

7. We were unable to verify the facts recorded by other observers in the ossification
of the limbs, as we were handicapped by the decalcifying action of formalin even in
weak solutions, so that our observations in this direction could not be considered
altogether satisfactory.

ILLUSTRATIONS.

PLATB I.

Fig. 1. Gentoo penguin, embryonic area, early stage.

Fig. 2. Gentoo penguin, within first 24 hours, showing head fold, medullary folds, and primitive streak.
This stage resembles that of the chick at 22 hours.

Fig. 3. Gentoo penguin, said to be 3 days. This stage of development resembles the chick at 24 hours.

Fig. 4. Gentoo penguin, said to be at 5 days. The stage of development resembles the chick at 26 hours.
The photograph shows specially the cephalic flexure, the hinder ends of the medullary folds enclosing the
bulbous extremity of the notochord, and the remains of the primitive streak.

Fig. 5. Gentoo penguin, said to be 6 days. Comparable to the chick at 40 hours. The photograph
shows the bulbous expanded posterior end of the notochord, and evidence of a segmentation of the neural
folds.

Fig. 6. Gentoo penguin, about 7 days.

PLATE II.

Fig. 7. Duck embryo of 5 days. Compare fig. 4 of penguin of same age.

Fig. 8. Duck embryo of 6 days. Compare fig. 5 of penguin of same age.

Fig. 9. Duck embryo of 7 days. Compare fig. 6 of penguin of same age.

Fig. 10. Gentoo penguin embryo of 12 days. Both upper and lower limbs have appeared on the surface
of the trunk

Fig. 11. Gentoo penguin embryo of 13 days.

Fig. 12. Duck embryo of 8 days. Compare character of limbs with those of penguin of 13 days
(fig. 11).

Figs. 13, 14, and 15. Successive stages of Gentoo penguin embryos at 15, 16 and 17 days, to illustrate
stages in the development of the sickle-shaped fore limb of the adult.

Fig. 16. Duck embryo of 12 days, in titu, to show how the embryonic attitude is maintained, and is not
dependent upon external pressure.

Fig. 17. Gentoo penguin embryo of 22 days, to show advancing change in the position of the feet.
Compare figs. 15 and 23, which show other stages in the development of the peculiar flat base of the adult.

III.

Fig. 18. Duck embryo of 13 days.

Figs. 19 and 20. Gentoo penguin embryo of 18 and duck embryo of 12 days, photographed to the same
scale, to show altering characters of the limbs, when the trunks are of the same size.
Fig. 21. Duck embryo of 14 days.

Fig. 22. Duck embryo of 15 days. Compare with fig. 15 of gentoo penguin embryo of the same age.

(BOY. soc. EDIN. TKAKS., VOL. ILVII., 243.)

58 EMBRYO PENGUINS OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

Fig. 23. Gentoo penguin embryo of 24 days. Notice the character of the fore limb, and the attitude of
the hind limb and tail.

Fig. 24. Gentoo penguin embryo of 25 days.

Fig. 25. Gentoo penguin embryo of 27 days, to show the stage of development reached before leaving
the shell.

Fig. 26. Photograph of fore limbs of gentoo penguin embryo of 22 days, to show ossification.

The clear portions of the shafts are composed of bone, and the dark extremities are of cartilage. Both
extremities of the radius and ulna are cartilaginous. Beyond them is a darkish area, in which are the
cartilaginous rudiments of the carpus.

Beyond that region are the metacarpal bones and phalanges.

(BOY. soc. EDIN. TRANS., VOL. XLVII., 244.)

Scot. Nat. Ant. Ex p. PLATE I. Vol. IV.

Drs WATERSTON AND GEDDES. — EMBRYO PENGUINS: SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

FIG. l.

FIG. 3.

FIG. 2.

FIG. 4.

FIG. 5.

FIG. 6.

Scot. Nat. Ant. Exp. PLATE II. Vol. IV.

Drs WATERSTON AND GEDDES. — EMBRYO PENGUINS : SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

FIG. 7,

FIG. 8.

FIG. 9.

FIG. 10.

FIG. 11.

FIG. 12.

FIG. 13.

FIG. 14.

FIG. 15.

FIG. 16.

FIG. 17.

MTirtane & Erekine. Edin.

Scot. Nat. Ant. Exp. PLATE III. Vol. IV.

Drs WATERSTON AND GEDDES. — EMBRYO PENGUIN'S : SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

FIG. 18.

FIG. 19.

FIG. 20.

FIG. 21.

FIG. 24.

FIG. 22.

FIG. 23.

FIG. 25.

FIG. 2G.

M-Faifene It Erskine. Edin.

PART V.
SEALS.

SCOTTISH NATIONAL ANTARCTIC EXPEDITION.
V.- ANATOMY OF THE WEDDELL SEAL

(LEPTONYCHOTES WEDDELL1)
(2) GENITO-URINARY ORGANS.

BY DAVID HEPBURN, M.D., C.M.,

Professor of Anatomy, University College, Cardiff (University of Wales).

Scottish National Antarctic Expedition : Observations on the Anatomy of the
Weddell Seal (Leptonychotes Weddelli). By David Hepburn, M.D., C.M.,
F.R.S.E., Professor of Anatomy, University College, Cardiff (University of AVales).
Part II.*

(MS. received December 4, 1911. Read January 8, 1912. Issued separately January 19, 1912.)

GENITOURINARY ORGAN-.

In my former contribution I gave a general summary of the animal under considera-
tion, and discussed in detail the peritoneal arrangements of its abdominal cavity and
the naked-eye anatomy of its alimentary organs. In the present paper I shall give an
account of the genito-urinary system.

The kidneys were situated on each side of the dorsal mesial mesentery. Each was
covered on its ventral aspect by the peritoneum forming the dorsal wall of the greater
peritoneal sac. The right kidney was quite free from contact with the liver and the
duodenum, while the left kidney was equally free from contact with the spleen. Both
kidneys were therefore situated well back towards the pelvic end of the abdominal
cavity. Each kidney measured 5 inches in the longitudinal diameter and 2 inches
in the transverse diameter. The hinder or caudal end of each reached a point two
inches from the pelvic inlet, which, as formerly described, was narrow and well defined
.by the course of the hypogastric (umbilical) arteries.

The surface of the kidney indicated lobulation, but the lobules were not separated from
each other. The hilum was placed ventro-mesially , and at its point of emergence from the
surface of the kidney the ureter was nearer to the caudal than to the cephalic end of the
organ. On opening up the hilum, the ureter was seen to result from the union of two main
tributaries, each of which, in its turn, was formed by the junction of several smaller root-
lets, which corresponded more or less closely in number to the number of the kidney
lobules. There was no distinct pelvis to the ureter, which was gradually formed by the
junction of smaller ducts in the manner indicated. Nevertheless, the widest point of
the ureter was found at the junction of its two main tributaries. The ureter and its
chief tributaries lay on the ventral (anterior) aspect of the renal vessels, and not on their
dorsal (posterior) aspect, as is the case in man.

The size of the ureter suggested a vessel about half the diameter of an average
human radial artery. The ureter followed a course along the dorsal wall of the abdomen
towards the pelvic inlet ; and half an inch beyond the termination of the abdominal
aorta, or, in other words, at the point where the common iliac artery divided into its
external and internal branches, the ureter crossed to the mesial side of the internal iliac
and hypogastric arteries, and continued its course along the margin of the pelvic inlet.
In this position the ureter and the hypogastric artery were both in their turn crossed

* Part I. was published in the Trans. Roy. Soc. Edin., vol. xlvii., pt i. (No. 3), 1909.
(REPRINTED FROM THE TRANSACTIONS OF THE ROYAL SOCIETY OF EDINBURGH, VOL. XLVTH., PP. 191-194.)

64 PROF. DAVID HEPBURN

by the vas deferens, which now assumed the mesial position to both of the others. Up to
this point the ureter had not entered the pelvic cavity, and about three-quarters of an
inch onwards, that is, in the direction of the tail, the ureter, still lying close to the pelvic
brim, entered the lateral aspect of the urinary bladder, travelling between the folds of
a lateral vesical mesentery or peritoneal ligament. Thus, as a consequence of the great
obliquity of the pelvic inlet, the ureter was able to reach the bladder by skirting the
pelvic brim, and at no point did it require to enter or sink into the interior of the pelvis.

The urinary bladder was placed mesially, and was attached to the ventral wall of
the abdomen by a ventral mesial mesentery composed of peritoneum, which, as formerly
described, closely invested the bladder except on its pubic aspect. The apex of the
bladder extended to the umbilicus, where it still presented an open lumen. There was
no obliterated part or urachus, and throughout its entire length it presented a uniform
calibre, suggestive of an empty portion of small intestine. Developmentally, it may be
said to represent an enlarged and patent allantois ; but as this animal was only two days
old at the time of its death, probably a sufficient period had not elapsed for the closure
of the umbilical end of the organ.

The hypogastric arteries were carried along each lateral aspect of the bladder,
suspended in peritoneal folds half an inch in width, so that these arteries were not in
contact with the wall of the bladder until they reached a point between 2 and 3 inches
from the umbilicus, where the peritoneal folds disappeared, and the arteries closed in
upon the sides of the bladder.

The length of the bladder from the umbilicus to the prostate gland was 10 inches.
The prostate gland lay close to and on the abdominal side of the symphysis pubis.

The interior of the bladder was lined by a mucous membrane, presenting numerous
rugosities, which to a large extent lay parallel to each other, and in the longitudinal
axis of the bladder. Towards the outlet the mucous membrane became comparatively
flat and smooth.

The orifices of the ureters were longitudinal oblique narrow slits 2 mm. in length and
5 mm. apart. The lateral margins of each of these openings were continued towards
the outlet as slight ridges for a distance of 10 mm. These ridges met in the mesial
plane, thus forming a mesial longitudinal ridge or uvula vesicae. The actual trigonum
vesicse was therefore a triangular area 5 mm. wide at its base and 10 mm. long on
each side.

The uvula vesicae was continued into the urethra, and became continuous with the
crista urethrse, which attained its greatest prominence 20 mm. from the apex of the
trigonum vesicse. The sinus pocularis was represented by a very small mesial aperture
opening on the distal side of the summit of the crista urethrae.

The prostate gland did not attract attention, and at first sight one would have
doubted its presence. Certainly in cutting into the urethra from its pubic aspect no
variation in consistence was detected. Still, there was a definite thickening of the pubic
wall of the urethra corresponding to the general position of the urethral crest. On the

(ROY. soc. EDIN. TRANS., VOL. XLVIII., 192.)

ON THE ANATOMY OF THE WEDDELL SEAL. 65

other hand, on the rectal or pelvic aspect of the urethra, and in relation to the urethral
crest, there was a mesial longitudinal thickening of firm consistence, from 5-6 mm. in
length. Into the hinder end of this denser part the vasa deferentia entered. The
pelvic portion of the urethra was therefore not surrounded by visible prostatic tissue at
its vesical end, and the prostatic tissue was not prolonged in relation to the urethra as
far as the sub-pubic pelvic wall, because, whereas the prostate was only from 5-6 mm.
in length, the pelvic urethra measured from 35-40 mm. long. No doubt the extreme
youth of the animal accounts for the primitive condition of the prostate, but it is
interesting to note that the part readily recognisable is the mesial longitudinal lobe.
A portion of the prostatic part of the urethra, along with the surrounding tissue, was
prepared for microscopic examination. Definite glandular prostatic tissue was revealed
in relation to the pubic and lateral aspects of the urethra. On the rectal aspect of the
urethra dense fibrous tissue was displayed. The two vasa deferentia were visible, each
quite distinct from the other, so that their close proximity and apparent fusion previous
to their entering the prostate on its rectal aspect was not a real fusion. The urethral
crest presented the section faces of the bifurcated end of the uterus masculinus
(Miillerian ducts).

Each testis was lodged in its own peritoneal pouch, which was situated to the outer
side of the pubic body, in the depression between the pubis and the head of the tibia.
These pouches were completely separated from each other by the keel-like projection
of the pubic symphysis, and thus they did not form any object comparable to a
scrotum. The testes had descended through the abdominal wall on the ventral aspect
of Poupart's ligament, i.e. through the inguinal canal, and not through the crural canal,
notwithstanding the novel position occupied by the testis in relation to the limb as a
whole. The tunica vaginalis testis was in open communication with the sac or cavity
of the abdominal peritoneum, and thus the whole condition might fairly be said to
resemble two imperfectly descended testes, although in this case there was no scrotum
into which they could have descended, nor was it possible for them to descend any
farther. Each testis was considerably flattened, and measured 25 mm. long by 14 mm.
wide.

No hydatids of Morgagni were visible. The epididymis presented a globus major
and a globus minor. It did not lie close to the testis, but was supported by a mesentery,
which at its deepest measured 10 mm. The vas deferens was similarly supported, and
therefore it presented itself clear from the epididymis at the distal end of the testis,
instead of lying close to it as far as the proximal end of the testis, as in man.

The vas entered the inguinal canal in the usual way and crossed the iliac fossa,
running superficial to the external iliac vessels and the hypogastric artery. Thereafter
it hooked round the hypogastric artery, and, passing to its mesial side, it proceeded
backwards, i.e. tailwards, towards the base of the urinary bladder, taking its place to the
mesial side of the ureter on its course. As the vas approached the proximal or pelvic
end of the prostate gland it came into such close contact with its fellow of the opposite

(ROY. soc. EDIK. TRANS., VOL. XLVIII., 193.)

<6'6 PROF. DAVID .HEPBURN. ON THE ANATOMY OP THE WEDDELL SKAL.

side that their adjacent walls became firmly blended together. This produced the
appearance of an enlargement common to both of them, but there was no dilatation or
ampulla on each one. There was no trace of seminal vesicles.

The penis was constructed on familiar lines. A strong, flexible cylindrical structure
was present in the body of the penis extending 7 mm. from the base of the glans
penis backwards. A portion of this structure was removed for microscopical examin-
ation. In transverse section it presented a circular outline, and was equally associated
with the two corpora cavernosa penis. Its resistance to the knife suggested young
bony tissue, and accordingly it was decalcified. Afterwards sections were cut out of
paraffin, mounted, and stained in hsematoxylin and eosin. Under the microscope it
presented the distinctive characters of cancellated bone, being more spongy towards the
centre of the section and denser towards the surface, where it was closely enveloped in
a fibro-vascular sheet of membrane, comparable to periosteum. Numerous bone-cells
were embedded in the developing processes of bone. No trace of hyaline cartilage could
be detected. No doubt this short cylindrical piece of young bone is comparable to the
much larger os penis of the walrus, as well as to the furrow-shaped and partly bilateral
os penis of the fox and the dog. The bulb on each corpus cavernosum penis was situated
in relation to the crus penis, and not on the penile portion of the organ. From the region
between the bulb of the corpus spongiosum penis and the rectum, i.e. corresponding to
the central point of the perineum, there were two parallel bands of tissue running forwards
'towards the distal end of the body of the penis. These were similar to muscular bands
which I have elsewhere* described in connection with the penis of the porpoise.
Probably these act as retractors of the penis. As in the case of the porpoise, a
microscopic examination of sections cut longitudinally and stained after Van Giesen's
method revealed unstriped muscular fibres, with fibrous tissue bundles. Since there
is no scrotum in the porpoise, whose testes are situated mtra-abdominal, and since in the
seal under consideration each testis occupied a recess placed under the integumentary
layers in relation to the inner side of the head of the tibia, it seems not unfair to
consider these non-striped muscular bands as being homologous to the tunica dartos
layer of an ordinary scrotum, more especially as the muscular fibres of the tunica
dartos are of the unstriped or involuntary variety.

* " The Anatomy of the Genito-urinary Apparatus of the adult male Porpoise," HEPBURN and WATERSTON, Trans.
Royal Physical Society, Edinburgh, 1902.

(ROY. .800. EDIN. TRANS., VOL. XL.VIII., 194.)

PART VI.

SEALS.

SCOTTISH NATIONAL ANTARCTIC EXPEDITION.
VL-ANATOMY OF THE WEDDELL SEAL

(LEPTONYCHOTES WEDDELLI}.

(3) THE RESPIRATORY SYSTEM AND THE MECHANISM OF

RESPIRATION.

BY DAVID HEPBURN, M.D., C.M.,

Professor of Anatomy, University College, Cardiff (University of Wales).

Scottish National Antarctic Expedition : Observations on the Anatomy of the
Weddell Seal (Leptonychotes Weddelli). By David Hepburn, M.D., O.M.,
Professor of Anatomy, University College, Cardiff (University of Wales). Part III.

(MS. received March 28, 1912. Read June 3, 1912. Issued separately July 18, 1912.)

THE RESPIRATORY SYSTEM, AND THE MECHANISM OF KESPIRATION.

In the specimen under consideration there were fifteen pairs of ribs, of which nine
pairs were vertebro- sternal. The costal cartilage associated with each of these was long
and very flexible. The articulation of the first pair of costal cartilages with the sternum
was effected by means of a short but strong band of fibrous tissue, which permitted
considerable freedom of movement and did not form a junction of the more or less rigid
character seen in man.

The chondro-sternal joints of the second, third, and fourth costal arches were of the
diarthrodial variety, each joint being divided into two separate cavities by an inter-
articular ligament. The fifth, sixth, seventh, eighth, and ninth chondro-sternal joints
presented diarthrodial joints without interarticular ligaments.

The sternum was long, narrow, somewhat like a four-sided rod, and divided into
segments (suggestive of vertebral centra) by amphiarthrodial joints which were
placed opposite the chondro-sternal joints from the second to the ninth. A suprasternal
tapering cartilage extended towards the head for a distance of two inches, while the
ensiform cartilage extended backwards to a similar distance and ended in a broad semi-
lunar expansion.

The intercostal muscles were well developed, being thick and fleshy, presenting little
or no fibrous intersection. They were arranged so as to present an external and an
internal muscle in each intercostal space, and the direction of their fibres was similar to
that seen in man ; but the fibres of the external muscle were continued between the
costal cartilages close up to the margin of the sternum without the intervention of an
intercostal membrane.

The triangularis sterni muscle arose from the deep surface of the sternum on its own
side of the mesial plane. It consisted of a number of slips, which were wide enough to
give the appearance of a complete sheet of muscle. These were attached to the sternum
from the level of the third costal cartilage backwards to the level of the ninth. The
fibres ran forward and outwards to be inserted into the deep surfaces of the costal
cartilages from the second to the ninth inclusive, and into fibrous bands which passed
from one cartilage to the other. The general line of insertion into the costal cartilages
was near to the series of costo-choudral joints, each of which, except that of the first rib,

(REPRINTED FROM THE TRANSACTIONS OF THE ROYAI SOCIETY OF EDINBURGH, VOL. XLVIH., PP. 321-332.)

72 PROFESSOR DAVID HEPBURN ON

formed a diarthrodial joint. On the ninth costal cartilage this muscle interdigitated
with the attachment of the diaphragm. This large, well-developed muscle was supplied
by a series of twigs derived from the intercostal nerves in relation to which it was
attached.

The sterno-mastoid muscle extended from the anterior end of the sternum and from
the side of its pointed suprasternal cartilage to the mastoid process. There being no
clavicle, this muscle appeared narrow.

The sterno-hyoid and sterno-thyroid muscles arose from the sternum under cover of
the previous muscle. They formed a thin continuous sheet which probably included
the omo-hyoid muscle along its lateral border in the vicinity of the hyoid bone. The
entire sheet was innervated from the hypoglossal nerve, and the insertion of fibres into
the thyroid cartilage and into the hyoid bone suggested the character of its constituent
parts. A thin band of muscle fibres occupying their usual position formed the thyro-
hyoid muscle.

There were two well-defined scalene muscles, both of which were situated on the
dorsal side of the subclavian vessels and cervical nerves, and may therefore be regarded
as the representatives of the scalenus medius and scalenus posticus muscles.

The musculus scalenus medius was inserted into the costal cartilage of the first rib
close to the costo-chondral articulation.

The musculus scalenus posticus was inserted into the lateral aspects of the third,
fourth, fifth, and sixth costal cartilages close to the costo-chondral articulations. At
each insertion the pointed attachment interdigitated with similar attachments of the
musculus obliquus externus abdominis, whose digitations extended to the cartilage of
the first rib.

Regarding the skull and the cervical column as providing the more fixed or rigid
attachment for the -scalene and sterno-mastoid muscles, it is fairly evident that these
muscles may act as elevators of the ribs and sternum by drawing them towards the head.

The diaphragm was well defined in all its parts, but its dorso-lateral portions were
very thin, and in the absence of a central tendon of the trefoil type it presented appear-
ances deserving detailed description, more especially in regard to the important position
occupied by this muscle in the mechanism of respiration. Its strongest part was the
mesial or vertebro-sternal element, which presented two well-marked, pointed crura
attached to the lumbar vertebrae. From this origin the muscular fibres passed in a
ventral direction on either side of the abdominal aorta until they reached the ventral
aspect of this vessel, where to a small extent their fibres intermingled ; but for the most
part the fibres of the right crus were on the ventral side of those of the left crus.

This distinction between the fibres of the two crura was maintained as they continued
towards the oesophagus, along the lateral aspects of which they passed, thereby forming
the cesophageal opening, which was practically in the mesial plane. A short distance
on the ventral side of the oesophageal opening the muscular fibres were inserted into a
circular tendinous ring placed slightly to the right of the mesial plane, and through this

(ROT. soc. EDIN. TRANS., VOL. XLVIII., 322.)

THE ANATOMY OF THE WEDDELL SEAL. 73

ring the inferior vena cava passed. From the ventral face of this tendinous ring strong
muscular bands extended to the deep face of the broad ensiform cartilage.

From each side of the dorsal segment of the fibrous ring surrounding the inferior
vena cava there extended a narrow tendinous septum in the dorso-lateral direction.
Neither of these septa reached the ribs, although the one on the right side was more
strongly marked than that on the left side. Into the dorsal faces of these septa there
were inserted muscular fibres derived from the lateral aspects of their respective crura,
as well as a small, feeble muscular slip from the ventral surface of the second last rib
(the 14th) near its head.

From the ventral aspects of the tendinous septa under consideration, a sheet of
muscular fibres passed outwards to be attached by slips or digitations into the ribs close
to their junction with cartilages from the 8th to the 13th inclusive. The digitation be-
longing to the 13th rib was attached just in front of the angle of the rib. This ventro-
lateral part of the diaphragm was very thin. Areolar tissue occupied the intervals
between digitations attached to the 13th and 14th ribs, and also between the 14th rib
and the lateral margin of the crus. The association of the diaphragm with the 15th rib
was so feeble as to be doubtful. Probably these weak places may be regarded as corre-
sponding to arcuate ligaments, although in the human sense these structures were
undefined. At any rate, these arched ligaments had no other representation, neither
could the slight intermingling of crural fibres on the ventral aspect of the aorta be
regarded as other than a very feeble median arched ligament. Altogether the dorso-
lateral development of the diaphragm was extremely feeble. Those muscular fibres
attached to the 8th rib were in close contact with the mesial part of the diaphragm
between the sternum and the fibrous ring enclosing the inferior vena cava.

From what has been described it will be evident that there was no central tendon of
the trefoil pattern, but in its place a vena caval ring, from which there extended two
dorsal-lateral septa, of which the right was the stronger marked. The shortness of the
left septum permitted a greater mingling of the fibres belonging to the left crus with
those forming the rest of the left half of the diaphragm.

The general arrangements within the chest cavity do not call for special discussion.
The two pleural sacs and the mediastinal interval followed the customary disposition.
It may, however, be noted that the mediastinal layers of pleural membrane were in close
apposition from the second segment of the sternum backwards to the hinder end of the
sternum, and that consequently the ventral or anterior section of the mediastinum was
a mere chink, in its sternal relations.

The lungs were extremely dark in colour, brown almost to black. They were quite
soft, but yielded no feeling of crepitation on pressure, so that they suggested a complete
absence of air.

When they were removed each was placed in water, and they sank as if solid. A
small portion cut from the lung also sank in water, so that this tissue had entirely lost
its buoyancy. It is not quite easy to account for such a complete absence of air from

(ROT. soc. KDIN. TRAMS., VOL. ZLVIII., 323.)

74 PROFESSOR DAVID HEPBURN ON

the substance of the lung. The animal is known to have lived for some days, because it
was killed by poisoning with hydrocyanic acid after an attempt had been made to rear
it by artificial feeding. The carcase was preserved by injecting an arsenical solution.
Neither of these processes would account for the absence of air from the lung tissue.
We must therefore assume that either the natural elasticity of the lung tissue has pro-
duced the condition noted, so that the expiratory apparatus of the animal is able to pro-
duce a practical deflation of the lungs, which is doubtful ; or that, partly owing to the
length of time they have been preserved and partly owing to the preservative solutions,
the air has practically all passed into solution and disappeared.

A portion of the lung was prepared for microscopic examination, and, notwithstanding
the number of years that have elapsed since the animal was embalmed, the different
tissues were easily recognisable, but to staining agents such as hsematoxylin and eosin
they reacted very slowly and not very satisfactorily.

The hyaline cartilage of the bronchioles was cellular, and very similar to the cartilage
in the ear of the mouse.

The lobules of the lung were very clearly defined by interlobular tissue, which was
continuous with the sub-pleural tissue, and throughout this tissue there was a well-
marked amount of elastic fibres.

All the air spaces were shrunken, i.e. collapsed, almost to the point of obliteration,
but they were free from exudation. The capillary blood-vessels in the walls of the
air spaces were crowded with blood corpuscles, which may have been the result of the
preservative injection.

There is some reason, therefore, for considering that the normal elasticity of the lung
in this seal was much greater than that of man, and that, consequently, the air would
be much more effectively expelled from the lungs of the seal during expiratory move-
ments.

Attention may be drawn to certain of the body muscles whose attachments and dis-
position were such as to add to their expiratory value. The panuiculus carnosus muscle
was a thin sheet enveloping the trunk from the hinder end of the abdomen to the face,
and on the face and head forming a cowl modified for facial or expression muscles in
relation to the various apertures in that region. The fore limbs were in effect pushed
through this axial sheet. The disposition of its fibres showed dorso-lateral and ventro-
lateral directions, separated from each other by a lateral aponeurosis, and attached by
aponeurotic fibres to the dorsal and ventral mesial lines such as may be seen in the
porpoise, but less distinct. The direction of the muscle fibres in the dorso-lateral section
was obliquely from before (cephalic) backwards (caudal), whereas in the ventro-lateral
section their direction was obliquely from behind forwards. The general effect of the
contraction of this sheet would be to expel the air from the very elastic and flexible
thorax, as well as to compress the abdomen.

The musculus obliquus abdominis externus showed no attachment to the ilium.
By one end it was attached through digitatious to the entire series of costal arches from

(BOY. soc. EDIN. TRANS., VOL. XLVIII., 324.)

THE ANATOMY OF THE WEDDELL SEAL. 75

the first to the fifteenth. Its fibres were directed obliquely backwards towards the
ventral mesial line, and, having given place to a thin aponeurotic sheet, many of these
fibres interlaced with those from the opposite side to form the linea alba.

The hinder part of the muscle, however, did not form any attachment to the ilium,
but as a muscular arch, equivalent to the ligament of Poupart, they were attached to
the ventral aspect of the body of the pubis. Near to the pubis a slit in the muscle
sheet served the purpose of an external inguinal ring, in which the spermatic cord was
situated. A muscle so attached could clearly act as a very powerful expiratory muscle
provided the glottis were open.

The musculus obliquus abdominis internus was attached dorsally to the lumbar
aponeurosis and to the crest of the ilium, while ventrally it was inserted into the hinder
borders of the last four ribs and also through its aponeurosis into the linea alba. The
greater proportion of its aponeurotic fibres passed ventrally to the rectus abdominis
muscle along with those of the external oblique, but a few of them blended feebly
with the aponeurosis of the transversalis abdominis muscle. This muscle (transversalis)
presented lumbar and iliac attachments as well as a series of digitations on the hinder
seven or eight ribs. Its mesial attachment to the linea alba was by an aponeurotic
sheet placed on the deep side of the rectus abdominis muscle. The hinder or inguinal
margins of these two last muscles were very closely, almost inseparably, blended
together, and both were much thinner than the external oblique muscle.

The rectus abdominis muscle occupied an abdominal sheath whose composition has
already been indicated. It was attached posteriorly to the body of the pubis, and
extended anteriorly to the first costal cartilage, to which, as well as to all the other
sternal cartilages, it was attached by tendinous slips. Here again we can see that this
muscle, acting from a rigid attachment to the pubis, may act as a powerful expiratory
muscle in association with an open glottis.

The lungs, beyond what has already been said, do not call for detailed description.
Each presented a great oblique fissure, and thereby an apical and a basal lobe. In
addition the right lung possessed a transverse fissure, and therefore a middle or ventral
lobe. Furthermore, the right lung had an azygos lobe on its mediastinal aspect in
relation to the margin between diaphragm and pericardium.

On several occasions I have had the opportunity of making a detailed examination
of the respiratory mechanism of mammals whose habitat is either partly or entirely
marine, and on each occasion I have been impressed by the remarkable flexibility of
their thoracic wall, with the associated peculiarities in the attachments of certain of the
muscles. Attention has already been drawn to some of these peculiarities in the
descriptions given above, and it is almost impossible to avoid the conclusion that
respiration necessitates a more flexible chest-wall in the case of mammals surrounded
by water than in those which are surrounded by air, apart from the fact that the
normal attitude of the latter may be horizontal, as in the case of quadrupeds, or vertical,
i.e. erect, as in the case of man.

(HOY. soc. KDIN. TRANS., VOL. XLvm., 325.)

76 PROFESSOR DAVID HEPBURN ON

On the other hand, this flexibility of the chest becomes not only a drawback, but
may be an actual source of danger when the marine mammal comes on shore either by
intention or as the result of accident. Thus it is well known that a cetacean dies when
it runs aground, not necessarily by starvation, but because it is suffocated, since the
flexibility of its chest-wall renders respiratory movements impossible under the superin-
cumbent weight of its body.

The leader of the Scottish National Antarctic Expedition made very careful observa-
tions on the attitude of seals when they left the water and resorted to the ice, and he
noted that they do not assume positions which would hamper their chest movements.
Thus they recline on the side when asleep so as to leave the movements of one side of
the chest unimpeded, while at other times their common attitude is to lie prone with
the chest raised off the ground by the short fore limbs. Considerations of this kind lead
to the conclusion that respiration, but more especially the act of inspiration, can be
seriously impeded or even rendered impossible by the weight of the animal's own body.
That free inspiratory movement of the chest-wall in man may be hampered by the
weight of his body may be readily observed in the case of an operatic singer who, by
the exigencies of his performance, is called upon to sing in the supine position ; for
although in this position he can fill his lungs sufficiently for ordinary respiration,
yet he cannot inspire deep enough for effective vocalisation. Clearly, therefore, the
respiratory mechanism is affected by the attitude of the individual as well as by the
surrounding medium, air or water, in which the animal performs the necessary
respiratory movements.

There can be no doubt that, whatever the natural attitude of the mammal may be,
or whatever its habitat, the ordinary movements of inspiration and expiration are carried
out with the minimum expenditure of effort consistent with the amount of air required
for each respiratory act. On the other hand, special circumstances may call for
additional or extraordinary efforts both as regards inspiration and expiration. The
discussion of respiratory movements is usually left, and by many observers considered
properly left, to the physiologist ; but as these movements are entirely dependent
upon a definite mechanism in which the muscular arrangements play an important
part, they cannot fairly be excluded from the province of the anatomist, and it is
from the standpoint of structure that I propose to offer some observations which
seem warranted by the conditions I have seen in the seal under consideration, as well
as in the porpoise.

It may be well in the first instance to deal with the lungs themselves ; and, as the
condition in which I found them has already been stated, it will only be necessary to
add that, except for the presence of the azygos lobe on the right side, they corresponded
with the human lungs so far as the number and arrangement of fissures and lobes was
concerned. There is no reason to suppose that during the act of inspiration they would
inflate in a manner different from the lungs of man. Now, among the many interesting,
elaborate, and ingenious attempts to explain the respiratory act, none is more suggestive

(ROY. soc. EDIN. TRANS., VOL. XLVIII., 326.)

THE ANATOMY OF THE WEDDELL SEAL. 77

than that of KEITH,* who approaches the discussion of the subject more as an anatomist
than as a physiologist.

It is not my intention to follow Professor KEITH in detail and offer a criticism of the
conclusions arrived at by him. At the same time, some of his statements appear to me
to overlook certain of the anatomical facts. Perhaps the most important fundamental
statement made by KEITH is in reference to the lungs when he says " the upper lobe is
normally expanded by one mechanism, the lower by another," and as a consequence he
insists that " the great fissure, which divides the upper from the lower lobe, is functional
in its significance." Supposing this view to be correct, it would follow that since there
is a third lobe in the right lung of man and a fourth or azygos lobe in the right lung of
a quadruped, with the fissures required for their delimitation, the mechanism for expand-
ing the right lung must differ from that required for the left lung. Further, as regards
the apical or upper lobe, KEITH maintains that because of the impressions of certain ribs
upon the lateral and anterior aspects of the upper lobe, but not upon " the dorsal surface
of the upper lobe," there is " a constant relationship between ribs and spaces " for that
part of the lobe which presents impressions, but a " downward and upward " movement
of the dorsal unmarked part, in which it follows the movement of the lower lobe, because
" the lower lobe and the dorsal part of the upper lobe are chiefly expanded by a dia-
phragmatic mechanism." The argument for a functional significance for the great oblique
fissure seems to me unnecessary if the substance of the apical lobe is to expand in two
different ways simultaneously, for, at least as far as the dorsal part is concerned, the
presence of the fissure does not seem to confer any advantage.

I am not disposed to maintain that the fissures of the lungs have no significance,
although to my mind it is rather structural than functional. Even " the obliteration
of the pleura! cavity by adhesions has so little apparent effect on the respiratory
movements that their presence cannot be detected during life," any more than the
obliteration of the lobulated character of the kidneys interferes with their functions.
After all, the outstanding requirement is that the lungs shall expand to the capacity
corresponding to the immediate muscular effort that is being performed, and naturally,
therefore, the capacity undergoes constant variation. With this end in view, I cannot
but think it is best to consider the muscular mechanism of inspiration as a whole, and
the muscular mechanism of expiration as a whole, since it is their co-ordinated and not
their individual action that we depend upon. Probably, in quiet ordinary breathing,
no animal, any more than the human individual, employs the full scope of its inspiratory
mechanism, and hence in man it has become customary to employ such terms as
" thoracic " and " abdominal " to indicate the character of the inspiratory effort which
is most noticeable in the female and in the male respectively. At the same time, there
is no record of this distinction in the inspiratory act among the sexes of the lower
animals, nor between the human sexes during infancy and early adolescence. It

* "The Mechanism of Respiration in Man," by ARTHUR KEITH, pp. 182-207, in Further Advances in Physiology,
edited by LEONARD HILL, published by Edward Arnold, London, 1909.

(ROY. soc. EDIN. TRANS., VOL. XLVIH., 327.)

78 PROFESSOR DAVID HEPBURN ON

appears to me, therefore, that the double mechanism which KEITH supports for the
expansion of the apical and basal lobes of the lung is more apparent than real, and that
the so-called types of breathing in man are rather the result of the erect attitude
whereby, from the natural configuration and diameters of the diaphragmatic section of
the thoracic cavity in the two sexes, it is with less muscular effort that the male expands
the lower part of his chest and the female expands the upper part of her chest in order
that each may obtain the amount of air necessary for ordinary quiet breathing. When
additional efforts call for more air, or when, as in the supine position, the easy movement
of the chest is impeded, then there is an immediate departure from the characteristic
method ; but I do not think that we must postulate a double inspiratory mechanism.

The key to the whole mechanism of inspiration is undoubtedly the part performed by
the contraction of the diaphragm. We cannot, therefore, overestimate the importance
of its attachments and structure ; nor must we forget that, like any other muscle, its
action is the result of the contraction of its fibres, whereby its attached ends are brought
more or less near to each other. The most favourable method of examining the diaphragm
is to consider the adult structure from the point of view of its development.

The first part of the diaphragm to appear in the embryo, and the part which may
be considered the most powerful in the adult, is its mesial or vertebro-sternal portion,
whose vertebral ends or crura arise from the lumbar vertebrae and constitute its axial
or fixed end. These muscular fibres having adapted themselves to the positions of the
abdominal aorta and the oesophagus, by a certain amount of intermingling of the fibres
from opposite sides of the mesial plane, become inserted into the central tendon, whose
shape varies from the trefoil tendon of man to the vena caval ring with its lateral septa
as seen in the Weddell seal. From the ventral aspect of these tendinous structures a
second set of muscular fibres extends to the deep surface of the lower or hinder end of
the sternum. The mesial part of the diaphragm is therefore in reality a digastric
muscle pursuing an arched course from the vertebral column to the sternum. The
arched character of its course is more pronounced in its dorsal segment, while in its
ventral or sternal segment the arched character is lost, being replaced by a straight or
flat course. This change in the curve of the two segments is due partly to the disposi-
tion of the abdominal viscera and partly because the vertebral attachments are some
distance farther tailwards than a point which would correspond with the hinder end of
the sternum.

By its contraction two results may follow : — ( 1 ) the lower (hinder) end of the sternum
is either drawn closer to the vertebral column or else prevented from being projected
ventrally (forwards) by other influences ; (2) the arched dorsal segment between the
vertebral column and the central tendon becomes more or less flattened, and in conse-
quence the adjacent abdominal viscera are pushed towards the ventral abdominal wall.
At the same time its restraining or bracing action upon the hinder end of the sternum
becomes correlated to the restraining action of the first costal arches upon the manubrium
sterni, whereby the manubrium sterni is maintained at a relatively fixed distance from

(ROT. SOC. BDIN. TRANS., VOL. XLVIII., 328.)

THE ANATOMY OF THE WEDDELL SEAL. 7$

the vertebral column. By reason of the somewhat rigid character of the first costal
cartilages in man, as well as from the fact that they are frequently encased in an ossified
shell, the restraining nature of the connection between the first pair of ribs and the
sternum is very well marked ; but even in the Weddell seal, where a short and powerful
fibrous ligament takes the place of the first costal cartilage, the manubrium is very
firmly retained in its relation to the backbone.

The next part to be added to the diaphragm developmental!}' constitutes its ventro-
lateral segments. In the adult these are composed of muscular fibres which arise from
the ventral and lateral aspects of the central tendon. From this position they extend
in a fan-shaped manner to be inserted into the deep surfaces of the costal arches by
digitations which correspond very closely in number with those ribs that do not reach
the sternum directly through their costal cartilages — that is to say, the false or vertebro-
abdominal series of ribs.

This thin sheet of muscle becomes more and more arched as its slips sink lower on
the series of ribs. When therefore it contracts, each digitation will either draw its own
particular rib nearer to the central tendon or else maintain the ventral end of its rib
at a more or less definite distance from the central tendon.

In this way the ventral ends of the false ribs are provided with temporary or inter-
mittent fixed points, fixation by the contraction of the diaphragm being substituted
for fixation by the sternum, as is the case with vertebro-sternal ribs. In fact, the series
of ribs could with eifect be classified as vertebro-sternal and vertebro-diaphragmatic.

The flattening of the arched surfaces of the diaphragm must increase the available
thoracic space, but under ordinary conditions the addition so provided cannot of itself
be very great, and only becomes important as the central feature of a larger movement.

Developmentally, the last part to be added to the diaphragm is also its weakest
part both in man and in the Weddell seal. This is the dorso-lateral segment, which
consists of muscular fibres forming a delicate sheet extending between the dorso-lateral
aspects of the central tendon and the ligamenta arcuata externa and interna, and
through these with the vertebral column on the one hand and the last rib on the other.
The arched course of these fibres in man must enable them to aid the flattening of the
dorsal parts of the diaphragm and thereby again assist in pushing the abdominal con-
tents in a ventral direction, but in the seal they are so feebly developed that the effect
of their contraction must be practically negligible. I do not doubt that contraction of
the diaphragm may produce some depression of the central tendon, more especially at
its dorsal side, but I doubt whether the depression of the central tendon can take place
on its sternal side or be so pronounced as a whole as to give the " piston action "
described by some observers. My reasons for holding this view may be shortly
summarised. The pericardial bag rests by its base upon the diaphragm, and, when a
central tendon of the trefoil pattern is present, the fibrous bag and the central tendon
are intimately united to each other, but the ventral surface of the pericardium is
attached to the manubrium sterni by a sterno-pericardial ligament which is described

(BOY. soc. M>nr. TRAMS., VOL. XLVIII., 329.)

€0 PROFESSOR DAVID HEPBURN ON

by MACALISTER* as strong and rounded. A similar ligament of weaker character
attaches the pericardium to the ensiform cartilage. These ligaments, especially the
former, would resist the traction of the pericardium in the abdominal direction, and so
resist the depression of the central tendon. Again, the normal liver presents indented
grooves corresponding to, and resulting from apposition with, the ribs which cover it,
and these grooves indicate a fairly constant relation between the liver and the ribs,
since they could not be formed by any plunging or piston action communicated to the
liver from the diaphragm. If, on the other hand, the flattening of the dorsal portion of
the diaphragm pushed the liver ventrally towards the ribs, and at the same time the
contraction of the ventro- lateral portions of the diaphragm drew the lower ribs towards
the liver or even maintained them in a position to resist the liver, then the liver mark-
ings would be at once accounted for. Further, STARLING t states that during ordinary
respiration the central tendon of the diaphragm is practically motionless, but that as
soon as respiration becomes laboured there is an actual downward movement of the
diaphragm, and that during laboured inspiration the breathing is mainly thoracic in
both sexes, "and the abdomen recedes with each inspiration." Apparently, therefore,
according to this observer, it is fair to conclude that a laboured inspiration, by calling
for more powerful action of the diaphragm, results not only in greater flattening and
depression of the dorsal segment of the arched diaphragm, but also in the lower ribs
being drawn closer to its central tendon than during ordinary breathing, which is just
what an examination of the muscular attachments would lead one to expect. I there-
fore arrive at the general conclusion that the diaphragm is the keystone in the inspira-
tory mechanism, and that its chief action consists in resisting the ventral (forward)
movement of the hinder (lower) end of the sternum and of those ribs which are not
directly articulated to the sternum. As a result of this controlling action the whole
series of ribs may participate uniformly in a general lifting or elevating movement
which characterises their position at the end of inspiration as contrasted with their
sunken or depressed position at the end of expiration. Such an elevation of the ribs
does not call for any rotation of their shafts, and indeed, from the nature of their
capitular and tubercular articulations with the vertebral column, rotation of the shaft
would be impossible. But the chondro-sternal as well as the articulations just
mentioned are from the nature of their ligaments adapted to the movements of eleva-
tion and depression of the rib as a whole, and even a small amount of such movement
at the vertebral end of a rib would tend to be magnified by the length and obliquity
of its shaft. In fact, the capsules of the costo-transverse articulations are sufficiently
long to permit of the gliding action necessarily associated with such elevation and
depression.

In the seal there are two muscles whose attachments are only readily comprehen-
sible when considered as part of the mechanism for depressing the ribs after they have

* MACALISTER, Text-book of Human Anatomy, 1889.

t E. H. STARLING in Schafer's Text-book of Physiology, vol. ii. pp. 276 and 280.

(BOY. soc. EDIN. TRANS., VOL. XLVIII., 330.)

THE ANATOMY OF THE WEDDELL SEAL. 81

been elevated. These are the musculus obliquus externus abdominis and the musculus
rectus abdominis. Both of these muscles are attached to the pubis, i.e. to the un-
yielding or rigid pelvis, and between them they provide slips or digitations of insertion
into nearly all of the ribs, even extending to the first. Further, the musculus obliquus
abdominis externus has no attachment to the ilium — in other words, both of these
powerful muscles were pubo-costal in their attachments. With a distended chest and
a glottis firmly closed so as to render the chest wall fairly rigid, these muscles by their
contraction could clearly compress the abdominal contents ; but in association with an
open glottis, of necessity they must pull their rib attachments towards the pubis — in
other words, they must act as depressors of the ribs and thus as powerful expiratory
muscles. Such a depressor action compels one to presume and to accept an elevated
position of the ribs during inspiration.

There is nothing in the mechanism which seems to require one mode of action for
inspiration by the vertebro-sternal ribs and another mode of action by the vertebro-
abdominal ribs. Of course, the first costal arch, from the nature of its sternal articula-
tion, is, even in the seal, capable of less elevation than those ribs whose shafts and
costal cartilages are longer and whose sternal articulations permit greater freedom of
movement ; but, in order to secure a uniform method of elevation throughout the series
of ribs, it is necessary to provide some more rigid line or point d'appui for the start of
the movement, and the combination of the first costal arch with the manubrium sterni,
together with their powerful scalene and sterno-mastoid muscles, provides such a line.
Moreover, in man the clavicle is added to this line by the sterno-mastoid and trapezius
muscles as well as by such ligaments as the costo-clavicular and the sterno-clavicular.
Again, it will be found that those ribs which elevate most readily are just those whose
heads are provided with an interarticular ligament passing between the head and the
intervertebral disc. Such a powerful structure would be unnecessary unless there
were a tendency for the head of the rib to be drawn away from the vertebral column as
the dorso- ventral and transverse thoracic diameters increased owing to the elevation of
the ribs in full inspiration. There is no ordinary form of inspiration which requires a
larger amount of air under regulated control than the inspiratory movement performed
by the trained singer, and one of the approved methods of obtaining this result aims at
the expansion of the large dorsal surface of the lungs by cultivating the upward move-
ment of the ribs in relation to the dorsal surface of the chest, as being not only easier
than, but preferable to, a forced action of the diaphragm. An examination of the
mechanism of respiration leads me to the conclusion that, in all forms of respiration,
this mechanism acts in the same way, but not, in all its parts, to the same extent, for
any particular respiratory act ; that ordinary and laboured respiration differ in degree
rather than in kind ; that the degree of respiration depends upon the amount of air
required for any particular form of exertion ; that the difference between the respiration
of the quadruped and the respiration of man results from differences in their attitude
^horizontal and erect), whereby each cultivates that form of chest movement which

(ROY. soc. EDIJJ. TRANS., VOL. xLvm., 331.)

82 THE ANATOMY OF THE WEDDELL SEAL.

requires the minimum of muscular effort. The differences between the adult human
male and female types of breathing are adaptations due to the avoidance of severe
muscular effort so long as a smaller effort will serve the purpose, and in my opinion
they result from the normal differences in the lower or diaphragmatic diameters of the
thoracic cavity in the two sexes. To a large extent these differences may be accounted
for by the width of the female false pelvis as compared with that of the male. Since
the abdomen proper (i.e. excluding the true pelvis) contains no organs which are not
common to both sexes, it follows that a wide false pelvis, by providing increased
accommodation in the lower abdominal regions, is naturally associated with a reduction
in the dimensions of the upper or diaphragmatic end of the abdomen. These conditions
make elevation of the sternal ribs more necessary in the female than in the male, whose
larger diaphragmatic thorax permits of ordinary breathing without the pronounced or
visible elevation of his sternal ribs, although their movement may become visible when-
ever the supply of air required calls for an extension of the elevating movement.

In either sex, a change from the erect attitude to the horizontal (e.g. to the supine)
position is usually followed by the introduction of more or less of the respiratory
features of the opposite sex, owing to the temporary interference with the amount of
rib movement in common use when the ribs are unobstructed.

(ROY. soc. EDIN. TRANS., VOL. XLVIII., 332.)

PART VII.
TUNICATA.

VIL-THE TUNICATA OF THE SCOTTISH
NATIONAL ANTARCTIC EXPEDITOIN.

BY W. A. HEKDMAN, D.Sc., F.R.S.,

Professor of Zoology in the University of Liverpool.

(WITH ONE PLATE.)

The Tunicata of the Scottish National Antarctic Expedition. By W. A. Herdman,
D.Sc., F.R.S., Professor of Zoology in the University of Liverpool. (With
One Plate.)

(MS. received January 8, 1912. Read February 19, 1912. Issued separately July 3, 1912.)

So far as regards number of individuals, and their size, this is one of the largest
collections of Ascidians brought back in recent years from Antarctic seas. It contains
almost exactly the same number of species of ASCIDIACEA (Ascidise Simplices + Asddiae
Composite) as the Discovery collection — viz. fifteen or sixteen — but whereas in the
latter collection nearly all the species were represented by single specimens, in the
Scotia collection most species can show long series of individuals — in all there are about
two hundred specimens, as against the thirty-three brought home by the Discovery.

The sixteen species in the present collection represent almost as many genera, and
half a dozen families. The systematic arrangement is as follows :—

ASCIDIACEA.

Family MOLGULID.E.

Paramolgula gregaria (Lesson).
Paramolgula horrida (Herdman).

Family CYNTHIIDJE.

Boltenia legumen, Lesson.
Fungulus-antarcticus, n. sp.
Halocij/tthia setosa, Sluiter.

Family STYELID.*:.
Styela lactea, Herdman.
Styda paessleri, Michaelsen.
Synstyela incriistans, Herdman.

Polyzoa opuntia, Lesson.
Goodsiria placenta, Herdman.

Family ASCIDIID.E.
Ascidia charcoti, Sluiter.

Family DISTOMID.E.

Colella pedunculata (Q. and G.).
Holozoa cylindrica, Lesson.

Family POLYCLINID.E.

Polyclinum complanatum, Herdman.
Amaroucium distomoides, Herdman.
Amaroudum sp.

It is interesting to notice how greatly some of these recent collections from the far
South differ from one another in the species represented. The following table — which
gives only the sixteen species in the Scotia collection — shows that only one form
(Halocynthia setosa) from that collection was also taken by the Discovery, whereas ten
species were taken by the Challenger, eight by the Hamburg Magellanic and South
Georgia Expedition, and five by the French Antarctic Expedition under Charcot. This
can be explained to some extent, at least, by the precise localities visited : the Scotia, the
OiaJlenger, and the Hamburg collections were largely made in the Magellan and Falk-
lands neighbourhood, while the other three collections were mainly from farther south.

(REPRINTED FROM THE TRANSACTIONS OF THE ROYAL SOCIETY OF EDINBURGH, VOL. XLVIII., pp. 305-320.)

88

PROFESSOR W. A. HERDMAN ON THE

" SCOTIA " SPECIES.

"DISCOVERY."

"CHALLENGBU."

CHARCOT.

"SOUTHERN
CROSS."

MAGELLAN
AND SOUTH
GEORGIA.

Paramolgula horrid a,

X

X

„ gregaria,

X

X

Boltenia legumen, .

X

X

Funr/ulus antarcticus,

("Scotia"

only)

Halocynthia setosa, .

X

X

Styela lactea, .

X

X

X

„ pnessleri,

X

Syiistyela incrmtans

X

X

Polyzoa opuntia,

X

X

Goodsiria placenta, .

X

Ascidia charcoti,

X

Oolella peduneulata,

X

X

X

Holozoa cylindrica, .

X

X

X

X

Polydinum complanatum.
A maroucinm distomoides,

}ln collection of Australian Museum, Sydney.

Amaroucium sp.,

Although only one of the Scotia Tunicata requires to be described as new to science,
several of the species are of considerable interest, and most of them add something to
our knowledge either in the characters and variation of the species or in distribution.
The one new species (Fungulus antarcticus) is a very remarkable form belonging to the
deep-sea genus Fungulus, known only from a single species obtained during the
Challenger Expedition between the Cape of Good Hope and Kerguelen Island.

This collection shows again what I remarked upon more than twenty years ago in
the case of the Challenger collection, that the Ascidian fauna of the far South is
characterised by the abundance and the large size of the individuals of a comparatively
few species. Halocynthia setosa and Holozoa cylindrica are the two largest species, the
one simple and the other compound, and both are represented by a large number of
specimens. I have, however, written on this matter, and also on the number of Antarctic
as compared with Arctic species, so recently in my report * upon the Discovery collection
that these matters need not be discussed further here.

Family MOLGULID.E.

Paramolgula gregaria (Lesson). (Plate, fig. 9.)

Cynthia gregaria, Lesson, Cent. Zool., p. 157.

Molgula gregaria, Herdman, Challenger Report on Tunicata, Part I., p. 73.

Locality. — Station 118, on hulks, Stanley Harbour, Falkland Islands, January
16, 1903.

There are over forty specimens of this species in the collection, ranging in size from
2 x 1*5 cm. up to 6 '5 x 5 cm. The majority are about 4 cm. in diameter. They have

* National Antarctic Expedition: Natural History, vol. v., "Tunicata," 1910.
(ROY. soc. EDIN. TRANS., VOL. XLVIII., 306.)

TUNICATA OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 89

the characteristic smooth test and translucent grey tint, and they also agree closely
in internal details with the Challenger specimens from shallow water off the Falkland
Islands.

MICHAELSKN has suggested that this species and Paramolgula gigantea (Cunningham)
are the same. No doubt they are related forms ; both belong to the restricted genus
Paramolgula, having only broad, ribbon-like longitudinal bars but no true folds in the
branchial sac (a fact I overlooked in drawing up my " Revised Classification of the
Tunicata" in 1891, as MICHAELSEN has pointed out), but I do not consider them as
identical. In addition to differences in the external appearance — the shape and the
condition of the test — the branchial sacs are not alike in detail, and the dorsal tubercles
differ widely. I give here a figure of the dorsal tubercle (Plate, fig. 9) of P. gregaria
from the Scotia collection for comparison with that of P. gigantea figured in the
Challenger Report.

LESSON figures his species with five lobes round each aperture, but that is no doubt
an error. The branchial aperture has six and the atria! four lobes.

Paramolgula horrida (Herdman) (?). (Plate, figs. 10 and 11.)

Locality. — Station 118, on hulks, Stanley Harbour, Falkland Islands.

I have very little doubt that this single large specimen (measuring 7 '5 cm. in length
and 5 '5 cm. in breadth) belongs to the same species as the specimen from "off the
Falkland Islands, 5-12 fathoms," which I described in the Challenger Report as Molgula
horrida. They both fall within the more modern genus Paramolgula, separated off
from Molgula by TRAUSTEDT because of the absence of true folds in the branchial sac.
As the Challenger description was drawn from a single specimen, and as this Scotia
specimen differs a little in detail, it may be well, in the interests of a fuller knowledge
of the species, to add a few of the characteristics of the individual before me.

The shape is irregularly ovate, and flattened, and the colour is a very dark
brown. The other external characters can be seen from the figure (Plate, fig. 10).
The Test is leathery and rough on the surface. It is thin but tough, and dark
but smooth and glistening on the inner surface. The Mantle is dark brown and
opaque. It is thick, but soft and not muscular, or at least the muscles do not form
obvious bands.

The Branchial Sac has on each side seven wide longitudinal vessels which look like
narrow folds. Between the distant wider transverse vessels, narrower intermediate ones
branch in all directions in a dendritic manner, so as to form rounded and oval and
variously shaped meshes in which the stigmata lie. The stigmata are also rather
irregular in arrangement, being in some parts in spirals and in other places side by side
in rows (see fig. 1 1 ).

The Tentacles are of different sizes, there being eight larger much branched, with
some smaller ones between.

(ROY. soc. EDIN. TRANS., VOL. XLVIII., 307.)

90 PROFESSOR \V. A. HBRDMAN ON THE

The Dorsal Tubercle has its long axis antero-posterior and its opening at the side.
The horns form two close spirals, both coiled inwards. Whether P. patagonica of
MICHAELSEN is also the same species as P. horrida is very doubtful. I am inclined to
regard it as distinct.

Family CYNTHIIDA;.
Boltenia legumen, Lesson.

Locality. — Station LI 8, on hulks, Stanley Harbour, Falkland Islands, about twenty
specimens ; and eight specimens from 6 fathoms, Port Stanley, February 2, 1904.

There are over two dozen specimens of this species in the collection, and they range
in size from I '5 to 7 cm. in greatest length of body — about the same range as in the
case of those in the Challenger collection. All of these specimens of B. legumen belong
to the " forma typica " of MICHAELSEN'S system * of subdivision of this species, and
agree in character with the Challenger specimens from the same locality. In some
cases the little bristles on the surface of the test are more abundant and more prominent
than in others, but there are all gradations between. This is evidently a very common
Ascidian in shallow water at the Falklands.

Fungulus antarcticus, n. sp. (Plate, figs. 15 to 18.)

A single specimen which clearly belongs to the rare and interesting genus Fungulus
was obtained at Station 301 from a depth of 2485 fathoms, on March 13, 1903, at
lat. 64° 48' S., long. 44° 26' W. ; temp. 31°'02. The genus was established in 1882 for
another solitary individual found in the Southern Ocean during the Challenger Expedi-
tion, at Station 147, between the Cape of Good Hope and Kerguelen Island, lat. 46°
46' S., long. 45° 31' E. ; depth, 1600 fathoms, on a bottom of Globigerina ooze. The
two localities are thus nearly 3000 miles apart, but agree in that both are in the far
south and in very deep water.

The Scotia specimen, although closely related to the Challenger Fungulus cinereus,
Herdman, cannot be placed in the same species. The general appearance and anatomy,
and especially the remarkable structure of the branchial sac, are the same ; but the
relation of the peduncle to the apertures and the details of structure are different in
the two forms. The description of F. antarcticus is as follows : —

The body is club-shaped (fig. 15), like a rounded knob about 1'5 cm. in diameter
on the summit of a short, stout peduncle, which is also about 1 -5 cm. in length and
from 4 to 6 mm. in thickness. The peduncle is continuous with the ventral edge of
the body, while the dorsal edge projects markedly. The surface is smooth and the

* " Die Holosomen Ascidien des magalhaeniseh-siidgeorgischen Qebietes," Zoologica, Bd. xii., Heft 31, Stuttgart,
1900, p. 109.

(ROY. SOC. EDIN. TRANS., VOL. XLVIII., 308.)

TITNICATA OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 91

colour pale yellowish grey. The branchial aperture is a little way along the ventral
edge of the anterior end, and appears to be bilabiate or elliptical rather than lobed.
The atrial aperture is in the middle of the dorsal surface, and is a square or four-lobed
opening. In F. cinereus, the branchial aperture was described as triangular, and the
atrial as bilabiate, but the figure of the former in the Cliallenger Report is not very
difierent from the figure now given (fig. 16) from the Scotia specimen.

The Test is thin and membranous, but tough. Under the microscope it is seen to be
finely roughened all over the outer surface. In minute structure the test agrees with
that of F. cinereus as described in the Challenger Report. The Mantle is very thin and
inconspicuous, but muscular. It is penetrated by numerous, very fine, closely placed
muscle bundles which, in the tubular extension of the mantle which occupies the hollow
peduncle, run longitudinally parallel to one another.

The Branchial Sac is remarkably delicate, and is, in fact, merely a very loose wide-
meshed net with folds at intervals where the longitudinally-running vessels are crowded
together (fig. 17). The transverse vessels are of two sizes, occurring alternately. The
looseness of the branchial sac and the minute undulations in practically all the muscle
bundles of the mantle give the impression that when alive and filled with sea-water the
animal had the power of expanding to a considerably larger size than it now shows.
Possibly the test when alive was of a gelatinous consistency and capable of being
dilated.

There are no spicules in the vessels of the branchial sac. The endostyle is narrow
but conspicuous ; there are no spicules in its wall. The branchial tentacles are few and
only slightly branched. The alimentary canal is relatively small, and is confined to
the posterior end of the left side close to the top of the peduncle (fig. 18). The stomach
wall has a number of close-set longitudinal folds.

The gonads are one on each side, rather long and irregular, with the narrower end
pointing to the atrial aperture (fig. 18).

This new species differs from Fungulus cinereus, Herdman, in the shape and pro-
portions of the body (see figures) and in the much paler colour of the test ; in the
details of position and shape of the branchial and atrial apertures ; in having the trans-
verse vessels of the branchial sac distinctly of two sizes ; in having a well-marked
stomach with longitudinal folds ; and in the length and shape of the gonads.

Halocynthia setosa, Sluiter.

This very striking and characteristic species was obtained by the Scotia in consider-
able quantity at the South Orkneys. It was originally described by SLUITER * from two
specimens obtained by the French Antarctic Expedition under Dr JEAN CHARCOT at " He
Booth Waudel, 40 metres " ; but as the figures in the report on the Charcot Expedition
did not seem to me to be characteristic, I gave a supplementary description, with figures.

* Bull. Miis. Hist. Xat. Paris, 1905, No. 6, p. 472 ; and Expe'd. Antarct. Fraiif. (Charcot), " Tuniciers," p. 40.

(BOY. soc. EDIN. TRANS., VOL. XLVIII., 309.)

92 PROFESSOR W. A. HERDMAN ON THE

of the species in my report upon the Tunicata of the Discovery Expedition."" The
Discovery obtained in all five specimens from the east end of the Barrier and the
neighbourhood of the winter quarters in M°Murdo Bay, in 10-20 fathoms.

The more abundant material obtained by the Scotia gives us still further informa-
tion in regard to the characteristics and variation of the species. Some of the Scotia
specimens, measuring up to 11 x 7 x 5 cm., are the largest yet obtained ; and some of
them show a short peduncle at the place of attachment, a feature not previously
observed. The characteristic spines on the test reach in some of these larger specimens
to a length of 21 mm. and a breadth of about 1 mm. at the base. In some of these
specimens the musculature of the mantle is remarkably strong, and consists externally of
circular siphonal sphincters, beyond which is an oval sphincter, of numerous fibres,
enclosing both siphons, while more internally radial muscles formed of exceedingly
stout and strong fibres run outwards from the base of the siphons. Connective tissue
permeated by fine fibres covers and unites all these various muscle bundles.

Of the six folds on each side of the branchial sac, the largest is the one next to the
dorsal lamina on each side, and the smallest is generally the third counting from the
dorsal to the ventral edge. The transverse vessels are of four different sizes, and the
stigmata are from nine to twelve in a mesh. One dorsal tubercle examined measured
4 '8 mm. from side to side and 3 '2 mm. antero-posteriorly.

The nerve ganglion is extraordinarily narrow and elongated, and may reach 9*5 mm.

*

in length, with two nerves diverging from each end which can be traced with the eye
round the sphincters of both siphons. The subneural gland is in the form of a thin
layer over the ganglion.

A strong band of muscle fibres lies under the dorsal lamina and extends from the
mantle into the wall of the branchial sac near the posterior end of the nerve ganglion.

Two of the specimens had Amphipods in the branchial sac. The specimens were
obtained as follows :—

I. Station 325,t 9-10 fathoms, Scotia Bay, South Orkneys, July 1903.

(1) 9x6x6 cm. (on a short peduncle).

(2) 9'5 x 6'5 x 3 cm. (very short peduncle).

(3) 8x5-5x4 cm.

II. Station 325, 9-10 fathoms, June 1903 ; temp. 29° F.

(1) 4 x 4'5 x 2'5 cm. (test only).

(2) 8 x 5'5 x 4 cm. (with a smaller one attached).

III. Station 325, 9-10 fathoms, August 1903 ; temp. 29° F.

(1) 6 x 5 x 4'5 cm. (also four empty tests).

IV. Station 325, 9-10 fathoms, May 1903 ; temp, about 28° F.

(1) 7'4x 5 x 5 cm.

(2) 6x4x4 cm. (also four empty tests).

* Report National Antarctic Exped. : Nat. Hist., vol. v., "Tunicata," London, 1910.
t The whole of Scotia Bay is termed Station 325 ; consequently, depths vary.

(ROY. soc. EDIN. TKANS., VOL. XLVIII., 310.)

TUNICATA OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 93

V. Station 325, 9-15 fathoms, April 1903 ; temp. 28° to 29° F.

About a dozen specimens ranging from 11x7x5 cm. down to 5'5 x 5 x 3'2
cm. (one empty test).

VI. Station 325, 9-15 fathoms, December 26, 1903 ; temp. 3l°'4 F.

(1) 4^x3x3 cm.

Family STYELID.E.

Styela lactea, Herdman. (Plate, figs. 1-8.)

Styela lactea, Herdman, Challenger Report on Tunicate, Part I., p. 156.
Styela flexibilis, Sluiter, Charcot Exped., "Tuniciers," p. 36.
(]) Cynthia verruco«a, Lesson, Ceitt. Zool., p. 151.

Localities. — (1) Station 118, on hulks, Stanley Harbour, Falkland Islands.

(2) Scotia Bay, South Orkneys, Station 325, February 2, 1904.

(3) Attached to Holozoa cylindrical, thrown up on beach. Scotia Bay, January
17, 1904.

(4) Station 118, shore pools, Port Stanley, January 1903. (Two elongated
specimens.)

The specimens from the Falkland Islands are about twenty in number, ranging from
little globular spiky balls (see figs. 3, 4) of 1 cm. in diameter to irregular barrel-shaped
masses (fig. 1 ) of 8 cm. in length and 5 to 6 cm. in breadth. The specimens from
Scotia Bay attached to the compound Ascidian Holozoa cylindrica, Lesson ( = Distaplia
ignota, Herdman), are small and globular, bristling with short pointed spikes, and of a
pure white colour (fig. 2) ; while the remaining specimen from Scotia Bay (February 2)
is much larger, roughly cylindrical in shape, less spiny, and of a duller colour (fig. 1).
Still, all transitions in shape and appearance can be found between the extreme forms,
so there can be no doubt that all belong to the one species, S. lactea, found by the
Challenger Expedition at Kerguelen Island, and by the Southern Cross Antarctic
Expedition at Cape Adare.

The largest Scotia specimens correspond closely with SLUITER'S S. jlexilnlis, found
during the Charcot Expedition at " He Booth Wandel." That species agrees in internal
characters with S. lactea so closely that I have no doubt that the two are the same,
and that S. flexibilis must be regarded as a synonym of S. lactea. It is, I think,
possible also that the Cynthia verrucosa of LESSON, found attached to Fucus on the
shores of Malonines Islands, Antarctic, which is figured as having five lobes round each
aperture, is really this same species. If so, the number of lobes shown by LESSON is, of
course, erroneous.

The following additional characters, taken from the larger Scotia specimens, may be
useful to compare with the descriptions of other specimens :—

Size 7 x 4 x 3'5 cm. Barrel-shaped, attached by flat area at posterior end about
3 '5 cm. in diameter. Colour pale creamy white with a pinkish tinge in places. Test

(ROT. HOC. EDIN. TRA.N8., VOL. XLVTII., 311.)

94 PROFESSOR W. A. HERDMAN ON THE

thin, leathery, raised at intervals to form little pointed tubercles, the larger of which
are echinatecl (fig. 5). Mantle muscular, with regular circular and longitudinal bands.
Branchial Sac with four large folds on each side. There are six to nine bars on a fold,
and four in the interspace. Dorsal lamina a broad plain membrane. There are about
thirty very long simple tentacles and some intermediate smaller ones. The dorsal
tubercle has both horns coiled inwards to form short spirals (fig. 8). There are two or
three long gonads on each side, and many endocarps. Fig. 7 shows the arrangement
of the alimentary canal.

In the smaller, more globular specimens the conical spiny tubercles on the test are
relatively more numerous and more closely and regularly placed (see figs. 2, 3, 4, and 6).
The Challenger specimen figured, from Kerguelen, was intermediate in size to the
larger and the smaller Scotia examples, and was smoother in character of test.

Styela paessleri, Michaeisen. (Plate, figs. 12 to 14.)

This species was described by MICHAELSEN in 1900 from specimens obtained in the
Straits of Magellan. The Scotia specimens from the Falkland Islands seem to be
rather larger on the whole, but agree in essential characters.

The following description, from the Scotia material, may be useful : — There are
about twenty specimens, varying in size from 1 cm. to 3 cm. in length by 1'5 cm. in
average breadth, obtained from Station 118, at the Falklands, depth 6 fathoms; and a
couple from Port Stanley, February 2, 1904, 6 fathoms.

The colour varies from a creamy white to a yellowish brown, and the surface of the
test is in most places closely wrinkled. The branchial sac has four folds on each side,
the largest being those adjacent to the dorsal lamina, with ten bars each, while the others
have usually six bars. Fig. 12 gives the plan of both sides of the branchial sac as seen
in section, with the number of bars and of rows of stigmata shown by the figures. The
folds have from five to ten bars, and there are from two to five (usually four) bars in the
spaces between. These numbers agree fairly well with those given by MICHAELSEN.
The transverse vessels are of three sizes arranged with regularity : 1 — 3 — 2 — 3 — 1, and
having a narrow horizontal membrane in addition crossing the meshes (fig. 13). Most
of the meshes are square, with five to seven stigmata in each. The extreme dorsal and
ventral meshes are more elongated transversely, and contain a greater number of stigmata.

The dorsal tubercle is of curious form (fig. 14), a simple crescent with the horns
anterior and having a globular excrescence in the concavity. The dorsal lamina is a
plain membrane. The tentacles are crowded and number about a hundred. They are
of two sizes, roughly fifty of each. MICHAELSEN records only sixty tentacles, but as
the specimens he examined were smaller than ours, the difference need not be regarded
as important.

Although some of the above characters do not agree precisely with those given by
MICHAELSEN, still the differences are not, I think, greater than what may reasonably be

(ROY. soc. EDIN. TRANS., VOL. XLVIII., 312.)

TUNICATA OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 95

ascribed to individual variation within the limits of a species. The dorsal tubercle is
perhaps the feature that shows most divergence, but MICHAELSEN himself remarks in
the original description that it is probable that other specimens might show a different
form of tubercle.

Polyzoa opuntia, Lesson, subspecies coccinea, Cunningham.

Goodsiria coccinea, Cunningham, Trans. Linn. Soc. Land., xxvii.
Several specimens of this common species were obtained at the Falklands :—

(1) Station 349, shore pools, Cape Pembroke, January 1903 to January 1904. One

large, lobed colony and a couple of small ones. This is part of collection
made on behalf of Scotia by Mr PEARSON, Cape Pembroke lighthouse-keeper,
during twelve months.

(2) Station 118, rock cod trap, Stanley Harbour, 3£ fathoms, January 1903. One

elongated colony, about 26 cm. in length

Goodsiria (Gynandrocarpu) placenta, Herdman.

Several specimens that seem to agree closely with this South African species were
obtained at the Falklands, as follows :—

(1) Station 118, Stanley Harbour, January 7, 1903. One small colony.

(2) Station 118, rock cod trap, Stanley Harbour, 3^ fathoms, January 1903. Part

of a large colony which probably measured 10 or 12 cm. across.

(3) Station 118, Port Stanley, 6 fathoms, February 1904. One colony measuring

about 10 by 5 cm.

Synstyela incrustans, Herdman.

(1) Allaeocarpa zschaui (Michaelsen).

Locality. — Station 118, on hulks, Stanley Harbour, Falkland Islands.

There are about a dozen colonies of this species, ranging in size from 1 or 2 cm. up
to 5 or 6 cm. in diameter. Most of them were adhering in masses along with the larger
specimens of Paramolgula gregaria.

In detailed characters these specimens agree well with the Challenger specimens of
Synstyela incrustans obtained in the Straits of Magellan, but they also agree with
MICHAELSEN'S description of Allceocarpa zschaui from South Georgia ; and when
mature, the Ascidiozooids show the male unisexual polycarps on the left, and the female
on the right-hand side of the mantle, which is a character of MICHAELSEN'S proposed
generic division Alloeocarpa. As, however, he names my species Synstyela incrustans
as the type form of Allceocarpa, and as he apparently does not in his system retain
Synstyela as a genus, but substitutes the name Alloeocarpa for it, I must point out

(ROT. soc. EDIN. TBANS., VOL. XLVIII., 313.)

96 PROFESSOR W. A. HERDMAN ON THE

that Synstyela, Giard, has the prior claim and must be retained as the name of the
genus, even when, as happens to be the case, our knowledge of the internal characters
has been increased and the definition added to since the genus was originally created.
Consequently I must regard MICHAELSEN'S Allceocarpa zschaui as a Synstyela, and
furthermore I h'nd myself unable to distinguish it as a species from S. incrustans of
the Challenger Report. In MICHAELSEN'S " Revision der compositen Styeliden oder
Polyzoinen," * where both species are described, in his table on p. 73 he distinguishes
them by the proportions of the oviduct and the number of internal longitudinal bars in
the branchial sac, as follows :—

S. zschaui having the oviduct broader than long, and having sixteen to seventeen

bars on each side ; and
S. incrustans having the oviduct longer than broad, and having twelve to fourteen

bars on each side of the sac.

Now, in the first place, with a soft, easily deformed structure like the oviduct it is
almost impossible to be sure of the true proportions ; and secondly, I find them varying
considerably in my specimens ; so that I cannot say they agree more in this character
with the one species than with the other. Then as to the number of longitudinal bars,
on dissecting out and mounting a branchial sac from a Scotia specimen I find the
number of bars to be fifteen on each side. According to MICHAELSEN, if it had sixteen
the species would be zschaui, and if it had fourteen it would be incrustans. Under
these circumstances, and as I find the specimens before me agree equally well with the
descriptions of these two species, 1 think there can be little doubt but that A. zschaui,
Michaelsen, is a synonym of Synstyela incrustans, Herdman.

Diandrocarpa monocarpa (Sluiter) is certainly not the same species as Synstyela
incrustans, although it is probably a Synstyela. The number of longitudinal bars in
the branchial sac is very much smaller than in the present species.

Family ASCIDIID^E.

Ascidia charcoti, Sluiter.

Locality.— Station 325, in shore pool, Scotia Bay, South Orkneys, February 2,
1904.

The single large Ascidia in the collection clearly belongs to SLUITER'S A. charcoti,
a species found by the Charcot Expedition to be abundant at " He Booth Wandel."
The Scotia specimen measures 8'5 x 5'5 x 2 cm., and was attached by a small area in
the middle of the left side. The branchial aperture has only seven lobes, a curious little
detail in which it agrees with SLUITER'S description. The atrial has the usual six lobes
characteristic of the genus. The test reaches a thickness of 2 to 3 mm., but has not
the red colour mentioned by SLUITER ; and the mantle is unusually thick and spongy

* Milteilungen aus dem Naturhistor. Museum,, xxi., Hamburg, 1904.
(ROY. soc. KDIN. TRANS., VOL. XLVIII., 314.)

TUNICATA OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 97

for an Ascidia. The branchial sac is also thick, and both mantle and branchial sac are
of a distinctly pinkish colour which may be the remains of the orange-red that SLUITER
records. There are twelve moderate-sized tentacles, and the horse-shoe shaped dorsal
tubercle is very large, reaching up to the base of the tentacles. It seems larger than in
SLUITER'S specimens, in which, however, the dorsal tubercle is recorded as being rather
variable.

SLUITER states that no teeth are present on the dorsal lamina ; but I find that in
the Scotia specimen the dorsal lamina has marked denticulations along its free edge,
amounting in one part to short tentacular languets. But still I have no doubt that
my specimen belongs to SLDITER'S species, and that the dorsal lamina must be regarded
as somewhat variable in character. The viscera on the left side of the body are
unusually large and opaque.

Family DISTOMID^E.

Colella pedunculata (Quoy and Gaimard).

T Sycozoa sigillinoides, Lesson.
1 Colella tenuiccudii, Herdman.
? Colella umbellata, Michaelsen.

One colony having a stalk bifurcated near the top and bearing two " heads " was
found at Station 346 on Burdwood Bank, 56 fathoms, on December 1, 1903, and presents
to some extent characters recalling all the species named above. In the branching of
the peduncle it is like MICHAELSEN'S C. umbellata from the Falklands ; in the general
appearance of the " head," however, it is more like QDOY and GAIMARD'S C. pedunculata,
found by the Challenger at the Straits of Magellan, the Falkland Islands. Kerguelen,
etc. The long slender stalk recalls the Australian C. tennicaulis ; and it is possible
that LESSON'S Sycozoa sigillinoides may be identical with one or more of these other
named forms. Both the "heads" are, unfortunately, in the single colony in a very
ragged condition — possibly dead when collected — so that the more minute characters
of the Ascidiozooids cannot be determined.

Holozoa cylindrica, Lesson. (Plate, fig 2.)

(?) igntitus, Herdman, Challenger Report, ii., 1886, p. 251.

Julinia amtralis, Caiman, Quart. J. Micr. Sci., 1894, p. 1.

Distaplia ignota, Herdman, Report on "Southern Cross" Tunicata, Brit. Mus., 1902, p. 197.

Holozoa cylindrica, Less.— Hartmeyer, in Bronn's Tier-Reichs, "Tunicata," 1909.

This large and apparently abundant Antarctic species was obtained by the Scotia
Expedition at the following localities : —

I. Station 346, Burdwood Bank, 56 fathoms, December 1, 1903. Seventeen
fraoments from 10 to 30 cm. in length by 2 to 4 cm. in diameter. All

O •_ »

in bad condition, soft and partly macerated, with many other animals,
Hydroids, Polyzoa, etc., entangled in the irregular, ragged surface.
(ROY. soc. EDIN. TRANS., VOL. XLVIIL, 315.)

98 PROFESSOR W. A. HERDMAN ON THE

II. Station 325, Scotia Bay, April 1903. One specimen, 30x4x3 cm., bad
condition.

III. Station 325, Scotia Bay, South Orkneys, December 6, 1903; temp. 29'8° ;

floating on surface. 80 cm. x 2 (tapering to 1) cm.

IV. Station 325, Scotia Bay, South Orkneys, December 26, 1903 ; temp. 307°.

(1) 85 (incomplete) x T5 (tapering to 1) cm.

(2) 75 cm. (incomplete) and two fragments.

V. Station 326A, Brown's Bay, South Orkneys, November 1903. Two specimens :

(1) 55 x 2 to 3 cm ; (2) 40 x 2 cm.

VI. Scotia Bay, South Orkneys, January 17, 1904; temp. 32"5° ; thrown up on
beach. One colony, 20 x 5 cm., with several Styela lactea attached ; in
bad condition ; most of Ascidiozooids lost.

VII. Scotia Bay, South Orkneys, January 3, 1904; temp. 31'5°; thrown up on
beach. Two very long specimens: (1) over 100x2 cm.; (2) over
150 x 2 cm.
VIII. Scotia Bay, South Orkneys, November 25, 1903; surface. Three small

colonies, 20 to 30x 1 to 1'5 cm.
IX. Scotia Bay, March 25, 1903. One small colony, 10x2 cm. ; bad condition ;

most of Ascidiozooids gone.

Most of these specimens are, unfortunately, in very bad condition, and were probably
dead or decomposing when collected. The Challenger specimens were in such a rotten
condition that it was impossible to determine even the genus. But from the rather
better material brought home by the Southern Cross Expedition I was able to determine
that the Challenger specimens — evidently the same species — belonged to the genus
Distaplia. What CALMAN described as Julinia australis in 1894 is again the same.

SLUITER, in his report on the Charcot Tunicata, thinks that " Julinia " may be
recognised as an independent genus because of the elongated form of the colony ; but
Distaplia clavata (Sars), from Arctic seas, although it does not attain to such a length,
has the same elongated form — and therefore it cannot be said that a Distaplia with this
habit of growth is unknown.

The colony found floating on the surface in Scotia Bay, December 26, and measuring
about 85 cm. in length, is the best preserved specimen in the collection, and I think
the best preserved that I have seen in any collection brought back from the Antarctic.
The colony, although soft, does not seem to be rotten. The Ascidiozooids are distinct
and large and closely placed throughout its length. Their exposed ends measure about
2 mm. across, and are of an opaque pale yellow colour, in contrast to the translucent grey
of the test in which they are embedded. Throughout the greater part of the colony the
Ascidiozooids appear to be in long meandering lines, but here and there one comes upon
a circular, elliptical, or more irregular group (fig. 2), reminding one of the arrangement
in a Botrylloides. Both ends of the colony are incomplete, and at the upper end the
Ascidiozooids appear to be dropping out of the test.

(ROY. soc. EDIN. TRANS., VOL. XLVIII., 316.)

TUNICATA OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 99

Some of the Ascidiozooids in this colony are the best preserved I have seen in all the
various samples of this species that have passed through my hands, and their anatomical
and histological characters agree in detail with the excellent account of " Julinia " given
by CALMAN. In fact, I can agree with CALMAN in every respect save that of bestowing
a new name on the genus. It is evident from his remarks that he recognised the close
affinity to DistapUa, and the only mistake he made was in not referring the species to
that genus.

I agree, however, with HARTMEYER* that it is practically certain that this form had a
distinctive generic name applied to it at a still earlier date. The " Holozoa cylindrica"
of LESSON (Voyage " Coquille," Zool., ii. p. 439 ; 1830) agrees in all the points that are
mentioned in the brief description with our form. It is said to have a " holothuriform "
body, cylindrica], with rounded ends, free and floating (which is apparently the con-
dition in which our form is usually picked up), of mucous appearance, with a whitish
fibrous centre composed of tubes coming from the ends of the animals ( = Ascidiozooids).
It was found " 30 leagues from Terre-des-Etats," at the southern extremity of America.
I notice that MICHAELSEN (Hamburger magalhaensiscJie Sammelreise, " Tunicaten,"
1907, p. 40) has also suggested with a (?) that LESSON'S Holozoa cylindrica is the same
as "Julinia" (or DistapUa) ignota.

Family POLYCLINID/E.
Polyclinum complanatum, Herdman (?).

The species was described t from a specimen obtained at Port Jackson, Australia.
The Scotia material was taken at Station 483, at the entrance to Saldanha Bay, on
May 21, 1904, from a depth of 25 fathoms. It consists of four fragments, cut probably
from the same colony, the largest of which measures about 6 cm. by 2. The colony
was apparently flattened, and had much the same shape and colour as the Australian
one. The Ascidiozooids also have the same type of structure. The post-abdomen is
rather longer than in the Australian specimens, but that is a matter that varies with
the reproductive condition. The specimens are, however, so fragmentary, and there
is so little that is distinctive, that I cannot be certain as to the identity of the species ;
but there is nothing in the microscopic details to negative the view that the Falkland
Islands specimens belong to this Australian species.

Amaroucium distomoides, Herdman (?).

I refer one large colony and a few small fragments in the Scotia collection to this
Australian species. J The original specimen came from Port Jackson; the Scotia

* In the new edition of the " Tunicata" of BRONX'S Tier-Reichs.

+ See HERDMAN, Descriptive Catalogue of the Tunicata oftlx Australian Museum, Sydney, N.S. W., 1899, p. 81. On
the plate (Pel. I. figs. 9-12) it is referred to as "Polyclinum depretsum."
| See HERDMAN, ibid,, p. 75.

(ROY. soc. KDIS. TRANS., VOL. XLVIII., 317.)

100 PROFESSOR W. A. HERDMAN ON THE

material is from Port William, Station 349, Falkland Islands, February G to 8, 1903,
6 fathoms. The large colony measures about 14 cm. by 3 cm., and is attached along
the length of a Laminaria-like Alga. The test is dark greyish brown, and the small
yellow Ascidiozooids show all over the surface as closely placed dots or streaks of a
lighter colour. In further details this specimen agrees well with the description of the
Australian one. One zooid was, however, noticed with an eight-lobed branchial aperture.
The stomach has longitudinal folds. The stigmata are large. The dark colour of the
test is due to dense crowding with small test-cells. The Scotia colony was evidently
taken at the reproductive season, as it contains abundance of embryos in various stages
of development up to the tailed larval stage ready to be set free.

Amaroucium sp. (?).

Some small colonies, a few millimetres to about 1 centimetre across, which were found
attached to groups of Styela paessleri and other Ascidians from the Falkland Islands,
belong to the genus Amaroucium, but may be only young colonies of some larger form
such as A. distomoides, or A. pallidulum obtained by the Challenger Expedition at
Port William.

It may be remarked in regard to the three last species of Compound Ascidians that
they require re-examination in the living state. Many of the Compound Ascidians
are scarcely determinable from the contracted and bleached specimens in preserved
collections. It may well be that one or other of the above Polyclinids had in
the living state a bright colour or some other characteristic appearance that is now
wholly lost.

THALIACEA.

(MS. received March 13, 1912.)

Family SALPID^;.

The very large collections of Thaliacea, which were obtained at the South Orkneys
and other Antarctic localities (some from under the ice), were found on examination
to belong entirely to the genus Salpa and to represent two species only ; and in fact
all the specimens, except a single one, are different conditions and sizes of the common
and widely distributed species, Salpa runcinata-fusiformis.

Salpa runcinata-fusiformis, Chamisso-Cuvier.

Station 432, surface, March 30, 1904 ; temp. 31'8°. Nearly one hundred specimens,
from 3 cm. to 6 cm. in length, all of the aggregated form, and many of the larger
ones showing echinated ridges on the test. Most of them showed embryos projecting
into the peri-branchial cavity, one in each.

Station 427, from coarse tow-net, March 26, 1904. About one hundred specimens,
from 3 cm. to 5 cm. in length. In other respects they resemble those from the last

(ROY. soc. EDIN. TRANS., VOL. XLVIII., 318.)

TUNICATA OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 101

locality, except that the nuclear mass has more of a canary-yellow tint in these, and
was a pinker colour in the others.

Station 325, trap-hole, surface, June 4, 1903, Scotia Bay, South Orkneys. A
dozen specimens, all rather large, up to 6 cm., and having some red pigment on
the nucleus.

Station 416, taken by trawl lowered to 2370 fathoms, but probably captured at
surface, March 17, 1904, lat. 71° 22' S., long. 18° 15' W. About forty specimens,
much smaller, up to 4 cm. in length at most, all with a pale yellow-coloured nucleus.

Station 422, vertical net, surface to 800 fathoms, March 23, 1904 ; temp. 31'1° ; lat.
68° 32' S., long. 12° 49' W. This jar contains a large matted mass of Salpse, Medusae,
Macrurous Crustacea, and small fish, also a large species of Sagitta. It looks as if it
had at one time become dry. The Salpse seem to be all of the aggregated form of this
species, and are of medium size.

Station 391, "Tunicate," water-bottle on sounding wire (depth of sounding, 2630
fathoms), February 27, 1904. A single, very large specimen of the solitary form,
fully 8 cm. in length, but rather damaged, with a chain measuring at least 4 cm.

Surface otter trawl, February 24, 1904. One specimen, aggregated form.

Station 325, " Doliolum," trap-hole, surface, June 1903, Scotia Bay, South Orkneys.
About twenty specimens of the chain form.

" Doliola ? ," while sounding, March 22, 1904. Three specimens.

Station 325, surface, June 1903, temp. 28'9°, Scotia Bay, South Orkneys. About
ten specimens.

Station 325, surface, June 1903, temp. 29°, Scotia Bay, South Orkneys. Three
specimens.

Station 430, vertical net, March 28, 1904, temp. 31 '0° F. About fifty specimens,
badly preserved, along with some Medusae.

Station 432A, surface, March 30, 1904, temp. 3T80. Haifa dozen specimens.

Station 325, surface, June 1903, temp. 29°, Scotia Bay, South Orkneys. One
damaged specimen.

Station 409, "Tunicate," vertical net, fathoms 0-100, March 5, 1904, temp. 30°.
One damaged specimen.

Station 391, evening, while sounding, February 27, 1904. One damaged specimen.

Salpa scutigera-confederata, Cuv.-Forsk.

Station 535, on Gulf weed, surface, June 27, 1904, "Doliolum." One badly preserved
specimen, which had probably been dried at one period, solitary form of the species.

(ROY. soc. EDIN. TRANS., VOL. ILVIII., 319.)

102 THE TUNICATA OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

ADDENDUM.

As this paper was going to press I received from Dr BKUUE a couple of tubes con-
taining the following :—

Polyzoa pictonis, subspecies patagonica, Michn. One poor specimen from shore,
Port Stanley, Falklands, January 1903. Station 118.

Lissamaroucium magnum, Sluiter. One colony about 3'5 cm. in diameter, trawled
from Station 346, 56 fathoms, December 1, 1903, Burdwood Bank.

Amaroucium sp. (?). One colony about 3 cm. in diameter from same haul as the
last species. Station 346.

EXPLANATION OF THE PLATE.

Fig. 1. Styela lactea, Herdman. Large barrel-shaped specimen. Nat. size.

Fig. 2. Part of large colony of Holozoa cylindrica, Lesson, with three small specimens of Styela lactea
attached. Nat. size.

Fig. 3. Globular specimen of Styela lactea, showing the positions of the branchial and atrial apertures.
Nat. size.

Fig. 4. Anterior end of similar specimen, showing the branchial aperture. Nat. size.

Fig. 5. One of the echinated spines of Styela lactea. Enlarged.

Fig. 6. Small, globular and very spiny specimen of Styela lactea. Nat. size.

Fig. 7. Alimentary canal of Styela lactea. Slightly enlarged.

Fig. 8. Dorsal tubercle and tentacles of Styela lactea. x 40.

Fig. 9. Dorsal tubercle of Paramolijula grer/aria, Lesson. Enlarged.

Fig. 10. Paramolgula horrida, Herdman, right side. Nat. size.

Fig. 11. Part of branchial sac of P. horrida. x 40.

Fig. 12. Diagrammatic plan of both sides of branchial sac of Styela paessleri, Michaelsen, supposed to
be cut through the endostyle and spread out : I. to IV., branchial folds. The number of bars on the folds and
in the interspaces is shown.

Fig. 1 3. Small part of branchial sac of Styela paessleri. x 40.

Fig. 14. Dorsal tubercle of Styela paessleri. x 40.

Fig. 15. Fungulus antarcticus, n.sp., from the left side. Nat. size.

Fig. 16. Branchial aperture of the same. Enlarged.

Fig. 17. Part of branchial sac of same species, from the inside. x 40.

Fig. 18. Dissection of Fungulus antarcticus, to show alimentary canal and gonads. A little enlarged.

(ROY. soc. EDIN. TRANS., VOL. XLVIII., 320.)

SCOT. NAT. ANT. EXP.

HKKDMAN : "ScoriA" TUNK ATA

Vol. IV.

At*

D.L.

H A H. del.

M-raiUne t EnUnc, Liti... E<iin.

PART VIII.
FISHES.

SCOTTISH NATIONAL ANTARCTIC EXPEDITION.
VIII -FRESHWATER FISHES FROM BUENOS AIRES.

BY R. S. CLARK, M.A., B.Sc.,

Scottish Oceanographical Laboratory, Edinburgh.
(WITH ONE PLATE.)

Scottish National Antarctic Expedition : Notes on some Fresh-water Fishes
from Buenos Aires. By R. S. Clark, M.A., B.Sc.. Scottish Oceanographical
Laboratory. Edinburgh.

(Handed in July 2, 1912. Issued separately August 12, 1912.)

The six fishes here described were brought home by Dr W. S. BRUCE in 1904.
They were taken, with one exception, from the Government Dry Dock at Buenos
Aires in January 1904, while the Scotia was undergoing repairs there, immediately
after her return from her first voyage to the Antarctic. All six have been previously
described, though one — Plecostomus laplatx, Eigenmann — only as late as 1907, three
years after the capture of the two Scotia specimens.

The system adopted in describing the species is that outlined by Dr GUNTHER in
the British Museum Catalogue ; ihefundamentum divisionis being the body-length —
tip of snout to root of middle caudal rays. On account of the considerable variations
shown by each species at different stages of its growth, the work of identification has
been rendered extremely difficult. In this connection, I tender my grateful thanks to
Mr C. TATE REGAN (British Museum) for his invaluable assistance.

The result of the following investigations has been to corroborate substantially the
statement put forward by EIGENMANN in Proc. Wash. Acad. Sciences, to the effect
that the fresh-water fish fauna of Buenos Aires is essentially Amazonian.

The species are : —

SILURID.E.

Pimelodus nigribarbis Boulenger.
Pimelodus albicans Cuv. and Val.

LORICARIID^E.
Plecostomus laplata? Eigenmann.

CH A RACID.E.
Prochilodus argenteus Agassiz.

Micropogon undulatus Linnaeus.

GYMNOTID.fi.

Rhamphichthys blochii Bloch and Schneider.

108 MR E. S. CLARK ON

Pimelodus nigribarbis Boulenger (PI. fig. 3).
1 Pimelodus valenciennis Lutken.

D. I, 6. A. 17. V. 6. P. I, 8.

This specimen is undoubtedly similar to BOULKNGER'S two type specimens from the
Oamapuam Eiver, and described by him in Ann. Mag. Nat. Hist., iv., 6th ser., 1889.
It differs from Pimelodus valenciennis, Lutken (Occ. Papers Cal. Acad. Sci., i.,
1890), in the lengths of anal and adipose dorsal fins. In valenciennis these are equal,
but in the Scotia specimen the adipose dorsal is five-sixths of the anal.

Length of specimen, 167 mms.

Pimelodus albicans Cuv. and Val.

Anus albidm, Val., Voy. d'Orb., 1847.

„ albicans, Guv. and Val., Hist. Nat. Poins., xv., 1840 (Buenos Aires).
Piramutana albicans, Giinther, Ann. Mag. Nat. Hist., 1880, vi., 5th ser. (Rio Plata).
Arius moroti, Val., Voy. d'Orb., vi., 1847 (La Plata).
Piramutana blofhii, Cuv. and Val.

Pimelodns albicans, Eigenmann and Eigenmann, Proc. Cal. Acad., i., 1890 (Rio Plata).
,, ,, Eigeumann, Proc. Wash. Acad. Sciences, March 4, 1907.

D. I, 6. P. I, 10. V. 6. A. 11.

EIGENMANN'S description in Proc. Cal. Acad., 1890, adequately covers the
characteristics of the Scotia specimen. There are no pterygoid teeth, and those on
the palate are reduced to two small separate patches on the vomer. The maxillary
barbel reaches the root of the caudal fin, " as in young specimens 6 to 8 inches long,
though it reaches only to the anal fin in specimens 10 inches long" (GUNTHER).

Length of specimen, 235 mms.

Plecostomus laplatse Eigenmann (PI. figs. 1 and la).
Plecostomus laplatse, Eigenmann, Proc. Wash. Acad. Sciences, March 4, 1907,

D. I, 7. A. I, 14.

The two specimens are certainly EIGENMANN'S laplatte, described by him in 1907
as a new species. The number of scutes in a longitudinal series is 31 + 1 caudal scute.
Smooth patches between pectoral and ventral are well marked. Head and abdomen
are covered with small dark spots, while anterior part of body has larger spots. Fins,
except caudal, spotted. Alternating pale and dark longitudinal bands on body.

Two specimens. Lengths, 283 and 420 mms.

Prochilodus argenteus Agassiz.

P. argenteus, Cuv. and Val., xxii. (Rio San Francisco, Brazil).
„ Giinther, B.M. Cat., v. (Brazil).

n_ i A

D. 11 (12). A. 10, Lat. line 45, lat. trans v. —.

Six excellent specimens were brought back by the Scotia. They average in length

SOME FRESH-WATER FISHES FROM BUENOS AIRES. 109

39 cms. The scales are iridescent, greenish above, and yellow below the lateral line,
while each bears a distinct cross-mark on its surface.

Micropogon undulatus (Linnaeus).

Perca undulata, Linnaeus, Syst. Nat., 12th ed., 1766.

Micropogon undulatus, Guuther, Chall. Rep. on Shore Fishes (Monte Video).

„ Giinther, Ann. Mag. Nat. Hist., 5th ser., 1880 (Rio de la Plata).

Gunther, B.M. Cat., ii p. 211.
,, „ Guv. and Val. (New Orleans).

,, „ Perugia, Ann. Mus. Civ. Stor. Nat. Geneva, x., 1890 (Monte Video).

„ ,, Berg, Anal. Mus. Nac. Buenos Aires, iv., 2nd ser., 1895 (Bahia

Blanco, Mar de Plata, Monte Video, Maldonado).

„ „ Evermann and Kendal, Proc. U.S. Nat. Mus., xxxi., 1907 (Argentine).

,, ,, Berg, Ann. Mus. Nac. Buenos Aires, iv., 2nd ser., i., 1895 (Emboco-

dura del Rio de la Plata, Monte Video).

7

D. X. I, 28. A. II. 8. Lat, line 52 + 3 caudal. Lat. transv. — .

15

One specimen was caught on the line in Rio de la Plata, while the other was taken
from the Government Dry Dock at Buenos Aires. The respective lengths are 27 and
31 cms. The comparative measurements of the Scotia specimens agree with those
given by EVERMANN and KENDAL in Proc. U.S. Nat. Mus., xxxi., where the specimens
are of the same size. The colour is silvery . There are dark grey spots on back, while
below the lateral line these have a tendency towards stria ticm. . ..

Rhamphichthys blochii (Bl. and Schn.) (PI. fig. 2).

Gymnotus rostratus, Bl. and Schn., 522, t. 106.
Rhamphichthys blochii, Kaup, Apod., p. 133, fig. 9.

„ rostratus, Mull, and Trosch.

Carapus rostratus, Cuv., ii. p. 357.
Rhamphichthys blochii, Gunther, B.M. Cat., viii. 5, 1860 (Para).

KAUP attributes the original of this species to BLOCK, but comments on the imper-
fection of the description. The number of anal rays is 365, as in the Scotia specimen,
though BLOCH gave 292. Anus accompanied by a genital papilla which lies behind
the vertical line through the posterior margin of the eye. Snout tubiform and slightly
more than half the length of the head. Small round spots on the head and body.
Irregular black lines and spots on the fins. Between the anal fin and the lateral line
runs an interrupted series of oblique bars.

Length, 296 mms.

EXPLANATION OF PLATE.

Fig. 1. Plecostomus laplatx, side view, reduced (about f ).
„ 10. „ „ ventral aspect, reduced (about f ).

,, 2. Rhamphichthys blochii, side view, reduced (about £).
,, 3. Pimelodus nigrilarbis, side view, reduced (about f ).

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PART IX.
SEALS.

SCOTTISH NATIONAL ANTARCTIC EXPEDITION,
IX.-ANATOMY OF THE WEDDELL SEAL

(LEPTONYCHOTES WEDDELL1}.
(4) THE BRAIN.

By DAVID HEPBURN, M.D., C.M.,

Professor of Anatomy, University College, Cardiff (University of Wales).
(WITH ONE PLATE.)

Scottish National Antarctic Expedition : Observations on the Anatomy of the
Weddell Seal (Leptonychotes Weddelli). Part IV. : The Brain. By David
Hepburn, M.D., C.M., Professor of Anatomy, University College, Cardiff (University
of Wales). (With One Plate.)

(MS. received June 18, 1912. Read December 2, 1912. Issued separately February 8, 1913.)

The material placed at my disposal for the purposes of this paper comprised the brains
of four adult specimens of the Weddell seal, in addition to the brain of the young
animal which has formed the subject of my former contributions.* The four adult
brains having been removed at the time the animals were killed, and preserved in a
solution composed of spirit (90 per cent.) 6 pints and formal (2 per cent.) 4 pints,
were, with one exception, in a firm and satisfactory condition for detailed anatomical
examination. The body of the young seal had been preserved with a view to ordinary
dissection, and therefore its brain was not in the firm state of the adult specimens ; but
as I had the opportunity of removing this brain from the skull, I was able to observe
the disposition of the dura mater to the hemispheres of the cerebrum and cerebellum.
While the dura mater presented, as a whole, its usual arrangements, it was noteworthy
that the falx cerebri did not act as a septum between the two hemispheres of the
cerebrum except to a very slight extent, and certainly for not more than one-third of
the distance between the vertex of the cerebrum and the dorsal surface of the corpus
callosum. As a result, in the region referred to the opposing mesial surfaces of the
two hemispheres lay not only in close apposition with each other, but their convolutions
were intimately adapted to each other. Similarly, the tentorium cerebelli only extended
a short distance between the cerebrum and the cerebellum, and, as the occipital ends of
the cerebral hemispheres fell considerably apart from each other, there was space for
the accommodation of the well-developed vermis of the cerebellum as well as for the
bulbous pineal body, which occupied a position upon its dorsal aspect. As I removed
the brain from the skull the stalk of the pineal body gave way, and probably the same
thing had occurred during the removal of the adult brains, for, while different lengths
of the stalks had been preserved, there was only one complete specimen of its bulbous
extremity. Looked at from the vertex, the general outline of the whole brain was that
of a four-sided figure with rounded angles, and the cerebral hemispheres concealed the
cerebellum except where the vermis was exposed between them at their occipital ends.
The frontal ends of the hemispheres were not rounded into frontal poles ; but, on the
contrary, they almost formed flat frontal surfaces. Similarly, the occipital ends were
rounded and not pointed to form occipital poles. There was a small amount of differ-
ence in the absolute size of the adult brains, and the largest specimen measured 120 mm.

* Part I., Trans. Boy, Soe. Edin., voL xlvii. p. 57, 1909. Part II., Tram. Roy. Soc. Edin., vol. xlviii. p. 191, 1912.
Part III., Trans. Pay. Soc. Edin., vol. xlviii. p. 321, 1912.

(REPRINTED FROM THE TRANSACTIONS OF THE ROYAL SOCIETY OF EDINBURGH, VOL. XLVHI., PP. 827-847.)

116 PROFESSOR DAVID HEPBURN ON

in its fronto-occipital diameter ; 115 mm. in its greatest transverse diameter at a point
well forward on the temporo-sphenoidal lobes ; and 71 mm. in vertical height, measured
from the pons varolii to the vertex of the cerebrum. . Thus, apart from the peculiarity
of its general outline in total size, it was only slightly less than an average human brain.
Throughout the anterior two-thirds of their extent the cerebral hemispheres were, as
already indicated, in very close apposition, and the falx cerebri only dipped into the
pallial or superior longitudinal fissure to a slight extent ; but in its posterior third this
cleft opened to form a wide interval, measuring 65 mm. in the transverse direction at
its hinder end and narrowing as it ran forwards towards the posterior end of the corpus
callosum. In the deep level of this interval the pineal body and the upper surface of
the vermis were visible, as well as part of the upper surface of the cerebellar hemispheres.
It should be stated that the backward extension of the occipital lobes of the cerebrum
carried them 2 mm. beyond the cerebellar hemispheres.

In its essential features the basal aspect of the brain conformed to current
descriptions of the mammalian brain ; but it presented many special points of interest, to
which reference will be made in the course of my survey.

I. CEREBRAL CONVOLUTIONS AND FISSURES.

Regarded as a whole, the cerebral convolutions (gyri) were large and well defined
from each other by deep, well-marked fissures (sulci), and yet many furrows not deep
enough to be regarded as sulci were seen crossing the surfaces of convolutions. In-
variably these shallow furrows were in the position of blood-vessels ramifying in the pia
mater, and it was clearly demonstrable that the furrows were produced by the blood-
vessels. In appearance they resembled the arterial grooves upon bony surfaces, and
their presence upon the surface of the brain suggested either arterial pulsation or resist-
ance to brain growth as their determining cause. Indeed, from the distinct character of
many of them it would not be difficult to credit these vessels with the possibility of
determining the position of new fissures in a rapidly expanding hemisphere. In their
chief and outstanding characters the two hemispheres corresponded with each other ;
but in the matter of intimate detail they presented a considerable amount of asymmetry,
although neither hemisphere could be said to be more elaborately convoluted than the
other.

The general plan of the convolutions and fissures was not simple or easy to determine.
In fact, the whole arrangement bore very little if any resemblance to that presented by
the brain of a typical member of the carnivora, e.g. the dog ; and this is somewhat
remarkable and unexpected when we remember that the seals are themselves carnivores
notwithstanding their numerous adaptations to an aquatic habitat. Partly for this
reason, and partly because my observations do not altogether harmonise with those of
MURIE* in his description of another seal (Otaria jubata, the sea-lion), nor with those

* MURIE, Trans. Zool. Soc. Land., vol. viii., 1874.

(ROY. 8OC. EDIN. TRANS., VOL. XLVII1., 828.)

THE ANATOMY OF THE WEDDELL SEAL. 117

of Sir WM. TURNER * in his account of the brain of the elephant seal, I propose to deal
at some length with the arrangement of the convolutions and fissures and the possibility
of dividing the cerebral surface into subordinate lobes, after the manner adopted in
describing the human brain.

The complexity of the convolution pattern of the brain of seals has led observers to
devise such an elaborate terminology for the description of the separate convolutions
and fissures that it is a matter of considerable difiiculty to correlate the different terms.
Consequently, bearing in mind the variations which specimens of these brains present
among themselves, as well as their divergence from the ordinary type of carnivore brain,
I have preferred to restrict the use of terms as much as possible, and to limit the attempt
to establish homologies to such characters as were fairly comparable to those presented
by the human brain.

1. The Lateral Surface of the Hemisphere. (Fig. 1.)

On this aspect the convolutions and fissures were well developed both as regards
their size and their numbers, and yet any underlying " pattern " resulting from the
disposition of the primary fissures was most elusive and difficult to decide. Fortunately,
there was no uncertainty with regard to the fissure of Sylvius (sulcus Sylvii). Its
commencement in relation to the locus perforatus anticus on the basal surface of the
brain, and its position between the orbital and temporo-sphenoidal parts of the hemi-
sphere on the same surface, fixed the position of its main stem without any doubt, and
so by its outer end it provided one fixed point from which to unravel the complexity of
the lateral surface. TURNER found this fissure traceable on the lateral aspect of the
hemisphere " upwards and backwards for 32 mm. on the side of the right hemisphere,
but not so far on the left." Nevertheless, for some time I found great difficulty in
deciding which, and how many, of the fissures upon the lateral surface were entitled to
be accepted as its direct continuations, although, as the dissection proceeded, the decision
arrived at in the first instance was verified as correct. My initial difficulty was increased
by the fact that in the lateral view of the hemisphere of the brain of the dog, as may be
seen in the figure given by WIEDERSHEIM and PARKER.^ the fissure of Sylvius is
represented as a " closed " fissure, i.e. one provided with " opercula," forming an " arcuate
gyrus " which surrounds the fissure on all aspects except the basal segment of the
fissure. Further, in the brain of the dog, this " arcuate gyrus " is repeated twice, so
that altogether on the lateral aspect of its hemisphere, to quote WIEDERSHEIM and
PARKER, "In carnivores, cetaceans, and ungulates, three gyri arch over the Sylvian
fissure, one above the other, and are separated by the so-called arcuate fissures." J
Certainly this was not the manner in which the convolutions and fissures were disposed
on the lateral aspect of the hemisphere of the Weddell seal in relation to the fissure of

* TURNER, Gludlenger Reports, vol. xxvi, Zoology : Report on Seals.

t WIEDERSHEIM and PARKER, Comparative Anatomy of Vertebrates, 3rd ed., 1907, p. 224.

I Ibid., p. 228.

(ROY. soc. EDIN. TRANS., VOL. XLVTII., 829.)

118 PBOFESSOR DAVID HEPBURN ON

Sylvius. On the other hand, in MURIE'S * paper already referred to, there is a fairly
close resemblance shown in pi. Ixxviii. fig. 40, between the brain of the sea-lion and
that of the Weddell seal under consideration (fig. l), so far as the general position of
gyri and sulci is concerned ; although, having verified my interpretation of the surface
appearances by dissection of the interior of the hemisphere, my conclusions differ
considerably from those arrived at by MURIE, and so far as carnivora in general are
concerned I am of opinion that at least the Weddell seal presents a very novel arrange-
ment of the fissure of Sylvius, but still one which is quite compatible with, and readily
explainable by, reference to the mode of its development from the embryonic to the adult
condition. As is well known, the Sylvian fissure, in the course of its embryonic
development, results from the more or less close apposition of those portions of the
cerebral cortex which, as derivatives from the orbital, frontal, fron to -parietal, and
temporal portions of the cortex of the hemisphere, and under the term " opercula,"
extend beyond so as to overshadow and gradually conceal from lateral observation that
portion of the cortex called the central lobe or island of Reil, and thus ultimately the
surface of the island of Reil may become completely hidden by convolutions which are
no longer upon the same superficial plane as those of the insula. Further, until these
" opercula " practically come into contact with each other, not only does the insula
remain more or less visible, but the lateral segment of the fissure of Sylvius is
represented by a gap or interval of varying width. Again, if the growth of the insula
kept pace with the growth of the surrounding " opercula," then the insular convolutions
would continue to present themselves upon the same superficial plane as that of the
" opercula," and thus instead of a fissure of Sylvius we should find in its place the sulcus
which limits and marks off the island of Reil from the surrounding cortex, viz. the
limiting sulcus (sulcus insulae). In other words, we should find the island of Reil pre-
senting or protruding between the " opercula " by whose apposition the fissure of Sylvius
derives its lateral characteristics.

In my opinion, that is the interpretation of the condition which is presented by the
brain of the Weddell seal. As a result there appear to be two sulci extending from the
basal stem of the fissure of Sylvius, and between them the greater part of the island of
Reil not only presents itself, but is to a large extent upon the same superficial plane as
that of the surrounding gyri.

The convolutions upon the surface of the insula were irregular, and neither upon
different brains nor upon the two sides of the same brain were they closely repeated ;
but I have given in fig. 1 a drawing of the brain in which they showed a tendency to
radiate from the basal end of the fissure of Sylvius, and I have done so because in the
human brain a radiating arrangement is their normal characteristic. From all this it will
be evident that the fissure of Sylvius as such is not represented on the lateral surface
of the brain of the Weddell seal ; but that in its place there is a vallecula, wide anteriorly
and narrower posteriorly, which is occupied by the convoluted surface of the island of

* MURIE, loc. cit., pi. Ixxviii., fig. 40.
(ROY. soc. EDIN. TRANS., VOL. XLVIII., 830.)

THE ANATOMY OF THE WEDDELL SEAL. 119

Reil, whose boundaries are indicated by the limiting sulcus which almost completely
separates the insula from the surrounding cortex.

In my opinion, this interpretation of the appearances is in conformity with the facts
elucidated by a dissection of the corpus striatum, as well as with the facts of develop-
ment, although I am not aware that it has hitherto been advanced by any of the
observers who have described the brain of the seal. Indeed, in his description of the
brain of the elephant seal, TURNER says : " I can make no definite statement as to the
presence of the island of Reil, unless the concealed part of the anterior limb of the
Sylvian fissure be regarded as representing it." Again, in reference to the brain of the
walrus, the same observer says : " I could not speak with any precision of the island of
Reil, unless the concealed part of the anterior limb of the sylvian convolution passed
deeply into the fissure and was concealed by the anterior limb of the supra- Sylvian
convolution, which for some distance therefore formed the anterior lip of the fissure of
Sylvius." (In the first of these quotations the reference is to the concealed part of the
Sylvian fissure, and in the second to the concealed part of the Sylvian convolution, but
probably this is by inadvertence.)

The Plates which illustrate the papers of MURIE and TURNER, if compared with
fig. 1 of the present communication, will show how much minor variation the brains of
this group of marine mammals may present, while to my own mind they emphasise
the interpretation which I have ventured to put forward. It is difficult to conceive a
brain of the dimensions of those under consideration without an island of Reil ; and as
this part of the convoluted surface of the hemisphere corresponds more or less exactly
to the surface aspect of the corpus striatum, the presence of the latter practically com-
pels us to account for the former.

My next endeavour was to determine which of the sulci could be accepted as the
fissure homologous to the fissure of Rolando (sulcus centralis), because of its importance
as a guide to the position of the sensory-motor area and its value as a dividing line
between the frontal and parietal lobes of the cerebrum. Reference may again be made
to the brain of the dog, in which the sulcus cruciatus is an outstanding feature, and to
WIEDERSHEIM and PARKER'S * description of the fissure, where the following occurs :
" Along the lateral surface of the hemisphere, the cruciate sulcus (the homologue of the
central sulcus or fissure of Rolando of primates) extends upwards to the pallial fissure."
Now, in the Weddell seal the cruciate sulcus is well marked ; but, as may be seen by refer-
ence to figs. 1 and 2, it is situated so far forwards that, if it be accepted as the homologue
of the fissure of Rolando, practically not only is there no frontal lobe remaining, but
the parietal lobe is carried forwards to a position in front of the basal limb of the
fissure of Sylvius, both of which contingencies are so serious as to compel us to doubt
whether the homology be correct in the case of this seal, in view of the importance of
the Rolandic area as a sensory-motor centre. For these reasons, therefore, so far as the
Weddell seal is concerned, I am driven to accept as the fissure of Rolando that fissure

* WIEDERSHEIM and PARKER, lot. cit., p. 228.
(ROY. soc. BDIN. TRANS., VOL. XLVIII., 831.)

120 PROFESSOR DAVID HEPBURN ON

whose lower end will be seen resting upon the fronto-parietal operculum of the insula,
and I have marked it by this name in fig. 1. In this respect my drawing and its
interpretation are more in agreement with MURIE'S * account of the sea-lion, although
in his drawing the fissure of Rolando is represented as much more extensive than it
appears to be in the Weddell seal.

TURNER describes the cruciate fissure of the elephant seal as seen from the front and
not from the norma verticalis, and states that " a large sigmoid gyrus was bent around
its outer end." To some extent this description would apply to the Weddell seal,
although in the latter the cruciate fissure was visible from the norma verticalis, but it
was much more effectively seen from the norma frontalis, while its outer end was
blocked by an arched gyrus (fig. 1).

I could nob find any satisfactory evidence of a homologue for the external parieto-
occipital fissure, and therefore no fixed indication of a limit between the parietal and
occipital lobes of the cerebrum on its lateral aspect, or between the occipital and
tempero-sphenoidal lobes on the same aspect, for the reason that these areas were freely
connected with each other by annectant gyri.

The Convolutions on the Lateral Surface.

The frontal lobe having been delimited in the manner described, its convolutions
resolved themselves into a pre-central (ascending frontal) ; the frontal contribution to
the opercula of the insula ; and two or three short convolutions running forwards from
the pre-central convolutions towards the sulcus cruciatus.

The elongation of these short convolutions in a forward, i.e. frontal, direction would
have the effect of forcing the sulcus cruciatus forwards and downwards towards the roof
of the orbit, and would thus bring the cruciate fissure into position as a kind of
boundary line between the frontal and orbital aspects of the frontal lobe. It appears
to me that the blunt frontal end of the brain of the Weddell seal is due in some
measure to the presence of convolutions, which in the human brain would be found in
relation to the roof of the orbit. Further, in the human brain there may sometimes be
seen a fissure which runs transversely from the pallial fissure across the frontal lobe
and close above the orbital margin of the hemisphere. In my opinion this is a fissure
which may fairly be regarded as corresponding with the sulcus cruciatus.

In the figure given by MURIE, and already referred to several times, there is, on the
frontal side of the fissure which is marked "Rolando," a convolution named in three
places as the antero-parietal convolution (AP) ; and I cannot but think that this was an
unfortunate term to introduce at such a place so long as the fissure of Rolando is
accepted as a boundary line between the frontal and parietal lobes of the highly
elaborated brain of man.

From the fissure of Rolando (fig. l), and beginning at a point about its middle, a
well-marked fissure ran backwards towards the occipital end of the hemisphere. This

* MURIE, loc. cit., pi. Ixxviii., fig. 40.
(BOY. soc. EDIN. TRANS., VOL. XLVIII., 832.)

THE ANATOMY OF THE WEDDELL SEAL. 121

fissure, which was deepest at its ends and shallowest about half way between them,
divided the parietal region of the brain into an upper and a lower lobule, and it might
quite fairly be termed the intra-parietal sulcus. Each of the lobules above and below
the intra-parietal sulcus presented in its turn a short and less defined fissure whose
course was roughly parallel to that of the intra-parietal sulcus, but neither of these
short fissures opened into the fissure of Rolando. Thus the frontal ends of the convolu-
tions both above and below the intra-parietal sulcus were united together, with the
result that the arrangement suggested an interrupted post-central (ascending parietal)
gyrus.

It has already been stated that there was no definite guide which could be selected
as a demarcation between the parietal and occipital lobes, and therefore I can only say
that, as a whole, the convolutions in the occipital region ran from behind forwards, and
more or less parallel to each other, to make connections with the parietal and temporo-
sphenoidal convolutions. One of these connections seems worthy of special notice. It
joined the hinder end of the island of Reil and the hinder end of the temporo-sphenoidal
operculum to one of the occipital convolutions. In this relation it should be
remembered that the Sylvian fossa (which ultimately becomes the posterior limb of
the Sylvian fissure in the primate brain) is shallowest in this region during the process
of its development.

The lateral aspect of the temporo-sphenoidal lobe, which provided one of the opercula
of the island of Reil, was situated below and behind the Sylvian fossa. It presented
two fairly well defined convolutions, an upper and a lower, separated by a definite
sulcus, with irregular sulci of smaller dimensions, suggesting the possibility of further
subdivision.

2. The Mesial Surface of the Hemisphere. (Fig. 2.)

This aspect of the hemisphere presented considerable elaboration and complexity as
regards the structures belonging to the pallium, but in the basal region it was simpler
and more easy of interpretation. As on the lateral surface, the convolutions and fissures
were large and well defined, although the determination of their homologies was a
matter of considerable difficulty.

The corpus callosum measured 5 cms. in length and 4 mm. in vertical depth over
the greater part of its length. The genu was 10 mm. long and 9 mm. in vertical depth,
while the vertical depth of the splenium was 5 mm. From the anterior end of the
genu to the frontal end of the hemisphere the distance was 2 cms., and from the posterior
margin of the splenium to the occipital end of the hemisphere the distance was 4 cms.
Therefore, as a whole, the corpus callosum was situated nearer to the frontal end of the
brain. The rostrum of the corpus callosum was very short, but the lamina terminalis
(lamina cinerea), extending from the rostrum to the optic chiasma, was a well-defined
object.

The callosal sulcus, which separated the dorsal surface of the corpus callosum and

(ROY. soc. EDIN. TRANS., VOL. ZLVIII., 833.)

122 PROFESSOR DAVID HEPBURN ON

the anterior aspect of the genu from the surrounding convoluted surface, commenced
at the locus perforatus anticus, which to a considerable extent encroached upon the
mesial aspect of the hemisphere and presented itself in front of the lamina terminalis
below the genu of the corpus callosum. Several shallow extensions of the callosal
sulcus, in relation to the anterior half of the corpus callosum, ran forwards and upwards
into the superincumbent convolution, thereby complicating the appearance of that gyrus.

The sulcus cruciatus was visible upon this aspect of the frontal lobe, and here it
divided into several branches, of which the hindermost was the longest.

There also appeared on this surface the fissure which I have accepted as the fissure
of Rolando, and it extended from the superior margin of the hemisphere downwards
and backwards to a point almost half way to the dorsal surface of the corpus callosum.

The calloso-marginal sulcus was much interrupted by the invasion of other fissures,
so that it was composed of not only the fissure on the dorsal aspect of the callosal
gyrus, but also of a branch from the cruciate sulcus anteriorly, and a branch from a
fissure situated posterior to the callosal gyrus (fig. 2).

The mesial aspect of the occipital lobe was reduced to comparatively small dimen-
sions in comparison with the size of the hemisphere, a condition which resulted from
the fact that occipital structures, which in a human brain of corresponding magnitude
would have been visible on its mesial face, were in this seal's brain turned to the
inferior or cerebellar aspect of the occipital lobe. For this reason there was very
considerable difficulty in selecting a fissure which could be regarded as homologous
with the internal parieto-occipital sulcus. As the result of a later dissection, which
determined the position of the calcarine fissure, I concluded that the fissure which is
immediately posterior to the callosal gyrus, and whose course is upwards and forwards
towards the supero-mesial border of the hemisphere (fig. 2), should be regarded as the
internal parieto-occipital sulcus. Apparently this is the splenial fissure of some authors.

The callosal gyrus started by rising gradually from the locus perforatus anticus
immediately below the genu of the corpus callosum. It ran forwards, and growing
larger as it proceeded it wound round the anterior end of the genu, forming several
well-marked folds situated between the callosal and cruciate sulci. Thereafter it passed
backwards in a straighter or less elaborate form above the posterior two-thirds of the
corpus callosum and between the callosal and calloso-marginal sulci. Posterior to the
splenium it turned abruptly towards the basal aspect of the hemisphere, constituting
what is known in human anatomy as the isthmus of the limbic lobe. Subsequently
(fig. 3) it curved along the infero-lateral aspect as the hippocampal gyrus, which
steadily expanded as it proceeded forwards to terminate in a wide flattened extremity
situated close behind the locus perforatus anticus, but separated from it by the basal
segment of the Sylvian fissure. In a later stage of the dissection the uncus was found
in connection with the hippocampal gyrus.

So far, therefore, as the essential elements which enter into its formation are
concerned, the limbic lobe in all its parts was fully represented ; and only at its frontal

(ROY. BOC. EDIN. TRANS., VOL. XLVIII., 834.)

THE ANATOMY OF THE WEDDELL SEAL. 123

end, where several definite sinuosities appeared, and at the widely expanded end of
the hippocampal gyrus were • there any marked deviations from the much simpler
appearances presented by the limbic lobe of the human brain.

3. The Inferior or Basal Aspect of the Hemisphere.

As may be seen by reference to fig. 3, the general appearance and the interpretation
of this surface were relatively simple in comparison with the other surfaces, except in
the occipital region, where again there was considerable complexity due to the fact that
so much more of the convoluted surface of the occipital lobe was directed towards the
tentorium cerebelli than towards the falx cerebri. with the result that objects which
appear on the mesial aspect in the primate brain were found upon the tentorial aspect
in that of the seal.

In the mesial plane the two hemispheres were divided from each other in the
frontal region by the pallial fissure as far back as the lamina terminalis, below and
behind which the optic chiasma was situated. The inter-peduncular space presented
the usual boundaries, viz. : anteriorly, the optic chiasma ; antero-laterally, the optic
tracts ; postero-laterally, the crura cerebri ; posteriorly, the pons Varolii. The structures
forming the floor of the space were the tuber cinereum, provided with a short
infundibulum to which the hypophysis cerebri was attached, this latter being a large
object in proportion to the size of other structures ; the corpora mammillaria ; and the
locus perforatus posticus. The occulo-motor nerves emerged from the mesial aspect
of the crura cerebri.

The basal surface of the frontal lobe was clearly defined posteriorly by the fissure
of Sylvius and the locus perforatus anticus. This surface presented the following
fissures : — the olfactory sulcus, which was occupied by the olfactory tract (fig. 3),
pursued a straight course from the locus perforatus anticus forwards towards, but not
quite up to, the orbital margin ; the rhinal sulcus commenced a short distance in
front of the Sylvian fissure and ran forwards in a curved manner, following the lateral
contour of the orbital surface, but separated from the margin by a convolution, then,
winding round the anterior end of the olfactory sulcus, it turned backwards between
the olfactory and pallial sulci, and terminated as a shallow groove upon the gyrus
rectus.

The convolutions on the orbital surface were the following : — the gyrus rectus,
situated between the olfactory sulcus and the mesial orbital border ; the posterior
orbital gyrus, forming the anterior boundary of the Sylvian fissure and the orbital
operculum of the island of Reil ; a triangular gyrus, occupying the space between
the olfactory and rhinal sulci ; and a long curved gyrus, situated between the rhinal
sulcus and the lateral margin of the orbital surface. The triangular and curved gyri
were both connected with the posterior orbital gyrus behind and with the gyrus rectus
in front, but otherwise they were separated throughout their length by the rhinal

(ROY. 80C. EDIN. TRANS., VOL. XLVIII., 835.)

124 PROFESSOR DAVID HEPBURN ON

sulcus. The general arrangement of their surface suggested the possibility of the
rhinal fissure being the foundation for the more elaborate fissures which characterise
the orbital surface of the higher brains.

The olfactory tract presented two distinct roots, separated from each other by a
large area of the locus perforatus anticus. Of these, the mesial root came into view
from the mesial surface in relation to the anterior end of the callosal gyrus of the
limbic lobe. The lateral root emerged from under cover of the expanded end of
the hippocampal gyrus. Closely adhering to the locus perforatus anticus, these
roots converged and fused to form the olfactory tract, which occupied and moulded
itself to the olfactory sulcus.

In all my adult specimens the olfactory bulb had been broken off, so that I am
not able to state its size, frontal relations, etc. ; but it was present in the young specimen
as an ovoid enlargement 17 mm. in length and 6 mm. in breadth. It turned upwards
upon the frontal surface of the hemisphere, to which it was closely applied.

Behind the fissure of Sylvius, the basal surfaces of the occipito-temporal lobes were
much more expanded in the lateral direction than is the case in the primate brain ;
and, as a consequence, convolutions and sulci which are not found on this aspect in
the human brain were visible in the brain under consideration. At the same time,
it presented sulci which do not occur in a human brain, and therefore it is not easy
to suggest a nomenclature for some of these sulci, nor to be quite certain that they
should be accepted as providing boundaries between the occipital and temporal sections
of the surface.

The dentate and collateral sulci, situated respectively on the mesial and lateral
aspects of the hippocampal gyrus, were distinctly indicated. Towards the hinder end
the collateral sulcus was interrupted by a bridging gyrus, behind which the sulcus
corresponded to the general position of the eminentia collateralis in the lateral
ventricle, as was afterwards revealed by dissection. Further, with the same part of
the sulcus, i.e. posterior to the annectant gyrus, just mentioned, other two well-marked
sulci communicated. Of these, one was directed backwards towards the occipital end
of the hemisphere, and the other diverged backwards and outwards towards the
infero-lateral margin of the hemisphere in its occipital area. Thus a large segment
of the occipito-temporal surface, situated between the collateral sulcus and the infero-
lateral margin of the hemisphere, was divided into three wedge-shaped gyri whose
bases were directed towards the infero-lateral margin and whose apices converged
towards the annectant gyrus above referred to. Indeed, this annectant gyrus connected
the anterior and the middle of the three wedge-shaped gyri with the middle portion
of the hippocampal gyrus. The posterior one of these three wedges presented a free
apex, but the surfaces of each of the three showed indications of further subdivision
by additional sulci.

The callosal and hippocampal gyri were united to each other by a narrow gyrus
which curved round behind the splenium of the corpus callosum and the crura cerebri.

(ROY. soc. EDIN. TRANS., VOL. XLVIII., 836.)

THE ANATOMY OF THE WEDDELL SEAL. 125

In the human brain this gjrus is named the isthmus of the limbic lobe, and I have used
the same term for its description in this account of the brain of the Weddell seal.

Posterior to the isthmus, a distinct deep fissure entered this region, i.e. the
basal aspect of the occipital end, as the continuation of a fissure well defined upon
the mesial aspect of the hemisphere. Upon the basal aspect it was cut off from
junction with the hinder end of the collateral sulcus by an annectant gyrus, whereupon
it turned abruptly backwards towards the occipital end of the hemisphere (fig. 3). It
appears to me that that part of the fissure immediately behind the isthmus should
be regarded as the continuation of the internal parieto-occipital sulcus (fig. 2), and
that its extension towards the occipital end of the hemisphere is the calcarine sulcus
(by some observers called the post-horizontal fissure). My reasons for this view will
be further elaborated in connection with the description of the posterior cornu of the
lateral ventricle, but meantime I may state that the calcar avis or hippocampus
minor was closely related to the position of the deep anterior end of what I have
named the calcarine sulcus. Resulting from appearances verified by dissection as
well as by transverse section of the posterior cornu of the lateral ventricle (fig. 4), I
feel warranted in concluding that the narrow gyrus which is situated on the lateral
aspect of the calcarine fissure and connected with the hippocampal gyrus must be
regarded as the lingual gyrus, while the larger gyrus situated on the mesial aspect
of the calcarine fissure and posterior to the internal parieto-occipital fissure (fig. 3)
must be regarded as the foundation for the cuneate lobule, which is found in a
corresponding position on the mesial surface of the human cerebrum.

Although the foregoing account shows that there was great deviation from the
convolution pattern characteristic of a typical carnivore brain on the one hand, and by
the human cerebrum on the other, yet the internal appearances exposed by dissection
underwent an entire change and became simplified to a remarkable degree. So much
was this the case that, in consideration of its size and with certain points of exception as
to the details, the various objects were as readily recognised as they are in a human brain.

II. DISSECTION OF THE CEREBRUM.

The method of procedure followed was that adopted in the dissection of the human
brain.

In the first place, the hemisphere was divided by a horizontal transverse section at
about 4 mm. distance above the mesial free surface of the corpus callosum, in order to
expose the white core or centrum ovale minus, which, considering the total size of the
hemisphere, was smaller than one expected. The reduction in the size of the central
white core could be explained by the depth of the sulci. Many of the sulci at the
frontal end were 2 cms. in depth, and at the occipital end some were 2 '3 cms. deep. As,
of course, all the sulci were bounded by a zone of grey matter, the general effect was a
reduction in the apparent size of the central white core.

(ROT. soc. EDIN. TRANS., VOL. XLVIII., 837.)

126 PROFESSOR DAVID HEPBURN ON

Another unexpected result was, that at the level mentioned, viz. about 4 mm.
above the corpus callosum, the section opened into the cavity of the lateral ventricle,
which therefore rose to a higher level than the mesial surface of the corpus callosum,
and consequently there must be a corresponding deviation from the horizontal direction
of those fibres of the corpus callosum which form the roof structures in relation to the
body of the cavity of the lateral ventricle. This upward extension of the ventricle,
taken in conjunction with the large size of the convolutions as indicated by the depth of
the fissures, shows that notwithstanding its superficial dimensions the brain of the seal
falls considerably short of a human brain of similar size as regards the amount of grey
and white matter.

In addition to what has already been stated with regard to the corpus callosum, the
following additional facts may be noted. The mesial faces of the two hemispheres were
so closely in apposition that opposing gyri practically interlocked with each other, and
therefore the dorsal surface of corpus callosum was entirely concealed. When this
surface was exposed it showed feeble striae longitudinales mediates and still feebler
laterales. The cingulum was present, but much smaller than the size of the surrounding
gyri led me to anticipate. The forceps major and/creeps minor were easily dissected
and were of characteristic appearance.

On removing the roof of the lateral ventricle and of its cornua I was impressed by
the apparent simplicity of the basal ganglia, which were large, and at the first glance
suggested strongly such appearances as one is familiar with in the human brain. Taking
into consideration the somewhat elaborate and intricate condition of the convolutions
of the pallium, the simple nature of basal objects was remarkable. The anterior or
frontal cornu of the lateral ventricle was very shallow. Its course was outwards and
forwards into the substance of the frontal lobe, where it terminated in a blind recess.
Its relations to the septum lucidum and to the caput of the nucleus caudatus were
similar to the arrangements seen in the human brain.

The middle or descending cornu likewise followed the human plan in its chief
features and direction. On its floor there were the choroid plexus, theflmbria hippo-
campi, and the hippocampus major terminating in the pes hippocampi. The choroid
plexus was continuous with the pia mater of the dentate sulcus, and thus, as in man, the
termination of this cornu was situated on the lateral aspect of the crus cerebri and closed
by the ependyma ventriculorum. The choroid plexus, however, was wider than in man,
and spread itself out so as to form a vascular sheet which separated the objects in the
roof of this cornu from the other structures on its floor. Further, the hippocampus
major and the fimbria, with the overlying choroid plexus, were pressed upwards against
the roof of the cornu, where they adapted themselves to a deep furrow which was
bounded mesially by the optic tract and laterally by the tail of the caudate nucleus.
Again, on its convex margin the hippocampus major was separated from the floor of
the cornu "by a deep fissure which almost completely detached this object from the floor
of the cornu. Indeed, the connection between the hippocampus major and the floor of

(ROY. soc. EDIN. TRANS., VOL. XLVIII., 838.)

THE ANATOMY OF THE WEDDELL SEAL. 127

the cornu was reduced to a narrow band in close relation to the concave margin of the
hippocampus. Consequently, in the brain of this seal the hippocampus major could not
be described as the reverse or ventricular surface of the sulcus hippocampi.

Again, the pea hippocampi terminated as a rounded end, only slightly wider than
the general body of the object and not expanded or notched as in man.

Th&Jimbria hippocampi occupied the concavity of the hippocampus major, but it only
spread over the surface of the hippocampus major for about a fourth of the width of the
latter. Both the concave and convex margins of the fimbria were free, so that it only
adhered to the surface of the hippocampus to a slight extent. So far as could be seen,
the fimbria became continuous with the lower end of the gyrus dentatus and the adjacent
part of the gyrus hippocampi close to the uncus.

The posterior cornu was not narrow and pointed towards its occipital end as in
man. Indeed, it appeared more like a wide backward extension of the middle cornu,
for at its commencement it was 2 cms. wide, and at this place the eminentia collaterals
appeared as a large well-defined elevation indented anteriorly by the convex face of
the hippocampus major, but these two objects were separated from each other by the
upward extension of the fissure already referred to on the floor of the middle cornu.
On the mesial aspect of the cornu, and above the eminentia collateralis, there were two
strongly defined convex ridges, the one above the other. Both of these ridges appeared
from under cover of the hinder end of the corpus callosum, with which they were con-
tinuous. The lower of the two was directed outwards and backwards. It descended
to the floor of the cornu, and ceased to be an elevated object immediately behind the
eminentia collateralis. As already indicated in an earlier part of nay description, this
elevation corresponded to the general position of the calcarine fissure on the inferior
aspect of the occipital lobe of the cerebrum, and for that reason I have regarded the
elevation just described as the calcar avis or hippocampus minor. The upper of the
two elevated ridges seen in the posterior cornu was the larger at its commencement,
but it narrowed down rapidly, and disappeared on the floor of the cornu behind the
calcar avis. This object may be taken as the bulb of the posterior cornu. Fig. 4 .
shows these two structures in relation to the calcarine fissure, and it will be observed
that the bulb of the cornu has a more direct relation to the calcarine fissure than the
calcar avis has. The posterior cornu extended backwards for a distance of 2 cms., and
terminated in a blind rounded extremity which, from the size of the eminentia collater-
alis, appeared to dip downwards. Certainly it showed no tendency to bend towards
the mesial surface of the occipital lobe.

The body of the lateral ventricle was roofed over, as already stated, by the tapetal
fibres of the corpus callosum. On its floor the following structures were noted :—

Anteriorly the nucleus caudatus, which was particularly well shaped ; to the mesial
side of its tailed part, there was the choroid plexus of the velum interpositum, and this
choroid plexus was spread out sufficiently to entirely conceal the tsenia semicircularis ;
behind the choroid plexus there lay the widely expanded lateral half of the body of

(ROY. 30C. KDIN. TRANS., VOL. XLVIII., 839.)

128 PROFESSOR DAVID HEPBURN ON

the fornix, which, although attached to the under side of the corpus callosum in the
mesial plane, nevertheless was spread outwards as far as the tsenia semicircularis, thus
forming a complete layer above the velum interpositum and the optic thalamus, of
which, indeed, no part was visible until the fornix was removed.

At the hinder and outer end of the optic thalamus, the fornix was raised from
below by a subjacent object so that it appeared as if the fornix itself contained a
rounded mass of material in the position just stated. However, this underlying rounded
projection was the corpus geniculatum externum of the optic thalamus, which would
have been visible in the floor of the lateral ventricle but for its concealment by the
expanded overlying base of the triangular fornix. The foramen of Monro was clearly
denned and occupied its customary position.

The fornix was remarkably well developed and of large size as compared with that
of man ; but, as in man, its body or central portion was triangular in shape and flattened
from above downwards. By its upper or callosal surface it was attached to the under
surface of the corpus callosum along a narrow mesial line which extended from Verga's
ventricle posteriorly to the septum lucidum in front. Elsewhere the cavity of the
lateral ventricle on each side extended between the corpus callosum and the fornix.
The lateral margins of the fornix were sharply defined and free. The deep surface of
the fornix rested upon the velum interpositum, but no vessels could be seen passing
between the two structures. The two anterior pillars of the fornix followed their
usual course towards the base of the brain, curving round in front of the foramen of
Monro. The two posterior pillars were wide like the body from which they started.
Each entered the descending horn of a lateral ventricle having its anterior margin
closely adapted to the concave margin of the hippocampus major, so as to form the
fimbria hippocampi in the manner already described. A closer examination of its
disposition now revealed a somewhat remarkable fact which had so far escaped observa-
tion— viz. the surface of the hippocampus major, although rounded and solid in appear-
ance, was now found to consist for the most part of the fibres of the posterior pillar of
• the fornix arranged somewhat like an incomplete hollow tube, within which there lay
concealed a much smaller ridge of grey substance representing the grey matter of the
hippocampus major, which became continuous with the isthmus of the limbic lobe at a
point below the splenium of the corpus callosum.

It is probable, therefore, that the longitudinal fibres of the posterior pillars of the
fornix are distributed to the hippocampus major ; to the hippocampal gyms with its
uncus, as well as to the gyrus dentatus. Further, such an increase of the amount
of grey matter in the hippocampus major as would deepen the sulcus dentatus would
also probably lead to the obliteration of the fissure on the floor of the descending horn
and to a thinning out of the fibres of the posterior pillar of the fornix, and thus produce
appearances which are characteristic of the brain of man without materially increasing
the total size of such a hippocampus major as is presented by the brain of the seal.

The velum interpositum was chiefly notable on account of its large choroid fringes

(ROY. soc. EDIN. TRANS., VOL. XLVIII., 840.)

THE ANATOMY OF THE WEDDELL SEAL. 129

and for its extreme thinness under the body of the fornix, where it covered the optic
thalami and formed one of the roof structures in relation to the mesial or 3rd ventricle,
for which it likewise provided the usual choroid plexuses. It transmitted numerous
vessels into the upper or dorsal surface of the optic thalamus, to which it was closely
adherent, but especially so at the hinder part.

The third ventricle was situated as usual between the optic thalami, and its most
noteworthy character was the large size of the middle commissure (fig. 2). The position
of its anterior and posterior commissures did not call for special comment, and the
structural arrangements and composition of its boundaries were not in any way peculiar.

The optic thalami formed large well-developed masses, and, as already described, no
part of their upper surfaces was visible within the lateral ventricles until the fornix
and velum interpositum were removed. When the upper surface of the optic thalamus
was fully displayed, it presented certain very interesting features. At its postero-
lateral end — that is, close to the entrance to the descending horn of the lateral ventricle,
but upon the upper surface of the optic thalamus — the corpus geniculatum externum
constituted a well-marked elevation which was related to the fornix as previously
explained. Along the mesial margin of its upper surface, a flattened ridge — the tsenia
thalami or stalk of the pineal body — ran backwards towards the anterior end of the
mesencephalon above the posterior commissure of the 3rd ventricle and the entrance
to the aqueduct of Sylvius, where it was joined by its fellow from the opposite thalamus,
and thus formed the peduncle of the pineal body.

The pulvinar was situated between the corpus geniculatum externum and the tsenia
thalami. It formed a flattened area which did not project backwards with an over-
hanging border as in man. The habenula was situated partly to the lateral side and
partly to the mesial side of the taenia thalami. In other words, the taenia thalami ran
across the surface of the habenula. Considered as a whole, the habenula formed a
narrow pyriform projection whose wider end was directed backwards and presented
itself on the lateral wall of the 3rd ventricle high up in the interval between the middle
and posterior commissures.

The corpus stnatum was displayed by making horizontal transverse sections from
the surface of the insula towards the mesial plane so as to include the caudate nucleus,
but it was not until the lower levels of the island of Reil were reached that definite
evidence of striation was observed. The grey substance of the surface convolutions and
that of the caudate nucleus were always distinctly seen, but it was only after the sections
had been subjected to the staining influence of a saturated solution of bichromate of
potash for forty-eight hours that the other grey masses were clearly visible.

The lenticular nucleus occupied its usual position on the postero-lateral aspect of
the head of the caudate nucleus. Its mesial border was convex and separated from the
caudate nucleus and the optic thalamus by the internal capsule. This band was quite
definite, but very narrow ; and it presented the characteristic anterior and posterior
limbs with an intermediate genu. The lateral margin of the lenticular nucleus in its

(BOY. soc. Ems. TRANS., VOL. XLVIII., 841.)

130 PROFESSOR DAVID HEPBURN ON

higher levels presented the ridges and depressions which are the characteristic of the
claustrura, and it was only after the sections had passed below the level of the general
mass of the lenticular nucleus that the claustrum was seen as a separate structure, with
a definite external capsule between it and the more deeply placed grey mass. Indeed,
the appearance of striation, which was directed forwards and outwards, was more
definite below the level at which the lenticular nucleus still retained its biconvex
outline and while the striated substance intervened between the claustrum and the
head of the caudate nucleus. The effect of this disposition of the grey and white
masses of the corpus striatum was to suggest that the differentiation of the external
capsule was incomplete and had not advanced to the stage of separating the claustrum
from the lenticular nucleus.

A band of white substance intervened between the cortical grey matter and wavy
margin of the claustrum, and, since the claustrum is usually regarded as a detached and
submerged portion of the grey cortex of the insula, it would appear the white fibres
which separate the grey cortex from the claustrum are developed earlier than those
which, in the higher brains, separate the claustrum from the caudate nucleus and are
known as the external capsule. In TURNER'S account of the elephant seal, it does not
appear that he submitted his specimen to this dissection. The grey nature of the tail
of the caudate nucleus was always distinct, and an extension of the sections through the
optic thalamus revealed quite plainly its grey substance, bounded laterally by the
posterior limb of the internal capsule. The grey matter, however, did not resolve itself
into the subordinate nuclei (anterior, mesial, and lateral) which characterises the
human brain.

The Pineal Body.— I was aMe to examine three specimens of this interesting object,
and in each case it presented widely different characters. Indeed, the differences were
so pronounced that they were not easy to reconcile and certainly not easy to explain.

In the brain which I removed from the skull of the seal which was two days old
at the time of its death, the pineal body was a large prismatic object resting upon the
vermis of the cerebellum and wedged into the interval between the occipital ends of
the cerebral hemispheres. It projected about 27 mm. behind the splenium of the
corpus callosum. The peduncle broke in the process of removal, but it was very short
and apparently just sufficiently long to permit the expanded part to clear the splenium.
The dimensions of the expanded, prismatic part were as follows : — greatest length,
27 mm. ; width, 18 mm. ; vertical depth, 12 mm.

In a second specimen, belonging to one of the adult brains, the peduncle was again
broken, but the expanded part still occupied its natural position. In this case the
peduncle was cylindrical and the expanded end was pyriform in shape, its measurements
being : — length, 20 mm. ; width, 15 mm. ; vertical depth, 9 mm. It showed no signs of
faceting by pressure from surrounding structures, as might have been expected,
supposing the reduction in its size as compared with the young specimen to have
resulted from the effects of preservative solutions. In the third specimen, also that

(ROY. soc. KDIN. TBAJSS., VOL. XLVJII., 842.)

THE ANATOMY OF THE WEDDELL SEAL. 131

of an adult brain, the complete object was in an undisturbed position. The peduncle
was very thin, flattened from above downwards, and measured 6 mm. in the transverse
direction. It was closely enveloped in the pia .mater, and extended backwards on the
vermis of the cerebellum to terminate in a disc-like expansion 12 mm. in width. The
discoid part was flattened upon its cerebellar surface, while it was slightly conical on
the opposite side. From the commencement of the peduncle to the extreme edge of
the disc it measured 25 mm., of which the peduncle represented 15 mm. and the disc
10 mm. Numerous vessels travelled between the pineal body and the pia mater.
These two adult brains had been preserved in precisely the same way, and therefore it
would appear as if the pineal body of the Weddell seal underwent a gradual reduction
in size subsequent to birth, but that the shrinkage is not accompanied by any marked
shortening in the total length of the object. Similar facts have been recorded by
TURNER in connection with the pineal body of the elephant seal, in which the
measurements were: — length, 16 mm.; greatest breadth. 8 mm.; greatest vertical
diameter, 6 mm. In two specimens taken from the walrus the dimensions were, in one
case, 30 mm. long and 18 mm. wide ; in the other case, 29 mm. long and 13 mm.
wide. There is thus satisfactory evidence that, so far as the seals are concerned, the
pineal body attains an unusual size as compared with other mammals ; although in the
case of Otaria jubata, described by MURIE, the size of this structure may not have been
so noteworthy as in the specimens above detailed, otherwise such a competent observer
could scarcely have confined his account of its size to the statement that it was
" relatively large."

The optic tract followed the usual course from the optic chiasma backwards and
outwards to wind round the lateral aspect of the crus c~-ebri. Thereafter — owing to its
relations to the hippocampus major, as already described — it became compressed into a
somewhat triangular band upon the under side of the thalamus, and sweeping past the
corpus geniculatum internum, with which it became closely associated, it continued its
course, spreading out certain of its fibres towards the pulvinar, but reserving a bundle
of considerable bulk for the corpus genicnlatum externum. So far as the eye could
judge, some of the fibres also reached the superior of the quadrigeminal bodies, but it
did not divide into the brachia which characterise its human arrangement.

III. THE MESENCEPHALON.

The mesencephalon presented the corpora quadrigemina on its dorsal aspect, and
each one of these was quite distinctly defined from the other by longitudinal and
transverse furrows. On its ventral surface the cnira cerebri were also well marked.
Latterly, the corpus geniculatum internum constituted a large oval elevation, larger
than either of the corpora quadrigemina and separated from them by a deep furrow
through which many vessels entered the brain substance. The aqueduct of Sylvius
(fig. 2) was a fairly wide canal, and was not reduced to a T-shaped chink as in man.

(ROT. SOC. EDIX. TRAXS., VOL. XLVm., 843.)

132 PROFESSOR DAVID HEPBURN ON

IV. THE HIND BRAIN.

(a) The Cerebellum.— As is usual among carnivores, the cerebellum possessed a
relatively large vermis in proportion to the size of the hemispheres. When examined
in longitudinal section (fig. 2), the relation of the vermis to the 4th ventricle and the
other constituents of the hind brain presented a great similarity to the corresponding
appearances seen in the human brain.*

The central lobe rested upon the superior medullary velum and possessed a lingula.
The culmen and declive were similarly recognisable, as were also the nodule and the
tonsil upon the inferior or ventricular aspect of the vermis. The pyramid, the tuber
valvidse, and the folium cacuminis were not so easily determined in the brain of the seal
as they are in the brain of man, because, whereas in the latter these structures are turned
towards the floor of the skull, in the former they were turned more towards the hinder
end of the vermis.

The hemispheres were small and practically impossible of the detailed subdivision
which is customary in the descriptions of the human cerebellum, and any attempt to do
so would introduce unnecessary risks of error. In a measure, the points of entrance of
the middle cerebellar peduncles from the pons Varolii provided a guide to what might
be regarded as the dorsal and ventral portions of the cerebellum. On this assumption,
the biventral lobe and the tonsil projected laterally some distance beyond any other part
of the hemisphere, while the flocculus formed a mass of considerable size which over-
lapped the middle peduncle from behind. If we accept the position of the middle
cerebellar peduncle as a sufficiently reliable guide from which to continue the great
horizontal fissure by means of which the upper and lower aspects of the human
hemisphere are located, then in the brain of this seal all that remained of each hemi-
sphere, in addition to the objects already mentioned, occupied the same aspect and was
directed towards the tantorium. Nevertheless, it was divided into two clearly defined
areas by a fissure which commenced at the point where the middle peduncle entered
from the pons Varolii. If, now, we name these lobes respectively superior -anterior and
superior-posterior, then all the parts of the cerebellum of the seal have been accounted
for. It may be noted that the part which I have just named the superior-anterior lobe
is reduced to a single folium in relation to the vermis, and it is this folium which is
named the folium cacuminis (fig. 2).

Compared with the human cerebellum, it would appear that whereas in the seal the
vermis and its subordinate parts are well developed, and the flocculus, biventral lobe,
and tonsil are produced on a large scale, the remainder of the hemisphere is much
reduced in proportion. On the other hand, in man the hemisphere proper has become
much expanded and thickened, with corresponding reduction in the size of the flocculus,
the biventral lobe, and the tonsil. In fact, a theoretical enlargement of the superior-
anterior and superior-posterior lobes of the hemisphere of the seal, accompanied by their

* Text-Boole of Anatomy, edited by CUNNINGHAM, 3rd ed., p. 512.
(ROY. soc. EDIN. TRANS., VOL. XLVIII., 844.)

THE ANATOMY OF THE WEDDELL SEAL. 133

expansion backwards as well as laterally, and a reduction in the size of the flocculus,
biventral lobe, and the tonsil, would be capable of producing a cerebellum with practically
the same superficial characters as that of man.

The pons Varolii was well developed, and measured 25 mm. from its anterior to its
posterior border in the line of its very definite basilar groove. The anterior and
posterior borders converged upon each other so rapidly, as they travelled outwards to
form the middle cerebellar peduncles, that the outline of the posterior border was
interrupted by the emergence of the large root of the 5th cranial nerve. As a result
of this arrangement, the greater part of this nerve-root made its appearance from the
side of the medulla oblongata between the olivary eminence and the pons, but of course
close up to the latter. In the elephant seal TURNER notes that the 5th cranial nerve
arose from the pons Varolii and not from the bulb, whereas, in describing the same nerve
in the walrus, he remarks that some fibres of the sensory root " passed backwards
between the facial and auditory nerves to the anterior and outer part of the medulla
oblongata."

The medulla oblongata was wide at its upper end, where it measured 40 mm. ; but
it narrowed rapidly towards the lower end, and instead of being conical it was markedly
flattened in the dorso-ventral direction. Its upper or " open " part was associated with
the 4th ventricle, while the "closed" or lower part contained the central canal in its
unexpanded condition. Its bilateral character was indicated by the anterior and
posterior medium fissures, the former shallow and terminating in relation to the
posterior border of the pons Varolii at the foramen of Vicq d'Azyr. On each side of
the anterior median fissure or groove the pyramid formed quite a distinct tract. The
point of emergence of the 6th cranial nerve was not between the pyramid and the
pons as in man, but from a flattened area situated external to the pyramid, so that the
nerve-stem made its appearance close to the mesial side of the large medullary root of
the 5th nerve and without any fibres of the pons intervening between them. The 7th
and 8th cranial nerves emerged from the side of the medulla oblongata close behind
the 5th and 6th nerves, but slightly nearer the dorsal or ventricular aspect of the bulb.

The olivary eminence was small and not so prominent as in man, but it distinctly
separated the 9th and 10th cranial nerves from the 1 2th or hypoglossal nerve.

The closed part of the medulla oblongata presented the general appearances and
proportions of the adjacent part of the spinal cord as regards its fissures and main
columns. Posteriorly, the funiculus gracilis with the clava at its upper end, the
funiculus cuneatus with its upper expansion, the cuneate tubercle, and also the tubercle
of Rolando were all definitely recognisable. They turned outwards in a common bundle
from a point immediately below the obex, and skirting the infero- lateral margin of the
4th ventricle they entered the cerebellum as the restiform body or inferior cerebellar
peduncle. The visible decussation of the pyramids began at a point 32 mm. from the
hinder margin of the pons Varolii, so that we may consider the total length of the bulb
to be distinctly less than its width at its upper end.

(ROT. soc. EDIN. TRANS., VOL. XLvin., 845.)

134 PROFESSOR DAVID HEPBURN ON

The rhomboid or ktli ventricle was distinctly lozenge-shaped, but neither in regard
to its size nor in regard to the detailed modelling of its floor was it so well marked as in
man. The floor presented a median furrow, as well as an inferior and a superior fovea
in relation to each quarter of the lozenge. Associated with the fovea inferior, the
trigonum hypoglossi and the trigonum vagi formed quite recognisable elevations. The
area acusticse was likewise a well-marked elevation on the floor, but its surface was
smoother than in man because of the absence of visible strise on its free surface. The
•eminentia teres was placed to the mesial side of the superior fovea, but it was prolonged
upwards as well as downwards by a longitudinal ridge which ran upwards along the
floor of the aqueduct of Sylvius in the one direction, and downwards to join the trigonum
hypoglossi in the other.

The obex was a distinct object in the roof of the ventricle in relation to its inferior
angle, and the ligula could be seen extending from it on each side.

SUMMARY.

In making a general summary of the naked-eye anatomy of the brain of the
Weddell seal, the features which have impressed me most and seem most deserving
of special reference are the following : —

1. Its angular appearances in association with its large size, suggesting that the
general fish-like outline of the entire animal has to a certain extent influenced the
shape of its skull, and thereby the shape of the brain within the cranium.

2. The size and elaborate ramification of the cerebral convolutions, together with
the considerable amount of asymmetry in the details of the arrangement of the
convolutions of the one hemisphere as compared with the other.

3. The width of the interval between the two hemispheres posterior to the hinder
end of the corpus callosum.

4. The marked departure from the arrangement of the cerebral convolutions in
such a typical carnivore as the dog.

5. The presence of those convolutions belonging to the island of Eeil upon the
same superficial plane as that of the surrounding convolutions which form the
opercula.

6. The definite and complete character of the limbic lobe.

7. The position of the calcarine fissure, and thereby of the visual area upon the
inferior aspect of the occipital end of the hemisphere.

8. The large size of the fornix, and particularly of its posterior pillars, in association
with a well-marked hippocampus major, of which the greater part is composed of
fornix fibres and only a small part of grey substance.

9. The long stalk and the large size of the pineal body and its position upon the
vermis of the cerebellum, in the open interval between the cerebral hemispheres.

10. The well-developed but, on the whole, simpler characters of the basal structures
as compared with the elaboration of the pallium.

(ROY. soc. EDIN. TRANS., VOL. XLVIII., 846.)

THE ANATOMY OF THE WEDDELL SEAL.

135

11. The relatively simple character of the mesencephalon.

12. The small size of those parts of the cerebellar hemispheres which in man
would constitute their main bulk ; and the large size of those objects in relation to
the vallecula, which in man would be relatively small.

13. The reduction in the posterior margin of the pons Varolii, which thereby
permits the bulbar root of the 5th cranial nerve to emerge directly from the side of
the medulla oblongata.

LITERATURE.

MURIE, Trans. ZooL Soc. Land., vol. viii., 1874.

WIEDERSHEIM and PARKER, Comparative Anatomy of Vertebrates, 3rd ed., 1907, p. 224.

CUNNINGHAM, Text-Bonk of Anatomy, edited by Cunningham, 3rd ed., p. 512.

Sir WM. TURNER, Challenger Reports, vol. xxvi., Zoology : Report on Seals (" Brain of Elephant Seal and

of Walrus," p. 89 et seg., plates viii., ix., x.). In connection with this Report a very full bibliography

is provided.

DESCRIPTION OF PLATE.

Fig. 1. The left cerebral hemisphere, natural
size.

F. Sy. Fissure of Sylvius.

F. Ro. Fissure of Rolando.
S. P-c. Pre-central sulcus.
S. Or. Sulcus cruciatus.

S. I-p. Intra-panetal sulcus.
/. R. Island of Reil.

Fig. 2. Mesial surface of the right cerebral hemi-
sphere.

S. Cr. Sulcus cruciatus.

F. Ro. Fissure of Rolando.

/. P-o. Internal parieto-occipital fissure.

Op. C. Optic commissure.

G. C. Corpus callosum.
Hy. C. Pituitary body.

P. Var. Pons Yarolii.
Pin. Pineal body.

Fig. 3. Basal surface of cerebral hemisphere.
Rli. f. Rhinal fissure.

F. Sy. Fissure of Sylvius.
S. Col. Sulcus colateralis.

C. f. Calcarine fissure.
P. o. Parieto-occipital fissure.
S. D. Sulcus dentatus.
0. T. Olfactory tract.
Op. N. Optic nerve.
Hy. C. Pituitary body.
Cr. C. Crus cerebri.

Is. Isthmus.
Gun. Cuneus.

G. L. Gyrus lingualis.

Fig. 4. Vertical transverse section through the
posterior horn of the lateral ventricle of
the right hemisphere, viewed from behind.
C. f. Calcarine fissure.
V. Lateral ventricle.

(ROY. soc. EDIX. TRASS., VOL. XLVHI., 847.)

SCOT. NAT. ANT. EXP.

HEPBURN : BRAIN OF WEDDELL SEAL.

Vol. IV

-p. Man..

I). II.-pl'llllt. ilrl.

ll'Fariane & Bnkiim K.lin.

PART X.

SEALS.

SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

X.-HISTOLOGY OF THE CENTRAL NERVOUS SYSTEM

OF THE WEDDELL SEAL

(LEPTONYCHOTES WEDDELL1).

BY HAROLD AXEL HAIG, M.B., B.Sc. (LOND.), M.R.C.S. (ENG.), L.R.C.P. (LoND.),

Lecturer on Histology and Embryology, University College, Cardiff.

(WITH TWO PLATES AND NINE TEXT FIGURES.)

Scottish National Antarctic Expedition : A Contribution to the Histology of
the Central Nervous System of the Weddell Seal (Leptonychotes weddellii).
By Harold Axel Haig, M.B., B.S. (Lond.), M.R.C.S. (Eng.), L.R.C.P. (Lend.),
Lecturer in Histology and Embryology, University College, Cardiff. Communi-
cated by Dr W. S. BRUCE. (With Two Plates and Nine Text Figs.)

(MS. received June 18, 1912. Read December 2, 1912. Issued separately February 17, 1913.)

CONTENTS.
PART I. (/) The Cerebellum.

(<i) The Spinal Cord.

(6) The Medulla Oblongata.

(c) The Pons Varolii.

The Mesencephalon.

(«) The Motor Cortical Area.

(g) The Optic Thalamus.

The Pituitary Gland.

PART I.

INTRODUCTORY AND PRACTICAL DETAILS.
The specimens submitted for examination were :

(a) Portions of the brain (labelled Specimen XXXI.).
(6) Portions of spinal cord (labelled Specimen XXIV.).

Both were in excellent condition as regards fixation and hardening, having been
preserved for many years in a fluid composed of formol and 95 per cent, alcohol (the
fluid was also injected into the cerebral vessels). They were, previous to histological
examination, submitted to the following processes : —

i. Comparatively thin slices were taken from various regions and placed for

twenty-four hours in absolute alcohol,
ii. Then transferred to acetone for twelve hours,
iii. Placed in xylol until permeated.

iv. Embedded in paraffin of melting-point 52° C. Sections were then taken
with an improved form of the Cambridge rocking microtome, and fixed to
slides by means of the albumen method.

Staining of the Sections.

The best stain for differentiating the grey and white matter was found to be
Weigert's method with acid fuclisin, used alone, or subsequent to treatment with
Delafield's haematoxylin. Other stains were tried (Bismarck brown with acid fuchsin
as counterstain, aniline blue-black, haematoxylin and orange-G., etc.), but acid-fuchsin

(REPRINTED FROM THE TRANSACTIONS Of THE ROYAL SOCIETY OF EDINBURGH, VOL. XLVm., PP. 849-866.)

142 MR HAROLD AXEL HAIG ON THE

alone appeared to give the best results. The Weigert-Pal method was given a trial,
but with negative results, owing to the special means of preservation and hardening
employed.* The following regions were studied histologically :—

(a) Mid-lumbar, dorsal, and cervical regions of the spinal cord.

(b) Medulla oblongata :

i. In the region of the pyramidal decussation.

ii. At the lower level of the olivary body.

iii. At the middle of the olivary body.

iv. At the upper level of the olivary body.

(c) Pens, lower, middle, and upper regions.

(d) The mesencephalon :

i. At the level of the posterior corpora quadrigemina.
ii. At the level of the anterior corpora quadrigemina.

(e) The optic thalamus.

(f) The motor region of the cortex of the cerebral hemispheres (precentral gyrus).

(g) The cerebellar vermis (mesial region).
(k) The optic chiasma.

(i) The pituitary gland.

(k) The uncinate gyrus of the hippocampus major.

I. HISTOLOGY OF THE SPINAL CORD.

A. Cervical Region. (Text-fig. 1.)

A transverse section across the upper region of the cervical cord shows a very
marked breaking up of the grey matter into four main groups in each half, viz. :—

(a) A posterior mass extending from the posterior grey commissure into the

posterior median column of Goll ; in this mass at the base small nerve-cells
are seen, and a few at the peripheral part near the surface of the cord. The
latter represent the beginning of the nucleus gracilis, which soon becomes
prominent even in the cervical region.

(b) A postero-lateral mass, into which the fibres of the posterior roots may be seen

to extend. In this mass small nerve-cells may be made out, but they are
very scattered and not at all conspicuous.

(c) An intermedio-lateral mass, at the central portion of which some large nerve-

cells are to be made out, these representing the cells of Clarke's column.

(d) An anterior mass (anterior cornu) of wide extent, and possessing four or five

groups of very large nerve-cells (motor cells of anterior cornu).

* The pituitary gland sections were stained with Delafield's haematoxylin, followed by eosin to differentiate.

(ROY. soo. EDIN. TRANS., VOL. XLVIII., 850.)

HISTOLOGY OF THE CENTRAL NERVOUS SYSTEM OF THE WEDDELL SEAL. 143

The postero-lateral and intermedio-lateral masses of grey matter are united by
strands of neuroglial tissue, which break up the lateral regions of the cord into a well-
marked reticular formation. The grey matter in general is characterised by its deeply
staining neuroglia fibres, which are of a somewhat coarse nature, and are apparently
less in number per unit area as compared with the supporting tissue of higher types
of cord.

H

rfc

fr.%.

K

m.

CL-K.

FIG. 1. — A transverse section of the spinal cord (upper cervical region).

a.f. Anterior median fissure.
p.f. Posterior median fissure.
p.r. Posterior root-bundles.
a.r. Anterior root-bundles.
f.g. Funiculus gracilis(posteriormediancolumn),

showing an indication of the beginning

of the nucleus gracilis.

p.h. Posterior horn of grey matter (beginning
of the substantia gelatinosa of Rolando).
i.l.h. Intermedio-lateral horn.
f.r. Format io reticularis.
a.h. Anterior horn.
p. in. Pia mater.

In the central region of the section the posterior grey commissure is of wide extent,
and a small amount of neuroglia is to be seen anterior to the central canal of the cord :
the latter is flattened antero-posteriorly, and is lined by a well-marked layer of
columnar ciliated epithelium.

The postero-median fissure is of about the same length as the anterior median
fissure, perhaps slightly longer ; but the cord has apparently been subjected to a certain
amount of mechanical pressure, causing flattening, and giving the whole section the
appearance of abnormal transverse as compared with antero- posterior diameter. The

(BOY. soc. KDIN. TRANS., VOL. xivra., 851.)

144 MR HAROLD AXEL HAIG ON THE

postero-median fissure, which normally is of greater extent than the anterior, would
thus be reduced in length.

The. white matter of the cord appears to consist of very fine medullated nerve-fibres :
in the posterior region, the columns of Goll (mesial) and Burdach (lateral) are very
well marked, the pia mater dipping into the cord and marking off these two wedges of
white matter. The pial septum, separating the postero-lateral column from the crossed
pyramidal and direct cerebellar tracts, passes almost to the postero-lateral horn of
grey matter.

The fibres of the anterior roots of the spinal nerves pass out through the white
matter in three or four main bundles : those of the posterior roots in one thick bundle
of fibres situated at the middle of the postero-lateral horn.

Comparison with higher type of mammalian cord (cervical region}.

The main points in the comparative histology are the following :—

(a) The early appearance of the nucleus gracilis, which in the human cord does

not appear before the medulla has -been reached.

(b) The extensive spreading out of the grey matter of the posterior and intermedio-

lateral horns.

(c) The wide disproportion between the transverse and the antero- lateral diameter

of the cord, due account being of course taken of any pressure which may
have arisen.

(d) The marked development of the formatio reticularis.

(e) The relatively enormous size of the motor cells in the anterior cornu : this is

perhaps not sufficiently emphasised in text-fig. 1, but is nevertheless a very
striking feature in the actual section. This feature is possibly related to
the highly developed powers of locomotion shown by members of the
seal family.

The proportion of grey to white matter is as 3 to 4 approx.

B. Dorsal region. (Text-fig. 2.)

The cord appears in this region to have been submitted to mechanical pressure
in the transverse diameter : for this reason, the central canal appears elongated
antero- posteriorly and somewhat distorted. The postero-median fissure is longer
than the anterior and reaches to the grey commissure ; the anterior fissure stops short
of the central grey matter, leaving a well-defined anterior or white commissure. The
antero-posterior diameter appears greater than the transverse. The grey matter is
divided in each lateral half into three main masses, viz. :—

(a) A posterior horn, in which are to be seen a few medium-sized nerve-cells.
(6) An intermedio-lateral horn, at the tip of which a group of medium-sized nerve-

(ROY. soc. EDIN. TRANS., VOL. XLVIII., 852.)

HISTOLOGY OF THE CENTRAL NERVOUS SYSTEM OF THE WEDDELL SEAL. 145

cells is situated ; the cells of Clarke's column are also seen, having been
displaced below the base of the posterior horn by mechanical pressure,
(c) A broad anterior horn possessing several groups of large nerve-cells.

The white matter in the posterior region shows but slight indication of a division
into postero-mesial and postero-lateral columns : the lateral columns are of relatively
wide extent.

71

n.

L./l

FIG. 2.— A transverse section of the spinal cord (mid-dorsal region). Lettering as in fig. 1, except :—

a.m. Arachnoid mater.
c.c. Central canal of cord.

n. Posterior roots.

Anterior spinal vessels.

The posterior nerve-roots are narrower than in the cervical region, but are very
well denned : the root-bundles lie just outside, and are seen cut across in the section.
The anterior roots pass out in three or four narrow strands.

The proportion of grey matter to white is as 9 to 20 approx.

Compared with higher types, the dorsal cord shows but few divergences : the
posterior horn of grey matter is rather shorter than in the human cord, and Clarke's
column of nerve-cells is but feebly developed. On the other hand, the motor cells in
the anterior horn are very well defined.

(ROY. soc. EDIN. TRANS., VOL. XLVIII., 853.)

146

MR HAROLD AXEL HAIG ON THE

C. Lumbar region. (Text-fig. 3.)

The three main subdivisions of each lateral crescent of grey matter can be dis-
tinguished, viz. posterior, intermedio-lateral, and anterior horns.

A few nerve-cells are present about the middle of the posterior horn, and a well-
marked group is to be seen at the base of the posterior horn (Clarke's column).
Groups also occur at the tip of the intermedio-lateral horn, and to the number of three
in the anterior horn, these being very obvious and the component cells very large.

H

n.

FIG. 3. — A transverse section of the spinal cord (lumbar region). Lettering as in figs. 1 and 2, except : —

d.m. Dura mater.
p.r.b. Posterior root- bundles.
a.r.b. Anterior root-bundles.

n. Bundles of the candii equina.

v. Large branches of the vertebral vessels.

The posterior or grey commissure is wide, and the anterior white commissure is
relatively broad in the antero-posterior direction : the central canal appears as a
four-sided space lined by an ependyma possessing an internal layer of ciliated
epithelium.

The posterior root-bundles are very large and number three or four on each side
posteriorly, lying between the dura mater and the pial investment of the cord ; the
anterior bundles are two in number on each side antero-laterally.

Of the white matter, the lateral columns are of fairly wide extent, as also are the
anterior columns ; there is no marked distinction between Goll's and Burdach's columns
posteriorly, although a small pial septum does show the superficial division.

(ROY. soc. EDIN. TRANS., VOL. XLVIII., 854.)

HISTOLOGY OF THE CENTRAL NERVOUS SYSTEM OF THE WEDDELL SEAL. 147

In the section figured the postero-median and antero-median fissures are of about
the same length, but in a normal specimen the posterior would be rather longer
than the anterior.

Outside the dura mater may be seen nerve-bundles representing a high origin of
the cauda equina : this feature is very marked in the seal, the cord ending in a
filament at a much higher level than in the human.

The proportion of grey matter to white is as 1 to 3 approx.

The chief features of comparative value are the relative shortness of the posterior
horn as compared with the human, and the presence of the surrounding bundles of
the cauda equina even high in the lumbar region of the cord. The motor cells of
the anterior cornua are, as in the dorsal and cervical regions, larger relatively than
those found in the cord of man.

II. HISTOLOGY OP THE MEDULLA OBLONGATA.
A. At the middle of the pyramidal decussation. (PI. I. fig. 1.)

Several marked points of difference are seen here as compared with the upper cervical
region : firstly, the transverse diameter of the section is twice the antero-posterior
diameter, and the anterior fissure is about twice the length of the posterior. The
central canal is approaching the posterior surface of the cord, but there still remains a
fairly wide grey commissure.

Indications of the beginning of a restiform body may be made out laterally, and the
intermedio-lateral horn of grey matter now forms a well-defined mass, known as the
substantia gelatinosa Kolandi (s.g.R.}.

The grey matter generally exists in relatively large proportion : the large motor
cells of the anterior horn are still very obvious, and in the postero-median and postero-
lateral columns are to be seen respectively the nucleus gracilis and nucleus cuneatus
(n.g. and n.c.).

The pyramidal decussation is a marked feature, fibres passing across from the
lateral column of one side to the anterior column of the opposite half, and to a certain
extent separating the grey matter of the posterior region from that of the lateral and
anterior regions.*

The proportion of grey to white matter is as 3 to 4 approx.

B. At a point just below the calamus scriptorius of the 4th ventricle.

(PI. I. fig. 2.)

In this region the central canal is fast approaching the floor of the 4th ventricle to
open out into that cavity. The pyramidal decussation is now no longer noticed, the
upper level having been passed.

* The decussation of the pyramids is of greater extent longitudinally than in the human medulla, being found
quite close to the calamus scriptorius at its upper level.

(ROY. soc. EDIK. TRANS., VOL. XLVIII., 855.)

148 MR HAROLD AXEL HAIG ON THE

The substantia gelatinosa Rolandi is now quite separate from the rest of the grey
matter and lies enclosed by white fibres which partly arch round it and partly form the
lower limit of the restiform body. The pyramids form marked protuberances anteriorly
in the middle line.

No trace of an olivary body is seen as yet, a feature which is of comparative value ;
the nuclei gracilis and cuneatus are well defined, the latter lying now well to the outer
side of the former.

Large nerve-cells are to be made out in the grey matter of the still visible represen-
tatives of the anterior cornua.

The proportion of grey to white matter is as 3 to 5 approx.

The transverse diameter of the section markedly exceeds the antero-posterior
diameter : this feature is now prominent throughout the whole extent of the medulla,
until, when the pons is reached, the disproportion becomes less obvious.

C. At the lower limit of the olivary body. (PL I. fig. 3.)

A median raphe has now appeared, and the central region is occupied by intercross-
ing strands of fibres (internal arcuate fibres), and just dorsal to the pyramids the tract
of the fillet may be seen.

The central canal has opened out on to the floor of the 4th ventricle, the latter being
covered by a layer of ependyma.

The restiform body is now becoming a more obvious feature, and outside this fibres
forming a well-marked covering, passing from the pyramid regions round the restiform
body towards the dorsal region.

Two well-defined nuclei have now taken the place of the nuclei gracilis and
cuneatus: these are respectively the nucleus of the 12th cranial nerve and the dorsal
nucleus of the 10th cranial nerve (n. 12th, n. 10th).

The olivary body is peculiar in that in section it shows an internal mass of grey
matter, loaded with rather large nerve-cells, having the form of a U-shaped fold, which
however does not possess any folds of the second order such as are to be seen in the
grey matter of the human olivary body. In some regions this olive is open centrally
(hilus), but in the lower regions takes the form of a closed oval of grey matter (o.n.).
The olive does not form a very marked external projection, and, moreover, especially in
the higher regions of the medulla, appears to be further removed from the pyramids. A
few of the internal arcuate fibres may be traced from the hilus olivse across the median
raphe to the opposite restiform body.

The rest of the grey matter is of a somewhat scattered aspect : the substantia
gelatinosa still forms an obvious mass laterally enclosed by the curved restiform body :
some of the masses of grey matter lying ventrally may possibly represent accessory
olivary bodies, but some certainly belong to the category of arcuate nuclei.

Just below the floor of the 4th ventricle in the middle line is a tract of fibres repre-

(ROY. soc. EDIN. TKANS., VOL. XLVIII., 856.)

HISTOLOGY OF THE CENTRAL NERVOUS SYSTEM OF THE WEDDELL SEAL. 149

senting the posterior longitudinal bundle : and ventral to this another band, not well
defined, which is the anterior longitudinal bundle.

D. At the middle of the olivary body. (PI. I. fig. 4.)

The main points in this region of the medulla are : the olivary nucleus, the restiform
body, the nuclei of the 12th and 10th cranial nerves, and outside both of these a group
of nerve cells which apparently represent the nucleus of the vestibular division of the
8th cranial nerve.

Other well-marked features are the issuing fibres of the 12th, 10th, and 9th nerves :
the first of these pass down through the internal arcuate fibres to emerge from the
medulla between the olive and a mass of grey matter which has become separated from
the substantia gelatinosa ( ? nucleus ambiguus) : the fibres of the 10th nerve pass
through the arcuate fibres between the restiform body and the fibres of the 12th nerve
to emerge at the inner edge of the restiform : the fibres of the 9th nerve are only seen
for a short part of their course and emerge further forward (see next section). The
pyramids are now very well defined, and the tract of the fillet and the posterior
longitudinal bundle form characteristic features.

E. At the upper level of the olivary body. (PI. I. fig. 5.)

In this section, a layer of grey matter appears spreading over the floor of the 4th
ventricle, quite distinct from the subjacent nuclear groups ; of the latter groups, two,
representing nuclei of the 9th nerve, are to be seen just below and external to the
very obvious posterior longitudinal bundle. The fibres of the 9th nerve are seen
issuing between the olivary body and the restiform body, whilst external to the latter
some fibres of the 8th cranial nerve are to be distinguished.

Several small groups of grey matter, just mesial to the olivary nucleus, may
possibly represent accessory olivary nuclei.

The anterior longitudinal bundle, the fillet, and the pyramids all form marked
features from above downwards ; and the fibres which enclose and arch round the
restiform body ( ? continuation of the external arcuate fibres) form also a point worthy
of notice.

III. HISTOLOGY OF THE PONS VAROLII. (PI. 1. figs. 6. 7, and 8.)

A section across the lower pontine region of the brain shows the 4th ventricle
closed over by the superior medullary velum, and the floor of the ventricle lined by a
well-marked ependyma ; a thick layer of grey matter lies subjacent to this, and, in the
middle line, the posterior longitudinal bundle is one of the most prominent features
of the section. In the middle of this region there is to be seen the formatio reticularis,
and ventral to this, a fairly wide trapezium makes its appearance ventral to the

(ROY. HOC. EDIN. TRANS., VOL. XLVIII., 857.)

150 MR HAROLD AXEL HAIG ON THE

trapezium, the pyramid bundles form two very distinct masses, and between them lies
grey matter representing the nuclei pontis ; two small masses of grey matter dorso-
external to the pyramid bundles may possibly represent superior olivary nuclei.
Pontine nuclei are also to be seen lying ventral and external to the pyramids.

The nucleus of the 7th cranial nerve lies at the outer and upper angle of the
formatio reticularis, and the issuing fibres of the 7th nerve pass down close to and
parallel to a wide bundle of fibres which arises (partly) from a nucleus close to the inner
side of the restiform body ; this bundle is the issuing root of the auditory or 8th
cranial nerve. Outside the restiform body and the above-mentioned nerves lies part
of the white matter of the cerebellar hemisphere (c).

Pontine fibres pass ventrally to the pyramid bundles, and form characteristic
parallel strands.

The mid-pontine region shows a very thick layer of grey matter beneath the
ependyma of the floor of the 4th ventricle, on either side of the middle line : beneath
this, the posterior longitudinal bundle forms a conspicuous band, the two lying close
together in the middle line. The tract of the fillet is also well defined, lying just above
and to the outer side of the pyramid bundle ; whilst the intercrossing fibres of the pons
alternate with parallel streaks of grey matter, breaking up the majority of the central
part of the section (PI. I. fig. 7) into a reticular formation of wider extent than that in
the lower pontine region.

The nuclei of the 5th cranial nerve (motor and sensory nuclei close together)
appear as a large group of nerve-cells just internal to the restiform body, which latter
is now known as the superior cerebellar peduncle, and the issuing fibres of the 5th
nerve are seen at the lateral part of the section just ventral to the white matter of the
cerebellar hemisphere.

The trapezium is represented by the dorsal portion of the reticular formation, and
can hardly be distinguished histologically from that, except by appropriate methods,
inapplicable in the case of the present material. The central bundle of the 5th nerve
forms a rather narrow band of fibres lying just beneath the grey matter of the floor
of the ventricle.

The pontine fibres which lie at the lower (ventral) aspect form very marked parallel
strands curving outwards towards the 5th nerve : and the grey matter just dorsal to
these fibres (nuclei pontis) are distinguished by their relatively wide extension laterally.

The upper pontine region appears in some respects very similar to the mid-region ;
the tract of the fillet is, however, becoming divided into two main portions, viz. the
intermediate fillet and the lateral fillet (PI. I. fig. 8 /).

The posterior longitudinal bundles are now slightly separated in the mid-line by
a small amount of grey matter, and the grey matter of the floor of the 4th ventricle is
not quite so thick as in the middle region of the pons.

The crossing of the 4th cranial nerves is seen as a band lying just above the
ventricle, forming a kind of roof to the cavity at this point ; the superior cerebellar

(ROY. soo. EDIN. TRANS., VOL. XLVIII., 858.)

HISTOLOGY OF THE CENTRAL NERVOUS SYSTEM OF THE WEDDELL SEAL. 151

peduncles are very distinct, and their decussation may be seen about the middle of
the section, although this does not form so obvious a feature as it does in the human
pons. A noteworthy feature is the absence of any sign of substantia nigra, as also is
the fact that the pyramid bundles are still isolated masses lying well in the grey
matter of the lateral regions. Pontine fibres are also well marked.

The anterior longitudinal bundle lies ventral to the posterior longitudinal bundle,
but is not a marked feature, since a large proportion of grey matter is present in the
mid-region. Below the decussation of the superior peduncles in the mid-line is some
grey matter, which, although ill-defined, would represent the so-called central nucleus of
the higher types.

The root-bundle of the 4th nerve appears just external to the grey matter of the
floor of the ventricle, and indications of the nucleus of the 4th nerve are to be made
out at the outer limit of the grey matter in the floor.

IV. HISTOLOGY OF THE MESENCEPHALON. (PI. I. figs. 9 and 10.)

A section across the mid-brain in the region of the posterior corpora quadrigemina
shows a structure very like that seen in the human mid-brain ; the aqueduct of Sylvius
is, however, much nearer the dorsal surface, and moreover is rhomboidal in shape.

Each corpus quadrigeminum possesses an outer coat of white fibres and inter-
mediate mass of grey matter, and an inner thin band of white fibres separating it from
the central grey matter round the aqueduct.

The reticular formation of the tegmentum is very distinct, and the posterior longitu-
dinal bundles show clearly just below the grey matter surrounding the aqueduct of Sylvius.

The nuclei of the 3rd and 4th cranial nerves are very well defined, lying mesially
just ventral to the lower angle of the aqueduct.

The substantia nigra forms a layer, containing some very large nerve-cells, lying
between the fillet tracts and the crustae ; prolongations from the substantia nigra pass
into the crusta and tend to subdivide it into two or more regions on either side.

There is an ill-defined mass of grey matter between the two crustee, representing an
inter-peduncular ganglion, and above this a tract of decussating fibres which form the
crossing (upper part) of the superior cerebellar peduncles. The decussating fibres of
the tegmenta are well marked in the mid region above the substantia nigra.

A section across the anterior corpora quadrigemina shows some divergence from
the human type ; the Sylvian aqueduct is still rhomboidal in shape, and is some dis-
tance from the dorsal surface. The fibres of the 3rd cranial nerve are seen issuing
through the posterior longitudinal bundles, and a portion of the section which in the
human is occupied by the red nucleus : the latter, however, cannot be markedly
distinguished as such in the section in question.

Each anterior corpus quadrigeminum possesses a very thin covering of white fibres,
grey matter in the intermediate region, and a narrow layer of white fibres internally ;

(ROY. soc. EDJN. TRANS., VOL. XLVIII., 859.)

152

MR HAROLD AXEL HAIG ON THE

the grey matter surrounding the aqueduct shows mesioventrally the nuclei of the 3rd
nerve, and ventral to the latter the posterior longitudinal bundles are fairly obvious.
The crustse are of less extent than those of the former section.

The posterior commissure of the brain shows dorsal to the aqueduct of Sylvius, and
is of moderate antero-posterior dimensions.

Points of comparative value in connection with the medulla, pons, and mid-brain
are as follows : —

(a) The disproportion between the transverse and antero-posterior diameters of the
medulla, the former being about twice or two and a-half times the length of
the latter.

(6) The relatively large proportion of grey matter in all these regions, much of it,
however, being composed of purely neuroglial tissue.

(c) The aberrant shape of the olivary nucleus and the outward displacement of the

whole olive.

(d) The marked development of the restiform body and its early distinction in a

comparatively low region.

(e) The late formation of the crustse, the mid-brain being reached before these are

well defined. Other minor points will be made out by reference to Plate I.,
and comparison of the figures with sections of normal human material.

V. THE HISTOLOGY OF THE OPTIC THALAMUS. (Text-fig. 4.)

The main features to be made out from a vertical (sagittal) section of the optic
thalamus consist mainly in the relative distribution of grey and white matter ; the
white matter occurs in two main masses — an external, thick superiorly and thinning off
towards the anterior aspect, and an internal oblique mass which divides the internal

FIG. 4. — A mesial sagittal section of the optic thalamus (semidiagrammatic). x 4 times.

w.f. Superficial layer of white fibres into which septa of neuroglia pass from the deeper grey matter.
n.n.n. Groups of large nerve-cells in the grey matter.

g.m. Central grey matter. The internal mass is not sharply differentiated into grey and white
matter : small nerve-cells appear scattered throughout the;former.

(ROY. soc. EDIN. TRANS., VOL. XLVIII., 860.)

'HISTOLOGY OF THE CENTRAL NERVOUS SYSTEM OF THE WEDDELL SEAL. 153

grey matter into two main groups, an anterior and a posterior. Just beneath the
superficial white matter several well-defined groups of nerve-cells may be seen (n.), and
posteriorly a group is also to be distinguished. Many small nerve-cells are scattered
throughout the masses of grey matter.

VI. THE HISTOLOGY OF THE CONVOLUTIONS OF THE PRECENTRAL GYRUS.

(Text-fig. 5.)

The cortex of the motor area presents a fairly typical structure, except that large
multipolar nerve-cells occur at a relatively deep level, forming a deeply staining layer

FIG. 5. — Semidiagrammatie vertical section of a part of the motor cortex cerebri (precentral convolution), x 6.

1. Plexiform layer.

2. Layer of small nerve-cells (granules).

3. Layer of large flask-shaped nerve-cells, the axons passing centrally, dendrons peripherally.

4. Layer of large pyramidal cells (comp. to Bett's cells of human motor cortex).

5. Deep layer of large pyramids, lying next to the white centre : there is much dense neuroglia in this layer.

lying next the fibres of the white centre ; in all, about four layers of nerve-cells may
be distinguished. The following layers are quite distinct :—

(a) A superficial layer formed of interwoven fibres (plexiform layer).

(b) A second layer of small nerve cells, the axons of which are not very obvious.

(c) A third layer of large flask-shaped cells not unlike the Purkinje cells of the

cerebellum, with axons passing centrally and dendrons peripherally.

(d) A fourth layer of rather large pyramidal cells, the axons passing centrally.

(e) A fifth layer of large multipolar cells, the axons passing in many directions ;

this layer contains much deeply staining neuroglia.

(ROT. SOC. EDIN. TRANS., VOL. XLVIII., 861.)

154

MR HAROLD AXEL HAIG ON THK

VII. THE HISTOLOGICAL FEATURES OF THE VERMIS OF THE CEREBELLUM.

(Text-fig. 6.)

A section across the lamellar of the vermis (or of the hemisphere) shows the typical
arrangement characteristic of higher mammals, viz. a white centre, an inner layer of
"granules" (small nerve-cells, the axons of which pass peripherally), an intermediate
layer of the large flask-shaped cells of Purkinje, the axons of which pass centrally, the
dendrons passing peripherally, and a superficial "molecular" layer, consisting of small

FIG. 6. — A portion of a vertical section of the vermis of the cerebellum, x 6.

g. Layer of granules (small nerve-cells and axons
of the Purkinje cells).

/. Fibres of the white centre.

m. Molecular layer.

p. Layer of large Purkinje cells.

nerve-cells, the dendrons of Purkinje cells and fibres derived from neuroglia cells, and
the " climbing" and " moss" fibres coming from the deeper layers.

The only point of comparative value is the relatively large size of the Purkinje cells ;
these are not only large, but exist also in greater numbers than are usually met with in
an equivalent area of the human type.

[The uncinate gyrus and optic chiasma present much the same features as the same
regions in the human type : no points of comparative value were made out in the
sections.]

PART II.
THE HISTOLOGY OF THE PITUITARY GLAND. (PI. II. and Text-figs. 7, 8, 9.)

The pituitary gland of the Weddell seal is a body of considerable histological
interest, inasmuch as all three portions are well developed, and in addition there occurs
upon its upper aspect a small portion of tissue the structure of which is not represented
in the human pituitary.*

The gland (text-fig. 7) is a large one, and is made up of three main portions, viz.

* It is possible that this structure is represented in some of the lower types by the sacci vasculosi found in
connection with the pituitary gland.

(ROY. soc. EDIN. TRANS., VOL. XLVIII., 862.)

HISTOLOGY OF THE CENTRAL NERVOUS SYSTEM OF THE WEDDELL SEAL. 155

a large anterior lobe, a narrow but well-defined intermediate part, and a posterior lobe ;
the hinder portion of the intermediate mass is bent at an angle upon the rest of that
part, so as to come to lie posteriorly between the lower limit of the posterior lobe
and the projecting posterior part of the anterior lobe. From the surface, the upper
part of the posterior lobe appears wedge-shaped and folded over the hinder part of
the anterior lobe .so as to cover about a third of the upper surface.

The atypical tissue on the upper surface takes the form of a small ovoid mass
folded upon itself, and the upper leaf appears to be continuous with the floor of the

Bin.

•mi

a.£

CoE
anb-l

FK;. 7. — A mesial sagittal section of the pituitary gland (Leptonychotes weddellii). (Semidiagrammatic. )

anl.l. Anterior lobe : A A, its front part ;

15 B, its posterior part (see Plate II.).
tnid.l. Intermediate portion.
post. I. Posterior lobe.

n. Neck of the infundibulum.
op.l. Optic tract.

x. Hypothetical mass on the superior aspect of

the infundibulum.
.SIM. Sinusoids.

col. Colloidal substance formed in the middle lobe.
b.v. Large blood-vessels in the posterior lobe (often

filled with colloid).

3rd ventricle ; its structure, however, in no way resembles either grey or white
matter, being made up of strands of conglomerate cells (syncytium) between which is
a considerable amount of connective tissue and some relatively large vessels (text-figs.
8 and 9). Below this body comes a layer continuous with the posterior lobe, then
the persistent cleft continues with the third ventricle and passing some way into
the " neck " of the posterior lobe, and below this again the lower part of the neck in
which a small amount of the above atypical tissue also occurs, but, as will be seen,
quite separate from the main mass.

(ROY. SOC. EDIN. TRANS., VOL. XLVIII., 863.)

156

MR HAROLD AXEL HAIG ON THE

The large anterior lobe appears to be made up of two distinct portions (histologically
distinct), viz. a front part composed of syncytial strands of cells, with well-defined nuclei
and a few intervening sinusoids, or rather capillaries, since the endothelial walls are

c.6:

71- L

FIG. 8. — Mesial section of the portion lettered x in fig. 7 (moderately magnified).

n.i. Neck of infundibulum.
p.l. Posterior lobe of pituitary.
s<. Strands of epithelial cells.
c.t. Connective tissue.

v. Large blood-vessels.
xl. Similar tissue to above seen in the
tissue above anterior lobe.

present (Plate II. B). The hind part of the anterior lobe is made up of fairly
large clumps of cells, the cell-outlines being quite distinct and the majority of them
having deeply stained cytoplasm of a somewhat granular character ; the nuclei are

SyrC

con.

FIG. 9. — A small portion of fig. 8 more highly magnified.

syn. Syncytial strand (st. of fig. 8).
con.t. Connective-tissue fibres and cells.

n. Nuclei of fibroblasts (lamellar cells).

large and distinct.* In this part many large sinusoids occur, filled with erythrocytes ;
a few of the above-mentioned deeply-staining cells occur in the front part of the anterior
lobe, but the majority in the front part are syncytial and the cytoplasm but lightly

* In these clumps, some of the cells possess much clearer cytoplasm, which is not deeply stained with eosin ;
in this respect the anterior lobe resembles that of the human pituitary gland.

(ROY. soc. EDIX. TKANS., VOL. XLVIII., 864.)

HISTOLOGY OF THE CENTRAL NERVOUS SYSTEM OF THE WEDDELL SEAL. 157

stained with eosin and clear. The intermediate portion is a typical syncytium of clear
protoplasm in which are embedded many large nuclei : colloidal substance occurs at
various points bordering on the anterior lobe (Plate II. C, and text-fig. 7, col.).

No sinusoids are present in the intermediate portion, there being, however, cleft-like
spaces between the strands of the syncytium. At the upper part of the gland the
intermediate portion seems to pass insensibly into the anterior lobe — in fact, almost
to blend with that lobe, and the above structure would thus lead to an inference
that the middle portion is a derivative of the front part of the anterior lobe, since
both these portions are syncytial in character. The posterior lobe (Plate II. D) is
made up of a large amount of neuroglia in which large nuclei occur (nuclei of
neuroglia cells), and also here and there small masses of colloid which are passing
through the lobe from the intermediate portion. The large vessels of the posterior
lobe (not sinusoids) are also occasionally seen to be filled with masses of colloid
in which are embedded small bi- or tri-lobed masses which are either leucocytes or
degenerating nuclei from the intermediate portion.

EXPLANATION OF PLATES.

PLATE I.

Serial sections of the medulla oblongata, pons, and mid-brain (Leptonycliotes weddellii).

1 . At the decussation at the pyramids :

S.G.R. Substantia gelatinosa Rolandi.

n.c. Nucleus cuneatus.
n.g. Nucleus gracilis.
py.'l. Pyramidal decussation.

2. Just above the pyramidal decussation :

Pi/. Pyramids.

Other lettering as in 1.

3. At the lower region of the olivary body :

C.R. Corpus restiforme.

/. Tract of the fillet.

o.n. Olivary nucleus.

.< a. Superficial arcuate fibres.

p.l.b. Posterior longitudinal bundle.

». XII. Nucleus of 12th cranial nerve.

n. X. Dorsal nucleus of 10th cranial nerve.

4. At the mid-olivary region :

i.a. Internal arcuate fibres,
n. IX. Nucleus of the 9th cranial nerve.
n.d. Part of Deiter's nucleus.

(ROT. soc. EDIX. TRASS.,

X. and XII. Issuing fibres of the 10th and

12th cranial nerves.
Other letters as iu preceding
figures.

5. At the upper olivary region :

IX. and VIII. Issuing fibres of the 9th and 8th

cranial nerves.
a.n. Arcuate nuclei.

6. Across the lower pontine region :

'•. 4- Cavity of the 4th ventricle.
VII. and VIII. Issuing fibres of 7th and 8th cranial

nerves.

D. VII. Nucleus of 7th cranial nerve.
p.n. Pontine nuclei.
f.p. Pontine fibres.
f.r. Formatio reticularis.
s.o. (?) Superior olivary nuclei.
bi. Trapezium.

7. Across the middle of the pons :

n. V. Nucleus of 5th cranial nerve (motor
nucleus).

VOL. XLVIII., 865.)

158 HISTOLOGY OF THE CENTRAL NERVOUS SYSTEM OF THE WEDDELL SEAL

V. Issuing fibres of 5fch nerve.

s.e.p. Superior cerebellar peduncle.

p.l.b. Posterior longitudinal bundle.

/. Tract of the fillet.

8. Across the upper region of the pons :

IV. Intercrossing fibres of 4th cranial

nerves.

n. IV. Root-bundles of 4th nerves.
s.p.d Decussation of superior peduncles

of cerebellum.
/. Fillet.

9. Across the posterior corpora quadrigemina
(mesencephalon) :

Sy. Aqueduct of Sylvius.
f.ret. Reticular formation.

p.l.b. Posterior longitudinal bundle.
s.c.p. Superior peduncles of cerebellum.
fify Intermediate and lateral fillet.

t.d. Decussation of trapezium.
i.p.g. Interpeduncular grey matter.
n. III., n. IV. Nuclei of the 3rd and 4th cranial

nerves.

p.p. Crusta (pes pedunculi).
s.n. Substantia nigra.
C.Q.p. Posterior corpus quadrigeminum.

10. Across the anterior corpora quadrigemina :

p.c Posterior commissure of brain.
C.Q.a. Anterior corpus quadrigeminum.
n. III. Nucleus of 3rd nerve.
/3. Issuing fibres of 3rd nerve.
/. Fillet.

PLATE II.

A. Portion of the posterior part of the anterior

lobe of the pituitary gland (Leptonychotes
weddellii). x 500:

g. Large granular cells, staining deeply

with eosin.

sin. Sinusoidal blood-spaces, some of
them containing erythrocytes,
and occasionally showing an
endothelial lining (n.ep.) indi-
cative of a true capillary.

B. Portion of the anterior part of the anterior

lobe of the pituitary gland, x 500 :

syn. Syncytium, including here and
there large isolated cells typical

of the posterior part of the
anterior lobe (g).
v. True capillaries.

C. Part of the pars intermedia of the pituitary

gland, x 500:

syn.p. Syncytium = fused spindle-shaped
cells with nuclei (n.) embedded
in the protoplasm.

col. Colloidal substance at the edge of
the pars intermedia.

D. Part of the posterior lobe of the pituitary gland :

n.f. Neuroglia fibres.

n. Nuclei of neuroglia cells.

col. Isolated colloid masses.

c»ll. Large mass of colloid in a vessel (•«.).

(ROY. soc. EDIN. TRANS., VOL. XLVIII., 866.)

Scot. Nat. Ant. Exp.

HAIG : CENTRAL NERVOUS SYSTEM OF THE WEDDELL SEAL — PLATE I.

n;3-

^VTL^lVSfetr^ 3

Vol. IV.

PY

Scot. Nat. Ant. Exp.

HAIG : CENTRAL NERVOUS SYSTEM OF THE WEDDELL SEAL — PLATE II.

Vol. IV.

syn.

B

n.

?)

^WfP^^i

' *' v J>'"V^/^** » ** »\ s"*-**-1 i^^^ '* '

I)

PAKT XI.

SEALS,

PART XL-MEASUREMENTS AND WEIGHTS OF

ANTARCTIC SEALS TAKEN BY THE SCOTTISH

NATIONAL ANTARCTIC EXPEDITION.

BY WILLIAM S. BRUCE, LL.D., F.R.S.E.,

Director of Scottish Oeeanographical Laboratory, Edinburgh.

(WITH TWO PLATES AND ONE TEXT FIGURE.)

Measurements and Weights of Antarctic Seals taken by the Scottish National
Antarctic Expedition. By William S. Bruce, LL.D., F.R.S.E., Director of the
Scottish Oceanographical Laboratory, Edinburgh. (With One Text-Diagram and
Two Plates.)

(MS. received February 18, 1913. Read March 17, 1913. Issued separately August 4, 1913.)

During the voyage of the Scotia in the Weddell Sea, and during her wintering at Scotia
Bay, South Orkneys, I measured as many of the seals taken as possible. I was usually
assisted in this work by Mr WILTON. When a seal was killed near the ship, and it could
be arranged, I also weighed them both in whole and in part, weighing the different viscera
as at a human post-mortem examination. The results of these investigations are now
summarised in tabular form, which makes it easy at a glance to make comparisons. In
addition to the measurements and weights of the true seals of the Antarctic regions,
those of a fine specimen of Otaria jubata (the Patagonian Sea-lion) secured at the
Falkland Islands are also given.

Leptonychotes weddelli (the Weddell Seal).

Of the true seals twenty-seven of the thirty-three specimens of Leptonychotes
weddelli taken were measured, and, except in a few cases, in great detail, and several
were weighed in whole or part. The longest Weddell Seal was a female measuring
129^ inches and weighing 908 Ibs., while the heaviest, also a female, measured 124 inches
and scaled 920 Ibs., or about 940 Ibs. counting loss of blood, etc., in cutting up. Both
these were secured towards the end of September. Another female, killed on 28th August,
was also of large size, but as it was killed three miles from the ship its weight could not
be taken, neither was it measured. The largest male secured was 117 inches long.

Eleven adult females were taken and six adult males ; six young females and ten
young males. It is doubtful whether this indicates any real proportion of sexes. It
may be due to the males going further afield than the females, especially just prior to
and after the birth of the young. Many hundreds if not thousands of these Weddell
Seals could have been obtained, but science did not demand that extensive slaughter
which sealing expeditions indulge in.* I made a point of not allowing more killing
than was necessary for food supply and for scientific purposes. Weddell Seals were
never seen in the pack, except in the vicinity of Coats Land and the South Orkneys.

It will be noticed that about 250 Ibs. of blubber may be obtained from a single
seal, i.e. more than a quarter of the weight of the whole animal. The enormous weight

* Fully 6(KX) seals of the four Antarctic species were killed by the Balcma in 1892-93. They were mostly Crab-
eaters and Sea-leopards. Upwards of 25,000 were killed by four Dundee whalers and one Norwegian whaler at
that time.

Thanks are due to the Executive Committee of the Carnegie Trust of the Universities of Scotland for defraying
the expenses of this paper and that entitled " Skulls of Antarctic Seals," recently published.

(REPRINTED FROM THE TRANSACTIONS OK THE ROYAL SOCIETY OF EDINBURGH, VOL. XLIX., PP. 567-577.)

164 DR WILLIAM S. BRUCE ON

of 50 Ibs. for the stomach and contents of one of these Weddell Seals shows the heavy
meal, which usually consist of fish (mostly Notothenia coriiceps), that seals can digest.

Stenorhynchus leptonyx (the Sea-leopard).

Only seven Sea-leopards were secured, three males and four females, the longest of
which was a female measuring 155 inches. On the Balsena in 1892-93 I obtained one
measuring 162 inches. Two males of average size, 128 and 134 inches, scaled respec-
tively GOG and 630 Ibs. They are thus by measurement longer and lighter animals
than the Weddell Seals, and they give this appearance. They are extremely lithe in
their movements, jumping great distances out of the water. They catch swift penguins
in the water. On the Balsena I frequently found stones in their stomachs, which are, I
believe, derived from the stones in stomachs of the penguins they devour.

Dr PIRIE during the voyage of the Scotia found sand in their stomachs, which is
probably derived from the same source. In four of the specimens obtained penguin
feathers and remains were found, and on one occasion a Sea-leopard was seen in Scotia
Bay to catch a penguin and take it under water. The penguin undoubtedly forms the
staple diet of these animals, and the size and nature of the teeth is indicative of such
diet. Only on one occasion did we see a young Sea-leopard, and this was early in
November in Brown's Bay, off Point Thomson, when Mr WILTON, Dr PIRIE, and I tried
to secure it, but failed on account of the rotten state of the ice.

Lobodon carcinophaga (the Crab-eating or White Antarctic Seal).

We only obtained five specimens of the Crab-eating Seal during the voyage of the
Scotia, although many of these were seen on the voyage southward in 1903 at a time
when we were unable to secure them. In 1892-93 the Dundee whalers secured a great
number of these Lobodons, and 1 have seen as many as forty on a single piece of ice.
This is probably the smallest of the four species of Antarctic Seals, although the Ross
Seal is of very similar dimensions. One very old male that we secured weighed 494 Ibs.,
but the other three were very much smaller and lighter than this animal. This animal
is one of the specimens that I have presented to the Royal Scottish Museum. The skin
of one of these animals was very badly scarred, and it is uncertain what is the cause
of these scars. They may possibly be due to the attacks of a grampus, but we have
no definite evidence of this. We had little evidence on the Scotia to tell us what was
the food of these Lobodons, but on the Kalxna I found the remains of fish and
crustaceans in their stomachs.

Ommatophoca rossi (the Ross Seal).

This is the rarest seal in the world, and the Scotia naturalists only obtained two
specimens of it, one male and one female, the skins of which have been deposited in the
Royal Scottish Museum, and the skeletons in the Anatomical Museum of the University

(ROY. soc. EDIN. TRANS., VOL. XLIX., 568.)

MEASUREMENTS AND WEIGHTS OF ANTARCTIC SEALS.

165

of Edinburgh. It is the very antithesis of the Sea-leopard ; instead of being long and
lithe, it is rather short and bulky, though of remarkably graceful form. It is charac-
terised by a very thick neck, which it has the power of dilating at will. The weight
of the female we obtained was 475 Ibs., and that of the male 400 Ibs., and probably
this represents the average weight.

On board th&J$ala0na Mr BURN MURDOCH and I obtained what was almost certainly a
very young Ross Seal, which we were bringing on board alive, but which the late Captain
FAIRWEATHER ordered to be killed. The skin was thrown into a heap with the rest, and
the carcase was left on the ice. The illusions of ignorance consequently robbed science
for the time being of an interesting discovery. The chief food of the Ross Seal is
cuttle-fish, and the stomach of one of those we secured had the beaks and remnants of
cuttle-fish in it. It must, therefore, be swift in the water to catch these swift inverte-
brates, but its teeth are of extremely small and delicate character, which is all that
is necessary for such soft diet. There were apparently fish scales in the stomach of one
of the animals, but these were probably the scales of small fishes.

Length- Tip of Nose to Tip of Tad.

Tota.1 Length -Tip of Nose to Tip of Hind

Flipper

NecK Girth
R>st to Occiput

Axillary
Girth-

Greatest
Girth

Anal

or

Least Girth

Anterior
or

Neck Girth

Diagram illustrating Seal Measurements.

Otaria jubata (the Patagonian Sea-lion).

Little need be said regarding this well-known species which, as already indicated,
is not Antarctic in distribution, keeping clear of the ice limit. There are a considerable
number of these animals round the shores of the Falkland Islands, where they live
among the thick tussock grass.

(BOY. soc. EDIN. TRANS., VOL. XLIX., 569.)

166

DR WILLIAM S. BRUCE ON

Leptonychotes weddelli

LENGTH.

GIRTH.

LIMBS.

EYE.

s

No.

DATE.

SEX.

AGE.

- ~

•H

O

A
H _;

1|U

*« £•

S SJS

&>

4^

§

"3
q

Fore Flipper.

Hind Flipper.

nd Hori-

ameter.

00

3
1

43 . ™

«

*>."&

°£

S "3

a

C3

£

0

3> CJ

<* rt £
fe a> JS « £P

t-i •

i. a).

* *3 g

^S

2

O

Is

3£

B

O

to

E c •")

5'" o
o <o *s

•^^

"

O

CO

J

O W

3 SO

S T3
1— 1 —

-2 1 3 ^4e

|a|OH

ii

X' H p^

13
s §
> «>

4>

t-^H

1903.

in.

in.

in.

in.

in.

in.

in.

in.

in

in.

in.

in.

in.

in.

VI.

Mar. 25

Female

Adult

1141

100

54

64

14- x 1

44-

VII.

VIII.

Apr. 10

Male

Young

79

67

401

47

19J

Hf

13}

12

10}

Lixf

3|

(base)

IX.

May 19

„

ii

781

671

37*

48

48

121

91

15^

13

10

f xl

31

(base)

Xa.

„ 30

Adult

X.

July 8

Female

Young

841

741

3H

431

541

...

13i

7|

7*

15

14*

*

20}

Frozen

I

XL

„ 12

Male

Adult

114

100

62

741

80

...

16

7

H

19

16

24

W

3f

XII.

,, 23

n

Young

981

881

49

60

651

iH

vj

M|

Hi

m

• • •

35

XIII.

„ 25

102

90

XIV.

,, 28

Female

Adult

La

ree

XVII.

Aug. 31

Male

Young

54

47

19

25

28

10f

4

6J

2|

7i (base)

XVIII.

„ 31

601

511

34|

341

n*

6*

7*

13j

12

13}

3}

10i (base)

XIX.

Sept. 2

...

»

64

55

241

29

34

"i

7

6

-,

13

H

3

(ROY. soc. EDIN. TRANS., VOL. XLIX., 570.)

MEASUREMENTS AND WEIGHTS OF ANTARCTIC SEALS.

167

(the Weddell Seal).

WEIGHT.

Total.

Liver.

Kidneys.

Lungs.

Heart.

Right.

Left.

Right.

Left.

Ib.

Ib. oz.

Ib. oz.

Ib. oz.

Ib. oz.

Ib. oz.

...

Ib. oz.

220

278
787

16 13

0 11|
21b.

0 10
9 oz.

1 7
3 12

1 3
3 1

1 4
3 4

...

5H
83

85

...

...

GENERAL RKMAKKS.

Shot by D. W. WILTON in Buchan Bay, South Orkneys. Skin
presented to the Natural History Museum, Marischal College, Uni-
versity of Aberdeen.

Killed by the cooks, FLORENCE and MURRAY, in Scotia Bay. Preg-
nant. Embryo and skull preserved. Embryo presented to the
University of Edinburgh.

Clubbed by R. DAVIDSON in Scotia Bay. Temperature 97°-4 F.
Skin poor, but preserved. Additional measurements : umbilicus to
penis, 1\ inches; penis to anus, 11| inches.

Stabbed by a seaman in Jessie Bay, South Orkneys.

Killed by a seaman near Ailsa Craig, South Orkneys. Skeleton
preserved, but the skull and several ribs were lost while macerating
in the sea.

Stabbed by D. W. WILTON in Scotia Bay. Skeleton almost complete.
Additional measurements : small intestine measured 60 feet, and large
intestine, 3 feet.

Stabbed by D. W. WILTON near Point Rae, Scotia Bay. Additional
weights: blubber, 250 Ibs.; skin, 80 Ibs.; carcase, 457 Ibs. Skin pre-
sented to Royal Scottish Museum. Part skeleton at Scottish Oceano-
graphical Laboratory.

Shot by J. H. HARVBT PIHIB in Scotia Bay. Skeleton preserved at
Scottish Oceanographieal Laboratory.

Stabbed by R. DAVIDSON in Scotia Bay. Skeleton preserved at
Scottish Oceanographical Laboratory.

Stabbed by D. W. WILTON near Point Rae, Scotia Bay. No
dimensions taken, but of large size. No embryo in uterus. It was
found that the right scapula had been fractured across the blade and
had united again at an angle. Skin presented to H.S.H. the Prince
of Monaco, for Musee Oceanographique de Monaco.

Killed by chloroform on 31st August. Age one day. Born about
30th August. Carcase presented to the Anatomical Museum of the
University of Edinburgh, and partly dissected for vascular system.
Skin at Scottish Oceanographical Laboratory.

Killed with potassium cyanide on 31st August. Age two days.
Born about 29th August. Injected and embalmed by Professor
DAVID HEPBURN'S method, by whom it has been dissected and
specially reported upon. Additional measurement : tail 4| inches in
length.

Killed with potassium cyanide on 2nd September. Age probably
three or four days. Young of No. XX. Carcase presented to Ana-
tomical Museum of the University of Edinburgh. Skin presented to
Glasgow Corporation Art Galleries and Museum.

(ROT. soc. EDIN. TRANS., VOL. XLIX., 571.)

168

Dll WILLIAM S. BRUCE ON

Leptonychotes iveddelli

LENGTH.

GIRTH.

LIMBS.

EYE.

0)

u

CO

OH .

.,

••-

No.

DATE.

SEX.

AGE.

H o

0 *

jj p |

•M

O

a,

0 t-

aj o _^

Z; '£ >>

CD _

4J

08

Fore Flipper.

Hind Flipper.

1-2

i-H 0)

— £

1

Is

1! 1

"t? *4-H

H ,J

o '3

CD
H§

M -" c3

o a a

* fflT=

1|3
if*

M

CO

S

5

6

O
v>

J

•§ &D

~ ro

"~" r—

Jw

"S '5 c
1 §-1

S <D

« te

3 rO
^**^ rrl

t-.

S tf-

^s

-« _ 0)

CO C3 01

J3.5 ^C

s g i

2 g's

-U r~. r^>

* a
afi

l-a
•fil

Oi O

Interorl:

H =

•<"

ajH^

W2H(X,

[> «

1903.

in.

in.

in.

in.

in.

in.

in.

in.

in.

in.

in.

in.

in.

in.

XX.

Sept, 2

Female

Adult

123

110

57

74

75

...

151

7J

...

19

17 '

...

4f

XXI.

„ 7

Male

Young

61*

54

23£

33

34J

I0|

5

7f

12

10i

14|

l»«l

4

9J (base)

XXII.

„ 8

66£

58i

20f

29|

31i

13

*i

7

13

10

15

Hxi

3

9 (base)

XXIII.

„ 20

Female

61

53^

18J

26

28

13

TJ

6

13

13

16

3

8 (base)

XXIV.

21

Adult

124

XXV.

7 7

„ 22

Young

66

58

26

28

12

7*

13

3

8J (base)

XXVI.

,, 24

•'

Adult

129£

115J

55$

72

77

m

Ti

7*

20

18J

19

...

1

XXVII.

Oct. 3

Male

Young

62

54

19

26

27

10

6

T|

14

11

5|

3

10 (base)

XXVIII.

3

Adult

XXIX.

„ 22

Female

1211

108

72

XXX.

„ 22

Male

" * 4

117

65

XXXI.

„ 22

Female

122

41

59

18

15i

23

H

(base)

12.5 (base)

XXXII.

„ 23

"

Young

79

67

...

48

48

...

15

81

81

15

l»i

201

5

* Stiff wlien measured, probably more.
(ROY. soc. EDIN. TRANS., VOL. XLIX., 572.)

.MEASUREMENTS AND WEIGHTS OF ANTARCTIC SEALS,
(the Weddell Seal) — continued.

169

WEIGHT.

Total.

Kidneys. Lungs.

Liver.

Right.

Left. Right. Left.

Heart.

GENERAL REMARKS.

Ib.
838

83

84

61

920

71

908

Ib. oz.

207

Ib. oz.

Ib. oz.

Ib. oz. Ib. oz.

Ib. oz.

Killed off Point Davis, Scotia Bay. Mother of No. XIX. Ad-
ditional weights : blubber, 250 Ibs. ; skin, including skull and flippers,
228 Ibs. Carcase, less head and flippers, 340 Ibs. ; estimated weight
of blood lost, 20 Ibs. Skin presented to Peterhead Museum. ?

Killed with potassium cyanide. About two days old. Skeleton
preserved. Skin presented to Natural History Museum, Marischal
College, University of Aberdeen.

Stabbed. Two or three days old. Carcase preserved. Additional
weights: 74 Ibs., and estimated blood lost, 10 Ibs The mother
attempted to kill this animal, and would probably have done so had
she not been prevented.

Captured alive in Scotia Bay. Probably twelve days old. This
animal was kept on board for ten days, and was fed by bottle with
condensed milk. It died of convulsions.

Stabbed by J. FITCHIB off Point Martin, Scotia Bay. About 20 Ibs.
for loss of blood in cutting up and stabbing should be allowed, making
the total weight nearer 940 Ibs. No young with it, and no embryo j
within. Skin presented to the Glasgow Corporation Art Galleries and
Museum. Skeleton at Scottish Oceanographical Laboratory.

Killed with potassium cyanide and hydrochloric acid on 22nd
September. One day old. Taken from mother 21st September.
Carcase preserved and skull partially severed to inject brain with
formalin. Skin presented to the Royal Scottish Museum.

Stabbed by J. MACDOUGALL off the Point Martin, Scotia Bay.
Mother of No. XXVII. Young captured and taken on board alive
at same time. Skin preserved.

Two to three weeks old (nine days spent on board). It died on
3rd October. The mother was No. XXVI. Presented to Coats
Museum, Paisley.

Killed by J. H. H. PIBIE while at camp off Graptolite Island, South
Orkneys. Skull only preserved.

Killed by R. DAVIDSON off Delta Island (near Point Rae), Scotia
Bay. Skin presented to Coats Museum, Paisley.

Killed by R. DAVIDSON off Delta Island (near Point Rae), Scotia
Bay. Skeleton preserved. Lower jaw fractured.

Shot by R. DAVIDSON.
brought on board alive.
Dundee.

A young cub was with this seal and was
Skin presented to Municipal Museum of

Six to eight weeks old. Shot by J. H. H. PIRIB in Jessie Bay. This
young seal was quite alone. The skin was handsomely marked but
was spoilt by giant petrels, and the skeleton by dogs.

(ROY. soc. RDIX. TRANS., VOL. XLIX., 573.)

170

DR WILLIAM S. BRUCE ON

Leptonychotes weddelli

No.

DATE.

SEX.

AGE.

LENGTH.

GIRTH.

LIMBS.

EYE.

Interorbital Space.

(^ .

'S~\ =-i

C™^ 0^

® r3

Cfi ^1J

.°

II 1

1?

o

O c3

a>
m
o

Anterior or .Neck
(6 ins. anterior
to Axillary).

b

jS

'H

Greatest.

"c8

h

O

Fc

fj3
OW

in.

re Flipper.

Hind Flipper.

Vertical and Hori-
zontal Diameter.

II

^ 13 o>

-5-2 =c
•*-a S ^

£ SH

.2 ct

O W

in.

in.

Stretch at
Terminal
Phalanges.

1903.

in.

in.

in.

in.

in.

in.

in.

in.

in.

in.

in.

XXXIII.

Oct. 29 Male

Young

764

67

30

49

...

11

6|

n

15

13|

11 (base)

3

XXXIV.

„ 29 Female

»

79

684

29

50

...

...

1H

7i

8

164

15|

8

lOJ(base)

••

3|

XXXV.

XXXVI.

XXXVII.
XXXVIII.

XL.
XLI.

XLIV.

„ 29

Nov. 23
„ 23
„ 23

1904.
Feb. 17

„ 17

Mar. 10

Male

Female
Male

»)

»
Female

»

Adult

»
M

II

))

72

62

29

51

1 O 1

1. £i~K

6

8

16

14

11 (base)

3*

...

Lobodon carcinopliaga

1903.

I.

Feb. 2

Male

Adult

97

88

51

531

154

,5j

134

.. .

IV.

,, 10

Female

»

89

814

254

49

53

16

7i (base)

HI

134

...

U+4

4|

V.

XV.

XVI.

1

Mar. 3

Aug. 1
August

Male

))
»J

Young

»)

109

98

-.

55

58

...

19

20

...

17

17

22

"*'

5

(ROY. soc. EDIN. TRANS., VOL. xux.. 574.)

MEASUREMENTS AND WEIGHTS OF ANTARCTIC SEALS.

171

(the Weddell Seal) — continued.

WEIGHT.

otal.

Liver.

Kidneys.

Lungs.

Heart.

Right. Left.

Right.

Left.

Ib.

lb.

lb. oz.

lb. oz.

lb. oz.

lb. oz.

lb. oz.

830

36

•2 7

2 7

5 8

4 12

4 2

(the Crab-eating Seal).

4 to

5 ewts.

300

(fina)

494

325

(circa)

0 14 I 0 14

Length =$lins.
greatest width = 31

1 11

1 6

1 12

REMARKS.

Shot by R. DAVIDSON in Scotia Bay. About two months old.
Additional measurement, from anus to penis, 13 inches.

Shot by R. DAVIDSON in Scotia Bay. About two months old. Skin
presented to Glasgow Corporation Art Galleries and Museum.

Shot by R. DAVIDSON in Scotia Bay. About two months old.
Additional measurement, from anus to penis, 12£ inches. Skin pre-
sented to Coats Museum, Paisley.

Shot at Point Martin, Scotia Bay.
' dimensions taken.

Skeletons preserved. No

Shot by R. N. R. BROWN on west shore of Scotia Bay. Skeleton
preserved.

Shot by D. W. WILTON on west shore of Scotia Bay. Skin and
skeleton preserved. Skin presented to Museum of Perthshire Society
of Natural Science, Perth.

Shot by W. S. BRUCB, 2 miles off Coats Land, 74° 1' S., 22° W.
Additional weights and measurements : stomach and contents (fish and
cuttle-fish), 50 Ibs.; spleen, 5 Ibs.; diaphragm, 4 Ibs. 6 ozs.; gut, 43 Ibs.;
and measured 111 feet. Skin presented to Royal Scottish Museum.

Shot by J. H. H. PIRIB at Station 152, 60° 32' S., 43° 40' W. Skull
preserved. Additional measurements : from anus to penis, 16f inches ;
from penis to umbilicus, 9 inches. Weight between 4 and 5 cwts.
Skin preseuted to Perthshire Society of Natural Science, Perth.

Shot by J. H. H. PIBIE at Station 189, 60° 05' S., 32° 10' W.
Skin and skull preserved. Additional weights and measurements :
anus to umbilicus, 22£ inches ; large intestine up to caecum, 40 inches :
oecum, 5x3 incites; small intestine, 122 feet; thymus, 2 ozs.; spleen,
6£ ozs.; pancreas, 6£ ozs.; blubber, 1 to 1 J inches thick. Skin presented
to the Royal Scottish Museum.

Shot by J. H. H. PIRIE at Station 283, 68° 24' S., 323 31' W. This
was an old animal, and consequently may be regarded as being of
full size. Blubber, 147 Ibs. ; skin, 47 Ibs. Skin presented to the
Royal Scottish Museum.

Killed near the ship in Scotia Bay. Skinned scarred.

Killed by J. H. H. PIRIE off Point Davis, Scotia Bay. Skin very
beautifully marked, but not preserved.

Skeleton secured.

(ROY. soc. EDIN. TRANS., VOL. XLTX., 575.)

172

DR WILLIAM S. BRUCE ON

Stenorhynchus leptonyx

LENGTH.

GlETH.

LIMBS.

EYE.

XI

No.

DATE.

SEX.

AGE.

fi

•* ~

QJ """"

§^

11

~ K

JN

"S
_•&

2rt
t_j

01
CO

O

Anterior or Neck
(6 ins. anterior
to Axillary).

I

Greatest.

~a

h

O

Fore Flipper.

Hind Flipper.

Vertical and Hori-
zontal Diameter.

Illterorhital Spac

O W

I!

Stretch at
Terminal
Phalanges.

& SJ3

^ -i

<U w

Stretch at
Terminal

Phalanges.

—

1903.

in.

in.

in.

in.

in.

in.

111.

in.

in.

in.

in.

in.

in.

in.

III.

Feb. 9

Female

Adult

160

138

50

68

68

28J

22|

13 J (base)

22

20

7 j (base)

8J

1904.

XLII.

Feb. 26

Male

132£

120

36*

60£

59

f)O 1

-. J-fi

Oil
2

37

133 (base)

8

XLV.

Mar. 18

)j

134

37*

49

30

f) ~[ 1

22^

26

14 (base)

XXXVIIIa

Jan 7

Female

138

XXXVIIB

7

136

XXXVIIIc

30

Male

XXXVIIW.

Feb. 5

Female

»

140J

Ommatophoca rossi

1903. ;

II.

Feb. 6

Female

Adult

...

90

49

52

54

. . .

16

8i (base)

16

14J

25

Hxl

6

41*

1904.

XLIII.

Feb. 28

Male

II

100^

101 (ven-

92 (ven-

38J*

55^

56J

17i

...

18|

18f

24

11 (base)

Hxf

6J

tral)

tral)

Oft

iria j'

ibata

1904.

XXXIX.

Feb. 6

Male

Adult

128J

...

62

66J

8H

(shoul-

...

35£

11

19

25

OQ 1
"2

14

...

...

ders)

* Neck immediately posterior to occiput.
(ROY. soc. EDIN. TRANS., VOL. XLIX., 576.)

MEASUREMENTS AND WEIGHTS OF ANTARCTIC SEALS.

173

(the Sea-leopard).

WEIGHT.

Total. Liver

Kidneys.

Right.

Left.

Lungs.

Eight.

Heart.

Left.

GEXBRAL REMARK*.

Ib. Ib. oz.

Ib. oz.

Ib. oz.

Ib. oz.

Ib. oz. Ib. oz.

606 ... 1 7

1 9

2 12

630

(the Ross Seal).

475
400

(the Patagonian Sea-lion).

Shot by J. H. H. PIRIE at Station 183, 59° 42' S., 34° 13' W.
Anus to umbilicus, 35 inches ; tail, 5 iuches. Skin and skull preserved
and presented to the Eoyal Scottish Museum.

Shot by J. H. H. PJBIB at Station 387, 65° 59' S., 33° 06' W.
Stomach empty ; no parasites in stomach ; gut not examined.
Additional measurements and weights : entire gut, 54 feet ; rectum,
2 feet ; penis and testes weighed, 1 Ib. 4 ozs.; weight of body, 317 Ibs.;
skin, flippers and head, 286 Ibs.; blood about 3 Ibs. Anus to penis,
22 J inches ; anus to tip of tail, 9 inches. Mouth, tip of chin to gape,
8'1 inches. Skeleton presented to the Anatomical Museum of the
University of Edinburgh. Skin presented to the Museum of the
Perthshire Societv of Natural Science, Perth.

Shot by J. H. H. PIRIE, Station 417, 71° 22' S., 16° 34' W. Penis
to anus, 22 inches : anus to tip of tail, 10 inches ; from eye to gape, 4£
inches ; from angle of lower jaw to gape, 7| inches ; penis to umbilicus,
9 J inches ; tip of lower jaw to umbilicus, 77£ inches. Cast of skull
made. Nothing was found in stomach, but there were penguin feathers
in the gut. Skin presented to the Royal Scottish Museum.

Shot by J. H. H. PJRIE on the Beach, Jessie Bay. Uterus empty.
Stomach contained much sand, many penguin feathers, and a few
nematodes. Changing coat.

Shot by J. H. H. PIRIE on the Beach, Jessie Bay. Uterus empty.
Gut, 92 feet. Stomach contained large quantity of shrimp-like
crustaceans, too far digested to recognise ; one set of penguin tail
feathers. Numerous nematodes and castodes in intestine.

Shot by J. H. H. PIRIK on the Beach, Jessie Bay. Stomach con-
tained large quantity of small crustaceans.

Shot by J. H. H. PIRIB on the Beach, Jessie Bay. Uterus empty.
Stomach contained penguin remains.

Shot by J. H. H. PIRIK at Station 165, 60° 06' S., 43° 00' W.
Tail (webbed), 5 inches. Skull and skin preserved. Skin presented
to the Royal Scottish Museum.

Shot by J. H. H. PIRIE at Station 393, 66° 21' S., 28° 30' W.
Entire gut measured 268 inches ; from tip of lower lip to umbilicus, 57i
inches : umbilicus to penis, 4J inches ; penis to anus, 21 inches ; anus
to tip of flipper, 17J inches ; mouth from tip of chin to gape, 3| inches.
Cuttle-fish beaksand~cuttle-fish,aud apparentlyscales of fishes instomach.
Gut full of tape-worms. Skin presented to Royal Scottish Museum.

1150

Shot on Lower Tussock Island, Port William, Falkland Islands,

(circa)

Station 349. Additional measurements: from anus to penis, 12 inches ;
from penis to umbilicus, 1 1 inches ; chest between fore flippers,
31 iuches. Skeleton in Scottish Oceanographical Laboratory. Skin
presented to Royal Scottish Museum.

(ROY. soc. EDIN. TRANS., VOL. XLIX., 577.)

Scot. Nat. Ant. />/•.

VOL. IV.

DR AV. S. BRTJCK: "Measurements and Weights of Antarctic Seals taken by the Scottish National Antarctic Expedition." — PLATE I.

[Photo by W. S. Bniet.

1. Leplonycholes weddelli (Weddell Seal).

[Photo by R. X. R. Broirn.
2. Lobfxion cardnophruja (White Antarctic Seal).

•J-l.:'nbyJ.B. n.Pirie.
3. Stenorhynehus leptonyx (Sea-leopard Seal).

[Photo by W. S. Bmir.

4. Ommatophoca rossi (Ross Seal).

Scot. Nat. Ant. Exp.

You IV,

DR W. S. BRUCE: " Measurements and Weights of Antarctic Seals taken by the Scottish National Antarctic Expedition ." — PLATE II.

[Phuto b<i Rowland Ward.

Olariajubata (Patagonian Sea-lion) in Royal Scottish Museum, from Tussock Island, Port William, Falkland Islands.

Weight, about 1150 pounds.

PART XII.

SEALS.

XII.-ON THE SKULLS OF ANTARCTIC SEALS:
SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

BY WILLIAM S. BRUCE, LL.D., F.R.S.E.,

Director of Scottish Oceanographical Laboratory.

(WITH FIVE PLATKS.)

On the Skulls of Antarctic Seals : Scottish National Antarctic Expedition.
By William S. Bruce, LL.D., Director of the Scottish Oceanographical Labora-
tory. (With Five Plates.)

(MS. received March 13, 1913. Read May 5, 1913. Issued separately June 27, 1913.)

Although the osteology of Antarctic seals has been very completely discussed,
notably by the late Dr J. E. GRAY, Sir WILLIAM TURNER, and Professor ROBERT
THOMSON, yet the literature regarding the subject is somewhat scattered. I have
therefore considered that it might be important from the point of view, as it were, of
an index to publish a complete series of photographs of a set of the skulls of seals
taken by the naturalists of the Scotia during the Antarctic voyage of 1902-1904.

The species considered are :—

1. Leptonychotes Weddelli (Gill) : The Weddell Seal.

'2. Stenorhynchus leptonyx (F. Cuvier) : The Sea-leopard.

3. Lobodon carcinophaga (Gray) : The Crab Eater, or White Antarctic Seal.

4. Ommatophoca Rossi (Gray) : The Ross Seal.

5. Otaria jubata (Forster) : The Patagonian Sea-lion.

The type collection of the seals' skulls taken by the Scotia naturalists are chiefly
housed in the Anatomical Museum of the University of Edinburgh, and in the Scottish
Oceanographical Laboratory ; those housed in the Anatomical Museum of the Uni-
versity of Edinburgh have been duly recorded in Sir WILLIAM TURNER'S excellent
descriptive catalogue entitled Marine Mammals in the Anatomical Museum of the
University of Edinburgh ; while Professor ROBERT THOMSON, of Cape Town Uni-
versity, has contributed a paper to the Transactions of the Royal Society of Edinburgh
entitled " Osteology of Antarctic Seals," which also appears in the Scientific Reports
of the Voyage of S. Y. "Scotia" volume iv.

Beyond this it is unnecessary to say more at the present time, except to refer to
the plates published herewith. It will be observed that the skulls of the species of
each seal are shown from every possible aspect :—

From (l) The posterior aspect.

(2) The anterior aspect.

(3) The lateral aspect.

(4) The inferior aspect without the lower jaw.

(5) The inferior aspect with the lower jaw.
(5a) The superior aspect of the lower jaw.

(6) The superior aspect.

(HEI'KINTEU FROM THE TRANSACTIONS OF THE ROYAL SOCIETY OK EDINBURGH, VOL. XLLS., VI?. 345, 346.)

180 DR W. S. BRUCE ON THE SKULLS OP ANTARCTIC SEALS.

The figures speak for themselves, and, as already stated, are intended to be an
index for the use of museums and for the use of naturalists exploring in Antarctic and
Subantarctic Regions.

1 have considered the measurements and weights of Antarctic seals in the flesh in a
separate paper ; the anatomy of the Weddell Seals taken by the Scotia has been fully
considered by Professor DAVID HEPBURN and Dr HAROLD AXEL HAIG ; while Dr R. N.
RUDMOSE BROWN has dealt with "The Habits and Distribution of the Seals of the
Weddell Sea."*

* Vide Scientific Reports of the Voyage of S.Y. "Scotia," vol. iv., parts ii., iii., v., vi., ix., x., xi., xii., an<l xiii.

(ROY. SOC. EDIN. TRANS., VOL. XLIX., 346.)

Scot. Nat. Ant. Exj>.

VOL. IV.

BRUCE: "Skulls of Antarctic Seals." — PLATE I.

LEPTONYCHOTES WEDDELLI.

Scot. Nat. Ant. Exp.

VOL. IV.

BRUCE: "Skulls of Antarctic Seals."— PLATE II.

STENORHYNCHUS LEPTONYX.

Scot. Nat. Ant. Erp.

VOL. IV.

BRUCE : " Skulls of Antarctic Seals." — PLATE III.

LOBODON CARCINOPHAGA.

Scot. Nat. Ant. AV/>.

VOL. IV.

BRUCE: "Skulls of Antarctic Seals."— PLATE IV.

OMMATOPHOCA ROSSI.

Scot. Nat. Ant. Exp.

VOL. IV.

BRUCE: "Skulls of Antarctic Seals." — PLATB V.

OTARIA JUBATA.

PAET XIII.

SEALS.

SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

XIII. -THE SEALS OF THE WEDDELL SEA:
NOTES ON THEIR HABITS AND DISTRIBUTION.

BY R. N. RUDMOSE BROWN, D.Sc., University of Sheffield ;

Naturalist, Scottish National Antarctic Expedition.

(WITH NINE PLATES.)

The Seals of the Weddell Sea : Notes on their Habits and Distribution. By
R. N. Rudmose Brown, D.Sc., University of Sheffield ; Naturalist, Scottish
National Antarctic Expedition. (With Nine Plates.)

(MS. received February 15, 1913. Issued separately May 20, 1913.)

During the voyages of the Scotia all four species of Antarctic seals were met with,
the Weddell Seal (Leptonychotes weddelli), the Crab-eating or White Seal (Lobodon
carcinopfwga), the Sea-leopard (Stenorhynchus leptonyx), and the Ross Seal (Ommato-
phoca rossi). We found, as all other Antarctic expeditions have found, that the
Ross Seal was the rarest of these four species and was very infrequently seen. The
Weddell Seal we found to be far the commonest, and to occur in enormous numbers
around the land. It is probably this liking of the species for the vicinity of the land which
has made it a familiar seal to most South Polar expeditions, since, wherever the ship may
winter, these seals are sure to be seen in large numbers in spring, while many stay
far from the open sea in the winter, utilising holes in the ice for access to the water.
Captain Amundsen found them numerous off the Ross Barrier. On some coasts,
however, the Lobodon is the commonest species. During the voyage of the Dundee
whalers to the east of Graham Land and adjacent seas in 1892-93, Dr W. S. Bruce,
who accompanied the Balsena, noticed that " the mottled Grey Seals were in greatest
abundance " ; " the greatest number I saw on one piece of ice at a time was forty-
seven." These seals, which Dr Bruce referred at the time to the Ross Seal, were really
Crab-eating Seals. The mistake was very pardonable considering the marked difference
between young and adult Crab-eating Seals, and the fact that at the time no living
person, except Sir Joseph Hooker, was acquainted with Antarctic seals, outside a very
few museum specimens. Dr Bruce had not, it need hardly be said, a reference library
of even the most modest dimensions on board the whaler in which he sailed, and indeed
the first photograph of an Antarctic seal was taken by Dr Bruce during that voyage-
one of a Sea-leopard, a reproduction of which is now published. Moreover, the
commercial nature of the expedition afforded no facilities for collecting, and what skins
were saved for the purposes of identification at home were taken by the captain and
cut up for leather, while the skulls that Dr Bruce had managed to collect were thrown
overboard as being an encumbrance on the ship. Under such circumstances it can
readily be understood that mistakes were liable to be made in the identifications such
as the cabinet zoologist at home would have no excuse for and would probably fail to
understand.

Exclusive of these four species of seals, there are two which, though not truly
Antarctic, yet approach the regions of pack-ice : these are the Sea-elephant
(Macrorhinus leoninus) and the Southern Fur Seal (Arctocephalus aitstraliy). The

VOL. IV. 185 1

186 DR R. N. RUDMOSK BROWN ON

Sea-elephant, which used to frequent Kerguelen, Heard, Gough, Tristan da Cunha, and
other subantarctic islands in huge numbers, is now rapidly becoming extinct. The value
of its oil was too great to suffer it to live in peace. But it is not yet a vanished species :
individuals are occasionally seen in various subantarctic islands, including Fuegia,
South Georgia, Kerguelen, Heard, the Crozets, Tristan da Cunha, Macquarie, and
Campbell Islands. During 1902 one was shot in Port William, Falkland Islands, and
early in 1903 another came ashore near Cape Pembroke lighthouse, Falkland Islands,
but escaped ; and there are other records of its occasional occurrence in the Falkland
Islands. During our visit to Saddle Island, South Orkneys, in February 1903, we sighted
what was probably a Sea-elephant, but could not get to close quarters. However, in
April 1904 a large male 13|- feet long came ashore on the beach in Scotia Bay. This
record is of interest since it is not generally considered that the Sea-elephant penetrates
the pack. In fact, the edge of the pack-ice probably limits the southern distribution
of the Sea-elephant, but variations in the distribution of the pack may mean a con-
siderable southward extension of its range in some years. However, it must be
recalled that the Discovery sighted one in MacMurdo Strait (77° 50' S.), and that
Ross in 1840-43 recorded it from the Palmer Archipelago and off Louis Philippe
Land in 65° S.

Of the Southern Fur Seal (Arctocephalus australis) we saw nothing at the South
Orkneys. It is almost certain that this seal has been exterminated at the South
Orkneys, where even in Weddell's time it was rare. " In the evening the boats
returned, having coasted these islands for fifty miles. They had found but one Fur Seal
and some Sea-leopards " (A Voyage towards the South Pole, London, 1825, p. 23). Off
Cape Dundas, South Orkneys, Weddell got two Fur Seals, but none to the westward.
At the South Shetlands in 1820 and 1821 Weddell says 300,000 Fur Seals were
killed. This kind of slaughter naturally led to its extermination, and recent expedi-
tions to the South Shetlands, as well as Norwegian, Chilian, and other whalers, have
looked for it in vain ; though the Swedish Antarctic Expedition found one, and secured
the skin, at Nelson Island, South Shetlands, in 1902. There are probably no Fur Seals
now living in true Antarctic seas. In South Georgia, where it was once found in
enormous numbers, it is extinct. At the South Sandwich group it may still rarely
occur : it certainly was found there in comparatively recent years. On the subantarctic
islands of New Zealand a few still exist.

One other seal which the Scotia secured may be mentioned here, although it is
not, and never was, an Antarctic species : that is, the Sea-lion (Otariajubata), of which
a large male was shot at the Falkland Islands in February 1904. There are several
herds of them to be seen at certain of the smaller islands of the Falkland group, notably
at the Tussock Islands in Port William. These are low-lying islands covered with thick
tufts of tussock-grass growing man-high. The Sea-lions love to lie among this grass,
and in consequence it is necessary to proceed warily, for they are formidable antagonists
and exhibit surprising agility in their movements on land. This seal well merits the

THE SEALS OF THE WEDDELL SEA. 187

name of Sea-lion, with its creditable mane and its habit of emitting a deep roar as it
rears itself on its fore-flippers. The one secured, a male, measured 10 ft. 8 ins. over
all, and scaled over 1200 Ibs.

The chief importance of the Scotia collections lies in the large number of skins and
skeletons brought home, as well as two injected specimens of young Weddell Seals.
The anatomy of the Weddell Seal and the skeletons of this and other species are dealt
with elsewhere in this series of volumes. The notes in this paper deal with the dis-
tribution and habits of the seals observed in the Weddell Sea. In addition to observa-
tions made by the naturalists of the Scotia, I have included various notes made by
Dr W. S. Bruce during the cruise of the Dundee whaler Balaena to the western Weddell
Sea in 1892-93, since these deal with the same area of Antarctic seas and supplement
the observations of the Scottish National Antarctic Expedition. It may be added that
a complete set of skins collected by the Scotia is now in the Royal Scottish Museum,
Edinburgh. Others have been distributed to the museums of Glasgow, Aberdeen,
Dundee, Paisley, Perth, and Monaco : a few are in the museum of the Scottish Oceano-
graphical Laboratory. The skulls and skeletons are in the Scottish Oceanographical
Museum and the museums of the universities of Scotland, the type collection being
in the Anatomical Museum of the University of Edinburgh.

Leptonychotes weddelli (Lesson) — the Weddell Seal.

Both at the South Orkneys and at Coats Land this was the commonest species we
encountered, and one which we had facilities for studying better than others. Fifteen
years ago it was one of the rarest of known mammals, despite its extraordinary abund-
ance in the Antarctic.

While it is very plentiful near land, it is never found in large numbers in the open pack
far from the shore : in this it contrasts with the Lobodon. No recent expedition in
any part of the Antarctic has found the Weddell Seal to be rare, and its previous rarity
was solely due to the want of expeditions for a long period, and the little careful
coastal exploration that had ever been conducted until the days of the Belgica. In
fact, the Weddell Seal haunts the coast and adjacent pack to such an extent that its
presence may almost be taken as an indication of the proximity of land.

While during the winter at the South Orkneys some of these seals undoubtedly
deserted the floe for the edge of the pack, many, on the other hand, remained. In every
winter month at the South Orkneys a certain number, principally males, were seen,
and numerous seal-holes were noticed in the ice. Dr Nordenskjold found them common
during the winter at Snow Hill (1902-1903), and Dr Charcot likewise records them as
occurring during winter around Petermann Island (1909), sometimes singly, sometimes
in groups. But none of these localities are far from the edge of the pack. However, this
species, of all Antarctic ones, goes furthest from the open water, and makes use of holes
in the ice for breathing purposes to an extent that other species do not.

188 DR R. N. RUDMOSE BROWN ON

The Discovery in winter- quarters in 77° 50' S. was some ten miles from the open
water, and yet Weddell Seals were present all winter in such numbers that Dr E. A.
Wilson did not look upon them as migratory. In that latitude blowholes are far more
numerous than holes by which the seals leave and enter the water, and Dr Wilson
ascribed this to the reluctance of the seal to face the low temperatures and darkness of
winter : the sea is certainly much warmer. Mr James Murray, of the Nimrod, says the
Weddell, which was the only common seal at Cape Royds, stayed in winter, though
several weeks might pass without one being seen. At the South Orkneys the Weddell
Seals only emerged on to the floe during calm, sunny days : in windy weather they were
seldom to be seen unless under the lee of a hummock. It must be remembered that
in the latitude of the South Orkneys, 60° 43' S., there is no continuous night in mid-
winter, and this may account for the appearance of this seal being more frequent in
winter than it is further south.

The Weddell Seal, like the Lobodon, is a true Antarctic species, but there are records
of stragglers to subantarctic lands and even beyond. The large herds of Sea-leopards
which the Challenger heard of at Kerguelen were possibly herds of Weddell Seals.*
But authentic specimens have been observed at Santa Cruz in Patagonia, Heard
Island, Kerguelen, and even New Zealand.

The Weddell is the most lethargic of all the Antarctic seals. After a good meal
they come up through the seal-holes on to the floe to lie asleep for hours. On sunny
and calm days especially are they to be seen. They sleep on their sides or almost on
their backs, every now and then giving a scratch with the uppermost fore-flipper.
When disturbed, they look at the aggressor with a sleepy stare and appear to take very
little interest in him, and in a few moments lazily drop to sleep again. It requires
somewhat drastic treatment to awake a Weddell sufficiently to make him move away,
and even then his movements are far from energetic. One can sit down on a Weddell
Seal and only cause him to open his eyes and blink : then once more he falls asleep,
unalarmed. Not only is the animal of a naturally slothful disposition (except at the
pupping season), but he has never learned to fear man in the seclusion of his polar
wastes, and he has no natural enemies on the land or ice. This fearlessness applies
equally to other Antarctic seals, and birds. So tame and lethargic is the Weddell
Seal that ammunition was never expended on one when it was desired to secure a
specimen : they were always stabbed with a long knife.

At the end of August these seals returned to the South Orkneys, and large numbers
of females then collected at the rookeries. For the previous month or two no females
had been seen, only males. The females had probably been away feeding in prepara-
tion for their long period of starvation. The rookeries are on the floe near to land, and
though they may be some little distance from the open sea when the young are born,
the breaking up of the floes generally puts them in near proximity to the water by the
time that the young are able to swim. On the other hand, the nearness of the rookeries

* Notes of a Naturalist on the " Challenger," H. N. Moseley, London, 1892, p. 174.

THE SEALS OF THE WEDDELL SEA. 189

to land would enable the mothers, in the event of the floe breaking up prematurely, to
save their young at an age when they cannot swim.

In the last days of August the first young were born. Each mother has one pup,
which at birth measures from 2^ to 3^ feet in length. They are born with their eyes
open. The young have soft grey downy coats, with no suggestion of the coarse hairs
of the adult skin, nor any of its mottled markings. They have the usual disproportion-
ate head, and their flippers, especially the hind ones, are well developed and out of pro-
portion to the rest of the body. The eyes are large, brown, and slightly bloodshot in
the white. The baby moves in the same way as its parent, by drawing itself forward
with the help of the fore-flippers, and then dragging onward the hinder part of the
body ; but naturally the movements are slow, and in following its mother any distance
it has to rest every few yards. The mother lies alternately on one side or another
while suckling the pup, and often seems to shelter it by lying to windward. Sometimes
a mother plays with her young as well as such clumsy animals can, and at times one
saw a mother presumably caressing her baby by pressing her nose against the young
one's side. The cry of the pup is something like the bleating of a lamb, but strangely
human at times.

The altruism of maternity awakens the mother at this season to an alert and fero-
cious animal who savagely resents any approach to the pup — a great change from the
sleepy placidity of other seasons. It was only rarely that a cowardly or indifferent
mother was met with. The capture of a young one, without the sacrifice of the mother,
consequently entailed a certain amount of strategy. The mother places herself between
the aggressor and her baby, and heroically endeavours to defend it. On one occasion
when the young had been secured the mother made good her escape by plunging down
a seal-hole, but at other times the mother remained on the floes, no doubt bewailing
her lot in the peculiar hoarse roar which is heard only at the breeding-season. On
another occasion a mother, on seeing our approach to close quarters, made no attempt
to drive us away, but instead attacked her pup, giving it several bites about the head
and neck. Finally she seized it in her mouth and shook it as a dog might worry a
rat. The pup was badly mauled, and would probably have been killed had we not
separated the two. The deep roar of the mother at this season replaces the hoarse
rattling cuckle, and occasionally is accompanied, in anger, by foaming at the mouth.
Males were not seen at the rookeries or on the floe until September 21st, that is, three
weeks after the pupping commenced, when one or two began to appear, but they
remained rare for some time. In no case was a mother with more than one pup noticed,
and by the end of September all the young were born. The mothers who were robbed
of their young sometimes remained on the rookeries for days before departing. At this
season they probably do not feel hunger, as they are thickly coated with blubber when
they reach the rookeries, and on this they have to subsist for about four or five weeks
until the young are ready to take to the water. No instance was noticed of a mother
leaving her young earlier and going into the sea to feed.

190 DR R. N. RUDMOSE BROWN ON

It was noticed that the umbilical cord is severed by the young seal breaking it at
birth ; it is not bitten through by the mother. A number of Sheathbills (Pagodroma
nivea) haunt the rookeries and feed on blood and excrement, while Giant Petrels
( Ossifraga gigantea) are in waiting at each birth for the delivery of the placenta. No
sign was noticed, though carefully watched for, of attempts on the part of these petrels
to attack the young seals.

In the first week of October the young seals began to take to the water, that is, when
about four weeks old. The pup gingerly slides off the ice into a seal-hole and, keeping
near the surface, stays in the immediate vicinity of the hole ; in some cases the mother
gently pushes it in. In a few minutes he clambers out again, when mother and young
greet one another joyfully. Several days later the young seal is left to shift for
himself. It is only at that period, when the young are yet a little chary of the water,
though well able to maintain themselves in it, that motherless pups are to be found.
The mothers may go off for a few hours' fishing and return, but very soon they finally
leave their youngsters. At the end of October all the young had left their mothers.
The males then begin to return, and from this time onwards were frequently seen, all
in a fat and sleek condition, and with the slothfulness characteristic of the Weddell
Seal. In the end of October and in November the males and females are often seen
together. The period of gestation is probably eight to nine months, as far as could be
judged from the size of embryos taken from females killed during the first half
of the year.

During the summer individuals with bad scars were more than once seen, though
far less frequently than in the case of Lobodons. One was seen (January 30th, 1904)
with a bare patch over the back of the skull about five inches in width, and with one eye
badly injured. Another was noticed (October 10th, 1903) in a badly scarred condition
and with his jaw broken. One with a bleeding sore on its flank was noticed trying to
force his way between two narrow ice-tongues, which might suggest that ice is a possible
cause of the scars. However, in all probability most of these injuries are caused by
fighting at the rutting season. It is quite possible that the females join in the fights ;
at any rate they are occasionally scarred, though less than the males. The relative
immunity from scars which the Weddell Seal exhibited compared with the Lobodon
agrees with the more peaceful, lazy nature of the former.

It has been suggested by more than one naturalist, writing on Antarctic seals, that
these scars are due to attacks made on the seals by the Killer (Orca gladiator), which is
common in southern seas around the edge of the pack and among loose pack. While
the smaller scars which form the majority are probably due to fighting, the Killer quite
likely may be responsible for some of the larger gashes and severer injuries. There is
certainly no direct evidence that the Killer of the southern ocean feeds on seals, but the
probability is great, and is strengthened by the fact thai the southern Killer is
probably identical with that of Arctic seas. The Arctic Killer or Grampus certainly
feeds on porpoises and seals, and has been known to attack the largest whalebone

THE SEALS OF THE WEDDELL SEA. • 191

whales.* The immunity that the Weddell Seal undoubtedly experiences from these
scars Dr Wilson attributed to its frequenting coastal waters and pack-ice, where it is
free from the attentions of the Killer, which stays in deeper and generally more open
water. To a certain extent, no doubt, this is true, but I believe that the lazy nature
of the Weddell Seal means fewer fights among themselves and so fewer injuries.

The young adopt the adult coat soon after taking to the water, and the adults
change their coats in midsummer, about January. The general colour is grey or olive-
green, often very silvery, mottled with dark or light spots. Dr Harvey Pirie, who had
opportunities to observe many cases at the summer station at Scotia Bay, and made a
study of the colouring, says : " In observing closely their coats one finds there is great
variation, no two being alike, and they seem to be in all stages of changing coats ; some
even have a complete old coat, of which the hairs can easily be pulled out. Then again
the colour varies greatly, and apart from the actual variations it differs according to
your point of view, i.e. whether you look from for'ard with the lie of the hair, abeam,
or from aft for'ard against the grain. AVhen looking aft, and to a less extent abeam,
there is a silvery sort of sheen which is quite awanting if they are viewed from aft. Some
few were of an almost uniform creamy colour with a yellowish tinge, sometimes a
greenish yellow, and in two cases a brownish yellow. In practically every one the
dorsal aspect is darker than the ventral. Mottling is practically always visible, but
where the old coat is complete it may be very indistinct. The amount and size of the
mottling also varies very much, and may take the form of small spots or of large
patches. Generally speaking, however, it comes to be light spots with a dark ground
dorsally, and dark spots with a light ground ventrally. The mottling is best seen
ventrally and about the flippers, as these are the first places to cast the old coat ; a broad
dorsal ridge is the last region to change. The coloration of the new coat is on an
average a dark slatey-grey in the dark parts and a yellowish white in the light, but the
exact shades vary very much in their intensity." (Rep. on the Scien. Res. of Voy. of
" Scotia" vol. iv., Zoological Log, p. 99.)

The food of the Weddell Seal consists entirely of fish, lamellibranchs, cuttlefish,
crustaceans, and holothurians. They seem never to attack penguins. Many attempts
which we made to induce them to do so failed. Dr Turquet, of the Frangais expedition,
writes of a Weddell Seal eating a shag after playing with it half an hour. But it is
probable that this was in reality a Sea-leopard. It is very doubtful if this seal is lithe
and quick enough in its movements to catch penguins, and hence it has never taken to
them as food.

The adult Weddell measures up to 9|- feet in length, and weighs as much as 900 Ibs.
or more.

This seal is sometimes referred to as Weddell's False Sea-leopard, owing to a remote
resemblance between the two ; but it would be wise to drop the name, since it has

* " On the History and Geographical Relations of the Cetacea of Davis Strait and Baffin's Bay," Robert Brown,
Proc. Zool. Soc., xxxv. pp. 533-556, and Arctic Manual and Instruction*, 1875, pp. 69-93.

192 DR R. N. RUDMOSE BROWN ON

helped to cause confusion in the past. The type and only specimen was for long on
exhibition in the Royal Scottish Museum, and a figure of this, taken from the stuffed
specimen, appeared in Weddell's book (A Voyage towards the South Pole performed
in the years 1822-24, James Weddell, London, 1825). That specimen was grotesquely
stuffed and a caricature of the real animal. It has now been removed from the public
galleries, unstuffed, and put away as a cabinet skin. A number of specimens from the
Scotia collections replace it.

Stenorhynchus leptonyx Cuvier — the Sea-leopard.

The Sea-leopard, without being common, was frequently seen at the South Orkneys,
in the pack, and about Coats Land. We secured three skins of adults, but no young.
It is a solitary animal and is never found in herds : three were seen together on one
occasion only, and we never saw more in company. In 1893, off Louis Philippe Land,
Dr Bruce noticed large hosts of Sea-leopards, but generally met with them in twos or
threes or solitary. At the South Orkneys it was absent throughout the winter 1903,
with the exception of three which were seen in August. Otherwise it disappeared from
March until February. In 1904 Senor Valette records Sea-leopards in Scotia Bay only
in November and December. Dr Charcot notes their disappearance from Petermann
Island in midwinter (1909), and records the last one in April.

It is impossible to say with certainty where the Sea-leopard breeds, but we have only
slight evidence that it does so at the South Orkneys. In this connection it is of interest
to note that Dr Bruce saw a young Sea-leopard on Point Thomson in Brown's Bay in
November (19th), and an attempt was made by him, Mr Wilton, and Dr Pirie to secure it.
Senor Valette records young Sea-leopards in Scotia Bay in the end of December (1904).
Wherever the rookeries of the Sea-leopards occur, the young are probably born in
September, and would be quite able to look after themselves and travel some distance
even before December. However, the probability of these young ones having travelled
from Graham Land is not great. It is far more likely that they were born at the
South Orkneys. Unfortunately, the sex of those specimens seen in August could not
be determined, as they did not emerge from the water. In any case they had not the
appearance of pregnant females. The males of no species of seal appear near the
breeding- places in early spring, and had these been males it would indicate that no
rookery existed in the vicinity. In July and August, Mr A. E. Felton informs me,
Sea-leopards are common on the beaches of the Falkland Islands, but he has seen
nothing of their young or their breeding-places.

No other expedition has been any more successful than the Scotia in locating
the breeding- places of Sea-leopards, and this may of course be accounted for by the
comparative scarcity of the species, on which all expeditions have commented. On
the Bal&na expedition, however, as noted above, Dr Bruce and Mr Burn-Murdoch
report seeing many. Dr Bruce estimates that the crew of the Balssna killed fully a
thousand during December, January, and February. The expedition of the Frangais

THE SEALS OF THE WEDDELL SEA. 193

appears to have seen only one, and that is not certain, though the Pourquoi Pas? in
the same region saw several. But I am inclined to think that the failure to discover
the breeding-places is due, in reality, to there being no large Sea-leopard rookeries.
This species is probably more or less solitary, even at the breeding-season, and thus
would largely escape notice.*

Dr Bruce and Dr Donald brought back several foetal Sea-leopards from the expedi-
tion of the Balsena and the Active in 1892-93, which were given to the Zoological
Department of University College. Dundee.

The only other foetal Sea-leopard known was obtained by Dr Charcot at Petermann
Island in March 1909, during his expedition in the Pourquoi Pas?

The Challenger found this seal on Kerguelen in midsummer, and bones of it at
Heard Island. But the large herds on Kerguelen of which Moseley speaks by report
were probably Weddell Seals. t It is not uncommon at the Falkland Islands, and has
been recorded from South Georgia and Fuegia as well as from such low latitudes as
Tasmania, Wellington, N.Z., and the Lord Howe Islands, so that it is not confined to
Antarctic waters. However, the more northerly of these must be looked on as records
of mere stragglers from the habitual haunts.

The Sea-leopard is the largest of all Antarctic seals, and may reach 12 to 14 feet
in length and weigh over 900 Ibs. In appearance it is comparatively graceful, with
its long slim body and its light grey coat generally touched with darker spots. Alone
among the seals of the Antarctic does this species threaten man, even without pro-
vocation. \Yith its formidable array of teeth, its powerful jaw, and its remarkable
agility, it is not an animal to be played with. Other species never show any resent-
ment to man except during the pupping season, and then only on provocation. When
on the floe the Sea-leopard lies characteristically on its stomach, with its head resting
on the ice, and rears itself on its fore-flippers and at once shows fight on the approach
of a man. It is usually alert, ready, and energetic in its movements, while it has not
the same love of lying on the ice which the lazy Weddell Seal exhibits, and seems to
go on to the ice principally for a rest, when out in mid-sea. The food of this seal seems
to consist chiefly of penguins, which it chases with great agility under the surface of
the water, and even catches on the ice. In one instance, in Scotia Bay, a Sea-leopard
was observed swimming noiselessly up to a piece of pack, near the edge of which was
standing a Gentoo penguin : suddenly, with a quick jump, the Sea-leopard raised himself
from the water, caught the penguin's leg in his jaws, and drew the bird away into the
sea. In one specimen the Discovery found the body of an Emperor penguin ; and
Dr Davidson, of the Morning, in another found the remains of a young Weddell Seal.
Dr Bruce found large quantities of stones in Sea-leopards' stomachs in 1892-93, which
were evidently from the stomachs of digested penguins. Feathers from the same

* It is probable that the " Sea-leopards" which Mr Borchgrevinck describes as breeding in Robertson Bay, Victoria
Land, were really Weddell Seals (First on the Antarctic Continent, London, 1901, p. 237).
t Notes of a Naturalist on the " Challenger," H. N. Moseley, London, 1892, p. 174.

VOL. IV. 2

194 DR R. N. RUDMOSE BROWN ON

source were also found in the Sea-leopards' stomachs. However, on occasions when
penguins are not available the Sea-leopard does not despise fish and crustaceans.
Probably in midwinter the Sea-leopards migrate northwards to the edge of the pack,
and so keep in touch with the penguins, which do the same.

Lobodon cardnophaga (Jac. et Puch.) — the White Antarctic Seal.

The White Antarctic Seal or " Crab-eater" is not uncommon at the South Orkneys,
and in some parts of the Antarctic occurs in large numbers. Dr Turquet thinks it the
commonest species in the regions west of Graham Land explored by the Francais,
where the Pourquoi Pas? also found it plentiful. Dr Bruce in 1892-93 found this
species in the greatest abundance of any off Louis Philippe Land, where the Balsena
killed some 4000 to 5000 of them. In the Bay of Whales, in the Ross Barrier,
Captain Roald Amundsen found it to be a common species. No expedition has
found it uncommon, although we did not see it as frequently as the Weddell Seal.
This, however, may be because numerous Weddell rookeries occurred near our
winter-quarters.

The Lobodon typically occurs on the pack-ice, often far from land, and it was in
these conditions that most of the specimens we saw were recorded. They are seldom
seen on shore. During February and March 1903, while the Scotia was cruising in the
pack of the Weddell Sea, scarcely a day passed without several being seen. They are
never found far from open water. The Lobodon is not as a rule a solitary animal :
generally several are seen together. Dr Bruce in 1892 recorded as many as forty-seven
lying on the same piece of pack-ice.

During the winter the Lobodons desert the fast ice. Not a single specimen was
seen by us at the South Orkneys from the middle of March to the 1st of August. In
the somewhat less severe regions west of Graham Land they seem to leave a little later.
The Discovery also found that the Lobodous left when the sea froze over in winter. In
the winter they apparently stay at sea among floating ice, for there seems to be no record
of them visiting such lands as the Falkland Islands, Kerguelen, or Macquarie Island ; but
stragglers have been reported from Patagonia, and as far north as the Rio de la Plata
and also the coasts of Australia.

We came on no breeding-places of the Lobodon, but found a number of young ones
during the summer. An individual captured on August 1st, which Russ, one of the
dogs, chivied out of a tide-crack on to the floe, was a young one, but evidently about a
year old.

The pupping season, according to M. Racovitza, is September, which would agree with
the apparent age of the young that we captured during the summer. There is little
doubt that they lived at or near the South Orkneys, though not in the bays we visited,
for numbers of them were seen towards the end of August, after which none seem to
have been noticed until November. In that month two old males were seen. In the

THE SEALS OF THE WED DELL SEA. 195

later summer they again became common, both adults, male and female, and young.
Not only did we fail to find any quite young Lobodons, but we were unfortunate in not
securing any pregnant females, and so cannot offer any contributions to the embryo-
logy of the species. The Belyica alone secured a very young specimen. However,
among our collection of skins are two of young males not more than one year old.
M. Racovitza maintains that they bear their young on the pack away from land ; the
number of young he saw during the Belgicas long drift in the pack would confirm this.

The Lobodon is a comparatively shapely seal, and never attains the grossness of the
Weddell. It swims rapidly and gracefully, while on the ice it is generally alert and
vivacious in its movements, and can travel much faster than the Weddell.

The young has a thick mouse-grey coat, often with a mottled appearance, but this
changes in the adult into a coarser coat, but beautiful creamy-white in colour, though
some are rather silvery-white and others remain mouse-grey. Dorsally it is darker
than on the ventral surface. However, the colour varies somewhat with the position of
the spectator. If looked at from behind forward, it is several shades darker than in the
reverse direction ; and this applies equally to the Weddell Seal. With increasing age
the coat seems to become whiter.

A peculiarity of the Lobodon is the scarred appearance it presents in midsummer.
In December this is most noticeable ; in January their condition improves, and in
February, as a rule, they are free from scars. On the other hand, the young specimen
we captured in August had several half-healed scars, and badly scarred Lobodons were
seen both in February and March. Yet the fact seems to be that these scars are in-
flicted in November and December as a rule ; this coincides with the rutting season,
when no doubt the males are badly wounded in fighting with one another. As far as
the observations of the Scotia naturalists go, the scars are confined to the males ; but in
1892 Dr Bruce comments on the females being as badly scarred as the males. The
Discovery seems to have noted the same. If that is so, the females must join in the
fight. But the deep abdominal gashes that the Lobodon so frequently bears can scarcely
be ascribed to this cause. The idea that they are due to attacks by Sea-leopards can be
dismissed as groundless and unlikely in the extreme. The likelihood of the Killer
Whale being responsible I have discussed in speaking of the Weddell Seal. Dr Wilson
pointed out how the pelagic habits of the Lobodon would make it more exposed to these
attacks than other seals, and hence the more frequent scars. On the other hand, how-
ever, it must be noted that the Lobodon is more agile than the Weddell, not only on
land, but also in the water, which should give it greater immunity than the Weddell.
This agility of the Lobodon we had ample opportunities to note during the many months
the Scotia spent cruising in the pack. Possibly, however, it is more than counter-
balanced by the continual presence of the enemy, which the Weddell Seal, in its shallow-
water haunts, largely escapes.

The principal food of the Lobodon is Euphausia, an index to its more pelagic habits,
and hence presumably its name of " Crab-eater."

196 DR R. N. RUDMOSE BROWN ON

This seal has also been referred to as Dumont d'Urville's Seal, since it was discovered
by that explorer, in analogy with the names of the Weddell and Ross Seals. The
appellation, though quite justifiable on these grounds, is a clumsy one, and is unlikely
to come into use.

A full-grown Lobodon attains a length of over 9 feet and a weight of 500-600 Ibs.
or more. A temperature of 99° F. was recorded in one by Dr Pirie.

Ommatophoca rossi Gray — the Ross Seal.

This seal was rarely seen, and. despite the frequent expeditions of recent years, still
remains one of the rarest of known mammals. It is the only one of the Antarctic
seals that is entirely confined to Antarctic seas, and which has never been recorded from
extra-polar regions. During the voyage of the Scotia the Ross Seal was only seen on
five occasions, and on four of these it was among the pack some distance from land.
The most southerly one was seen about the latitude of the Antarctic circle, midway
between the South Orkneys and Coats Land, in the month of March. In December
1903 an old male with very worn teeth was observed in Scotia Bay, and several
were noted there in the course of the following and subsequent years. During the
first year (1904) of the Argentine occupation of Omond House Mr R. C. Mossman
captured a young female of this species, about 5^- feet in length and estimated at
about six weeks, near Cape Burn-Murdoch, Scotia Bay. This is particularly
interesting, since it is the first young specimen of the Ross Seal ever brought home,
and the only one known up to the present time. In 1892, Dr Bruce and Mr W. G. Burn-
Murdoch captured what Dr Bruce is now certain was a young Ross Seal. As soon as
the boat came alongside of the Bal&na, however, Captain Fairweather ordered the
animal to be taken out on to the ice arid skinned. The carcase was left on the ice, and
the skin heaved aboard as one of the 6000 skins taken home to Dundee. Prof. T. W. E.
David speaks of two young Ross Seals in February on the Drygalski Ice Barrier
Tongue in Victoria Land, but he does not give their age. Under the circumstances
neither could be preserved.* Previous records of this seal in the Weddell Sea are
those of Ross and Weddell. Dr Bruce, in 1892, refers to great numbers of Ross Seals,
but he admits that he was wrong in his identification in this case. Solitary individuals,
as in the Weddell Sea, have been recorded by practically all recent expeditions from
other parts of the Antarctic, but everywhere the Ross Seal is very rare. Dr Turquet,
of the expedition of the Franqais to the west of Graham Land, is not sure that he saw
any, but some of his colleagues certainly heard the characteristic cry. The collection
of skins, however, of this expedition seems to contain one of the Ross Seal. In the
same region the Pourquoi Pas ? seems to have sighted but two. M. Racovitza, of the
Belgica, comments on their rarity ; and the Swedish expedition in the Antarctic on
the east and north of Graham Land during their twenty-one months' stay saw none.

* Tlw Heart of the Antarctic, vol. ii. p. 208, London, 1909.

THE SEALS OF THE WEDDELL SEA. 197

Mr Mossman, on board the Uruguay (1905), saw a Ross Seal at the South Shetlands in
Pendulum Cove, Deception Island.

Of the breeding-habits of this seal nothing whatever is known, but in all probability
the young are born on the pack and not near land. From the rarity of the species
it is almost certain that they do not collect in rookeries at that season.

Their movements in winter are equally mysterious, for they seem to leave the ice-
bound region at that season. Presumably they migrate to the edge of the pack, but
there is no record of a Ross Seal having been seen on any subantarctic island.

The Ross Seal feeds principally on cuttle-fish, but remains of fish and Euphautda
were found in their stomachs. A large cephalopod, over 6 feet in length, was captured
in Scotia Bay in 1904 at the Argentine Meteorological Station, which from its beak
appeared to be of the same species as that on which the Ross Seal feeds.

The colour of this seal is a dark grey with lighter stripes or elongated spots, chiefly
on its sides, while its ventral surface is of a uniformly lighter tint, sometimes almost
white. Several peculiarities distinguish it in a striking way from other Antarctic seals.
It is probably the most shapeless of them all ; as M. Racovitza says, " the form of
the quadruped has almost disappeared." The enlarged size of the fore-flippers is
very noticeable, but the most remarkable feature of all is the thick neck due to a
great development of the larynx. This gives the animal an extraordinary appearance.
Furthermore, this seal is generally very fat, and the head is much dwarfed by folds
of fat about the neck. The distended larynx acts as a resonator, and hence the
explanation of the curious loud cry, which is quite characteristic and unmistakable.

Both on ice and in the water the Ross Seal is far more agile than the VVeddell, and
can be alert and active when the occasion demands it ; but, unconscious of danger, they
allow one to approach near without showing signs of fear. The agility of this seal may
account for its feeding habitually on cuttle-fish, in contrast to the slow, deliberate
Weddell, which has to be content with less active prey. Moreover, its quickness in the
water may be an adaptation to a pelagic mode of life which it seems to follow. Large
scars are seldom seen on the coat of this seal, and Dr Trouessart attributes this to the
greater agility of the Ross Seal, which enables it to escape the attacks of the Orca, to
which he ascribes such wounds. Certainly the Ross Seal is a very powerful swimmer,
but in view of the small number seen it is difficult to generalise regarding the scars.
In any case, the small scars which this seal occasionally bears are quite in keeping with
its feeble dentition, and would point to battles over females being the real cause.

The Ross Seal attains a maximum length of about 8 feet 6 inches ; we measured
none larger than this, and found most of them somewhat shorter. But Mr Bernacchi,
on the Southern Cross expedition, speaks of one nearly 11 feet. The Scotia secured
two skins and one complete skeleton. The temperature in one case was 96°'2 F. ; in
another case Seiior Valette recorded 98°'6.

198 THE SEALS OF THE WEDDELL SEA.

LIST OF REFERENCES.

ANDERSSON, K. A., " Das hb'here Tierleben im antarktischen Gebiete," Wissen. Erg. Schwed. Siidpolar-Exp.

(1901-3), Bd. v. 2, Stockholm, 1905.

BARRETT-HAMILTON, G. E. H., "Seals," Res. du voyage du S.Y. " Belgica" (1897-99), Anvers, 1901.
BARRETT-HAMILTON, G. E. H., "Seals," Report on the "Southern Cross" Collections, London, 1902, and The

Antarctic Manual, London, 1901, pp. 209-224.
BROWN, R. N. RUDMOSB, MOSSMAN, R. C., and PIRIE, J. H. HARVEY, The Voyage of the " Scatia," Edinburgh,

1906.
BRUCE, W. S., "Notes on Seals and Whales seen during the Voyage to the Antarctic, 1892-93," Report

Brit. Ass. Adv. Sci. Nottingham, 63, p. 807, 1893.
BRUCE, W. S., "Animal Life observed during a Voyage to Antarctic Seas" (read 17th January 1894),

Proc. Roy. Phys. Soc. Edin., vol. xii. part 2, pp. 350-354.

BRUCE, W. S., in From Edinburgh to the Antarctic, W. G. Burn-Murdoch, London, 1894, chapter xix.
BRUCE, W. S., "Antarctic Seals," Knowledge, 1893, December, vol. xvi. pp. 221-223.
BRUCE, W. S., and others, Life in the Antarctic, Glasgow, 1907. (A collection of sixty photographs of

vertebrate life, including several of Seals.)

LIOUSVILLB, J., in Rapports preliminaires sur les travaux executes dans I'Antarctique (1908-1910), Paris, 1910.
MURRAY, JAMES, in The Heart of the Antarctic, E. H. Shackleton, London, 1909, vol. ii., Appendix I.
RACOVITZA, E., "La vie des animaux et des plantes dans PAntarctique," Bull. Soc. Roy. Beige Geog., 1900,

pp. 177-230.
TROUESSART, E. L., " Pinnipedes," Exped. antarctique francaise (1903-5): Documents scientifiques, Paris,

1907.
VALETTE, L. H., " Viaje a las Islas Orcadas Australes,'' Ann. del Minis, de Agric., Sect, de Zoologica, iii. 2,

Buenos Aires, 1906.

WILSON, E. A., "Mammalia," (British) National Antarctic Expedition (1901-04), ii., London, 1907.
WILTON, D. W., and others, " Zoological Log of the ' Scotia,' " Report on the Scientific Results of the Voyage of

S.Y. "Scotia" (1903-4), iv. 1, Edinburgh, 1908.

THE SEALS OF THE WEDDELL SEA.

EXPLANATION OF PLATES.

PLATE I.

Fig. 1. Sea-leopard (Stenorhynchus leptonyx) (length = 13 ft. 6 in.), on board the s.s. Balxna, off Graham
Land, 1892. Mr W. G. Bum Murdoch seated. (This is the first photograph ever taken of a Sea-leopard.)
Photo by W. S. Bruce.

Fig. 2. Sea-leopard on The Beach, Jessie Bay, South Orkneys. Photo by J. H. H. Pirie.

PLATE II.

Fig. 1. The Lobodon or Crab-eater Seal (Lobodon carcinophaga). Adult with unmottled coat in fore-
ground. Young behind, showing mottled coat. W. Smith del. and photo.

Fig. 2. The Lobodon or Crab-eater Seal. Young, showing mottled coat. W. Smith del. and photo.

PLATE III.

Fig. 1. Ross Seal (Ommatophoca rossi) and Dr Pirie on board the Scotia. Photo by W. S. Brace.
Fig. 2. Weddell Seal (Leptonychotes weddelli), male, off Coats Land, Antarctica. Photo by W. S. Bruce.

PLATE IV.

Fig. 1. Weddell Seal on ice in Ellessen Harbour, Lewthwaite Strait, or Spencer's Straits. Photo by
W. S. Bruce.

Fig. 2. Weddell Seal (male) on beach of Mossman Peninsula, Scotia Bay. A usual attitude during
sleep. Photo by W. S. Bruce.

PLATE V.

Fig. 1. Weddell Seals on beach of Mossman Peninsula, Scotia Bay. Photo by W. S. Bruce.
Fig. 2. Weddell Seal (waking up) on beach of Mossman Peninsula, Scotia Bay. Photo by W. S.
Bruce.

PLATE VI.

Fig. 1. Weddell Seal off Coats Land. Photo by W. S. Bruce.

Fig. 2. Sea elephant Seal (Macrorhinm leoninus) at South Georgia. Photo lent by T. E. Salvesen.
Fig. 3. Weddell Seals on beach of Mossman Peninsula, Scotia Bay. Photo by W. S. Bruce.
Fig. 4. Weddell Seal and young, about three days old, Scotia Bay, South Orkneys. Photo by
W. S. Bruce.

PLATE VIL

Fig. 1. Weddell Seal (Leptonychotes weddelli) and newly-born young, Scotia Bay, South Orkneys.
Photo by W. S. Bruce.

Fig. 2. Weddell Seal, with young two or three weeks old, Scotia Bay, South Orkneys. Photo by
R. N. R. Brown.

Fig. 3. Young Weddell Seal, one day old. Photo by W. S. Bruce.

Fig. 4. Young Weddell Seal, two days old. Photo by R. N. R. Brown.

THE SEALS OF THE WEDDELL SEA.

PLATE VIII.

Wedclell Seals in Scotia Bay, South Orkneys.
Figs. 1-4. Adults.

Figs. 5-6. Mothers with young a few days old.
Fig. 7. Young, several weeks old, landing on beach.
Fig. 8. Young about one week old.

Photos by W. S. Bruce

PLATE IX.

Weddell Seal (mothers and young). The young are only a few days old. Photos by W. S. Bruce.
Nos. 7 and 8 show the mother and young playing with each other. The white spots in eyes of the
young of No. 8 are frozen tears.

SCOT. NAT. ANT. EXP.

Brown : Seals of the Weddell Sea. Plate I.

VOL. IV.

[Photo 6y W. S. Bruce.

1. Sea-leopard (Stenorhynrhux If pt onyx), (length, 13 ft. 6 in. ) on board the s.s. Jlalfma, off Graham Land, 1892.
Mr W. G. Burn Murdoch seated. (This is the first photograph ever taken of a Sea -leopard.)

[Photo by J. H. H. Pirie,

2. Sea-leopard on the beach, Jessie Bay, South Orkneys.

SCOT. NAT. ANT. EXP. VOL. IV.

Brown : Seals of the Weddell Sea. Plate II.

•

[II*. fmifh, del. and p/r.

1. The Lobodon or Crab-eater Seal (Lobodon carcinophaga). Adult, with unmottled coat, in foreground.

Young behind, showing mottled coat.

[II-. Smith, drl. and phot.,.

2. The Lobodon or Crab-eater Seal (young, showing mottled coat).

SCOT. NAT. ANT. EXP.

Brown : Seals of the Weddell Sea. Plate III.

VOL. IV.

[I'Koto by If. S. Brute.

1. Ross Seal (Ommatophoca rossi) and Dr Pine on board the " Scotia."

[Photo by Jr. S. Brvct.

2. Weddell Seal (Leptonychvtes weddelli), male, off Coats Land, Antarctica.

SCOT. NAT. ANT. EXP. VOL. IV.

Brown : Seals of the Weddell Sea. Plate IV.

[Photo by W. S. Bnct.
1. Weddell Seal on ice in Ellessen Harbour, Lewthwaite Strait, or Spencers Straits.

2. Weddell Seal (male) on beach of Mossman Peninsula, Scotia Bay. A usual attitude during sleep.

SCOT. NAT. ANT. Exp. VOL IV.

Brown : Seals of the Weddell Sea. Plate V.

1. Weddell Seals on beach of Moss man Peninsula, Scotia Bay.

[PKoto by W. S. Bnta.

[Photo by FT. S. Brux.
2. Weddell Seal (waking up) on beach of Mossman Peninsula, Scotia Bay.

SCOT. NAT. ANT. EXP.

VOL. IV.

Brown : Seals of the Weddell Sea. Plate VI.

[Photo liy ir. S. Eruce.

I. Weddell Seal off Coats Land.

[Photo lent by T. E. Salvcsen,
2. Sea Elephant Seal (Alacrorhinus leoninws) at South Georgia.

[Photo by W. S. Brute.
3. Weddell Seals on beach of Mossman Peninsula, Scotia Bay.

[Pkoto by W. S. Brtux.

4. Weddell Seal and young, about three days old,
Scotia Bay, South Orkneys.

SCOT. NAT. ANT. EXP.

Brown : Seals of the Weddell Sea. Plate VII.

VOL. IV.

v,-.

[PAoto by jr. .*. .Bri/<v.

1. Weddell Seal (Leptonychotes wedflelli) and newly-born young,
Scotia Bay, South Orkneys.

[Photo by H. .V. r.. Broirn.

2. Weddell Seal, with young two or three weeks old,
Scotia Bay, South Orkneys.

[Photo by ir. .S.

3. Young Weddell Seal, one day old.

[Photo by K. X. R. Drovn.
4. Young Weddell Seal, two days old.

SCOT. NAT. ANT. EXP.

Brown : Seals of the Weddell Sea. Plate VIII.

VOL. IV.

5.

[Photos by W. S. Bract.

Weddell Seals in Scotia Bay, South Orkneys.
(1-4. Adults ; 5-6, Mothers, with young a few days old ; 7, Young, several weeks old, landing on beach ; 8, Young about one week old.)

SCOT. NAT. ANT. EXP.

Brown : Seals of the Weddell Sea.

VOL. IV.

Plate IX.

1.

5.

[Photos by ir. S. Bruce.

Weddell Seal (mothers and young). The young are only a few days old. Nos. 7 and 8 show the mother and young
playing with each other. The white spots in eyes of the young of No. 8 are frozen tears.

PART XIV.
BIRDS.

PART XIV.-ORNITHOLOGY OF THE SCOTTISH
NATIONAL ANTARCTIC EXPEDITION.

SECTIONS I. TO XIII.

BY W. EAGLE CLARKE, F.R.S.E., F.L.S., Keeper of Natural History in the Royal
Scottish Museum; L. N. G. RAMSAY, M.A., B.Sc., Universities of Aberdeen
and Cambridge; R. N. RUDMOSE BROWN. D.Sc., University of Sheffield;
Naturalist, Scottish National Antarctic Expedition; and WILLIAM S. BRUCE,
LL.D., F.R.S.E., Director of the Scottish Oceanographical Laboratory, Edinburgh.

(WITH SEVEN PLATES AND TWO MAPS.)

OKNITHOLOGY OF THE SCOTTISH NATIONAL
ANTARCTIC EXPEDITION.

Section I. — Introduction.

By WILLIAM S. BRUCE, LL.D., F.R.S.E.,
Director of the Scottish Oceanographical Laboratory, Edinburgh.

THAT the ornithological results of the Scottish National Antarctic Expedition have
proved to be of considerable importance is because they were attained by the united
efforts of the officers and crew as well as the scientific staff of the Scotia. The keenest
observer on board, the one who would first detect a bird, not seen before, at a great
distance and be able to give a very exact description of its form and plumage, was our
second mate, the late Mr Robert Davidson. To him are due many of the excellent
records of the voyage. I spent many an hour on deck with Mr Davidson making notes
of birds that we observed together, and also notes of other birds he had seen during
his watch. Very often he would pass the word below that there was a new bird about
that we had not seen before, which brought the naturalists out on deck. To Mr D. W.
Wilton also great praise is due for his energy and pluck in being ever ready to be
lowered away in a boat in any sea a boat could live in. He was even ready, had he
been encouraged, to undertake many more risky boat expeditions than he did. Wilton
was usually accompanied by two seamen, and most often by the young Shetlander
Johnnie Smith. While the Scotia was lying during sounding, trawling, and vertical
tow-netting, and other operations, Wilton was almost always in the dinghy or gig
securing birds for the Scotia collections, and was responsible for many of the best
Scotia records. On one occasion, when the Scotia was trawling in one hundred
fathoms under the lee of Gough Island, Wilton got too far off the land and was
blown far out to sea in very rough weather, out of sight from the ship — a venture
which, but for the seaworthiness and the skilful handling of the boat, and level-headed-
ness of Wilton and his companions, might have ended in disaster. He was quite
unconcerned, however, when we picked him up after the trawl came on board, and
with his gun had secured a large part of the fine collection of birds that we collected at
Gough Island. In winter quarters there were more chances for everybody, and during
the absence of the Scotia refitting at Port Stanley and Buenos Aires, Dr .1. H. Harvey
Pirie specially distinguished himself by plucky rock-climbing, making many valuable
additions of eggs and young to the Scotia collections. Dr R. N. Rudmose Brown
was also assiduous in his observations at various penguin rookeries, and he and Pirie

VOL IV. 203 1

204 ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

secured the most complete series of penguin embryos in unbroken sequence from the
day of laying to hatching. These excellent embryological preparations form the basis
of the valuable investigations of Dr Waterston and Dr Geddes.*

On many occasions we "fished" for birds with baited hooks and triangles, but,
contrary to my previous experience on board the Balsena in 1892-3, we did not secure
many birds by this method. On the Balsena I have seen as many as thirteen
albatroses taken by the seamen in a day. On the same voyage I caught many birds
in this way myself, but, during the voyage of the Scotia, although we would some-
times hook a bird and bring in to within an arm's length, it invariably got clear before
being hauled aboard.

The consideration of the ornithology of the voyage has been based primarily on
geographical distribution ; the birds of every region passed over by the Scotia having
been considered separately. The first place that any special collection of birds was
made was St Paul's Rocks, where, in spite of a heavy swell running, the Scotia
naturalists secured two out of the three species that are known to breed there. Here
Dr Pirie had a narrow escape, when he was washed off the rocks, on to which he
jumped from the boat, into a sea swarming with sharks. So numerous were the
sharks that we caught several at the ship with ease, although our attention was
taken up at the time with testing our sounding gear and with kite-flying. At the rocks
fishing was tried from the boat, but each time a fish was hooked it was swallowed by
one of the sharks that were swarming round the boat. Little was then secured till
the Falkland Islands were reached, on account of our forcing as quick a passage
south as possible to the region we were specially fitted out to explore. At the
Falkland Islands my attention and the attention of the officers and crew had to be
given almost entirely to ship's business, but the naturalists managed to make several
excursions in the vicinity of Stanley Harbour. On two occasions I accompanied
excursions to Cape Pembroke, where I had to set up and subsequently inspect the
meteorological equipment at the lighthouse ; this made an opportunity for field
naturalist's work in that region. I also conducted a special excursion to the Tussock
Islands, while the Scotia was coaling, when we secured a Sea-lion and quite a number
of interesting birds. To Mr Thomson and Mr Coulson, of the lighthouse, is chiefly
due our collection of Falkland Island birds' eggs.

Very detailed and important work was done in ornithology at the South Orkneys,
in the Weddell Sea, and at Gough Island, and these collections have been examined
and described in three excellent reports by Mr W. Eagle Clarke in The 76zs,t which
are now reprinted. The type collection of the birds from these regions, including
males and females, types of every stage of plumage obtained, and a large and complete
series of eggs, I have presented to the Royal Scottish Museum ; while a further typical

* Idem, pp. 32-58.

t The Ibis, Eighth Series, vol. v., April 1905, pp. 247-268 ; Eighth Series, vol. vi., January 1906, pp. 145-187 ;
and Ninth Series, vol. i., April 1907, pp. 325-349.

INTRODUCTION. 205

series of the whole collection has been housed in the Scottish Oceanographical
Laboratory. I have presented several of the new and rarest species to the British
Museum, while the museums in Scotland have also benefited.

I have to thank Mr Eagle Clarke for his important contributions. I have also to
acknowledge the valuable work Mr L. N. G. Ramsay has done in putting in excellent
order the ornithological collections as a whole, and in giving such a complete account
of the birds seen and captured during the whole voyage. Dr R. N. Rudmose Brown
gives an important contribution on the " Life and Habits of Penguins," a brief summary
of the results of his own excellent observations and those of the other naturalists of the
Scotia. Free use has been made by the various authors of the bird notes in the
Zoological Log of the " Scotia " (Rep. on the Sci. Res. of the Voyage of the S. Y.
"Scotia," IV. i.), and also of the journals kept by the other naturalists and myself
during the voyage. The ornithological report, therefore, is a very complete one, and
the sum of the united efforts of all on board the Scotia.

Section II. — St Paul's Rocks, and the Voyages to the Falkland Islands,

and to Buenos Aires and Back.

By L. N. G. RAMSAY, M.A., B.Sc.

THE Scotia reached St Paul's Rocks on December 10th, 1902. "These isolated rocks
rise up in mid- Atlantic just a few miles north of the equator. They are only about
a half-mile in circumference and some sixty feet above sea-level. They have been
visited and described by Darwin, Sir James Ross, and the naturalists of the Challenger ;
but, geologically speaking, they are still a puzzle and of great interest. . . . We were
very anxious to secure some rock-specimens, but our hopes of effecting a landing were
frustrated by a heavy swell which intensified the strong westerly equatorial current ;
there was a rise and fall of about ten feet, and every now and again waves breaking
in spray almost right over the summit of the rocks." * Dr Pirie, who actually set foot on
them, was washed off into the sea, swarming with sharks, and was rescued with difficulty.
The rocks form the breeding-place of multitudes of boobies (Sula sula), and noddies
of two species (Anous stolidus and Micranous leucocapillus), which appear to nest all
the year round. Although the Scotia naturalists found landing was impossible, plenty
of birds were seen on the rocks, and some of the boobies were found to be nesting, the
young ones being nearly white in colour. Three boobies (an adult and two immature
birds) were obtained, and three adult noddies (Anous stolidus). The smaller tern,
Micranous, was not obtained.

From St Paul's Rocks the Scotia continued her voyage in a south-westerly direction,
passing within 50 miles (roughly) of Fernando Noronha, in the neighbourhood of which
a gannet (a white species — probably Sula cyanops or S. piscator) was seen.

On December 14th the Brazilian coast was sighted in 6° S. lat., and thenceforward
the ship made a direct passage for the Falkland Islands, her course gradually diverging
from the receding coast-line of the South American continent.

" Boobies " are logged as having been seen on various dates up to December
26th, in 30° S. These, with a few unidentified petrels and others, were the only
bird-life met with till 30° S. was approached, after which petrels began to be more
numerous.

The area of ocean between the River Plate and the Falklands was thrice traversed,
the two subsequent occasions being when the Scotia visited Buenos Aires to refit in
December 1903, returning to Port Stanley in January 1904.

* J. H. Harvey Pirie, The Voyage of the " Scotia," p. 40.
207

208 ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

The bird-notes of these three trips may be summarised together.

The Wandering Albatros (Diornedea exulans] on the first voyage was met with
for the first time in 35° S. On all three trips it was logged at intervals from this
latitude southward to near the Falklands.

The smaller species of albatros (Diomedea and Thalassogeron) were simply re-
corded as " Molliemauks " ; none were logged on the first trip ; on the second (December
1903) some were seen daily from 48° S. to about 34° S., while on the third they were
only encountered once, in 50° S., when six were seen.

Few Sooty Albatroses (Phaebetna spp ) were seen north of the Falklands, these birds
being logged from about 47° and 50° S. lat. on two of the three trips.

Wilson's Petrel (Oceanites oceanicus) was first encountered in 30° S., and thence
to the Falklands was noticed frequently, except on the last voyage, when it was
only once logged, and that was doubtful. On the first voyage they became more
numerous as the Falklands were approached, being noted as " very plentiful " on
January 4th (47° S.). On January 26th, 1904, " Stormy Petrels, like Wilson's, only
more white on their breasts and wings," were observed (43° S.). These may have
been the white-bellied Cymodroma grallaria, which was obtained later on, in the
South Atlantic. This species has lighter wings than Wilson's, and the inner under-
wing coverts are white.

"Blue Petrels" (Prion spp.) and Nellies (Ossifraga gigantea Gm.) were logged on
the first two days after the Scotia left Port Stanley in December 1903, and there is also
a doubtful record of the latter for 43° S. (January 26th).

On December 28th, 1902, three petrels were shot, the skins of two of which are now
in existence, in 33° S. lat. They prove to be CEstrelata incerta (SchL), a little-known
species, which has a wide distribution in the southern oceans : — *

(a) $ , December 28th, 1902. 33° 5' S., 48° 48' W. Length (in skin) about
18 ins. ; wing 1275 ins. ; bill (gape to tip) 2'0 ins. ; tarsus 1*75 ins. ; middle
toe 2'4 ins. ; inner toe 2'1 ins.

(6) ?, December 28th, 1902. Same lat. and long. Length about 17 ins.; wing
11 '9 ins. ; bill 175 ins. ; tarsus 17 ins. ; middle toe 2-2 ins. ; inner toe 1'85 ins.
Notes made on the spot describe the beak as black, the pupil blue, the iris sepia-
brown, the legs yellowish white, the lower parts of the toes and webs black.

In the dried skin the beak is black, with a greyish-white mark near the tip
on both upper and lower halves. Tarsi and toes yellow ; outer side of outer toe, and
distal portions of toes and webs, blackish.

The wings, mantle, lower back, rump, and tail are blackish brown, shading into
lighter brown on upper back, neck, sides of breast, and head ; a dark brown patch in
front of and above the eye ; middle of throat and of foreneck whitish ; middle of breast
and belly white. Flanks greyish brown, under wing- and under tail-coverts dark brown.
Faint lighter margins to scapulars and upper tail-coverts.

* Qodman, Monograph of the Petrels, 1910, p. 195.

ST PAUL'S ROCKS, AND VOYAGES TO THE FALKLAND ISLANDS, ETC. 209

The description of several petrels seen on the first voyage (December 24th to 26th)
corresponds with the above specimens, except that they are described as having
" a small white marking on the under and upper surface of the wing." This apparently
refers to another species, perhaps one of those species of (Estrelata with white on the
bases of the primaries.

Another species of petrel was seen in similar small numbers from December 29th to
January 5th. This was about the same size, but the under parts were entirely white,
the wings almost black, " verging into grey on the rest of the body, with a white ring
round beak and tail."

Several petrels of one or other of the two darker kinds were seen daily from
December 24th to January 3rd on this first voyage (26° to 45° S.), after which the
species with under parts wholly white was the only one observed.

Section III. — The Falkland Islands.
By L. N. G. RAMSAY, M.A., B.Sc.

THE Falkland Islands were visited on three occasions by the Scotia, namely : January
6th-January 26th, 1903 ; December 2nd-December 9th, 1903 ; January 3 1 st-February
9th, 1904. Dr Bruce had also previously visited these interesting islands in December
1892 and February 1893.

As the Scotia neared them for the first time, " in the cold grey dawn the
low, bleak, treeless, but grassy land looked very like some parts of the north of
Scotland."

" Vast stretches of undulating moorland, interspersed with peat-bogs and swamps,
constitute the general type of country, unless where the land rises into rocky ridges, or
higher into jagged hills. In place of heather, the ' diddle-dee ' (Empetrum rubrum),
a close ally of our blaeberry, grows in prolific abundance, clothing the moorland for
miles and miles, and each in its own particular haunt various kinds of sedges flourish.
The absence of trees on the islands is very noticeable." *

Although there was much to keep the staff occupied during these visits, apart from
scientific pursuits, a number of excursions were made, and a considerable number of
birds were collected.

The Scotia's collection of birds' skins includes twenty species, represented by sixty-
five skins, from the Falklands. Several other species were also observed. All have
been recorded as occurring in the Falklands by previous writers.

Tachyeres cinereus (Gm.). — Two specimens of the flightless "Steamer-duck" or
" Loggerhead " were brought home. They were seen in numbers on all three visits.
In December 1903 some of the birds had families of ducklings with them.

A number of the eggs of this bird are included in a collection of birds' eggs which
Mr Thomson and Mr Coulson, jun., of Cape Pembroke lighthouse, kindly presented to
the naturalists of the Expedition. These eggs measure 3 '25 by 2 '25 inches, and are
pale creamy buff in colour.

Chloephaga, hybrida (Mol.). — These birds were also seen on all three visits, and
seven skins are in the collection (three adult males, one immature male, and three
females).

The eggs, if the specimens are correctly named, are hardly distinguishable in size and
colour from those of the Loggerhead.

Nycticorax tayazu-guira (Vieil. ). — Two specimens of the " King-quawk," as it is

* R. N. Rudmose Brown, The Voyage of the "Scotia," p. 196.
VOL. IV. 211 2

212 ORNITHOLOGY OP THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

locally called, were obtained, an adult on January 12th, 1903, and a young bird of the
year on February 1st, 1904. A description of the latter is appended :—

Measurements: Length (skin) about 24 ins.; wing 1175 ins.; culmen 27 ins.;

tarsus 3 '2 ins.

Back, scapulars, and wing-coverts nearly uniform brown with a rusty tinge. Some
reddish-yellow and white tips among the lesser wing-coverts. Bastard wing
and primary coverts tipped with white. Quills and tail dark grey with a rusty
tinge. Feathers of head, neck, and under parts dusky brown with broad white
and yellowish-white shaft- streaks, crown blackish-brown. Bill blackish at tip
and along culmen, remainder yellow-brown. Feet dark brown (in dried skin).
The adult is a male. The bill and interior of mouth were noted as black, the
iris red ; its stomach was full of fish.

This bird was rather rare in the islands, according to the accounts of the islanders,
though Dr Bruce previously obtained it when he visited the Falklands in 1892.
Phalacrocorax magellanicus (Gin.). — Twelve skins are in the collection.
Five of these are adults. These were shot on January 7th, 1903 (one), and February
1st, 1904 (four). There is no trace of a crest, and black of the throat and foreneck is
being replaced with the white of the winter plumage to a varying degree.

(a) 7.1.03. Has a small white spot on the chin, but the feathers between the

rami of the mandible are black.

(b) 1.2.04. Ditto, but a small white cheek-spot also.

(c) 1.2.04. As (b).

(d) 1.2.04. Indistinct white patch on throat, white cheek-spot.

(e) 1.2.04. Feathers between mandibular rami and chin pure white, throat mottled

white, white cheek-spot.

Six full-grown young birds with brown down still adhering to the feathers of head
and neck were obtained at Tussock Islands in Port William on February 1st, 1904, where
numbers of them were seen. Three of these are glossy brownish black on head, neck,
upper parts, and thighs, with the belly greyish brown. The rest are similar except as
to the belly. One has this blackish brown ; one has it nearly white, only the tips of
some of the feathers being blackish ; and the last shows white, but to a much less
marked extent.

Phalacrocorax albiventer (Lesson). — One adult was obtained at Cape Pembroke in
January 1903.

Buteo erythronotus King. — An immature bird was obtained on January 21st, 1903.

Milvago australis (Genl.). — " The hawk known as the Johnny Rook " is logged as
having been seen during the Scotia's first visit to the Falklands. It is quite a familiar
bird in the vicinity of Port Stanley.

"Turkey-buzzards" (Cathartes sp.) were seen during the third visit.

Larus glaucodes Meyen. - - One adult skin is in the collection, obtained on
February 1st, 1904, at Tussock Islands, where many were seen. In life, this gull has

THE FALKLAND ISLANDS. 213

a "beautiful rosy blush" on its breast plumage, which has disappeared, however, in
the skin.

Larus dominicanus Licht. — These birds were observed on all three visits, and
were found breeding in numbers on the Tussock Islands in February 1904, a number
of eggs, and two young birds, being obtained there on the 1st. One of the young is a
chick in the down, the other older, with quills half-grown.

Sterna hirundinacea Less. — This species was also found breeding on the Tussock
Islands on February 1st, 1904, a half-fledged chick being obtained on that occasion. An
adult bird was also secured on February 8th, at the same place.

Megalestris antarctica (Less.). — Skuas were seen duiing the Scotia's second and
third visits.

H&matopus ater (Less.). — One adult was obtained at Cape Pembroke, during the
Scotia's first visit.

H&matopus leucopus (Garn.). — Four skins, all adults, are in the collection, obtained
on the Scotia's first and third visits.

Eudromias modesta (Licht.). — One skin is in the collection.

Gallinago paraguayise (Vieil.). — Of this species also there is one specimen.

Tringa fuscicollis Vieil. — Five skins of this winter migrant from the far north are
in the collection. Four of these were obtained in January 1 903, the remaining one on
February 2nd, 1904.

JEgialitis falklandica (Lath.). — Four specimens, two adult, two immature, were
obtained, in January 1903 (label of one immature bird lost).

Anthus correndera Vieil. — Two specimens were obtained on February 5th, 1904.
They both show the long wedge of white on the penultimate pair of tail-feathers,
characteristic of old birds of this species. The plumage of one is very much abraded,
the pale margins of the feathers on the back being worn away so as to leave it nearly
uniform dark brown. The tail especially has suffered, some of the feathers being
reduced to less than half their original breadth. There is an almost complete absence
of the sandy tint in the plumage of this specimen, as compared with the other.

The latter has apparently just moulted, the feathers being very fresh and new, and
the broad margins to those of the upper parts are of a very warm sandy buff colour,
while the whole of the lower parts are tinged with the same hue.

Muscisaxicola macloviana (Garn.). — Six specimens of this species were brought
home, two of which were obtained in January 1903, and four in February 1904. The
average wing measurement is almost 4*4 ins.

[Turdus falklandicus (ex Quoy et Gaim.). — A thrush logged as having been seen
during the Scotia s first visit to the Falklands was presumed to belong to this species,
which is confined to these islands.]

Trupialis militans (L.). — An adult accompanied by a full-grown young bird were
seen by Messrs Wilton and Brown one evening during the Scotia's second visit
(December 1903), and one or two also on the third.

214 ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

The collection of eggs referred to above includes three of this species. They
measure 1 '5 x 1 '1 ins., and are white, irregularly spotted and blotched with pale vinaceous
grey and chocolate-brown.

Phrygilus melanoderus (Q. et G.). — Many of these bright-coloured little birds were
seen flitting about the shore. Three specimens were obtained, two adult males and a
female, on January 12th, 1903.

Ossifraga gigantea (Gm.). — Giant Petrels or " Nellies " were seen during the Scotia's
second visit to the Falklands, and one also during the last visit.

Catarrhactes chrysocome (Forst.). — One example of this penguin was obtained at
Cape Pembroke on January 5th, 1903. It is an adult female.

Spheniscus magellanicus (Forst.). — Seven specimens (six adults and one young bird)
are included in the collection. All are from the Tussock Islands, February 1904.

The young bird measures (in the skin) about 19-20 ins. in length ; flipper 6 '5 ins.,
culmen 17 ins. It is therefore only about half-grown, as the adult measures 28 ins.
A quantity of brown down still adheres to the feathers of the median line of the crown
and the whole of the hind neck.

The plumage of the whole of the upper parts is of a uniform blue-grey appearance,
like that of Pygoscelis in fresh plumage, the feathers being tipped with blue and with
black shaft-streaks. The under parts are entirely white, with the exception of an ill-
defined dark grey band 1^-2 inches wide, corresponding in position to the neck-band
of the adult ; sides of head white mixed with grey. Underside of flipper much as in the
adult. The ridges on the base of the bill are already quite marked ; the groove between
culminicorn and latericorn is much more conspicuous than in the adult.

The haunts of the Jackass Penguin, on William Islands, were visited during the
Scotia's third stay at Port Stanley. This penguin was found to inhabit burrows vary-
ing from a foot to a couple of yards in length among the roots of the tussock-grass
(Dactylis cxspitosa), and, " with caution, one could see it sitting at the mouth taking
the air, but at the least alarm away it rushes into the safety of this underground
dwelling. It has a curious habit of sitting far back in its burrow eyeing the intruder,
with its head turned sideways, now on one side now on another. This penguin derives
its popular name from the very close similarity between its cry and an ass's bray."

During the Scotia's voyage from Buenos Aires to Port Stanley, penguins were
encountered on January 28th, in 45° 31' S., or about 350 miles from the Falklands.
Others were seen the next two days, before Port Stanley was reached. To the south,
penguins were met with in about 57° S., 55° 30' W., on November 30th, 1903, and again
on the next two days as the ship approached the Falklands.

* R. N. Rudmose Brown, The Voyage of the " Scotia," p. 200.

Section IV. — Voyages between the Falkland Islands and the

South Orkneys.

By L. N. G. RAMSAY, M.A., B.Sc.

ACROSS the tract of sea lying between the Falkland and the South Orkney Islands,
about 600 nautical miles apart, the Scotia made three voyages : southwards in January-
February 1903, northwards in November-December of the same year, recrossing
southwards again in February 1904.

The voyages averaged about seven days' duration ; the weather conditions were
sometimes clear and calm, sometimes stormy, with violent winds and snow showers ; no
ice was sighted north of the immediate neighbourhood of the South Orkneys, except on
the first voyage, when bergs were met with in some numbers during the southern half
of the journey, and the pack was encountered about 60° S. lat., unusually far north.
Bird life was fairly plentiful on all three trips, chiefly albatroses and petrels.
Diomedea exulans was noted on all three trips, chiefly towards the Falklands
(seven were seen on one day about 53C 20' S.), but they were also seen daily up to
58° S. on the first voyage.

" Molliemauks " (a general name for the smaller albatroses, except Phabetria) were
seen in some numbers throughout the greater part of the area. Two specimens were
obtained on December 1st, 1903, at the Burdwood Bank, which lies some 90 miles south
of the Falklands. These prove to be Thalassogeron culminatus (Gould), both adult.

The following notes were logged on December 1st, referring to these two albatroses
and a third which was probably not preserved : —

" They differ in colour especially about the head, some being much lighter than

others. The bills also differ in colour, the three we secured yesterday showing

this. Two of them have a yellow culmen and yellow under the mandible, while

the third is almost entirely grey. The inside of the mouth is yellow. The legs

are of a pale grey, almost white in parts, mottled with darker grey towards the

outer end of the webs. The grey has a distinct lavender tinge, much the same

as the feathers. Eyelid is black as well as the cere. At lower angle posteriorly

is a patch of white feathers, which in flight looks like a white eyelid. The eye

has a dark olive-green iris, while the pupil is greenish-black." In the dried skin

the feet are yellowish grey (Zoological Log of the "Scotia" p. 61).

Several "Grey-headed Molliemauks," recorded on February 12th and 13th, 1904

(58°-60° S.), may be of this species, or Thalassogeron chlororhynchiis. Several of the

" Real Molliemauk (Yellow-billed Albatros) " seen on February 12th of the same year

215

216 ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

were probably Diomedea melanophrys Temm. Sooty Albatroses were also observed
in fair numbers over the greater part of this tract of sea. The notes made by Dr Bruce
and the other naturalists at the time make it probable that Phcebetria cornicoides was
present as well as P. fuliginosa. On November 30th, 1903, a "Sooty" having "the
characteristic head and eye, but white underneath like a Mollie," may have been the
light-bodied P. cornicoides. Also on February 1st, 1903, Booties and " Molliemauks grey
and greyish black with no white at all " were logged. The latter would seem to have
been Phcebetria cornicoides.

Wilson's Petrels (Oceanites oceanicus) were seen every day on the first voyage, and
one was shot on February 2nd. On the two later voyages they were more irregular,
being logged as " plentiful " on two days, while on some days none were seen.

"Blue Petrels" were found in plenty throughout the area, except in the neighbour-
hood of the Falklands. Owing to the difficulty of distinguishing between the true
"Blue Petrel" (Halobsena ccerulea) and the "Whale-birds" (Prion spp.) on the wing,*
there are no records for these voyages in which the species is certain. However, it may
be supposed that the "Blue Petrels" logged were Banks' Whale-bird (Prion banksi,
Gould), f

A dead Blue Petrel was picked up on November 28th, 1903, which is stated to be
P. banksi in a footnote in the Zoological Log of the "Scotia" (published 1908), on
Mr Eagle Clarke's authority.

Cape Pigeons (Daption capensis] were plentiful, on the whole, numbers being seen
near the South Orkneys ; sometimes several were caught with landing-nets. On some
days, however, none were logged, on others few. On the return voyage to the Falklands
they were present up to the day on which Port Stanley was reached.

In the Zoological Log for November 29th and 30th, 1903 (58° and 57° S.), "Cape
Hens" are mentioned as being seen (one on the 29th and a few on the 30th) "for the
first time since we were in these latitudes last." On February llth and 12th, 1904
(55°-58° S.) a petrel "about the size of Larus dominicanus or a little larger, and
completely dark," was logged. These records may refer to Majaqueus tsquinoctialis,
commonly known as the "Cape Hen," and answering to the above description pretty
well. There is no record in the log of the first voyage which could refer to this species.

The above, with a few other unidentified petrels, were all the birds which were
apparently roaming at large on these seas when traversed by the Scotia. In the
neighbourhood of the South Orkneys a number of other species which were nesting on
those islands were encountered, viz. : —

Ossifraga gigantea, Priocella glacialoides, Pagodroma nivea (these were met
with when the ship entered the pack on February 2nd, 1903) Chionis alba, Sterna
hirundinacea, Megalestris antarctica, and Pygoscelis antarctica and P. adelise.

* The white tip to the tail of Halobsena is a sure mark of distinction : Wilson, (British) Nat. Ant. Exped., vol. ii.
Zoology, p. 104.

t See Mr Eagle Clarke's papers on South Orkneys and Weddell Sea under these species.

VOYAGES BETWEEN THE FALKLAND ISLANDS AND THE SOUTH ORKNEYS. 217

Penguins were seen on November 30th, and December 1st and 2nd, 1903, in
57°-52° S., as noted in a former section.

NOTE. — I take this opportunity of placing on record some observations of the bird-
life of the seas between the Falkland Islands and Graham Land, taken from the log
kept by Dr Bruce when naturalist on the whaler Balama on her voyage of 1892-3, in
the months of December and January.

A remarkable incident of the voyage occurred on December 16th, 1892, when birds,
especially Cape Pigeons (Daption capensis], but also Blue Petrels (? Halobiena) and
" Molliemauks," thronged literally in myriads over the sea, throughout the day, forming
a spectacle never to be forgotten, as Dr Bruce informs me. The previous and following
days these species were not observed in any large numbers. "Blue Petrels'"' were seen
in small numbers until the Baliena was approaching Graham Land, but were not seen
in the vicinity of this land or the adjacent islands. Daption capensis continued to be
present throughout the voyage.

Pagodroma nivea was observed from December 20th onward, after ice had been
encountered.

Ossifraga gigantea was also recorded (including several white examples) in the
neighbourhood of Graham Land, also Chionis alba and terns of some species. Several
large albatroses (Diomedea exulans) were seen two days after the Bal&na left Port
Stanley for the south.

Stormy Petrels (? Oceanites oceanicus) were present in small numbers throughout
the voyage.

An account of the Penguins observed in the vicinity of Erebus and Terror Gulf,
on the voyage of the Balzena, is given in an article by Dr W. S. Bruce entitled
"Antarctic Birds, I.," in Knowledge, xvii. pp. 208-210 ; and of the sister ship Active,
by C. W. Donald, M.B., in the Proc. Roy. Soc. Edin., 1895, pp. 170-176.

Section V. — On the Birds of the South Orkney Islands.*

By WM. EAGLE CLARKE, F.E.S.E., F.L.S., Keeper of the Department of
Natural History of the Royal Scottish Museum.

THE South Orkneys are a group of over a dozen islands situated between 60° and
61° S. lat., and 43° 3' and 47° W. long. They lie some 600 miles S.E. by E. of the
Falkland Islands, about 500 S.W. of South Georgia, and 200 E. of the South Shetlands.
They were discovered by Powell in 1821, and were visited by Weddell in 1823, by
Dumont D'Urville in 1838, and by Larsen in 1893. The descriptions furnished by
these explorers were, however, meagre in the extreme, and until the visit of the
Scottish expedition the South Orkneys remained among the least-known lands lying on
the fringe of the South Polar Sea. t

So far as their ornis is concerned, only two species of birds, and one of these prob-
lematical, have hitherto been alluded to — namely, the Ringed Penguin (Pygoscelis
antarctica), of which a specimen was obtained on Weddell Island by D'Urville, and a
Crested Penguin (Catarrhactes) described by Larsen.

The Scotia visited the islands on her voyage south in February 1903 ; subsequently,
having completed her first Antarctic cruise, she returned to the archipelago towards the
end of March and went into winter-quarters, remaining there for eight months, during
which period much valuable geographical and zoological work was accomplished.

Only two of the islands are of considerable size — namely, Coronation Island, which
is the most westerly, and Laurie Island, the most easterly.

Laurie Island, where the Scotia wintered, was the main scene of the labours of the
expedition, and it is almost entirely upon observations and collections made during
eleven months' residence there that the following contribution is based. The length of
this island is about 12 miles, its maximum breadth 6 miles, and its area fully 30 square
miles. The interior is lofty, and several of the summits reach to an altitude of from 2000
to 3000 feet. A number of deep bays run inland from north to south, separated by
narrow rocky peninsulas or steep lofty mountain -ranges, and cause the island to have a
very remarkable outline. All the valleys are choked by glaciers, and what little exposed
rock is visible is precipitous in the extreme. Here and there on the lower slopes and
at sea-level are a few acres of more or less level ground. In winter the whole island
and even the faces of the precipitous cliffs are covered with snow, which does not com-
mence to disappear till October and November (the late spring and early summer months) ;
but then many patches of moss-covered ground are laid bare, some of them bearing soil

* Reprinted, with slight verbal alterations, from The Ibis, Series VIII., vi., 1906, pp. 145-187.
t See The Voyage of the " Scotia," 1908.
VOL. IV. 219 3

220 ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

from six to ten inches deep. Except this vegetable- mould, there is little soil anywhere.
The rocks, various kinds of gray wacke, are mostly covered with lichens, especially Usnea,
which, with various species of moss, form the entire terrestrial flora of the island.

Concerning climatic conditions, Mr Mossman informs me that, in spite of their low
latitude, the climate of the South Orkneys is essentially polar. One of the most
powerful factors in determining the temperature of the air over this region is the cold
Antarctic current which carries streams of ice and numerous icebergs to a latitude
corresponding with that of the northern part of England. The mean annual tempera-
ture, based on nearly two years' observations, was found to be 22°'7 F., the means of
the seasons being: summer 31°'4, autumn 22°7, winter 13°-7, and spring 23°'3. The
most remarkable feature was the low and equable summer temperature, which rarely
rises above 37° or falls below 25°.* In winter, owing to the freezing up of the sea to
the south, the islands are virtually on the edge of a continent, and the temperature at
that season is thus characterised by great variability, the range of the thermometer
frequently exceeding 60° in twenty-four hours. If the wind is in the south, very low
temperatures, as low as 40° below zero F., are recorded ; but with a change of wind to
the north the thermometer may rise, even in the depth of winter, above the freezing-
point. Summer is characterised by almost continuously overcast skies, and the finest
and clearest weather occurs in winter. Owing to the large amount of cloud which
hangs over the islands in summer, the temperature is much the same as at places ten
degrees further south. The snowfall is excessive, the sunshine is very deficient, and
strong gales are frequent.

The first landing on the archipelago was effected at Saddle Island, which was
fortunately clear of ice, on February 4th, 1903. Here the explorers were met by a host
of Ringed Penguins (Pygoscelis antarctica), which had a large " rookery," where many
young and some eggs were found. Cape Petrels or " Pigeons " (Daption capensis),
Sheathbills(C/M0ms alba), and Skuas (Megalestris antarctica) were also nesting, and
specimens of both young and old were obtained. Gulls (Larus dominicanus), Giant
Petrels (Ossifraga gigantea), and Shags (Phalacrocorax atriceps) were observed on
the adjacent islets and rocks, and were apparently nesting there.

From Saddle Island the Scotia sailed for the far south, and, having made a success-
ful voyage in the southern waters of the Weddell Sea, the Expedition returned to
the South Orkneys on' March 21st. This was followed by a quest for suitable winter-
quarters, during which Leathwaite Strait and the east side of Coronation Island were
explored, and, finally, on March 25th, a bay, afterwards named " Scotia Bay," on the
south coast of Laurie Island, was selected.

It was now autumn, and the birds were beginning to emigrate in search of more
genial winter-quarters to more northern latitudes, or, in the case of some species, the
nearest open water to the archipelago, wherever that may have been. Even in mid-
winter (June and July) Laurie Island was not devoid of feathered inhabitants, for the

* Subsequent data slightly alter these values, but not as much as a degree — EDITOR.

ON THE BIRDS OF THE SOUTH ORKNEY ISLANDS. 221

following birds were observed more or less frequently, though not abundantly : — Snowy
Petrels (Pagodroma nivea), Giant Petrels (Ossifraga gigantea), Gulls (Larus doniini-
canus), and Sheathbills (Chionis alba). The Skuas (Megalestris antarctica) and the
Ringed Penguins (Pygoscelis antarctica} departed during the last days of April, and
were followed by the Cape Petrel (Daption capensis) and the Adelie and Gentoo
Penguins (Pygoscelis adelise and P. papua).

The first spring immigratory movements took place in October, when Cape Petrels,
Adelie and Gentoo Penguins, Skuas, and Terns (Sterna hirundinacea) arrived in the
order indicated, the last-named at the very end of the month. These were followed in
November by Wilson's Petrel (Oceanites oceanicus), the Ringed Penguin, and the
Silver Petrel (Priocella glacialoides).

With the return of spring the explorers were busy and journeys were undertaken in
various directions, while a camp was established on the northern shore of the island,
which \vas productive of excellent ornithological results, but had unfortunately to be
abandoned at an interesting period on account of the break-up of the ice.

After having been icebound for eight long months, the Scotia was liberated on
November 23rd, 1903, and immediately departed for the Falklands and Buenos Aires
to refit ; but a party under the charge of Mr Mossman, the meteorologist, and Dr
Harvey Pirie, the medical officer and geologist, was left to carry on the various observa-
tions and investigations and to make collections throughout the summer months. It is
to the assiduous labours of Dr Pirie that we owe most of our knowledge of the bird-life
of the island during this most interesting part of the year, and he has earned the best
thanks of ornithologists for the vast amount of valuable work which he accomplished.

During the summer bird-life was extremely abundant. Rookeries of the three
species of penguin (Pygoscelis) were numerous on the low rocky shores and less steep
cliffs on various parts of the coast. Some of these bird-cities contained several millions
of inhabitants, and their daily life presented scenes so remarkable as to be almost
beyond description. The cliffs and their screes were the home of several species of
petrels, which resorted in great numbers to the ledges and crannies for nesting-sites, and
the shores were the abode of the gull, the skua, and the tern.

The Ringed Penguin, hitherto regarded as being nowhere an abundant species, was
found to have its metropolis at the South Orkneys, where the summer population on
Laurie Island alone was estimated at not less than one million birds.

The finding of the eggs of the familiar Cape Petrel (hitherto unknown to science)
and of the chicks and young of the Ringed Penguin and the Snowy Petrel, the remark-
able extension of the known range of the Macaroni Penguin (Catarrhactes chrysolophus)
and of Fregetta melanogaster (which was undoubtedly breeding), were also among the
results of the summer's work. Eggs of Wilson's Petrel, the Sheathbill, the Blue-eyed
Shag (Phalacrocorax atriceps), and other well-known Antarctic species were also
obtained, some of them in great abundance. The collection of bird-skins, too, was
largely augmented.

222 ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

The series of bird-skins is one of the most important ever made in the Antarctic
Seas. It comprises one hundred and forty-three specimens, representing sixteen out of
the eighteen species now known to frequent the islands and their immediate vicinity ;
while the eggs number several thousands. Many of the skins afford additional informa-
tion concerning little-known phases in the plumage of several species, and enable me to
describe for the first time the young or immature stages of others, such as the Ringed
Penguin, Shag, Snowy Petrel, and so forth.

Add to the above slight summary of the bird- work accomplished the innumerable
notes on and accounts of the nidification and other habits of not a few little-known
species, and the investigations of their periods of incubation and the dates of their
arrival at and departure from their summer haunts, and we have an outline of the
ornithological results obtained by the Scottish National Antarctic Expedition at the
South Orkneys — results of the first importance, and meriting the full recognition, the
sincere thanks, and the most hearty congratulations of all interested in our favourite
science.

On the return of the Scotia from the Falklands, the members of the Expedition,
save Mr Mossman and another, embarked, and the South Orkneys were finally quitted
on February 22nd, 1903, for the southern shores of the Weddell Sea — the Antarctic
Continent, the then-discovered Coats Land.

In the preparation of this contribution I feel conscious that I have laboured under
one very great disadvantage, namely, that of not having been a member of the Expedi-
tion, a circumstance which must naturally result in unavoidable shortcomings. Much
valuable information, both written and verbal, has, however, been placed at my
disposal, including the official Zoological Log and full and interesting notes from the
private diaries of Dr Bruce, Dr Pirie, Dr Rudmose Brown, Mr Wilton, and Mr
Mossman. Mr Mossman, at the request of the Argentine Government, spent a second
winter and summer at Laurie Island, engaged in meteorological and magnetic work, and
he has most kindly supplied me with some additional information on bird-life made
after the departure of the Scottish Expedition. To all these friends I desire to express
my deep indebtedness and my sincere thanks. My friend Mr Norman B. Kinnear has
also earned my acknowledgments for his assistance in classifying records.

I propose to conclude this section of my contribution by instituting a comparison
between the avifauna of the South Orkneys, that of the nearest regions lying to the
south and north of them, and that of the Antarctic Continent.

Before proceeding to do this it will be well to remark that the avifauna of the
South Orkneys, as at present known, comprises 19 species; of these 13, perhaps 15,
are natives, i.e. breeding birds.

Turning our attention first to the south, and comparing the avifauna of the
Orkneys with that of the South Shetlands, which lie to the south and west, we find a
remarkable similarity between the ornis of the two archipelagoes. This similitude is no

ON THE BIRDS OF THE SOUTH ORKNEY ISLANDS. 223

doubt due to analogous climatic and other conditions influencing both animal and
vegetable life. As regards their bird-life, the two groups are practically identical, the
Orkneys only claiming one nesting species which does not occur in the Shetlands,
namely, the petrel Fregetta melanogaster ; while I am not aware that the latter group
possesses a single native species not found in the Orkneys.

When, however, we come to extend our ornithic survey to the nearest northern
land, South Georgia Island, the result is entirely different. Here we and that while
there are 9* native birds common to both, South Georgia has at least 12 t which do
not breed in the Orkneys, while the latter isles can only claim 3 which do not summer in
Georgia, namely Pygoscelis adelife, Phalacrocorax atriceps, and Sterna hirundinacea.
When we examine and compare the climatic conditions prevailing at these two stations
we have the key to these marked differences. In South Georgia, though only six
degrees north of the South Orkneys, the mean summer temperature, Mr Mossman
informs me, is 8°'8 higher, while autumn, winter, and spring are respectively 11°'6,
15°'9, and 10°7 warmer. At South Georgia the lowest temperature recorded was 9°'9
above zero, while at the South Orkneys 40° below zero has been registered. In South.
Georgia no less than 15 species of flowering plants (phanerogams) are known : in the
South Orkneys not one.

Extending our survey in like manner to the far south, and comparing the avifauna
of the South Orkneys with that of the Antarctic Continent, we find that the latter has
only 3 native birds not summering in the Orkneys, namely, the stately Aptenodytes
forsteri, Megalestris maccorrmcki, and Sterna macrura antistropha Reichenow. The
birds common to both are also 3 — Pygoscelis adelife, Oceanites oceanicus, and
Thalassceca antarctica.

The final instalment of the ornithological results of the Scottish National Antarctic
Expedition will be devoted to the birds of the Weddell Sea, southwards of the South
Orkneys, and Coats Land.

Pygoscelis antarctica (Forst.).
Pygoscelis antarctica Cat. B., xxvi. p. 634.

The Ringed Penguin is an uncommon bird in collections, and has hitherto been
regarded as not an abundant species anywhere within the somewhat limited area
in which it occurs ; while its phases of plumage were little known except in the
adult state.

Now, thanks to the work of the Scottish Expedition, we know that the species
is extremely abundant at the South Orkneys ; while the collections brought home

* These are Pygoscelis antarctica and P. papua, Fregretta melanogaster, Pagodroma nivea, Ossifraga gigantca, Daption
caperuis, Larus dominicamis, Megalestris antarctica, and Chionit alba.

t Apttnodytes patagonica, Catarrhactes chrysolophus (possibly a breeder at the South Orkneys), Pelecanoides exsul,
Garrodia nereis, Majaqueus cequinoctialu, Prion desolatus, Diomcdea exulans, Phabetria juliginota, Sterna vittata georgice,
"Phalacrocorax carunculatus Gm. (P. albiventer Less.),* Querquedula eatoni, and Anthus aniarcticus.

224 ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

enable me to describe all the stages of plumage from the newly hatched chick to the
mature bird.

Although not nearly so numerous as its congener P. adeliae, yet next to that species
it was the most abundant of all the birds found at the South Orkneys, where the total
number resorting to Laurie and Saddle Islands for the summer is estimated at over
one million.

This species was first seen by the Expedition on February 2nd, 1903, in lat. 60° S.,
to the north-east of the archipelago. Here a party was observed, some of which were
sitting on an iceberg, others on the water. Two clays later the first landing on the
islands was effected at Saddle Island, where the explorers met with a vast concourse of
these birds, and a number of specimens, young and old, and some eggs were secured.
The rookery at this island is believed to be tenanted by not less than 50,000 birds.

On her return to the South Orkneys in the autumn after the first voyage to the
Weddell Sea, the Scotia encountered Ringed Penguins off the east coast of Coronation
Island on March 23rd. On March 26th she went into winter-quarters in Scotia Bay,
•Laurie Island, and there these birds were observed until April 28th, on which date the
last of the autumn emigrants were seen. They were entirely absent during the whole
of the winter ; and the earliest of the spring immigrants were noticed on November 2nd.
On the following day a few more arrived, and the first bird to land walked straight up
to a small moraine, picked up a stone, and laid the foundation of its new nest. After
this date the immigrants were observed in varying numbers, many arriving on
November 9th.

When walking over hard surfaces to reach their nesting-grounds, it was noticed that
the birds maintained an erect position, marching in column of route ; but when they
came to soft snow they assumed a prone attitude and propelled themselves by means of
their legs ; when ascending a slope, or being chased, they brought their flipper-like
wings into play, using them either alternately or synchronously ; and when descending
from any height they tobogganed.

Eight rookeries were found on Laurie Island. The largest of these were at Cape
Robertson and Ailsa Craig, each of which contained many thousands of nests. There
were two other rookeries almost equal in size, but the rest were smaller, each tenanted
by a few hundred birds, and situated at different places on the coast. On some of the
off- lying Rudmose and Murray Islands there were jumbled rookeries of Ringed Penguins
and Shags (Phalacrocorax atriceps). Dr Pirie tells me that at Cape Robertson and
Ailsa Craig the inhabitants of the great bird-cities were solely composed of the ringed
species. At Cape Robertson, the birds occupied a strip of the coast about half a mile
long extending over the low rocky foreshore and up the gently rising cliffs behind,
until the farthest-inland birds must have been a couple of hundred yards from the sea,
and at a height of two or three hundred feet above it. He reckoned that on an average
there would be about a nest to each square yard, and there could not have been much
less than a quarter of a million birds. Here Dr Pirie and two companions took 1000

ON THE BIRDS OF THE SOUTH ORKNEY ISLANDS. 225

eggs in a very short time on December 12th. Dr Pirie had under more continuous
observation a small congeries which had taken up its abode amidst the Adelie rookery
at Point Martin, Scotia Bay. Here they constituted a small foreign element on fairly
high ground, with their congeners higher up as well as all over the ground between
them and the sea. Although massed together, they apparently get on with their
neighbours as well (or as ill) as with each other. The nests were poor affairs composed
of a few pebbles, varied occasionally by the bones of deceased ancestors. Woe betide
the inhabitant of these cities and villages that strays beyond the boundaries of his or
her domain ; then the beaks of all the penguins around dart out at the intruder and
soon drive it back to its own territory. There was one point where the path up the
cliff was very narrow, and here a constant stream of penguins of both species used to
go up and down on their way to the water. Right on the track were some nests of the
ringed species, and how these birds managed to hatch their eggs is a mystery, for all
day long they were incessantly engaged in pecking at the passers-by, who, though often
in a hurry, frequently stopped and retaliated. The cry of this bird is harsher than that
of the other species, and during the breeding-season it is active and always on the move,
though at other times it is solemn and phlegmatic in temperament. Its pugnacious
disposition made a visit to the rookery a painful adventure, for the protection of long
sea-boots did not always suffice. This bird is a good strategist, and believes in getting
in the first blow. Dr Pirie has seen one take a run of several yards, jump, and fasten
on to an intruder above his boots, at the same time lashing out vigorously with its
flippers. When on the beach or ice-foot mingled with other species, the Ringed
Penguins seemed always to take the lead in entering the water. They took the lead,
too, in repelling the attacks of the Samoyede dog "Russ." He could outmanoeuvre
any Adelie or Gentoo Penguin, but he had frequently to retire before the onslaughts
of the present species, which would face up to him and sometimes deliberately attack
him. It was decidedly the " boss," and jockeyed both the Adelies and the Gentoos.

Occasionally three eggs are laid, usually two, and sometimes only one. A con-
siderable number of eggs were obtained, and these vary in size from 770 cm. x 5*40 cm.
to 6'95 cm. x 5'50 cm. A small egg, one of a clutch of three, measured 4'43 cm. x 3*92
cm. The average weight of fresh eggs was 3 '5 6 oz. The eggs seem to be little known.
They vary in shape, some being almost perfectly oval, others more elongate in form and
narrower at one end. In colour the majority of those in the collection are of a uniform
very pale greenish white, with a thin coating of a chalky nature, such as is found on
the eggs of cormorants (Phalacrocorax) and other birds.

The first chicks were found on January 7th, 1904, and appeared to be about two
days old ; but this was not at the rookery where the first eggs were laid. The young,
though hatched considerably later than those of their congeners, seemed to develop more
quickly, and by February 1 1th some of them were beginning to show the characteristic
black ring.

In the autumn of 1904 Mr Mossman saw this species for the last time on April 26th.

226 ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

The first bird seen in spring was noted on November 14th, and the first eggs of the
season were found on November 27th.

The collection of skins contains only thirteen specimens of the Ringed Penguin,
but these represent the species in all stages of its plumage, and include a magnificent
albino example.

The following is an account of the various stages of plumage, most of them hitherto
unknown, passed through by this species :—

Chick (Laurie Island, January 7th, 1904). — The newly-hatched chick differs
somewhat remarkably from that of its congeners, since it lacks the dark or black head
characteristic of P. adelise and P. papua and is entirely clad in silky-white down,
except in the lower part of the abdomen, where it is partially naked. Bill black. Feet
yellowish. (It would have been more correct to figure this little bird in a nest rather
than erect, but such a mode of treatment would not have shown it to advantage. )

Young in Down (Saddle Island, February 4th, 1903). — There is a great change from
the plumage of the chick to the full-grown young in down. The latter is densely
clothed in short down resembling fur, the upper parts of which are mouse-grey, passing
into pale whitish grey on the head and cheeks ; the hind-neck is tipped with white and
the lores are blackish. The under surface is drab-grey, paler in the centre of the
abdomen ; the chin and throat are blackish. Bill black. Feet yellowish. Wing 4'9
inches. Culmen 1'4 inches. Tail-feathers 1*5 inches.

Young in Down and Feathers (Eillium Island, February 22nd, 1904). — Has blue-
grey feathers on the lower back, tail, sides of the back, and on the edge and tip of the
wing ; a band of blackish feathers on the crown and hind-neck ; lores feathered black,
and the rest of the upper surface covered with mouse-grey down. Under parts with
pure white feathers on the abdomen, lower breast, and chin ; upper breast and neck in
whitish down with a dusky band across the throat, under which the characteristic black
ring or bridle is in evidence. Wing 6'3 inches. Culmen 1'35 inches.

Immature Birds in First Plumage resemble the adults, from which they only differ
in having the back almost entirely blue, i.e. showing little black. Here, again, this
species differs from its congeners, which have more or less pronounced colour-characters
associated with their first plumage.

Adults. — The old birds on their arrival in spring (November) have the blue and
black of the upper surface very bright in tint, but as summer advances (February) the
blue fades and the black assumes a brownish hue. In February, too, some are in deep
moult, the under down shows through the scanty covering of contour-feathers, the
feathers on the wings are ready to drop off in patches, and the birds are quite tailless.
In March and April the new plumage has been assumed, with the exception of the tail-
feathers, which are still quite short, and yet these are the first to be assumed by the
otherwise downy young. The wing in the adult males measures from 7'1 to 7 '5 inches,
and in females from 6 '7 5 to 7 '2 inches.

The average weight of seven adult males taken on the 4th of February 1903 was

ON THE BIRDS OF THE SOUTH ORKNEY ISLANDS. 227

9'1 Ibs., the smallest scaling 7 Ibs. and the largest 1175 Ibs. Eight females averaged
8'65 Ibs., the smallest being 675 Ibs. and the largest 10 Ibs. On February 9th, 1904,
Dr Pirie got specimens weighing as much as 17 Ibs.

The albino is an adult female, and the plumage is entirely pure white with a silky
gloss. The bill is black and the feet are orange. It was obtained on the south beach
of Scotia Bay on the 2nd of February 1904.

Pygoscelis adelise (Hombr. et Jacq.).

Pygoscelis adelix Cat. B., xxvi. p. 632.

Thanks to the researches of the Expedition, the northern range of this truly
Antarctic species has been considerably extended, and the South Orkneys and their
neighbourhood now mark the extreme limits of its ascertained distribution at all
seasons of the year.

The Adelie or Black-throated Penguin is no doubt a resident bird in the archipelago,
for it was observed there all the year round, though only occasionally during the winter
months, which were probably spent on the open sea in the vicinity of the islands.

This bird was first noticed in lat. 60° 30' S. and long. 43° 40' W. on February 3rd,
1903, when the Scotia was nearing the South Orkneys. A number were then observed
swimming after the ship in company with P. antarctica, while others were seen on
the ice, either lying down or squatting. Those walking on the floes presented a comical
appearance, their gait resembling that of an " old salt" just ashore after a long voyage.
In jumping from the water on to the ice they made remarkable leaps of several feet,
but were not always successful, and fell back into the sea. The species does not appear
to have been observed at Saddle Island, but at Laurie Island it was the most abundant
of all the penguins, and its numbers during the summer were estimated at not less
than five millions.

Though a few were noticed throughout the winter of 1903, it was not until October
7th (October 8th in 1904) that the birds commenced to return to their summer haunts.
Over forty were then observed in Scotia Bay, most of them engaged in climbing up the
rocks into the old rookeries as if they had come to stay. They were all in plump
condition and travelled quickly, most of them moving on their bellies at full speed.
On the 10th large bodies were making their way from the open water, and on arriving
at the shore clambered up the rocks at once and made for the rookery. A party of
these birds, accompanied by some Gentoos, was met en route, and as soon as the Adelies
observed the intruders they hurried ahead, moving quickly on their bellies, to meet the
strangers, and on arriving moderately near they stood up, threw back their heads, and
loudly screeched defiance ; but they retreated on being approached, scuttling off in the
prone position at full speed, followed by the more timid Gentoos.

Mr Mossman noted that in the springs of 1903 and 1904 the first great arrivals of

penguins took place immediately after the last cold snap of the season.

VOL. IV. 4

228 ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

At Laurie Island and its off-lying islets no less than fourteen rookeries of Adelie
Penguins were discovered. The largest of these was located on the Ferrier Peninsula,
which for several miles was simply alive with these birds and some Gentoos, the former
being not less than two millions in number. Another vast colony was on Graptolite
Island, and there were smaller though still extensive rookeries on the west side
of Scotia Bay, on Delta Island, Point Rae. and on Watson and Pirie Peninsulas,
with numerous lesser settlements on other parts of the coast and on various small
islands.

The favourite sites for these communities were on plateaux where small stones
abounded, and these were sometimes occupied up to 500 feet above sea-level. As the
season advanced these rookeries became indescribably dirty, being masses of mud with
pools of filth, and the birds themselves became correspondingly defiled.

At the rookery in Scotia Bay the first signs of nest-building were noted on October
10th. By the 20th nearly all were paired, and the appearance of an unpaired bird
gave rise to a fearful commotion, every bird trying to get a billful of feathers from the
unhappy one, while all the penguins in the vicinity raised their voices and screeched
their loudest. The appearance of such wanderers, too, generally resulted in a free fight
among those around.

The nests were heaps of stones deliberately collected one by one from far and near,
even from under the snow. They were hollowed in the centre, and lined with some
bones of their departed brethren, or with dropped tail-feathers when procurable. Some
of the birds sat on the snow until it was thawed down to the stones beneath, and then
set to work to form an irregular hollow in which to lay their eggs. A number of the
nests became covered with snow, in some cases a foot deep, and several were deserted
in consequence. The birds are accomplished thieves, and start their knavish tricks as
soon as nest-building commences, but do not entirely desist when the young are
hatched, though they then practise them to a lesser extent.

Three eggs are sometimes laid, but two is the usual number, and not infrequently
one only. The first egg of the season was found on October 29th, 1903. On the
31st, 739 were gathered on Delta Island, which was covered with these birds. Between
November 2nd and 10th, 2075 eggs were taken for domestic use, and as late as the 21st
a number were obtained from a small rookery in Scotia Bay. The sitting bird incubates
in a procumbent position, the mate standing erect by her side. The period of incuba-
tion was ascertained to vary from 31 to 33 days.

This species is very bold as compared with the Gentoo, and attacks fiercely anyone
who enters the rookery. The birds had always to be forcibly evicted from their nests
if the eggs were wanted. It was quite a business to go through a rookery unless attired
in long sea-boots, and even then the birds sometimes got at the intruder unawares,
taking a running jump and fixing on his legs above the boot, whence they were not
easily shaken off. Other penguins passing the nests came in for violent assault, and
some were seen bleeding, while others were literally pecked to death. An angry bird

ON THE BIRDS OF THE SOUTH ORKNEY ISLANDS. 229

ruffles the feathers on the back of its head and neck, draws back its head, and glares
viciously with eyes and bill wide open. When the old birds leave the nest to go down
to the water to bathe, it takes them a long time to make up their minds to enter the
sea, and a whole crowd collects and walks up and down the ice-foot. They lean over
the edge, as if about to dive, and then retire again and run off to another point to go
through the same performance. When one makes a plunge a number of others immediately
follow. After they dive they roll over and over in the water, and wash themselves
thoroughly with the aid of their feet, gradually getting rid of the red dirt with which
they are bespattered and smeared. On leaving the water they have to jump about
four feet to reach the rock or ice. They often attempt to do this in places which are
too high, and fall back into the water.

The first young were found on December 6th, but probably some of these were
hatched on the 4th. Many were seen on the llth. On the 18th a mother penguin
was observed feeding her chicks. She bent her head until her bill was inclined about
45°, with the lower mandible uppermost, and the chicks sucked in the semi-digested
food brought up, taking it from the hollow between the rami of the upper mandible.
When the young were older they were fed from the beak. Some young under a
fortnight old were found to have already a small geological museum of pebbles in their
stomachs. By January 7th, 1904, the young were beginning to lose their down. The
rookeries at that date were in a greater state of filth than ever, and the stench was
almost unbearable. On February llth not a single old bird was in the rookery or
in the bay, and only a very few young were seen. They had evidently all gone out
to sea.

In 1904, Mr Mossman informs me, the first spring immigrants were noted on
October 8th, followed by several hundreds on the 14th, after which they were
continually arriving at the rookeries. On November 2nd the first egg was found, and
the first chick emerged on December 12th.

The collections contain forty-five specimens, in all stages of plumage, from the
South Orkneys ; also a large number of eggs.

As the various stages of plumage of this species have been carefully worked out
from the material collected by the Southern Cross Expedition, very little remains to
be said on the subject. I would remark, however, that of the thirty-four adult
specimens before me, obtained at all seasons, not one resembles the figure of the adult
bird on plate vii. of the Southern Cross collections. In all the South Orkney specimens
of this handsome species there is much less blue on the back, where black is the pre-
dominant colour, and the head and throat are almost entirely black, the feathers of the
head being merely tipped with blue.

Immature birds show more blue and less black on the upper surface than adults.
Some obtained in February, and presumably about a year old, have the chin entirely
black, and the throat a mixture of black and white. And these same birds vary in the
extent of the black apical spot on the under surface of the wing : in some it is developed,

230 ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

in others it is practically absent. This black apical spot cannot be regarded as a sign
of maturity, as some young birds have it more developed than certain adults — indeed,
one white-chinned example has this spot more pronounced than any other specimen in
the collection.

A fine albinistic male was captured on the south beach at Laurie Island on January
llth, 1904. The plumage of its upper surface is cream-coloured, washed with pale
brown on the hind-neck and crown ; the tail, wings, and under parts are white, except
the chin and throat, which are brown and indicate that the example is an adult ; the
bill and eyes were normal in colour ; and the feet pale on both surfaces.

A series of measurements taken in the flesh, and of the weights, revealed the fact
that there was great diversity in the size of the adults. The males varied in total
length from 28 to 33 '1 inches, and their wings from 7'1 to 7 '7 inches ; the females from
27 to 30 '8 inches, and their wings from 7 to 7 '4 inches.

As regards weight, it would seem that by the end of the nesting-season the weight
of birds of both sexes had run down to a low ebb, indicating, perhaps, that they had
been drawing on the stores of fat laid up since the previous autumn. In April males
ranged from 7 '25 to 10 Ibs., while in October the lightest bird scaled 11 "5 Ibs. and the
heaviest 14'1 Ibs. Females in April ranged from 6 to 8 Ibs., and in October from 9 to
13 Ibs. These results were based upon a large number of specimens.

The temperature of this species was found to be as high as 106° F.

Pygoscelis papua (Forst.).
Pygosceiis papua Cat. B., xxvi. p. 631.

The Gentoo Penguin, which nears the southern limits of its range at the South Orkneys,
was only found in small numbers as compared with its congeners, the total number
at Laurie Island being estimated at 100,000 birds. It was confined to four or five
rookeries, in which it nested in company with P. adelise.

This species was first met with by the Expedition at Saddle Island early in February,
though no mention is made of its nesting there, and later in the season it was observed
off Coronation Island on March 23rd. At the winter-quarters in Scotia Bay many were
observed departing late in March and early in April, and the last of the autumn
emigrants went north on April 25th. Not all of them departed, however, for a few
were occasionally seen during the winter months of May, June, and July. They
increased in numbers during the third week of August, and the spring return move-
ments set fairly in by September, on the 25th of which month they appeared at the
rookery on Cape Dundas, while numbers arrived from the north as late as November 5th.

On October 18th many were observed on their way from the open water to a big
rookery, in company with P. adelise. On reaching the shore they at once clambered
up on to the rocks forming the breeding-ground. Here the Gentoos occupied the
lower and less favourable sites, and formed a ring, as it were, round the Adelies. This
was, no doubt, due to the fact that many of them wandered about aimlessly for some

ON THE BIRDS OF THE SOUTH ORKNEY ISLANDS. 231

time ere commencing nesting, and thus allowed all the more desirable sites to be occupied
by the other species.

The nest was a much better and larger structure of stones than that of its congeners,
P. adeliae and P. ctntarctica, being from seven to eight inches high, and containing
some old tail-feathers and a few bones. The birds, too, were cleaner than the rest all
through the season. They were, however, great thieves, so far as pilfering nesting-
materials was concerned.

On November 14th many of the nests became snowed up, and some of the birds
sat in more or less deep holes in the snow ; many, too, were completely covered.

The first eggs were laid on November 6th. They were usually two in number,
frequently only one, never three. In shape they were very uniform, and rounder than
those of P. adeli&. The period of incubation was found to vary from thirty-one to
thirty-five days.

The birds are somewhat timid ; a few of those incubating were bold enough to peck
at human intruders, but the majority ran off their nests when approached. They were
a little more courageous after the young were hatched, but even then some of them
deserted their chicks without making any pretence at protection. They, however,
fought fiercely among themselves, using both wings and bills, giving some hard smacks
and sharp bites.

The young birds did not commence to lose their down until February llth; but
on one or two the white band across the crown had already begun to show itself, and
the neck to darken in colour.

The collection of skins from the South Orkneys contains specimens in all stages of
plumage, and many eggs were also obtained.

The newly hatched chick is clad in silky down, and is of an olive-grey tint on the
upper surface (darker, nearly black, on the head), but lighter beneath, and the bill is
bluish grey. This stage soon gives place to a darker coat of down, to the tips of which
the paler down of the first coat is attached for a time. In this second coat of down,
the upper parts, including the head, are slate-grey, the plumes of the back having pale
tips, and the under surface is white. When only a few days old, the bill begins to
assume the orange tint characteristic of the adult birds.

The adults obtained in February are in faded plumage, and late in that month and
during March and April had either moulted their tails or had that appendage only in
an incipient stage of growth.

Eighty specimens of mixed sexes, weighed on April 28th, 1903, varied from 8 '5 to
13'75 Ibs. Of these, the heaviest male scaled 1375 Ibs., the heaviest female 12'5 Ibs.

Catarrhactes chrysolophus Brandt.
Catarrhactes chrysolophus Cat. B., xxvi. p. 641.

The presence of this species at the South Orkneys is an interesting discovery,
since it indicates a considerable extension in its hitherto-known range, for there was

232 ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTAKCTIC EXPEDITION.

no reliable evidence before that the Macaroni Penguin had occurred south of the
Falklands and South Georgia in the western Antarctic seas, or of Heard Island in
their eastern waters.

Whether this species is an annual visitor, having breeding-grounds in the South
Orkneys, must remain an open question ; but it would seem not unlikely that such is
the case, perhaps on some of the unexplored islands of the archipelago.*

Five specimens were obtained in 1904 in the penguin rookeries on Laurie Island.
These were mostly captured singly towards the end of summer, and are regarded as
being more or less immature birds. Two of them, namely those last obtained, are
decidedly younger than the rest, and have the merest indications of yellow feathers on
the sides of the crown, and also small bills. The remaining three are considered to be
not fully adult, and have well-developed tufts of golden-yellow (not orange) plumes,
but are otherwise mature in plumage and in the dimensions of their bills.

The two younger birds are, there can be little doubt, birds of the year, and their
presence seems to point to the South Orkneys being their native land, for it is difficult
to believe that birds only a few weeks old could have accomplished the rough sea-
passage of 600 miles from their nearest known breeding-station at South Georgia.

The three older specimens, though not fully adult, are probably about a year old,
though whether this species breeds at such an age is uncertain.

None of these birds were observed in the autumn of 1903, when the Expedition
arrived, and their appearance in the summer of 1904 was a surprise to the explorers.

The first specimen, a male, was captured on January 7th in a big penguin rookery at
Scotia Bay, where it was found amongst a crowd of P. adelise. Ten days later a female
was secured in exactly the same place ; and on the 29th of January another male, just
below where the previous captures had been made. These three were the not fully adult
birds alluded to, and the place where they were taken was so frequently visited that
it is thought to be most unlikely that they could have been bred there without
being detected.

One of the younger birds was taken higher up in the same rookery on December 29th,
and the other was captured on the beach on February 6th. Both were males.

Regarding the age of these specimens I was somewhat uncertain, and sought the
assistance of my friend Dr A. E. Wilson, of the National Antarctic Expedition, who
had had considerable personal experience with the allied Catarrhactes schlegeli, and
whose aid it is a pleasure to acknowledge.

In connection with the occurrence of this species at the South Orkneys, it is well to
recall the fact that Capt. C. A. Larsen, of the Norwegian sealer Jason, informed
Dr Donald,f of the whaler Active, that he saw a rookery of Crested Penguins on the
South Orkneys. These birds he described as being intermediate in size between the

* A Crested Penguin, supposed to be of this species, was found by Mr Paulsen on November 29th, 1907,
"occupying the same nest with one of the 'Kinged' species."— Mossman, Scottish Geographical Magazine, xxiv.,
1908, p. 354.

t Of. Proc. Roy. Phys. Soc. Edin,, xii., 1894, p. 335.

ON THE BIRDS OF THE SOUTH ORKNEY ISLANDS. 233

Emperor and Adelie Penguins, and as having a yellow patch under each eye [? the
yellow angle of the gape] and a red superciliary crest three or four inches long.
This might well be regarded as a glorified description of fully adult examples of the
present species.

It is probable, too, that this is the species of Catarrhactes observed and obtained
by the Swedish Expedition on Nelson Island, one of the South Shetlands, which was
thought to belong to C. chrysocome. Most unfortunately the specimens were lost with
the wreck of the Expedition (cf. Lonnberg, Wissen. Erg. Schivcd. Siidpolar-Exped.,
Bd. v. Lfg. 5, p. 3).

Aptenodytes forsteri G. R. Gray.
Aptenodytes forsteri Cat. B., xxvi. p. 626.

The Emperor Penguin claims mention for the South Orkneys on the strength of the
following incident :—

On November 21st, 1903, two sailors reported having seen, by some open water at
the mouth of Scotia Bay, a large penguin, which was "three times the size of an
Adelie," but having black feet and a bill like a Gentoo, though with no mark on the
head like the latter species.

The bird unfortunately escaped into the water when the men attempted to capture
it. There can be no doubt about the size of the bird having been correctly described,
as the Adelies were close at hand for comparison. The two men who reported these
facts were among the most careful and trustworthy of the crew, and the conclusion
come to at the time was that the bird seen by them was an immature Emperor
Penguin.

Note. — As reported by Mr Mossman,* two Emperor Penguins were captured by
Mr Paulsen, of the Argentine Government Meteorological Station at Scotia Bay, South
Orkneys, on May 4th and 31st, 1907, respectively, and individuals visited the islands
also in the years 1905 and 1906, usually during the autumn and winter. — EDITOR.

Oceanites oceanicus (Kuhl).

Oceanites oceanieus Cat. B., xxv. p. 358.

Wilson's Petrel is a common summer visitor to the archipelago, and though not
nearly so abundant as either the Cape or Snowy Petrels, yet resorts in thousands to
Laurie Island to nest on the cliffs of its remarkably extensive coast-line. It was also
observed at Saddle Island during the short visit of the Expedition on February 4th,
1903, and was probably breeding there.

In the autumn of 1903 it was last seen on March 23rd, as the Scotia was approaching
the islands from the south, on her first voyage from the Weddeli Sea. It was never
observed during the winter months, and did not appear until late in the spring, namely

* Scottish Geographical Magazine, xxiv., 1908, p. 354.

234 ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

on November llth,* being the last of the summer visitors to arrive, On the 23rd the
ice broke up and many birds arrived, including numbers of this species. After this
date it was constantly under observation, for several dozens took up their abode in the
cliff above the observatory, where, on December llth, the first egg was obtained.

There was no attempt at nest-making ; the egg was simply laid in a hollow in the
earth in narrow clefts and fissures in the face of the cliffs, under boulders, and some-
times under stones on the screes sloping from the foot of the precipice, at heights vary-
ing from 20 to 300 feet above sea-level. It was often placed far in, and this and the
fact that the hole was so narrow made the egg difficult to procure. Some of the eggs
were laid at such a distance from the entrance that a spoon had to be lashed to a long
bamboo in order to reach them. The searchers could hear the low whistle uttered every
few seconds by the sitting bird, but on reaching the spot whence it seemed to proceed
the sound would appear to come from an entirely different direction. The dog
" Russ " proved to be a great aid in work of this kind, for he easily detected the bird's
presence by his keen sense of smell. When caught on the egg the birds brought up a
reddish fluid, which issued both from the mouth and nostrils. In addition to the low
whistle, these petrels had a harsh screaming chuckle. These noises they kept up almost
continuously after dark, especially on still nights.

They appear to return year after year to the same nesting-places, for both eggs and
dead young birds of previous seasons were numerous in the tenanted holes containing
the fresh eggs. This fact indicates that a very serious waste of life takes place in some
seasons, if not annually. It may be accounted for by the late arrival of the bird at its
breeding-stations, which, coupled with the lengthened period of incubation character-
istic of all petrels, results in winter setting in ere the eggs are hatched, or the young,
which develop slowly, are old enough to leave the nesting- holes. Another, and
perhaps more probable, explanation is that the disasters noticed were due to a succes-
sion of cold summers, which are actually known to have occurred. None of the eggs in
the summer of 1903-4 had been hatched when the Expedition left the islands on
February 21st. These facts would seem to indicate that the South Orkneys lie at the
extreme limits of possible breeding for Wilson's Petrel. Indeed, for many individuals
of this species, perhaps all, during some seasons the climatic conditions place the islands
distinctly beyond that range ; though it breeds further south, most likely with similarly
disastrous results.

From 7 to 1 1 p.m. these birds flitted about the cliffs and over the head of Scotia
Bay in great abundance, and in striking contrast to their habit in the daytime, when
only occasionally was one to be seen on the water, though there were probably many at
sea off the islands.

The nest figured was situated at the bottom of a crack in the rock, about four

* This and other species appear to be remarkably constant as to the times of their appearance and departure at the
South Orkneys. As an instance of this, it is interesting to note that Mr Mossman observed the first Wilson's Petrel
in the spring of 1904 on November 12th.

ON THE BIRDS OF THE SOUTH ORKNEY ISLANDS. 235

inches wide and two feet deep. It was the only one found which was open enough to
permit of a photograph being taken, and then only under particular circumstances
as to time. At about 7 a.m. the sun shone for a few minutes directly into the crack,
and it was during those moments that this unique picture was secured.

Eight eggs average 33'7 x 24 mm. The largest is 36 x 24 mm., and the smallest is
32 x 23 mm.

Fregetta melanoyaster (Gould).
Cymodroma mdanogaster Cat. B., xxv. p. 364.

On December 5th Dr Pirie discovered a pair of unknown petrels. He heard a low
whistling sound proceeding from a crevice in a rock on the east side of Uruguay Cove,
Laurie Island, and about fifteen feet above the sea, and on climbing up found what he at
first thought to be a pair of Wilson's Petrels, and managed to secure the female. Two
eggs, badly broken in the endeavour to capture the birds, were found near the mouth
of the crack — one of them obviously of a previous season, the other deeply incubated.

On examining the captured bird it was at once evident that it was not a specimen
of Oceanites oceanicus, for it had entirely black feet, had white on the under surface,
the feathers of the back slightly edged with white, a longer and more hooked mandible,
and strongly upturned nasal tubes. On the return of the Expedition, I found this bird
to be an example of Fregetta melanogaxter — the Black-bellied Storm-Petrel.

The dimensions of the egg secured were 3 '60 x 2 '5 5 cm., and correspond with those
of F. vnelanogaster, from the Falklands and Kerguelen, in the British Museum collec-
tion. The locality was again visited in the hope that the escaped bird might be found.
It was not there, however, nor were other individuals of this species observed elsewhere
in the islands.

The occurrence of this species is one of the most interesting ornithological dis-
coveries made by the Expedition. It implies a remarkable extension in its known
range, and removes the doubt which has hitherto overshadowed (cf. Antarctic Manual,
p. 228) the record of its having bred at South Georgia, as mentioned by Pagenstecher
(Die Vogel Siid-Georgiens, p. 18, 1885), in the southern summer of 1882-1883.

Thalassceca antarctica (Gm.).

Thalatsceca antarctica Cat. B., xxv. p. 392.

A few examples only of the Antarctic Fulmar were seen at the South Orkneys; but
it is thought by the members of the Expedition that it may possibly have bred on the
east side of the Ferguslie Peninsula, along with the Cape and Snowy Petrels, in the
summer of 1903.

A number of these birds were seen in the previous autumn when the Scotia
was between Saddle Island and Cape Bennet, the northern limit of Powell Island, on
March 23rd, 1903. Several were again observed on the following day in Leathwaite
Strait, between Coronation Island and Powell Island.

VOL. IV. 5

236 ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

On June 1st, when winter was well advanced, Dr Bruce noticed one flying round the
Scotia ; and another is believed to have been seen at the open water in Scotia Bay
on August 17th.

There are no South Orkney specimens in the collection, but a number had been
obtained in the Weddell Sea before the Expedition arrived at Laurie Island and went
into winter-quarters there.

Priocella glacialoides (Smith).
Priocella glacialoides Cat. B., xxv. p. 393.

The Slender-billed Fulmar, or Silver Petrel, was observed in the summer of 1903,
in MacDougall Bay, on the north coast of Laurie Island, on November 4th. After this
date examples were occasionally seen about the cliffs on the north side of the island
during November and December, and it is considered highly probable that a few pairs
were nesting there. The breeding-places of this bird, however, still remain to be
discovered.

This species was first observed during the previous autumn, when a number came
under notice between Saddle Island and Cape Bennet, the north end of Powell
Island, on March 22nd, 1903 ; and again on the following day when the Scotia was
in Leathwaite Strait, between Coronation Island and Powell Island, in search of winter-
quarters. It was also seen in numbers off the north-west end of Coronation Island
on February 14th, 1904.

It had been seen commonly, and specimens obtained, in the Weddell Sea just prior
to the date of the above observations.

Pagodroma nivea (Gm.).
Pagodroma nivea Cat. B., xxv. p. 419.

The Snowy Petrel of Cook was not only an abundant summer bird, but was by far
the most numerous of the few species which remained for the entire winter at the
South Orkneys.

In summer it frequented the high precipitous sea-cliffs which formed its breeding-
haunts, and where, during the nesting-season, some 20,000 birds were estimated to
be present on Laurie Island alone. It was never seen on the hills at the head of the
ice-sheets.

It was also found at Saddle Island and was nesting there. The single eggs were
laid under rocks, in caves, and in holes and crevices on the steep cliffs facing the sea,
at heights ranging from a few to several hundred feet above the water. The nests were
rough primitive structures, and consisted of a few stones or a little earth. They were
less accessible than those of the Cape Petrel, and mostly isolated ; but in one cave
under Mount Ramsay a dozen or more eggs were taken. This bird does not fly off' when
its nest is approached, but retreats a little, and ejects an oily fluid at the intruder,
uttering all the while shrill cries.

ON THE BIRDS OF THE SOUTH ORKNEY ISLANDS. 237

The first eggs were obtained on December 2nd, but were not quite fresh. By the 4th
all the birds seemed to have laid, and eighteen eggs were found, most of them in a cave
from twenty to twenty -five feet above sea-level. The cave was thickly carpeted with
the dung, and the nests, unlike the rough examples outside, were'all well formed in the
dung and had a few feathers in them. Some were placed as much as forty feet from
the entrance, where it was almost dark. In 1904 the first eggs were observed on
November 25th (Mossman).

Young birds were found on January 28th, 1904, but the parents were not present
with their chicks — not an unusual circumstance during the daytime with certain birds
of this order. When discovered, these chicks uttered the same harsh notes as arc
characteristic of the old birds. Their stomachs were found to be crammed with
crustaceans.

The young bird does not seem to have been described. One about one-third grown,
and captured on January 28th, 1904, is clad in long fluffy down which almost conceals
the feathers appearing on the wings and tail. The down is of a lavender-grey tint on
the back and chest, darker on the head, and dull ivory-white on the abdomen.

Ossifraga gigantea (Gm.).
Ossifraga gigantea Cat. B. xxv. p. 422.

The Giant Petrel was present at the Station all the year round, but was very much
less numerous during the winter months. There was a decided falling off in May, but
the lowest ebb was reached in June and continued until September, when the summer
birds of this species commenced to arrive. During the nesting-season it was estimated
that about 5000 were on Laurie Island alone, and when one remembers the savage
nature and almost insatiable appetite of these giants, it is easy to realise what a terrible
scourge they must have been to the penguins, upon which and their eggs and young
it was their one aim to gorge themselves to repletion.

They were to be seen everywhere in the summer-time, but their rookeries were
confined to the north and east coasts. Three of these rookeries were visited, two of
which, namely those on the Watson Peninsula, contained two hundred nests each,
while the third at Cape Geddes comprised only about one hundred. One of the larger
colonies was situated on bare rocky ground from 300 to 400 feet above sea-level, and
the other on a moraine at an elevation of from 250 to 300 feet. The nests consisted
of great piles of small angular stones, and were about two feet in diameter. The third
and smaller rookery was on a low strip of ground between a cliff and the shore, and
was close to the sea ; the nests were similar to the others. Although these contained
no eggs on November 3rd, yet the birds allowed a close approach, one of the parents
sitting on the nest, the other usually standing close alongside.

The first eggs were laid on November 4, but four only were found on that date.
On the 19th, however, eighty were obtained, all single specimens, except in two

238 ORNITHOLOGY Of THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

instances where two were found, probably laid by as many females. The birds had to
be pushed off' the nests ere the eggs could be taken, for very few flew away of their own
accord. They showed no fight when evicted, and usually sat down a yard or two away ;
nor did they shoot oil from their nostrils, but they vomited the contents of their
stomachs, not as a mode of defence, but to get rid of ballast in order to take wing.
They resorted to the same lightening process when chased. Unfortunately, the weather
conditions and those of the ice did not permit of these rookeries being again visited, so
that the period of incubation could not be ascertained nor the capture of young be
effected.

The average length of 80 eggs was 10 '38 cm. and the breadth 6 "5 7 cm.

This species was observed on Saddle Island, and was thought to be breeding on the
adjacent rocks.

The heavy toll ruthlessly demanded from the penguins was very manifest on visit-
ing their rookeries. Here abundant remains of recently killed young penguins, in the
shape of clean-picked skins and bones, were lying all around, while the gorged feathered
giants were either waddling about or sleeping off the effects of their orgies on the
neighbouring snow-slopes. They were observed to feed on dead seals, and during the
winter resorted to the ship's refuse -heap in search of scraps of meat. They were very
bold when in want of food, and one swooped down close to the cook and tore a piece
of flesh off a dead penguin.

The proportion of birds in pure white plumage in the rookeries was not more, per-
haps less, than two per cent. The colour of the birds ranged from very dark brown
through all shades of chocolate, and from grey through light grey and mottled white to
white. Some of these facts indicate interbreeding between the two forms and, perhaps,
between their offspring and typically coloured birds and others. Dr Pirie thinks that
they interbreed, because he has no recollection of seeing two white birds together on
the nesting-grounds.

Four specimens in the collection are from the South Orkneys, and two of these are
of the white form. The weight of these birds varied from 7 '25 to 10 Ibs.

Daption capensis (Linn.).

Daption capensis Cat. B., xxv. p. 428.

Although the Cape Petrel or " Cape Pigeon " is one of the most familiar birds to
voyagers in the southern oceans, and one, too, that has been known since the days of
Dampier (that is to say, since the closing years of the seventeenth century), yet the eggs
remained entirely unknown until December 2nd, 1903, when Dr Pirie took the first
specimens at the South Orkneys.

The three nests from which eggs were then obtained were placed on open exposed
ledges of cliffs on the west side of Uruguay Cove, Laurie Island, at heights of from
twenty to a hundred feet above sea-level. The nests were composed of a few small

ON THE BIRDS OF THE SOUTH ORKNEY ISLANDS. 239

angular fragments of rock and a little earth, and contained single eggs, which were quite
fresh. When approached, the sitting birds ejected an evil-smelling reddish fluid com-
posed of the semi-digested remains of crustaceans of the genus Euphausia. It was
extremely disagreeable to the collector to receive it in his face when peering over a
ledge, and the odour of it was found to cling to clothes for a very long time. The birds
can squirt this fluid with great precision for a distance of six or eight feet. They did
not leave their nests readily, and even allowed themselves to be captured while sitting.
The pure white eggs seemed very large for the size of the bird.

On December 3rd three more eggs were obtained. There were six nests on the ledge
where they were found, but three of them were empty. On the following day about
two dozen eggs were taken on the cliffs under Mount Ramsay, and on the 5th some
fifty eggs were found on the cliffs on the east side of Jessie Bay. The birds seemed
to be of a sociable nature, for several were frequently found nesting near to each other
on the same ledge, but isolated nests were not uncommon.

The work of collecting the eggs of this species proved to be such an unpleasant
business, owing to its nasty methods of defence already alluded to, that a long ski-pole
was used. With this the birds were pushed off their nests, and the eggs secured with-
out the captor being defiled. When thus removed they took short flights, and then
alighted near the nest. Both birds were often found sitting side by side (one on the
nest and the mate close alongside) and cooing and clucking to each other, though not
to the same extent as during the month previous, when courtship was in full swing.

On December 12th more eggs were procured from the locality in which they were
obtained on the 5th, and the nests robbed on that day, though still empty, were covered
by sitting birds. On January 13th, 1904, a fresh egg marked on December 2nd was
found chipped, so that the period of incubation was not less than forty-two days. On
January 18th a chick five days old was taken for a skin, and young birds were still in
down on February 5th, after which date the state of the ice did not permit of further
observations being made ere the Expedition left for the far south.

It was noted that before laying its eggs this petrel sits close on the nest for about
a month, and it was also observed that it entirely disappeared from its nesting-haunts
for some ten days before the first eggs were laid.

The eggs vary from oval to elongate-ovate in form. Taking two extreme forms, I
find their dimensions to work out as follows : — Oval type, 56'5 x 43 mm. ; elongate-
ovate type, 67'2 x 43'3 mm. The average of a large number of specimens is 62'35 x
43'11 mm. The length varies from 56'5 to 67'2 mm., and the breadth from 46'5 to
40'5 mm.

In 1904 the first eggs were laid on December 3rd, or one day later than in the
previous year (Klossman).

The numerous nests found were placed either on ledges of cliffs, or, though these
were few, in hollows in the earth and among small stones on steep scree-slopes, and all
were quite open. These are noteworthy facts, for the nests (containing young) found

240 ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

previous to the discoveries of the Scottish Expedition were obtained in burrows and
grottoes on the island of Kerguelen. There is little doubt that the Cape Petrel breeds
at South Georgia, and Mr Mossman tells me that he saw it in numbers off Deception
Island, one of the South Shetlands, in the height of the nesting-season.

This species is a summer visitor to the South Orkneys. In the autumn of 1903 it
was only once seen after April 21st, on which date a flock was observed flying north,
and was entirely absent during May, June, July, August, and September. The first of
the spring immigrants was seen on October 1st, but the bird was not noted again until
the 23rd, after which date it became frequent.

About 20,000 resort to Laurie Island for nesting purposes, and they are found in
hundreds all round the coast. In Uruguay Cove alone there were over one hundred
accessible nests, and many others were out of reach. They also nest on Saddle Island,
where both young and old were obtained on February 4th, 1903, and are doubtless
abundant throughout the other islands of the archipelago, which may be regarded as a
metropolis of the species.

They were never observed flying over the land, but were to be seen on the wing in
front of the cliffs (not wheeling high over them, like Pagodroma nivea) or sailing over
the sea.

The chick in down, five days old, taken on January 18th, 1904, is slate-grey above,
and paler and sooty on the under surface.

A young bird obtained at Saddle Island on February 4th, 1903, has the head and
body clad in down, with feathers developing on the wings and scapulars. The down on
the upper surface is sooty (darker on the head and cheeks) and paler and greyish on
the under parts. The wing-quills, the largest of which are 2 inches in length, are black,
some of them with the inner webs white towards the base. The feathers of the scapulars
are black and white. There are no signs of tail-feathers. Wing 8 inches.

The mature birds from the South Orkneys and the Weddell Sea present two types
of plumage. The first of these, which perhaps represents old birds in weathered dress,
were captured towards the end of summer (in February) ; and in them the dark portions
of the plumage are blackish with a brown cast, the head alone being black ; the feathers
of the mantle have whitish bases ; and the marginal and lesser coverts show less white
than in the next form. In the second type the dark portion of the plumage is slate-
black, and the bases of the feathers of the mantle are dusky. Specimens in this phase
were obtained early in the autumn (late in March), and are either in new or first
plumage. A male captured on the nesting-ledges on December 3rd, 1903, is intermediate
in plumage between these two forms.

Prion banksi Gould.
Prion banksi Cat. B., xxv. p. 434.

This " Whale-bird " fairly claims a place in the avifauna of the South Orkneys on
the strength of specimens seen off Coronation Island, within the territorial waters of

ON THE BIRDS OF THE SOUTH ORKNEY ISLANDS. 241

the archipelago, on November 27th, 1903, the day on which the Scotia left her winter-
quarters to proceed to the Falklands to refit.

It had been frequently observed on the outward voyage of the previous year, but
fell off rapidly in numbers as the pack-ice was entered, and ceased to be noted some
sixty miles ere the South Orkneys were reached. It was also seen, and specimens were
obtained at sea, to the eastward of the group during the early days of the first voyage
in the Weddell Sea.

Sterna hirundinacea Less.
Sterna hirundinafta Cat. B., xxv. p. 52.

The White-rumped Tern was first observed by the Expedition at Saddle Island, on
February 4th, 1903. It was not found to be an abundant species at Laurie Island,
where only some two or three hundred spent the summer, nesting in small scattered
colonies of about a dozen pairs, and also in isolated pairs.

In the spring of 1903 the first terns of the season were observed on October 21st,
but it was thought that some had been heard two or three days before. The bird had
been absent from the island since the 25th of March of the previous autumn.

The nests were mere hollows, lined with a few small fragments of stone, on the tops
of small rocks, or on raised beaches and small screes, and were always quite close to the
shore. The nests in the colonies were placed fairly close together, and often in proximity
to those of Lams dominicanus. When not surprised on their nests, these birds usually
betrayed the whereabouts of their treasures by hovering over them and screeching
loudly.

The eggs were one or two in number. The first were found on November 14th, and
from that date onwards they were observed until January 15th. In 1904 Mr Mossman
records the first eggs observed on November 27th.

The earliest chicks were obtained on December 25th, and by February 7th young
were noted as having lost all their down.

A few adult specimens and a chick are included in the collections ; also a number
of eggs, averaging 473 cm. x 3 '3 4 cm.

This species is also a summer visitor to the South Shetlands. But according to
Reichenow the South Georgian bird is a subspecies of Sterna vittata, which he has
named Sterna vittata georgite (Om. Monatsber., xii. p. 47); while the same authority
has described (I.e.) the tern of the Antarctic Continent as a race of the Arctic Tern,
Sterna macrura antistropha.

Larus dominicanus Licht.
Lariis dominicanus Cat. B., xxv. p. 245.

This Black-backed Gull has a remarkably wide latitudinal distribution, ranging as it
does from 1 0° S. in the South Atlantic to within a few degrees of the Antarctic Circle.
It was not a very abundant species at the South Orkneys, and the numbers visiting

242 ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

Laurie Island as a summer resort did not exceed some three hundred birds. It was also
observed at Saddle Island in the late summer, and had apparently been breeding there.

The chief nesting-haunts on Laurie Island were at Point Davis on the south coast
and Uruguay Cove on the north. At each of these places about a dozen nests were
found. Elsewhere it was found less abundantly, mostly in isolated pairs, all round
the coast.

The Southern Black-backed Gull was one of the few species that was observed all the
year round, for some of them braved the severities of the winter, and were seen more
or less frequently in the neighbourhood of the Expedition's winter-quarters at
Scotia Bay.

The return of the spring immigrants commenced in mid-October. The birds were
seen pairing on November 3rd, and the first eggs were laid on November 15th (on the
23rd in 1904 (Mos#man)). The first young are mentioned under the date of December
26th, and are described as being then about a week old. Fresh eggs marked on
December 3rd were found chipped on the 28th, indicating an incubation -period of about
twenty-five days. Young still in down were observed as late as January 30th, 1904.

The nests were placed on raised beaches, small screes, and rocks within a few yards
of the shore. The nest was a well-built structure of seaweeds, mosses, lichens, and
feathers ; and was usually surrounded by great quantities of limpet-shells, this mollusc
being evidently a favourite food of the bird. The eggs were usually two in number,
but sometimes three were found, and occasionally only one.

On April 15th Messrs Bruce, Pirie, and Wilton saw an entirely white gull, re-
sembling in all other respects this species, of which it may have been an albino
specimen ; and on September 21st an almost white example of this gull was seen, in
which the wings and upper surface were much lighter than usual.

The collection contains skins of this species in various stages of plumage, and a
number of eggs.

Megalestris Antarctica (Less.).

Megalestris antarctica Cat. 13., xxv. p. 319.

About five hundred Antarctic Great Skuas spend the summer on Laurie Island,
taking up their quarters in the vicinity of the penguin-rookeries, where they revel
among the eggs and young of their neighbours. They were also observed nesting in
similar situations on Saddle Island.

During the southern autumn of 1903 they were seen daily until the 28th of April,
on which date the last bird of the season was met with. They were entirely absent
during the winter months, and the first spring immigrants were noted on October 16th.
These were followed by a few others on the 26th, after which they gradually became
abundant.

On November 22nd two skuas, presumably males, were observed fighting fiercely
while a third was looking on, evidently an interested spectator. The birds fought with

ON THE BIRDS OF THE SOUTH ORKNEY ISLANDS. 243

bills and claws for nearly an hour, when one of them became quite exhausted ; and then
the victor flew off with his bride, and the vanquished was ruthlessly torn to pieces and
devoured by a Giant Petrel, which had been an interested spectator of the fight.

The first eggs were laid on December 2nd, and young birds a week old were found
on January 29th. By February llth, dark feathers were appearing on the wings and
sides of the breast of these youngsters.

The period of incubation was not precisely ascertained, but was believed to be
about six weeks.

In the spring of 1904 the skuas returned on October 21st; and the first eggs were
found on November 27th (Mossman).

The nests were usually placed on the tops of mossy rocks, or on plateaux from 100
to 400 feet above the sea, and consisted of well-made hollows in the moss, while teased-
out fragments of moss formed the lining. Occasionally nests were found on the tops
of moraines, and were then hollows in the earth lined with lichens.

The eggs were two in number, and on these the bird sat very close, her mate usually
remaining near at hand. When the nest was approached the owners screamed defiance,
and if the eggs were wanted the sitting bird had to be forcibly ejected from the nest —
not a very pleasant proceeding, as the sentinel bird wheeled above and dashed at the head
of the intruder, though never actually striking him. When a dog, however, appeared
upon the scene both birds swooped down on it, and sometimes struck it with their
wings. The nests were surrounded by many shells of eggs and remains of young
penguins. The young — pretty little masses of light brown down — soon wander away
from the nest, and are most difficult to detect among the moss, which they closely
resemble.

These birds were to be seen incessantly hovering over the penguin-rookeries, and
swooping down ever and anon at the sitting birds to snatch their eggs or young. On
such occasions the penguins combined in screeching at the harpies, but to little purpose.

Many were about the house all the summer, being attracted by the remains of
penguins thrown out by the cook. Nearly one hundred were observed around a seal's
carcase ; while dead Giant Petrels, and even deceased members of their own species,
did not seem to come amiss as food.

Numerous specimens were obtained at the South Orkneys. These vary in colour,
though mature and obtained at identical periods. Specimens captured in November,
soon after their arrival on the nesting-grounds, were of two types. One had the
ground-colour of both upper and under surfaces dark, being of a deep blackish brown,
rather paler below, and showing comparatively few light markings on the mantle and
scapulars ; indeed in some specimens the back is practically uniform. The other type
is less numerously represented in the collections, and is much paler (drab) generally,
except on the head ; while the feathers of the interscapulary region and under surface
have grey-buff margins. In these light- coloured birds the yellow streaks on the neck

are much more numerous and pronounced than in the darker birds ; and they agree
VOL. IV. e "

244 ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

with the form described by Saunders (Brit. Mus. Cat. Birds, xxv. p. 320) as inhabiting
the Falklands, except that they are not smaller in size than the ordinary dark form,
their wings measuring 16 '6 5 inches, as against 16 to 17 inches in the last-mentioned.
The Falkland Islands bird has recently been described by Lonnberg ( Wissen. Erg.
schwed. Siidpolar-Exped., Bd. v. Lfg. 5, p. 8, 1905) as a sub-species under the name
of M. antarctica falklandica.

It is of interest to remark that one of these light birds was observed to be mated
with one of the dark examples.

Megalestris maccormicki (Saund.).
Megalestris maccormicki Cat. B., xxv. p. 321.

Mr Mossman informs me that a specimen of McCormick's Skua was procured by the
Argentine naturalists at Laurie Island on November llth, 1904, and is in their
collection of birds. Mr Mossman saw this bird in the flesh and examined it, and he tells
me that it was quite different from the skuas, light or dark, which bred at the South
Orkneys. This South-Polar bird has not hitherto been obtained so far north as the
South Orkneys, indeed I believe not outside the Antarctic Circle.

Chionis alba (Gm.).
Chionis alba Cat. B., xxiv. p. 710.-

This Sheathbill, the " Paddy " of the explorers, was an abundant species, and
though chiefly a summer visitor to the islands, yet wintered in small numbers at
Scotia Bay, being attracted by the refuse cast out from the ship. Some wintered away
from the ship at the seal-haunts on the north side of the island.

In the summer it was present in all the penguin and shag rookeries, as many as
two hundred haunting some of the larger colonies. Altogether it is believed that from
2000 to 3000 of these birds passed the Antarctic summer of 1903 on Laurie Island
alone. Adults and young birds were present in considerable numbers at Saddle
Island on the occasion of the Expedition's visit early in the previous autumn, namely
on February 4th, 1903.

It was also fairly abundant around Scotia Bay in March, but towards the end of
April, when the temperature approached zero, the numbers fell off considerably, and
when winter conditions became fairly established only some twenty or thirty remained,
and for many days during that drear season were the only living creatures observed.
These pensioners eked out an existence on the refuse odds and ends which were daily
thrown out from the Scotia. One of the birds became very tame, and for a number
of days in succession visited the ship, remaining all day either in the fo'c'sle or
in the galley.

ON THE BIRDS OF THE SOUTH ORKNEY ISLANDS. 245

Late in September and during the first half of October many returned to their
summer-quarters, and their numbers greatly increased when the ice broke up on
November 23rd.

The first eggs were found on December llth, when eleven (two clutches of three,
two of two, and a single egg) were taken, but some of these proved to be considerably
incubated.

A nest found on December 3rd was on a ledge under an overhanging rock, and was
composed of small stones and penguins' tail-feathers. Five nests were found on the
llth in the large penguin-rookery in Scotia Bay : four of these were on the fringe of
the colony and quite low down, being only from ten to twenty feet above sea-level, and
placed in crevices of rocks or underneath boulders on the moraine ; while the other
nest was under a large boulder about one hundred feet up the moraine, and right in
the midst of the penguins. These nests were mainly composed of the shells of
penguins' eggs, bones, and feathers, and a number of limpet-shells. The position of
the nest is not difficult to detect, for one of the birds generally sits on a rock close by.
The eggs are usually three in number.

Fresh eggs marked on December llth hatched on January 7th, an incubation period
of twenty-eight days. The newly-hatched young are clad in brown down and show
conspicuous bare patches ; they are not by any means pretty objects like the young
penguins and skuas. On January 29th white feathers were beginning to develop under
the down of these chicks ; and by February llth the down had nearly all disappeared.

These birds were found to be very tame and unwilling to fly ; indeed, some of them
would not take wing when pushed with a stick, and most allowed an approach to within
striking distance ere they walked leisurely away.

In the penguin-rookeries they were to be seen perched in prominent places, on
the look out for dead birds or broken eggs. They are very bold, and one was observed
to abstract an egg from under a sitting shag, which was somewhat disconcerted at
having its photograph taken for the first time. Sheathbills were seen to revel in
garbage of every description, including the excrement and placentae of seals. Crustaceans
were found in the stomachs of some of those dissected.

The young bird figured is about one-third grown. The wings, scapulars, and
flanks have white feathers with a little down. The head, sides and back of
the neck, lower part of the back, and abdomen are clad in grey down mottled
with brown.

The temperature of an adult bird, taken on March 26th, 1903, was found to
be 107°'3 F.

The collection contains a few skins of adults and the young bird described, also a
small number of eggs. The latter are elongate-ovate in shape, and in colour white
boldly blotched with greyish black or dark brown and liberally freckled with the same
tints. They do not vary much in size, and measure from 54 to 58 mm. in length by
37 to 39 mm. in breadth.

246 ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

Phalacrocorax atriceps King.
Phalacrocorax atriceps Cat. 15., xxvi. p. 390.

It had long been known that a species of Phalacrocorax nested in the icy regions
of the Antarctic, for Ross found a " Cormorant " breeding at Louis Philippe Land, and
saw innumerable examples at Cockburn Island on January 6th, 1844. The specific
identity of these Antarctic Shags remained somewhat uncertain until the Scottish
Expedition finally settled the matter at the South Orkneys in 1903.

The Blue-eyed Shag, as the explorers termed this species, was present all the year
round in the archipelago.

In summer it was numerous, but avoided the main islands and sought nesting-
places on small islets or rocks off the coast of Laurie and Saddle Islands, where it
was estimated that about two thousand five hundred pairs were breeding, and
where some of the rookeries contained as many as two hundred nests. In winter it
was much less numerous, but the species never escaped observation for many days
in succession.

On August 2nd a flock containing several thousands, probably early immigrants,
was seen from the Scotia as she lay in her winter-quarters.

The first eggs were obtained on November 8th, on a small islet some forty feet high
situated off the north coast of Laurie Island, where a few Ringed Penguins were also
nesting. The nests were well-built structures composed of seaweed, moss, lichens, and
feathers. Some of the birds were still engaged in nest-building, and were diving and
bringing up masses of seaweed in their bills, while others were busily engaged in
picking mosses and lichens off the rocks. They were great thieves, even worse than
the penguins, for when the more timid of the nest-builders retreated on the approach
of the explorers, the bolder birds immediately carried off the momentarily deserted
materials for their own use. A few only of the nests contained eggs, mostly one apiece,
though some had two ; and the conclusion was arrived at that the birds had only just
commenced laying. Many of the nests were on rocks, some of them in the sites of
previous years ; while others were on pinnacles of ice, having been built on snow which
had gradually thawed away all round the nest, but not at its base. The sitting birds
were very confiding, and allowed themselves to be stroked on their nests. On the
following day, November 9th, another nesting colony was found on a small bare islet.
Here many of the nests contained three eggs, and the rock between them was in a
terribly unsavoury condition. The usual number of eggs was two, but three were not
unfrequent. The eggs varied in size from 51 to 67 mm. in length, and were 41 mm.
in breadth.

It was unfortunately impossible to visit these rookeries later in the season ; and
there were none within reach of Scotia Bay, though some birds rested every night on
the rocky islets in the bay in December, where no signs of their nesting were
apparent.

ON THE BIRDS OF THE SOUTH ORKNEY ISLANDS. 247

There are a number of skins in the collection in both adult and immature plumage,
and a considerable number of eggs.

Adult males shot in September have the crest well developed, the feathers being
from 1'5 to 1'75 inches long. A male shot in December has a much shorter crest, and
others of the same sex obtained in February are devoid of these ornamental plumes.
The September specimens are much more brilliant in plumage than the rest of the
adults, and also have the nasal caruncles more developed. The white dorsal patch
varies much in size, even in adult males obtained at the same season ; and in one
captured in December it is represented by a narrow band of white blotched with black
across the middle of the back. The culmen of adult males varies from 2 '2 to 2 '5
inches, and the wing from 11 '8 to 12'1 inches. Weight 6'5 Ibs.

The bird in first plumage has not, I think, been described. One obtained in
December has the upper surface hair-brown, tinged with green on the back, where the
feathers have narrow margins of lighter brown ; the head and hind-neck show a few
darker feathers ; outer scapulars and tail whitish, the shafts of the latter dull light
green ; primaries and secondaries dusky with a faint greenish tinge ; wing-coverts
edged with dull white, with a narrow buff-white alar band below the marginals ; under
wing-coverts brown ; under surface white ; thighs brown. These young birds show no
signs of a white dorsal patch. Slightly older birds obtained in February have their
upper plumage a mixture of brown and metallic feathers, and one specimen shows slight
indications of a white dorsal patch ; the central tail-feathers are blackish with white
shafts, and the rest of the plumage is as in the younger bird.

Section VI. — The Life and Habits of Penguins.

By R. N. RUDMOSE BROWN. D.Sc., University of Sheffield; Naturalist,
Scottish National Antarctic Expedition.

EARLY in the spring the penguins begin to return to their rookeries from the open sea
or the edge of the pack to the north, where they have spent the winter. They are sea-
birds, although they spend a considerable time at the rookeries every spring. By the
second week in October the flood tide of returning birds had set in at the South
Orkneys, but previous to this there had been many stragglers. From year to year the
time of return has proved to be much the same, whether the season is open or not.
Even if great stretches of land ice have to be crossed the penguins are not deterred.
They hurry onwards through water and over ice, perhaps even crossing necks of land,
to reach their favourite rookeries. On some days a steady stream of them passed the
Scotia from the north over the Beach across the fast ice in Scotia Bay to the large
rookeries on its western side, especially round about Cape Martin. Generally they
marched erect, but some in their hurry adopted the prone attitude and scurried ahead
of the others. Seldom did a penguin deviate from the line of march ; everyone knew
its goal and made straight for it. There can be little doubt that the same birds return
year after year to the same rookeries. Possibly the one-year birds return to their birth-
places, or may be they simply follow older birds. The erect mode of progression is slow
and ungainly, but the prone attitude is the reverse ; propelling themselves forward on
their bellies by aid of feet and flippers, they can move over the floe considerably faster
than a man on ski can travel, certainly over five, probably more than six miles an hour.
By the third week of October the rookeries around Scotia Bay were filled with a
noisy, excited crowd of birds, preparing for mating and nesting. As a rule, the rookeries
prove to be places that are clear of snow relatively early in the spring. Sometimes
they are at sea-level, but more generally several feet above, even as much as 100 feet
in places. At some rookeries we found birds as high as 300 feet. There are few suit-
able sites that are not occupied during the breeding season, and often enough the late-
comers or less fortunate ones are crowded out from the bare ground on to the snow,
where they are forced to build their nests. Sometimes it happens that, on the arrival
of the birds, parts or all of the rookery are still covered with snow. That makes no
difference to the activities of the penguins, unless, perhaps, they voice their dissatis-
faction and so add to the noisy clatter that always reigns on the rookeries. However,
by the time the eggs are laid the nest is clear of snow, through the sun's action, unless a
particularly heavy snowstorm leaves the rookery several inches deep for a day or more.

249

250 ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

The Adelia or Black-throated Penguins (P. adelise) arrived first in Scotia Bay.
About a fortnight after them arrived the slower, less energetic gentoos (P. papua), and
a few Ringed Penguins (P. antarctica) straggled southward to their haunts around
Saddle Island about the same time (see "Zoological Log of the Scotia," Rep. Sci.
Res. Voyage S. Y. Scotia, IV., and " Birds of the South Orkneys," ibid., Section V.).

As soon as the birds arrive at the rookeries the mates are chosen, and this involves
much display and showing off on the part of the male. He stands erect, drawing him-
self up to his full height, with head thrown back and neck craned forward, and slowly
and impressively waves his flippers several times, emitting a long, loud cackle. This over,
he resumes his normal, somewhat squat appearance, and looks round to see what
impression his charms and powers have had on the onlooking females. This process
repeated a few times generally results in a couple being paired off, and nest-building
then begins.

One often sees what is clearly a demonstration of affection on the part of a couple.
The two stand facing one another, and stretching forward cross their beaks, and proceed
to sway from side to side in unison, uttering the while a shrill, harsh cry. At other
times the two sit facing one another with a look of the smuggest complacency on
their faces.

Within a week of the arrival of the penguins nest-building is in full swing in the
rookeries. The first-comers get the best positions, already clear of snow, while the
later arrivals have to be content with less favourable sites. Nest material is not plenti-
ful, and the Black -throated Penguin almost entirely confines itself to the use of stones ;
the gentoo generally adds to his stones any stray feathers and bones which he can
find. Male and female take an equal share in the nest-building. The bird walks to
and fro collecting pebbles in its beak, and carries them to the chosen spot one by one,
there depositing them in a little heap. He is quite energetic over the work and,
though a penguin can walk but slowly on his feet, he often travels ten yards or more
for a stone. It is most amusing to watch their cunning in acquiring stones, for a
penguin never walks far for a stone if he can steal one near by. Since the nests are
seldom more than a couple of feet apart, theft is a relatively simple matter. A penguin
sets out to fetch a stone and notices a nest near by unwatched for the moment. He
approaches with the obvious intention of stealing a stone, when at that moment the
rightful owner, himself probably coming from a raid, returns. The intending culprit,
walking on with a most innocent air and with a look as if the thought of theft had
never entered his head, passes on to pick up an unclaimed pebble farther away. But
given a better opportunity he will not fail next time.

Often, of course, the culprit is caught in the act, and then an angry fight with beaks
ensues, and blood flows and feathers fly before peace is restored. In fact, the penguin
is a most pugnacious bird, and on every rookery there are always several fights in
progress, while neighbours continually snarl threateningly at one another. Not
infrequently one sees an unpaired male — very probably a one-year bird whose powers

THE LIFE AND HABITS OF PENGUINS. 251

had been unavailing to encharm a mate — running about the rookery in a forlorn and
battered state, pecked at by every bird it passes, and only too glad to escape to safety
beyond the rookery. Penguins fight almost solely with their beaks, dealing an
occasional blow with their flippers. But such blows in the case of an Emperor Penguin
are most powerful. A harsh cry accompanies every peck the penguin gives his enemy,
so that it is easy to imagine that with courtship and fighting a rookery is anything but
quiet. Wherever penguins nest, except in the very bad weather, there is an incessant
cackle that requires only a few hundred birds to become deafening at times. The nest
does not take long to complete, for it is merely a small heap of stones a few inches
high, scooped out in the middle into a slight hollow.

Towards the end of October, at the South Orkneys, the eggs of the Adelia Penguins
were laid. As a rule there are two in each nest, but occasionally three. The pug-
naciousness of the birds then increases, and they show much courage in defending
their eggs, and later their chicks, from the attacks of enemies. The skua (Megalestris
antarctica) is the bird for which the penguins must always be on their guard. As
soon as the eggs are laid skuas hover continually over the rookeries, and if one spies
an unguarded egg it swoops down immediately, seizes the egg, and carries it off to a
distance in order to suck the contents. On a skua flying low over the rookery all the
penguins in the vicinity combine in a shrill uproar in the hope of scaring it away, but
seldom with much success, and scores of eggs are thus destroyed during incubation. So
great, in fact, is the harvest of the skuas, that for a few weeks these birds congregate
near the rookeries and live on little else than eggs of penguins.

The presence of the Expedition introduced the penguins to a new and more per-
sistent enemy. Thousands of eggs were collected for food. This presented no
difficulty : the birds were simply pushed off the nests and the eggs picked up. But
the Adelia Penguin did not submit to this robbery without a protest. It would
rush at the intruder and bite him savagely, sometimes even jumping up and trying to
bite his arm. The bite of an angry penguin is not soon forgotten, so it was well to be
protected with long leather sea-boots. Occasionally a robbed bird would follow a man
several yards, attacking him courageously all the time, but generally they stayed near
the plundered nest and bewailed their lot. In most cases another couple of eggs
were laid very soon.

Two weeks after the Adelias had begun to lay, the first Gentoo Penguin's egg was
obtained. The Gentoos, who had been a fortnight later in arriving, inhabited the same
rookeries as the Adelias, but owing to their coming later had to be content with the
more outlying or otherwise less favourable sites. They had a small rookery of their
own on the west side of Scotia Bay, less than a mile from the ship. This afforded
excellent opportunities for the collection of dated eggs for embryological study. The
Gentoos are much more timid than the Adelias, and seldom made a stand when we
approached their nests, but bolted in a body, leaving their eggs at our mercy. This

certainly facilitated our work, if it lessened the interest and excitement. Scarcely ever
VOL. IV. 7

252 ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

did a molested Gentoo stand firm and show fight. Later on it was noticeable, however,
that an occasional Gentoo would defend its chick, fighting with flippers rather than beak.
In fact, their timidity and peacefulness seem to result in their being unable to hold
their own in competition with the Adelias for the rookeries. They build fairly large
nests, and use, in addition to pebbles, bones and tail-feathers. Some of the latest
arrivals, however, laid their eggs in mere holes in the snow. Their eggs frequently fall
a prey to the rapaciousness of the skuas. In all their movements the Geritoos show less
activity and alertness than the Adelias, and are altogether less interesting birds. The
cry of the Gentoo is strangely like the bray of an ass — so much so, in fact, that the
closely allied Falkland Island species (Spheniscus magellanicus) gets the name of the
Jackass Penguin.

The Bridled or Ringed Penguin (P. antarctica) occurs in large numbers at Saddle
Island, Nigg Rock, and Eillium Island, but around Scotia Bay and the shores of Laurie
Island in general no rookeries were discovered. This is more pugnacious than either
of the two common species, and a Ringed Penguin has been known more than once
deliberately to attack a man on the floe and only to retreat after receiving several hard
blows. The quagh of the Ringed Penguins is harsher than that of the other species.

For the first few weeks after their arrival the penguins live on their thick coats of
blubber and abstain altogether from fishing. After the eggs are laid, however, the male and
female take turns at making excursions to the nearest open water in search of food. The
one that remains behind looks after the eggs. In fine weather the penguin sits upright
against the eggs, partially enveloping them in a ventral groove in its coat of feathers.
In windy weather or a blizzard the bird lies prone and completely covers the eggs.
Sometimes in a blizzard the birds are almost snowed up, but the eggs rarely suffer.

A large rookery of penguins, such as the one at Graptolite Island, may contain several
million birds. At Route Point Dr Pirie estimated there were at least 200,000 Ringed
Penguins in one rookery. As rookeries fringe all Antarctica and its bounding islands,
wherever a site offers, it can be realised how enormous is the wealth of penguin life.
The nests occur at intervals of a few feet, and this gives a strangely regular appearance
to a rookery. If viewed from the right point of view, the birds seem to be arranged in
long streets. Some of the photographs reproduced in this volume illustrate that
symmetry.

About the middle of December the Adelia and Gentoo chicks were hatched, those of
the Ringed Penguin some weeks later. For a study of the rookeries and the habits of the
penguins at this season I am largely indebted to Dr Harvey Pirie, who was one of the
party left at the South Orkneys while the Scotia went to Buenos Aires to refit. The
young Adelias are covered with a dark, almost black down, the Gentoos and Ringed
Penguins of a light grey, the former becoming darker before the down is cast, the
latter always darker on the dorsal surface. The chick is fed by the adult bird.
Dr Pirie, describing the mother feeding her youngster, says : " She bends her head down
till her bill is inclined at about 45°, with upper mandible lowermost. Then the chick

THE LIFE AND HABITS OF PENGUINS. 253

sucks up the serai-digested food, brought up from her stomach, out of the hollow
between the rami of the upper mandible." The young have voracious appetites, and
grow rapidly. It was a funny sight to see a poor emaciated parent being chased by two
fat chicks as big as herself, demanding loudly to be fed, with cries of Mad, Mad, like a
young lamb. The young birds have several enemies, and the death-rate must be high.
While skuas do not neglect the chicks, most of their depredations are on the eggs.
Giant Petrels (Ossifraga gigantea), on the other hand, are greedy for young penguins.
On the outskirts of a rookery there are always a number of these birds lying asleep
after a gorge, or waddling to and fro in a state of repletion, unable to fly.

When about a month old the chicks begin to lose their down, and at this time
present a very bedraggled appearance, for the rookeries by then are like quagmires.
The birds are caked all over with mud and dirt, and the stench is abominable. When
the wind was in the right quarter a rookery at that season could be smelt more than a
mile away.

The casting of the down commences on the breast and under parts of the body,
neck, and flippers, and then on the dorsal aspects, the last parts to cast being generally
above the root of the tail and at the base of the flippers, and finally the crown of the
head. When the moulting is complete the young birds rapidly take to the water, and
then the parents leave them to shift for themselves. They are about six weeks old
when this happens, and the parents show no reluctance to abandon their young ; in fact,
they occasionally do so rather prematurely. Clumps of ten or twenty very forlorn-
looking youngsters may often be seen on the rookeries, left by the adults, but a little
timid of trusting to their own resources. In their defenceless state the young birds pay
a heavy toll to the giant petrels, skuas, and gulls.

The Sheathbill probably never attacks the chicks nor does it seize eggs, but haunts
the rookeries on the look out for offal.

Once their parental duties are over the old birds quickly fatten. Early in February
they had altogether lost their emaciated look, and were stout and prosperous again,
scaling some sixteen or seventeen pounds. By the middle of that month they had
deserted the rookeries and put to sea — their normal habitat. In the water the penguin
has only one enemy, the sea-leopard, which has been seen to come up alongside a floe,
seize a penguin in its huge jaws, and sweep down with its prey. At other times the
Sea-leopard chases the penguin in the water as the bird darts torpedo-like through the
sea before its enemy.

Further notes on the habits of penguins, more especially with regard to nesting,
will be found in the previous paper by Mr Eagle Clarke (pp. 223-233). Mr Clarke's
notes were compiled from observations supplied by the leader and other naturalists
of the Scotia.

Section VII. — On the Birds of the Weddell and Adjacent Seas, Antarctic Ocean.*

By WM. EAGLE CLARKE, F.R.S.E., F.L.S., Keeper of the Department of
Natural History of the Royal Scottish Museum.

THIS instalment of the ornithological results of the voyage of the Scotia deals with
the bird-life observed in the Antarctic Ocean southwards of the 60th parallel of south
latitude, and between the meridians of 12C and 45° of west longitude ; in other words,
the Weddell Sea and the waters fringing it on the north. Previous to the voyages
of the Scotia, the Weddell Sea had only been visited by Captain Weddell, during
his marvellously successful cruise in 1823; by Morrell later in the same year; by
Sir J. C. Ross, who traversed its waters in 1843 ; and lastly, by Dr Otto Nordenskjold,
who penetrated to its western portion in 1902.

Dr Bruce made two voyages into the Weddell Sea ; the first in the late southern
summer of 1903, and a second and more extended cruise in the same season of 1904.
On both these occasions a high southern latitude was attained, and on the latter the
Antarctic Continent within this area was discovered and named " Coats Land," after
Mr James Coats and Major Andrew Coats, in recognition of their public-spirited
liberality towards the Expedition. During these voyages birds received much attention,
and specimens were collected whenever opportunity afforded, chiefly while the Scotia
was engaged in taking soundings, or while trawling operations were in progress for the
capture of deep-sea organisms. As no landing was effected on terra firma, the birds
were observed or obtained either amid the ice-fields or on the open sea. The collection
thus formed comprises seventy-four specimens, and is one of great value and interest,
for it is not only the first ever made in the region, but adds greatly to our knowledge
of the geographical distribution of bird-life in the Antarctic Seas.

The first of the above-mentioned voyages commenced on the 4th of February 1903,
on which date the Scotia sailed from Saddle Island, one of the South Orkneys. It was
the intention of the explorers to follow a south-easterly course, as Weddell had done
in 1823, but the pack-ice, in places from fifteen to twenty feet thick, compelled them
to proceed in an easterly direction, hugging, as it were, the 60th degree of south latitude.
On February 14th, when some distance off Southern Tbule Island, the southernmost of
the South Sandwich group, the edge of the pack was fortunately found to trend
towards the much-desired south.

During the run from the South Orkneys towards the South Sandwich Islands the
following birds were either obtained or came under notice : — Banks's Whale-bird

* Reprinted, with slight verbal alterations, from The Ibis, Series IX., i., 1907, pp. 325-349.

255

256 ORNITHOLOGY OP THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

(Prion banksi), Wilson's Petrel (Oceanites oceanicus), Cape Petrel (Daption capensis),
Giant Petrel (Ossifraga gigantea), Silver Petrel (Priocella glacialoides), Snowy Petrel
(Pagodroma nivea), Hutton's Sooty Albatros (Phcebetria cornicoidcs), Ringed Penguin
(Pygoscelis antarctica), Sheathbill (Chionis alba), and several " Molliemauks."

From noon on the 15th of February until the 18th, when the Antarctic Circle was
crossed, the Scotia had a fine run southwards in a sea clear of pack-ice. On the 19th,
however, the edge of the pack was again encountered, and an E.S.E. course was
steered until the 22nd, when, in 70° 25' S. lat. and 17° 12' W. long., or a little short
of Ross's furthest south in this region, the ship became firmly beset. Later in the day,
however, the Scotia managed to free herself, but, the temperature having fallen to 13° F.,
it was resolved to retreat to the South Orkneys in search of winter-quarters.

The birds observed between the Sandwich group and the Antarctic Circle were
Wilson's Petrels, " Blue Petrels " (? Prion or Halob&na *), Giant Petrels, Hutton's
Sooty Albatroses, Cape Petrels, Snowy Petrels, Silver Petrels, Antarctic Skuas (Mega-
lestris antarctica), Antarctic Petrels (Thalassceca antarctica), and Terns (Sterna sp.).

Between the Antarctic Circle and the furthest south (lat. 70° 25' S., long. 17° 12' W.),
the same species came under notice, with the addition of Emperor and Adelie Penguins
(Aptenodytes forsteri and Pygoscelis adelias).

On the return voyage a north-easterly course was steered, the Antarctic Circle was
recrossed on March the llth, and the first southern voyage of the Scotia was completed
on the 21st, on which day she reached the South Orkneys.

The only bird worthy of note obtained during the final stages of this cruise was a
McCormick's Skua (Megalestris maccormicki).

Having spent the winter of 1903 at Laurie Island, the Scotia proceeded to the
Falklands and Buenos Aires to refit, and returned to the South Orkneys on February
14th, 1904. On the 22nd, the Expedition bade adieu to the Orkneys, and the Scotia
commenced her second and most important voyage to the Weddell Sea. In contrast
to the preceding season, the sea was almost free from pack-ice and a south-easterly
course was possible. At the beginning of March the old track of the previous year was
crossed, and a few days afterwards the Scotia broke her southern record of 1903, and
that of Ross in 1843. Skirting, more or less, the open pack for 300 miles, the ship
met with no obstacle to her progress southward, whereas in 1903 all had been im-
penetrable ice. On reaching 72° 18' S., 17° 59' W., however, a lofty ice-barrier
effectually barred further progress towards the south, and here Coats Land was
discovered. This barrier was traced for a distance of 150 miles to the south-west,
when, on the 7th of March, the ship was caught in a north-easterly blizzard and became
locked in heavy pack-ice. On the gale abating two days later, it was found that the
Scotia had been driven into a bight in the ice-barrier off Coats Land in 74° 1' S.,
22° 0' W. No open water was in sight, and with the temperature down to zero the pack

* Both Prion banksi and Halobcena ccerulea appear under the name of " Blue Petrels " in the Zoological Log,
and when specimens were not obtained it is impossible to say which species is intended.

ON THE BIRDS OF TJIE WEDDELL AND ADJACENT SEAS. 257

soon froze up. For several days there was no change, and the lateness of the season,
and the possibility of not being liberated, made it imperative that preparations should
be made for wintering. On the 13th, however, under the influence of a south-west
wind, the pack began to break up and the ship was once more afloat. Off this newly
discovered land Emperor Penguins were abundant, and twenty were captured ; while
Adelie Penguins, Giant Petrels, McCormick's Skuas, Arctic Terns (Sterna macnira),
and Snowy and Silver Petrels were swarming all around. The birds previously observed
during the voyage, in addition to the species named, were Cape Petrels, Wilson's
Petrels, Blue Petrels (Haloljsena cserulea), Mutton's Sooty Albatroses, and Bauks's
Whale-birds, the last-mentioned being only observed north of the Antarctic Circle.

The Antarctic summer was now fast drawing to a close, and the Scotia having
narrowly escaped the miseries of wintering in the pack, turned her prow towards the
north on the 17th of March, her immediate goal being the remote Gough Island in the
middle waters of the South Atlantic Ocean. In the run northwards to the Antarctic
Circle, the birds logged were Emperor and Ade"lie Penguins, Antarctic, Silver, Snowy,
Cape, Blue, and Wilson's Petrels, Button's Sooty Albatroses, Arctic Terns, and a new
petrel to the fauna of the South Polar Ocean, namely (Estrelata brevirostris. The
species noted between the Circle and 60° S. lat., which was crossed in about 12° W.
long., and between March 27th and April 2nd, included most of the species named,
the absentees being the two penguins, the Snowy and Antarctic Petrels, and the
Arctic Tern. On the other hand, an additional petrel, from the description probably
M'ljaqueus asquinoctialis, was encountered.

It may be interesting here to notice that the Expedition added no less than four
birds — namely, Sterna macrura, Phabetria comicoides, Halobiena cxrulea, and
(Estrelata brevirostris — to the short list of nine species previously known, according
to Mr Howard Saunders in the Antarctic Manual, to have occurred south of the
Antarctic Circle.

A specially important ornithological feature of these voyages of the Scotia was the
presence in the Polar Sea of a number of species of petrels far beyond the southern
limits of their breeding-areas. This seems to indicate that at the close of the southern
summer numbers of Hutton's Sooty Albatroses (P. comicoides), Cape Petrels(Z>. capensis),
Giant Petrels (O. gigantea), Antarctic Petrels (T. antarctica), Silver Petrels (P. glacia-
loides), Blue Petrels (//. cserulea), and (Estrelata brevirostris cross the Antarctic
Circle and sojourn among the polar ice ere they retreat northwards to pass the winter
in more genial oceanic resorts. It is possible, however, that some of these visitors to
the far south are non-breeding birds, and, if so, they may have spent the entire summer
there. The Tubinares are, as is well known, great wanderers, but these very remark-
able southern incursions are, perhaps, to be explained by the extraordinary abundance
of food to be found in the waters of the far south in the summer and autumn, which
allures some of the birds further and further towards the Pole, until the great ice-
barrier, which almost girdles the Antarctic Continent, arrests their further progress,

258 ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

since at its base the food-supply entirely ceases. This, too, explains why our familiar
Arctic Tern (Sterna macrura) passes the southern summer (our northern winter) amid
these ever-icy seas.

Full particulars of the distribution of all the species collected or observed by the
Expedition south of 60° 0' S. during the voyages that I have thus briefly described will
be found in the systematic portion of this contribution, which is chiefly based upon the
information afforded by the Zoological Log of the " Scotia," kept by Mr Wilton, though I
am also indebted to Dr Bruce for extracts from his private diary relating to this
portion of the expedition under his leadership.

Mr Kinnear has again earned my acknowledgments for the aid which he has
rendered me in classifying the numerous records.

I shall have occasion to refer to the following works :—

WEDDELL. — A Voyage towards the South Pole, performed in the Years 1822-1824. By JAMES WBDDELL,
Master hi the Royal Navy. (1825.)

Ross. — A Voyage of Discovery an/I Research in the Southern and Antarctic Regions during the Years
1839-43. By Capt. Sir JAMES CLARK Ross, R.N. Vol. ii. (1874.)

SAUNDERS (1). — Catalogue of Birds in the British Museum. Vol. xxv. Gaviae. By HOWARD SAUNDERS.
(1896.)

SALVIN.— Catalogue of Birds in the British Museum. Vol. xxv. Tubinares. By OSBERT SALVIN. (1896.)

CHUN. — Aus den Tiefen des Weltmeers. Von CARL CHUN. (1900.)

RAOOVITZA. — La vie des Animaux et des Plantes dans V Antur clique. Par EMILE G. RACOVITZA. (1900.)

SAUNDERS (2).— Antarctic Manual for the use of the Expedition of 1901. By HOWARD SAUNDERS. (1901.)

SHARPE.— Report on the Collections Of Natural History made in the Antarctic Regions during the Voyage
of the "Southern Cross" : Aves. By R. BOWDLER SHARPE, LL.I). (1902.)

ANDERSON. — " Das hbhere Tierleben im antarktischen Gebiete." Von K. A. ANDERSON. Wiss.
Ergebn. d. schwedischen Sudpolar-Exp. 1901-1903. Bd. v. Lfg. 2. (1905.)

NORDENSKJOLD. — Antarctica, or Two Years amongst the Ice of the South Pole. By Dr N. OTTO
NORDENSKJOLD and Dr JOB. GUNNAR ANDERSON. (1905.)

LONNBERQ. — "Die Vb'gel der schwedischen Siidpolar-Expedition." Von EINER LONNBERG. Wiiis. Ergelm.
d. schieed. Sudpolar-Exp. Bd. v. Lfg. 5. (1905.)

VANHOFFEN. — "Bericht iiber die bei der deutschen Sudpolarexpedition beobauhteten Vb'gel." Von Prof.
Dr E. VANHOFFKN. — J own. fur Urn., 1905, pp. 500-515.

BROWN, MOSSMAN, and PIBIE. — The Voyage of the " Scotia." By Three of the Staff (1906). An excellent
and graphic account of the work of the Scottish National Antarctic Expedition.

Aptenodytes forsteri G. R. Gray.
Aptenodytes forsteri Cat. B., xxvi. p. 626.

The occurrence of this bird as a visitor to the South Orkneys was considered
probable in 1903 (Ibis, 1906, p. 166), and the fact has since been fully confirmed by
the Argentine naturalists, who, Mr Mossman informs me, captured two specimens on
Laurie Island in March 1905. This establishes a record for the northern range, namely
60° 44' S., of this species.

On the first Antarctic voyage, in the early southern autumn of 1903, this species
was not observed until latitude 69° 46' S. (20° 58' W.) was reached. Here a male,
weighing 64 Ibs., was shot on February 21st, whose stomach contained cuttle-beaks,

ON THE BIRDS OF THE WEDDELL AND ADJACENT SEAS. 259

fishes, and three small gneiss pebbles. On the following day an individual was captured
on the ice in 70° 25' S., and was brought on board alive. This bird soon became
reconciled to its strange surroundings, and with remarkable equanimity paced up and
down the deck of the Scotia, every now and then giving utterance to a musical cry.
On the return passage to the South Orkneys for winter-quarters, another male was
obtained in 67° 10' S. (39° W.), but none were seen further north.

On the second voyage, in 1904, this species was not encountered till 72° 18' S.
(17° 59' W.) was attained on March 3rd, on which day three examples were seen and
secured. In 73° 30' S. (21° 28' W.) Emperor Penguins were in abundance on the water,
but only one was observed on the ice, which was captured. In 74° l' S. (22° W.),
while the Scotia was for several days fast in the pack off Coats Land, until March 14th,
an extraordinary number of these penguins were seen on the ice around the ship, and
many were captured with a view to providing food for the Expedition should it be com-
pelled to winter in the pack. While the ship was thus incarcerated, Piper Kerr played
on his pipes to one of the captive penguins which was tethered on the ice, but it is said
that neither rousing marches, lively reels, nor melancholy laments seemed to have any
effect on this lethargic and phlegmatic bird : it was absolutely indifferent to them all !

Some of the individuals captured weighed close on 80 Ibs., and it was all that a man
could do to lead one of them up to the ship. With their beaks they bit fairly hard,
while with their long flipper-like wings they dealt severe blows.

None were seen after the Scotia escaped from the ice and proceeded northwards.

Pygoscelis antarctica (Forster).

Pygosedis antarctica Cat. B., xxvi. p. 634.

Ringed Penguins were abundant in the South Orcadian waters, and were seen at sea
during the easterly run towards the South Sandwich group in February 1 903. After-
wards, when the Scotia was proceeding southwards, these birds were met with at
intervals until latitude 69° 39' S. (22° 58' W.) was reached on February 20th. On the
return passage to the Orkneys they were met with in 67° 39' S.

For the second voyage there were but few records, but it is said that several
individuals were seen by one of the sailors on March 18th (1904), when the Scotia was
in latitude 71° 22' S. (16' 34' W.). The only other observation is for April 1st, on
which date some were noticed on the water in 60° 33' S. (12° W.)— the last the Scotia
naturalists were to see of a bird to our knowledge of which they have added so much.

Pygoscelis adelise (Hombr. et Jacq.).

Pygosedis adelix Cat. B., xxvi. p. 632.

The Adelie Penguin — another native of the Antarctic Continent — was not observed
during the first voyage into the Weddell Sea until the latitude of 69° 46' S. (20° 58' W.)

was attained on February 21st, 1903, the day on which the Expedition made the acquaint-
VOL. iv. 8

260 ORNITHOLOGY OF THK SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

ance of the Emperor Penguin, which shares with this species the distinction of being
the most southerly representative of its order. On the following day, when the Scotia
reached her furthest south (71° 25') for the season, the birds were very abundant; and
on the return passage to the Orkneys were seen almost daily in considerable numbers.
On one occasion some twenty to thirty were observed seated on an iceberg from forty
to fifty feet high, to obtain a footing on which others were seen jumping out of the
water on to the berg's precipitous, slippery sides, and holding on where Dr Bruce
believes no other bird or mammal could. They took advantage of the wash of the sea,
but often had to try again and again ere they succeeded in landing on the lowest
ledges of the berg.

During the 1904 voyage this bird was not logged until March 6th, when a few
examples were noted in 73° 30' S. (21° 38' W.). A few more were observed on the
northward passage, but not beyond 61° 25' S. (12° 47' W.).

Three adults captured on February 23rd, 1903, were still in moult, having only
partially assumed their new coats of blue-black tipped with steely blue. The tempera-
ture of these specimens was found to be 102°-103C F.

Oceanites oceanicus (Kuhl).

Oceanites oceanicus Cat. B., xxv. p. 353.

After the nesting-season this little petrel becomes a great wanderer on the face of
the ocean. As such it was almost daily to be seen in numbers, both at sea and among
the ice, throughout the voyages of the Scotia — being observed as far south as 72° 22' S.,
while northwards it was present in abundance off Gough Island (40° 19' S.).

It was one of those birds which followed in the wake of the ship, probably on the
look out for scraps cast overboard from the galley. On February 7th, 1903, in lat. 60°
35' S. and long. 39° 44' W., a great number were observed around a dead whale, picking
up morsels of fat that fell from the bills of a host of Giant and Cape Petrels, which
were regaling themselves to repletion on the blubber of the defunct leviathan. On one
occasion "a flock" was observed resting on the water in 61° 22' (42° W.).

The Belgica obtained it in the pack in 70° S. (87° W.) ; the Southern Cross found
it breeding on Victoria Land ; and Dr Wilson saw it off the Great Ice Barrier in 78° S.,
some seventy miles from the nearest open water ( Voy. of the " Discovery" ii. pp. 482-3).

Thalassceca antarctica (Gm.).

Thalassoeca antarctica Cat. B., xxv. p. 392.

The Antarctic Petrel was first encountered by the Expedition on February 17th, 1903,
in latitude 64° 18' S. (23° 09' W.). Afterwards it was much in evidence in the vicinity
of and amidst the great polar ice-fields. During the first voyage this bird was seen
almost daily, sometime in flocks, its latitudinal range varying from 62° to 70° S., and
its longitudinal from 16° to 44° W.

ON THE BIRDS OF THE WEDDELL AND ADJACENT SEAS. 261

It was equally frequent during the second voyage, in 1904, and often followed the
ship all day. When off Coats Land in 74° 1' S. no less than twenty were shot on
March 14th, while the Scotia was locked in the pack. Just previously to this, namely
on March 5th, when in 72° 31' S., thousands of this species were seen in company with
McCormick's Skuas, Giant and Snowy Petrels, and Arctic Terns. On the northward
voyage it was not observed beyond 68° 26' S. (10° 11' W.).

As already recorded (The Ibis, 1 906, p. 1 69), a few came under notice at Laurie Island
in the summer of 1903-4, and it was thought not unlikely that the bird was nesting
there. A number were also seen near Saddle Island, another of the South Orkney
group, on March 22nd, 1903, in about 60° S., the most northerly point at which this
species was observed during the voyages of the Expedition. A number of specimens
were obtained in the month of March during the Antarctic voyages of 1903 and 1904,
and these vary in their plumage. Some are more or less faded (drab) iu colour and
abraded in feather, and are, no doubt, unmoulted birds ; others are evidently freshly
moulted adults or young of the year ; in these the head, back, scapulars, lesser wing-
coverts, quills and tip of the tail are slaty black, and the under parts purer white.
Specimens in moult are intermediate between these two forms, and make clear the
relationships of these phases in plumage as regards adult birds. Some of these moult-
ing birds lack rectrices, their tails being represented by the long under tail-coverts only.
The feet in life have the tarsus and outer toe greyish, the other toes and the webs paler
and washed with yellow.

Priocella glacialoides (Smith).
Priocella glacialoides Cat. B., xxv. p. 393.

The Silver or Silver-grey Petrel was observed during the voyages to and from the
Weddell Sea, between the latitudes 44° 30' and 71° 22' S., and the meridians of 9° 43'
and 42° 30' W. The extreme southerly range here indicated has only been exceeded,
I believe, in the case of the observations made by Dr Wilson during the National
Expedition under Captain Scott, when the bird was seen in the Ross Sea ( Voy. of the
"Discovery" ii. p. 481).

This petrel was first met with on the 9th of February, when the Scotia was skirting
the pack-ice in 58° 57' S. and 33° 34' W., or about midway between the South Orkney
and South Sandwich groups; but it was not observed beyond 63° 54' S. on the 1903
voyage. On the second voyage it was noted as high as 71° 22' S., on March 18th, 1904,
when one was seen in company with several other species of petrel and Arctic Terns.

It does not appear to be an abundant species in the Weddell Sea ; but it was
observed in numbers during the northward voyage, when latitude 60° S. was reached,
and was last seen when the Scotia was nearin^ Gough Island.

O O

As stated on page 236 of this volume, it was occasionally seen in the summer of
1903 about the cliffs at Laurie Island, where it was considered probable that a few
were nesting. I mentioned (I.e.) that its breeding-haunts remained unknown. Since

262 ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

then, however, I have learned that the Swedish Expedition under Nordenskjold found
it nesting at Cape Roquemaurel, on the west coast of Louis Philippe Land (Anderson,
t.c., p. 43).

A coloured drawing made by the artist to the Expedition shows the maxilla and
mandible tipped with black, the middle portion of the bill pale flesh-coloured, and the
base and nares pale cobalt-blue. The feet are pale flesh-coloured, the webs washed with
yellow, and the claws black. The iris is dark brown, and the pupil blue-black.

Pagodroma nivea (Gm.).
Pagodroma nivea Cat. B., xxv. p. 419.

The Snowy Petrel being a native of the Antarctic Continent, it is not surprising to
find that it was one of the most abundant and most frequently observed species which
came under the notice of the Expedition in the Weddell Sea. During both the southern
voyages of the Scotia, there was hardly a day on which this beautiful bird was not met
with — often in great numbers. It was most abundant in the neighbourhood of the
pack, and became less numerous as soon as open water clear of ice was entered.

The extremes of latitude between which this bird was met with during the voyages
of the Scotia were from 59° 44' to 74° l' S. (off Coats Land, the Ultima Thule of the
Expedition). Longitudinally it occurred from 12° 49' to 43° 40' W. On the voyage
from Coats Land northwards to Gough Island it was not observed beyond 68° 32' S.
(12° 49' W.).

The Snowy Petrel was sometimes seen in flocks around the ship, and often followed
it all the day long. It was seen to capture fish at or near the surface of the water.

CEstrelata brevirostris (Lesson).
CEstrelata brevirostris Cat. B., xxv. p. 409.

This species is new to the avifauna of the Antarctic Ocean, where it was discovered
by the Scottish Expedition on March 20th, 1904, in 69° 33' S. and 15° 19' W., during
the northward voyage from Coats Land towards Gough Island. On that day, while
the Scotia was engaged in trawling for the capture of various forms of marine life,
Dr Pirie left the ship in a boat for the purpose of obtaining specimens of birds, a
quantity of which, including terns and various species of petrels, were in the vicinity.
While thus employed he came across numbers of a petrel that he had not seen before,
and succeeded in shooting a male, which on examination I found to belong to this
species. Afterwards this " New Petrel," as it was logged, was observed on five different
days down to March 25, when it was seen in 65° 58' S. and 11° 24' W.*

The habitat of this interesting addition to the fauna of the South Polar seas has
hitherto been supposed to be confined, according to Salvin (op. cit., p. 410), to the
South Atlantic and South Indian Oceans, and he mentions specimens as being in the
British Museum Collection from Kerguelen (where it breeds) and Tristan da Cunha.

* See also infra, Section VIIL, p. 276.

ON THE BIRDS OF THE WEDDELL AND ADJACENT SEAS. 263

Vanhoffen (t.c., pp. 506 and 508) records it as seen so far south as 59° during the
voyage of the Gauss from Kerguelen towards Wilhelm Land ; and on the return
voyage northwards from that part of the Antarctic Continent it was again met with
200 nautical miles south of Heard Island, or in about 57° S.

The feet of the specimen obtained by Dr Pirie are, according to a coloured drawing
made at the time of its capture, purplish grey and the claws black ; while the iris is
dark brown and the pupil blue-black.

Ossifraga gigantea (Gm.).

Oxsifraga gigantea Cat. B., xxv. p. 422.

The Giant Petrel was seen everywhere and almost daily during the Antarctic
voyages of the Scotia, even in the highest latitudes reached during each year's cruise,
namely 70° and 74° S. ; and yet its breeding-grounds are all, so far as we know them,
to be found north of the Antarctic Circle.

Perhaps some of these petrels which frequent the far south in late summer and
autumn may be either non-breeding or immature birds which have spent the summer
there ; while others may proceed south at the close of the nesting-season, as may also
young birds. These are points which at present do not admit of satisfactory solution.

Off Coats Land, in 74° 1' S. and 22° AV., from the 9th to the 13th of March 1904,
when the Scotia was fast in the grip of the pack, a number of these birds were seen, as
were also many of McCormick's Skuas, Antarctic and Snowy Petrels, and Arctic Terns.
White examples are only alluded to in the Zoological Log during these voyages as
being seen on four occasions, all north of 61° S.

As regards the food of this bird as a marine species, a great host of individuals,
including white examples, were feeding on the carcass of a dead whale in 60° 03' S.
(39° 44' W.) on February 7th, 1903, and the stomach of one shot contained crustaceans.

Daption capensis (Linn.).

Daption capensis Cat. B., xsv. p. 428.

The Cape Pigeon was one of the most abundant species observed by the Expedi-
tion during its two Antarctic voyages. It was seen almost everywhere, both at sea
and amid the ice, as far south as 71° 50', though it was only found in small numbers in
the high latitude mentioned.

The presence of this (with other petrels) in great numbers in the Weddell Sea, far
to the south of its breeding-haunts, in the late summer and in autumn, has already
been alluded to (p. 257), and a possible explanation of its remarkable incursions amid the
South Polar ice, ere it moves northward to reach its accustomed oceanic winter-quarters,
has been offered.

All the specimens obtained in the Antarctic Ocean during these voyages in the
autumns of 1903 and 1904 have the dark portions of their plumage of a fresh slate-

264 ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

black tint, instead of the faded brown presented by the examples obtained at the South
Orkneys during the breeding-season. This is, with little doubt, attributable to the
fact that the birds were either adults recently moulted, or young in their first plumage,
or both.

Whenever the Scotia stopped for the purpose of taking soundings, these petrels
settled on the water on the look out for scraps of food, and so tame were they that
specimens were often captured by simply scooping them out of the water by means
of a large landing-net. Birds thus taken were sometimes liberated on the ship's deck,
where they showed their entire inability to escape, the Scotia not being long enough to
afford them a sufficiently extended run to enable them to rise on the wing : it was the
same with the Giant Petrels.

When in pursuit of food at or near the surface, the Cape Petrels were observed to
plunge downwards into the water after the manner of terns.

On February 7th, 1903; in 60° 35' S., 39° 44' W., a vast number, along with Giant
and Wilson's Petrels, were observed feeding on the floating carcass of a dead whale.

Ross (I.e., ii. p. 191) saw this bird off Victoria Land, on January 14th, 1841, in
71° 50' S., or in precisely the same latitude in which Dr Bruce made his southernmost
observation.

During the voyage of the Southern Cross it does not appear to have been noted
beyond 65° 3' S. (Sharpe, t.c., p. 157); but Vanhoffen (t.c., p. 507) observed it right
down to the winter-quarters of the Gauss, namely to the Antarctic Continent (Wilhelm
Land) in 66° 2' S., 89° 38' E. The Swedish Expedition (Anderson, I.e., p. 46) also
observed it near their southern limit, namely in the pack-ice east of Graham Land
in 64° 30' S.

Halobzena cserulea (Gmelin).
Halobxna cxrulea Cat. B., xxv. p. 431.

Under the collective name of " Blue Petrels." both this species and at least one of
the Whale-birds (Prion) were confounded by the Scottish explorers — a pardonable
error also made long years ago during Cook's voyage in the Antarctic Seas, and, no
doubt, often since repeated. Fortunately, however, a number of specimens of these
"Blue Petrels" were secured, and afford authentic information regarding both this
species and Prion banksi in the seas visited by the Expedition.

The data accompanying the skins of H. cserulea, the Blue Petrel proper, enable me
to extend the distribution of this species far to the south of all previous records of a
reliable nature. Salvin (t.c., p. 431), the monographer of the petrels, gives its range as
being between 40° and 60° S., or practically where Cook left it in the latter half of the
eighteenth century ; and it has no place in the bird section of the Antarctic Manual. From
the Scotia, specimens were captured as far south in the Weddell Sea as 69° 33' S., and
others, believed to be of this species, were observed as high as 71° 28' S. It probably
occurs even beyond the limits indicated, for I think there can be little doubt that this

ON THE BIRDS OF THE WEDDELL AND ADJACENT SEAS. 265

was the bird which Weddell met with on February 18th, 1823, a little further to the
west, in 73° S., where he tells us (op. cit., pp. 35-6), " the sea was literally covered with
birds of the Blue Petrel kind." Ross (t.c., p. 359) also mentions a "Blue Petrel" as
seen in the Weddell Sea in 67° 06' S. and 8° 35' W. Banks's Whale-bird (Prion
banksi), the other "Blue Petrel" of the Expedition, was not obtained beyond 66° S.,
and I am not aware of any reliable record of its occurrence within the Antarctic Circle.

"Blue Petrels" appear very frequently in the Log of the Scotia during the two
southern voyages, but H. cxrulea was not obtained north of 64° 29' S., and the ten
specimens in the collection were secured between that latitude and 69° 33' S., and
longitudes 12° 49' W. and 35° 29' W. All but two were obtained south of 68°, and in
the month of March (1903 and 1904). On February 25th, 1904, two specimens of this
species, and one of Priori banksi, were captured in 64° 29' S. and 35° 29' W.

Weddell (op. cit., p. 144) mentions the Blue Petrel as occurring at the South
Shetlands, but later explorers do not mention Halobsena cserulea for that group or for
the Antarctic regions proper. It would seem that this species is local in its far southern
range, and is a specially characteristic bird of the Weddell Sea. It was not seen at the
South Orkneys during the summer, nor was it encountered at sea in the vicinity of
that archipelago.

In some of the specimens in the collection the white feathers of the forehead show
their dark bases, and thus the front presents a mottled appearance. The bill in freshly
killed examples was cobalt-blue, except the nares and culmen, which were black. The
feet were cobalt-blue, the webs pale flesh-coloured, the claws black.

Pi-ion banksi Gould.

Prion banksi Cat. B., xxv. p. 434.

Banks's Whale-bird (and perhaps others of its genus*), as has already been stated,
when treating of Halob&na cserulea, was logged during the Antarctic voyages of the
Scotia as a " Blue Petrel." Here, however, the specimens collected with such praise-
worthy diligence again come to our aid, and enable us not only to distinguish between
the two species on important occasions, but also to extend the southern range of this
bird from 60° S. (fide Salvin, t.c., p. 434. and the Antarctic Manual) to 66° S.t

The first specimens, a male and female, were procured on February 9th, 1903, when
the Scotia was off the edge of the pack-ice in 59° 42' S. and 34° 13' W., or about mid-
way between the South Orkneys and Southern Thule, the most southerly of the Sandwich
Group. These were the only examples obtained during the Antarctic voyage of 1903.

* Prion desolat its appears (Antarctic Manual, p. 231) to reach the edge of the Antarctic Circle, having been ob-
tained by the Challenger at the ice-barrier. It breeds at Kerguelen. This bird was not obtained by the Scottish
Expedition.

f In the Liverpool Museum, however, there is a specimen which is believed to have been obtained by Dr J. Hooker
off Victoria Land in 70° S. This example is recorded, along with ^EgialitisfaUclandica, Nettionflavirostre, and Podicipes
ealipureus, as new to Antarctica in the Bulletin of the Museum (ii. p. 48). None of these species have come under the
notice of later observers within the Antarctic Circle.

266 ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

On the second voyage four (two males and two females) were captured in the Weddell
Sea, or its confines, on February 25th and 26th, 1904, in 64° 29' (35° 29' W.) and
65° 59' S. (33° 06' W.).

This species does not appear to have come under the notice of other recent Antarctic
explorers, but a species of Prion is recorded by Vanhoffen (t.c.) as having occurred off
Wilhelm Land, or just without the Antarctic Circle, on March 18th, 1903, and one was
also noticed between that part of the Antarctic Continent and Kerguelen Island where
P. desolatus is known to breed.

The bill and feet of the specimens secured by the Scottish Expedition are described
as being bluish grey, and the iris as brown.

Phoebetria cowiicoides Huttori.
Phcebetriafuliginosa Cat. B., xxv. p. 453; Chun, t.c., pp. 167, 220; Voy. of "Scotia," pp. 181, 232.

It is a matter for surprise that two such genuinely distinct species as P. cornicoides
and P.fuliginosa should have collectively passed for a considerable number of years
under the name of the Sooty Albatros. The Scotia collection of birds has been the
means of calling attention to the claims of Captain Hutton's so-called variety, described
in 1867, to full specific rank. Now there remains the important but at present almost
impossible task of unravelling the tangled skein involved in defining the geographical
distribution of the two species. Here the Scotia collections again lend a helping hand,
for they enable me to say that all the birds obtained and seen in the far south belonged
to Hutton's species, and that it was only when the waters of the South Atlantic were
approached that Gmelm's fuliginosa appeared upon the scene.

Specimens of both birds were collected, and the species under consideration was the
only one obtained in the Antarctic Ocean, where it was observed as far south as
69° 46' S.

During the first Antarctic voyage, in 1903, this bird was noticed on eight occasions
between February 14th and March 21st, in latitudes varying from 59° 33' to 69° 46' S.,
and in longitudes from 20° 58' to 27° 32' W. ; and on the second voyage, in 1904, it
was logged for south of 60° on ten days between February 23rd and April 1st, in latitudes
ranging from 60° 30' to 67° S., and longitudes between 10° 42' and 41° 55' W. On
February 25th, when in 64° 29' S. and 35° 29' W., six individuals were sailing around
the Scotia at the same time. North of 60° it was encountered as far as Gough Island
(40° 19' S.), as related below (Section IX., p. 289).

Previously to the researches of the Scotia Expedition, this albatros had not been
recorded within the Antarctic Circle, though Ross (t.c., p. 359) observed a "Sooty
Albatros" in the Weddell Sea in 67° 06' S. and 8° 35' W. on March 1st, 1843, which
most probably was of this species.

No specimen of P. fuliginosa was obtained or observed by the Expedition beyond
58° S., though it is certain that this albatros does attain to a higher degree of southern

ON THE BIRDS OF THE WEDDELL AND ADJACENT SEAS. 267

latitude. " Sooty Albatroses" were frequently logged during the voyages of the Scotia
between the Falklands and the South Orkneys, and are believed to have been P. corni-
coides, but no specimens were captured.

Having made some incursions into the literature of Antarctic ornithology in the
preparation of these papers, perhaps it may be well to allude to these researches so far
as they concern P. cornicoideg,

Captain Hutton told me, in 1905, that this species is the common form in New
Zealand, and that it breeds at the Auckland and Antipodes Islands at the end of
October ; and Lonnberg in his Contributions to the Fauna of South Georgia, i. p. 71
(1906), mentions it as breeding on cliffs at that island.

Chun (i.e., p. 167) informs us that during the voyage of the Valdivia, " Diomedea
fullyinosa, die aschgrauen Albatrosse mit schwarzlichen Kopfe," was met with about
midway between the Cape of Good Hope and Bouvet Island on November 20th, 1898,
and (p. 220) that the Smoke-grey (" rauchgraue ") Albatros was seen almost daily from
Bouvet Island onwards. There is no mistake as to the bird meant, for reference is
made to an excellent figure of P. cornicoides.

Bernacchi (To the South Polar Regions, pp. 316, 317) says that during the month
of January 1900 an albatros, which he describes as "generally sooty, paler on the
shoulders and under surface," was occasionally seen in the pack-ice.

Vanhoffen (t.e.) has a number of references to Phcebetria during the voyage of the
Gauss, as seen between Kerguelen and the Antarctic Circle, but he evidently did not
realise or recognise that there were two species of the genus.

Anderson (op. cit.) makes no allusion to Phcebetria under " Sturm vogel " in his
records of the " Hohere Tierleben " of the Swedish Expedition ; but Lonnberg (i.e., p. 6),
in his account of the birds seen during the relief expedition, mentions that several
examples of P. cornicoides were seen between Cape Horn and South Georgia, and that
it was last seen in 61° 42' S. and 57° 35' W.

No Sooty Albatros was obtained by the Southern Cross Expedition, though, as I
have said, Bernacchi saw this bird ; nor does Dr Racovitza (t.c.) mention either species
in his account of the animal life observed during the voyage of the Belgica.

Note. — For further discussion of these species in the light of more recent knowledge,
see Section VIII. of this volume, "'Scotia Rise' Region."-— EDITOR.

Diomedea exulans Linn.

Diomedea exulans Cat. B., xxv. p. 441.

Single examples of this species were seen on four occasions, namely : — March 20th,
1903, when the Scotia was in 61° S. and 43° 20' W. ; February 24th, 1904, in 63° S.
38° W., and on March 27th and 30th, in 67° S. 11° W. and 61° S. 13° W. respectively.

" Molliemauks " were logged on February 5th and 6th, 1903, when the ship was
proceeding eastwards from the South Orkneys and was a good way off the edge of the
pack-ice. This was from 60° to 61° S., and about 43° W. (Vol. IV. Part I. pp. 8 and 9.)

VOL. IV. 9

268 ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

Sterna macrura Naum.

Sterna macrura Cat. B., xxv. p. 62.

Sterna hirundinacea Bruce and Wilton, Scot. Geogr. Mag., 1904, p. 128; Pirie and Brown,
op. cit., 1905, p. 26.

In the Antarctic Manual (p. 223) Mr Saunders tells us that there is ample evidence
that terns are found in large numbers in the South Polar regions, and even within the
Antarctic Circle, and that Webster found terns at the South Shetlands, whence the
Dundee whalers brought back specimens which were referable to the well-known South
American species Sterna hirundinacea. Mr Saunders then expresses the opinion that
it may be reasonably assumed that all the terns found southwards of America are of
this form, and this is quite correct so far as the breeding species are concerned. No
tern is, however, known to breed within the Antarctic Circle, nor have, I believe, any
specimens hitherto been obtained to the south of 66°.

When the Scotia sailed from the South Orkneys she left Sterna hirundinacea
behind her. Other terns were met with, often in considerable numbers, and specimens
were fortunately obtained in widely scattered portions of the Weddell Sea. These,
strange to say, I found to belong to the most northern representative of their genus,
namely, to Sterna macrura, the Arctic Tern ! Thus this familiar bird to British
ornithologists would seem to have the most extensive latitudinal range to be found
among vertebrate animals, since it is now known to occur from 82° N. to 74° 1' S. It
was doubtless the species seen by McCormick in 76° 52' S. in Ross Sea, off Victoria
Land, in the 'forties of the last century, and also the bird noted by him in the ice
between 65° and 66° S. and 158° W.

During the Antarctic voyages of the Scotia terns frequently came under notice, and
specimens of Sterna macrura were obtained between 64° 29' and 72° 18' S. latitude
and from 12° 49' to 35° 29' W. longitude. They were often observed in considerable
numbers, and are logged for March 5th, 1904, as being seen in thousands in 72° 31' S. ;
while from the 9th to the 13th of the same month, when off Coats Land, in 74° ]' S.,
22° 0' W., many were seen along with McCormick's Skuas, Giant, Antarctic, and
Snowy Petrels, when the Scotia was locked in the pack.

It seems very remarkable that the far-off ice-fields of the South Polar Ocean should
be visited during the northern winter season by this boreal species. That it is only
a winter visitor does not admit of doubt, for the bird certainly does not breed there ;
nor is any other tern, so far as we know, a native of the Antarctic Continent. During
the southern summer (the northern winter) there is an extraordinary abundance of
marine life, especially of surface-swimming crustaceans, and so this elegant bird is
tempted to seek retreats which can otherwise only be regarded, even in summer, as
inhospitable in the extreme.

The finding of this tern in the seas off the South Polar Continent must be regarded
as one of the most important ornithological discoveries made by the Expedition, for.

ON THE BIRDS OF THE WEDDELL AND ADJACENT SEAS. 269

as has already been stated, no terns appear to have been previously captured within
seas girdled by the Antarctic Circle.

The Scotia collections include some interesting specimens. An adult female
obtained on March 23rd, 1904, in 68° 32' S. and 12° 49' W., has already assumed full
breeding-plumage, and shows no signs of moulting. Another (a male) is assuming its
summer hood, leaving the head a mixture of black-and-white feathers ; this specimen
still retains the dusky upper wing-coverts of youth. In addition to gaining the black
head for the first time, it exhibits further evidence of moulting, inasmuch as neither
the primaries nor the rectrices are quite fully grown, the first primary being still
shorter than the second by about half an inch. I am inclined to think that we have
here a bird about twenty-one months old.

The series also includes two immature examples in the plumage known as the
S. portlandica stage. These have the forehead and crown nearly white, the rest of
the head blackish, the lesser wing-coverts conspicuously dark, and the bill and feet
black. They are in deep moult so far as their primary -quills and tail-feathers are
concerned, but apparently not otherwise. Some of their primaries are only three
inches long.

The Gauss obtained a tern off Wilhelm Land, in 66° S., 89° 38' E., on February
18th, 1903, which Reichenow (0m. Alonatsber., xii. p. 47) described as a new sub-
species of the Arctic Tern under the name of Sterna macrura antistropha. This is
said to be " very like S. macrura, but with the bill darker, not cinnabar or poppy-
red, but carmine at the base and blackish towards the tip ; lower edge of mandible
longer, 21-22 mm., in macrura only 16-19 mm. Feet not yellowish red to cinnabar-
red, but dusky carmine-red, webs blackish. Tarsus somewhat longer, 15-17 mm." If
described from skins, that fact might account for the subtle differences in colour, or
if such differences really exist they might be due to the season (the winter for
S. macrura). The Scotia specimens do not exhibit the peculiarities attributed to this
subspecific form, except that one of them agrees with it so far as the dimensions of the
mandible and tarsus are concerned.

[Gulls.

No gulls were observed during the Antarctic voyages of the Scotia except Lams
dominicanus in the vicinity of the South Orkneys. In the Antarctic Manual (p. 232)
it is stated that this gull was obtained in 64° 18' S., and that a specimen of Lams
scoresbyi, also in the British Museum, was obtained in the vicinity of the South
Shetlauds in 64° 55' S. Dr Bruce, who was naturalist on the Balsena, from which
vessel the birds in question were obtained, tells me that these examples were un-
doubtedly captured at the Falkland Islands, and that they were skinned by him.]

270 ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

Megalestris maccormicki Saunders.

Megalestris maccormicki Cat. B., xxv. p. 321.

Megalestris antarctica Pirie and Brown, Scot. Geogr. May., 1905, p. 26.

This species was not distinguished from the Antarctic Skua (M. antarctica) during
the active work of the Expedition, and hence it is only possible to discriminate with
certainty between the two species by reference to the specimens obtained. No doubt,
however, all the skuas seen south of the Antarctic Circle belonged to the species named
after Dr McCormick.

There are only two skins of this skua in the collection brought home by the
Scotia. The first of these, an adult, was procured on March 10th, 1903, in 66° 40' S.
and 40° 35' W., or the lowest latitude in which this species has, I believe, been obtained,
except the example alluded to by me above as captured at the South Orkneys by the
Argentine observers in 1904.

The second example, an adult male, was shot alongside the ship on March 9th, 1904,
in 74° S. and 22° W. The Scotia was then fast in the pack and about two miles off
the Ice Barrier at Coats Land. Many of these birds were then present, as were also
numerous Giant and Snowy Petrels and Arctic Terns (Sterna macrura).

The Antarctic Skua (M. antarctica) did not occur with certainty beyond 62° 49' S.
(38° 12' W.).

Chionis alba (Gmelin).

Chionis alba Cat. B., xxiv. p. 710.

This bird, the only one found in the regions treated of that may be regarded as a
terrestrial or, to speak more correctly, a semiterrestrial species, was several times
observed at sea during the first voyage of the Scotia. It does not, however, penetrate
into the Weddell Sea, but was met with when the ship was running eastwards from the
South Orkneys towards the South Sandwich group in the late summer of 1903.

During this voyage the White Sheathbill was observed on three occasions : — First,
on February 6th, in lat. 60° 10' S. (42° 35' W.), when the Scotia was a good way off
the edge of the ice. Here this species, along with Banks's Whale-bird, Cape, Wilson's,
and Snowy Petrels, followed in the wake of the ship during the day. Again, on
February 8th, in 59° 44' S. (36° 40' W.), or about midway between the Orkney and
Sandwich groups and 300 miles from land, Sheathbills were observed along with the
birds already mentioned. Finally, it was noted at sea to the southward of the Orkneys
on March 21st in about 61° S. — its furthest south.

In the Liverpool Museum (Bull. Liverpool Mus., ii. p. 48) there is a specimen of this
bird which is said to have been shot on the Antarctic Continent in 78° S. by Dr Gunn,
who, strange to say, was afterwards proved never to have been there ! (see The Ibis,
1895, p. 165, and Antarctic Manual, p. 234, footnote).

Section VIII. — "Scotia Rise" Region: the Voyage from the Weddell Sea

northward to Gough Island.

By L. N. G. RAMSAY, M.A., B.Sc.

THIS section deals with the observations made by the Scotia naturalists and the
specimens obtained during the period between April 1st, 1904, and the ship's arrival at
Gough Island. An irregular northerly course was followed, chiefly between the 8th
and 1 3th meridians of west longitude.

The voyage lasted twenty days, and some 1200 or more nautical miles of ocean were
traversed. Very rough weather was experienced for most of the way. A cluster of a
dozen bergs seen on April otb was almost the last ice encountered.

Bird-life was rather plentiful throughout the voyage, in the form of numerous
albatroses and petrels — of which eighteen to twenty species were seen — and a few
penguins and terns and (towards Gough Island) skuas.

Hutton's Sooty Albatroses (Phcebetria cornicoides) were about the ship in some
numbers the whole way (the Scotia had first met them, after leaving the Antarctic Seas,
in 67° S. lat.).

P. fuliginosa was not met with till 55° S. was reached. These two species, as
stated by Mr Eagle Clarke in the preceding section, had not been recognised as distinct
previous to the voyage of the Scotia, although the form cornicoides had been described
long before as a variety of P. fuliginosa. The difference between the two was sufti-
ciently striking, even when seen only on the wing, to make Dr Bruce believe that they
must be specifically distinct His diary for April 4th records that another species of
" Sooty," different from the " blue-billed " form (cornicoides), by which they had been
accompanied for a week past, was seen, thus : " A Sooty Albatros with a yellow instead
of a blue stripe on its beak, and the streak is broader, the plumage altogether more
uniform and darker in colour than that of the blue-billed form, whose head alone is
very dark."

After this date, fuliginosa was seen in about equal numbers with cornicoides until
Gough Island was approached, but for 70 miles or so south of the island the "yellow-
billed " species was the more numerous.

The Scotia's collection includes ten specimens of Phcebetria, three of which are
P. cornicoides, the remaining seven P. fuliginosa. The labels of one of the former
and of two of the latter have unfortunately been lost, but all the others were obtained
on the part of the Scotia's voyage at present under consideration, except one of the
specimens of P. fuliginosa, which was obtained at Gough Island. The dimensions of

271

272 ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

these specimens do not indicate any very marked difference in size between the two
species. They may be tabulated thus :—

Culmen .
Tarsus ....
Midtoe ....

P. fuliginosa (7 examples).

P. cornicoides (3 examples).

Average.

Extremes.

Average.

Extremes.

4-45
3-2

4-8

4-8-4-1
3-0-3-4

5-0-4-5

4-05
3-1
4-75

4-2 -3-9
3-15-3-1
4-8 -4-6

It will be seen that there is a distinct difference in the length of the culmen, which
is shorter in P. cornicoides. This may, or may not, be characteristic of the species.
The sexes were not determined. The primaries are damaged or being moulted in most
cases, so that it is impossible to compare the length of wing. In plumage, all the
specimens of P. cornicoides can be picked out at a glance by the very distinctly
lighter grey colour of the neck, back, breast, and belly (the two species in this respect
may be compared to the Hooded and Carrion Crows, although the contrast is not quite
so marked) ; the scapulars and wing coverts of P. cornicoides also have a slightly
greyer tinge than those of P. fuliginosa. In other respects the plumage is quite
identical in the two species. The bills in the dried specimens are similar in size
and form, but the groove on the mandible is dull yellow \nfuliginosa, greyish black
in cornicoides.

Diomedea melanophrys was first logged on April 8th, in 52° 30' S., and again the
next day, and later, on the 12th, 15th to 17th, 19th, and around Gough Island on
April 21st and 23rd.

The Scotia's collection includes four skins of D. melanophrys :—

(a) No. 275 (obtained in April on the voyage north to Gough Island). Bill

entirely orange-yellow. Phase 1.
(6) No. 292, April 30th, 1904 (probably killed April 29th, 40° S., 2° 30' E.).

Bill orange-yellow, tip dusky brown ; head white. Phase 3.
(c), (d) The labels have been detached ; on April 27th, 1904, " two Mollies with
horn-coloured bills " shot, may refer to these, (c) Bill pale yellow, tip
greyish yellow ; head white. Phase 2 or 3. (d) Bill pale yellow, ridge of
hook brownish ; head white. Phase 2 or 3.

Dr E. A. Wilson * gives the following table of what he came to the conclusion were
various plumage- stages of D. melanophrys :—

Phase 1. White head and neck, orange-yellow bill, deep orange tip.

2. „ „ lemon -yellow „

3. ,, ,, ,, ,, dusky-brownish tip.

4. „ ,, dull yellow bill, blackish tip.

5. Grey head and neck, dusky brownish bill, darker tip.

* (British) National Antarctic Expedition: Natural History, vol. ii., 1907, pp. 111-1 13.

"SCOTIA RISE" REGION. 273

Of these birds he says :—

" Some were pale grey all over the head and neck, others had merely a broad or
narrow collar of grey incomplete below. The size also varied considerably, but all had
the bill of D. melanophrys, dusky yellow and always with a darker tip. We considered
these grey-headed and grey-necked individuals, if the bill was broad, to be the young
of D. melanophrys. There was no difficulty in distinguishing the grey-headed D.
melanophrys from the grey-headed Th. chlororhynchus and Th. culminatus, since the
colour of the bill in the two latter is much more clean-cut and distinctly black and
yellow than the dusky-brownish or yellowish bill of the immature D. melanophrys.

" Until October 19th we saw only immature birds, but on that day the white-headed
adult, with lemon-yellow bill and orange tip, appeared. After this we saw each day
for a while only adult birds, but on October 28th we had the following together : white-
headed adults with lemon-yellow bill and bright orange tip ; white-headed birds with
bright yellow bill and dusky tip, or with dull yellow bill and blackish tip ; and grey-
headed birds with dusky brownish bill and darker tip.

" Throughout the first half of November we saw one or other of these forms almost
daily, and came to the conclusion that they were simply age-changes, and that the grey-
headed were the immature."

The Scotia's specimens belong to phases 1 to 3 according to the above arrangement.
It does not appear that the Scotia encountered any birds in the grey-headed immature
stages of this species, although this cannot be definitely affirmed. It is significant,
however, in view of the curious way in which the Discovery sometimes for long periods
encountered only white-headed adults, at other times only grey-headed young birds,
and at others, again, all the phases together. The different varieties of " Molliemauks "
seen on the present section of the Scotia's voyage were described fairly clearly in the
Zoological Log, but it is possible that grey-headed stages of D. melanophrys were not
distinguished from the grey-headed or grey-collared birds which will now be described.

From April 9th onwards a distinct form of Albatros was noticed, which was logged
as the " Black-billed Albatros." It is thus described on the above-mentioned day in the
Zoological Log : — •" Mr Bruce saw a new albatros of the size and shape of D. melano-
ph rys, and of the same colour except the back of its neck, which was dark with a dark
ring complete or almost complete round about it, and a black beak.'' This form, after
being observed on April 9th in 51° S., was not again met with until the 16th, on which
day and on the ensuing days up to the 20th April it was again encountered. It was,
however, not observed in the immediate vicinity of Gough Island, nor subsequently.

In his account of the Discovery's birds, Dr Wilson (loc. cit., p. 113) says: — "In
the Atlantic between Punta Arenas and the Falkland Islands we occasionally saw one
or two of the typical adults, the last on July 27th, when D. melanophrys disappeared
entirely, and its place was taken by a form which we had before this hardly seen at all,
a bird in every respect the same in shape and size as D. melanophrys, but with a grey
ring always round the neck and the bill always quite black " ; and again in a footnote

274 ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

to the foregoing quotation: "This bird appears to agree with the ' Mollymauk '
(Thalassogeron sp. inc.) mentioned by Mr Eagle Clarke amongst the birds of Gough
Island (see p. 287 of this volume, ' The Birds of Gough Island '). Those that we
saw were evidently adult. They had the bill entirely black and the head white,
shading on the occiput, or sometimes on the hind-neck, into grey, which deepened on
the sides of the neck to form a well-marked grey collar incomplete on the fore-neck.
The feet were rosy pink. In other respects, as in size, the bird closely resembled
D. melanophrys. We saw it several times in March from 55° S. lat. northwards as
we came up to the Auckland Islands from Wilkes Land. We saw nothing of it in the
South Pacific; but in July we found it again in the South Atlantic between 30° and
40° S. lat. as we came north from the Falklands in 1904."

It appears highly probable that the " Black-billed Albatros " of the Scotia was the
same bird as that described by Dr Wilson in the extracts just quoted. Also, as he
suggests, it is possible that the Gough Island specimen (see Section IX., p. 287) is
likewise of this species. This specimen has, in the meantime, been submitted to
Mr Godman and other eminent authorities, and in his great work on the petrels
the former says : — *

" Mr Eagle Clarke, in his paper on the ' Birds of Gough Island,' procured by
the Scottish National Antarctic Expedition, mentions (The Ibis, 1905, p. 265) a bird
which he was unable to identify with any known species. . . . Mr Rothschild, to
whom he referred it for examination, recognised its resemblance to Thalassogeron
carteri, but noticed that it differed in having the toes nearly '75 in. shorter, and
also the hind-neck washed with grey like the back, instead of being white (Bull.
Brit. Orn. Club, xiv. p. 6). Mr Eagle Clarke has now kindly lent me this specimen
for examination, and I find that the short foot is occasioned by the absence of a
phalanx in two of the outer toes on both feet, but in other respects it agrees very
well with Thalassogeron carteri. There are, however, some minor points worthy of
notice, viz. : — the extension of the dark colour of the back to the hind-neck as mentioned
above, and a slight difference at the base of the culminicorn ; there is also a grey tinge
on the face and crown ; these characters, however, appear to me of trivial importance.
I ascribe the absence of a joint in the toes to a malformation, or an individual peculiarity,
and I am the more inclined to do so as the feet look out of proportion to the size of the
bird. Moreover, I am not aware that any other member of the order Tubinares is
without the full complement of toe-bones ; consequently I place this bird, for the
present at all events, under Thalassogeron carteri."

Accordingly, we may say that it seems probable that the " Black-billed Albatros "
of the Scotia's log, as well as the similar birds mentioned by Dr Wilson, may belong
to the species Thalassogeron carteri.

From April 10th to 19th inclusive (49° 30' to 43° S.) a grey-headed species of
albatros with dark bill and yellow culmen, and otherwise resembling Diomedea

* A Monograph of the Petrels, 1907-1910, ]> 3(51.

"SCOTIA RISE" REGION. 275

melanophrys, was seen daily (except on one day). They were recognised as belonging
to the same species as the two albatroses secured at Burdwood Bank in December 1903
(Thalassogeron culminates, see Section IV., p. 215), and were logged as "Burdwood
Bank Albatroses." None were seen in the neighbourhood of Gough Island or
subsequently on the voyage to the Cape of Good Hope. It is, however, equally if
not more probable that these birds may have been Thalassogeron chlororhynchus.

Diomedea exulans appeared irregularly on this voyage. Several were met with
on April 3rd, when one was shot, and again on the 7th, the 10th, llth, and 18th;
on the 19th very many were seen (nearing Gough Island, about 43° S.), but next day
only some young birds are logged. On the 21st, when the Scotia was lying to off
Gough Island, many were flying about her all day long.

Priocella glacialoides, after having been unusually numerous on March 31st, was
seen in decreasing numbers for the next three days (up to about 57° S.), and not again,
except a single example on April 17th (44° 30' S.).

" Blue Petrels " were logged nearly every day during this part of the Scotia's voyage,
chiefly up to 55° S., after which none were seen for several days. From 51° to 46° S.
one or two were seen daily, and then they began to appear in larger numbers. Many
were seen about 43° S., and also in the neighbourhood of Gough Island.

As no specimens of these " Blue Petrels " were obtained in this region, it is not
possible to say with certainty whether they were Halobasna csendea, or species of
Prion, but it may be regarded as probable that most were of the latter genus
(see Section VII., under Halobasna). The only "Blue Petrel" obtained at Gough
Island was a Prion, P. vittatus (Gm.) (see Section IX. of this volume, p. 286).

A fair number of Cape Pigeons (Daption capensis) were seen up to 55° S.,
thereafter one or two per day occasionally up to 43° S.

Giant Petrels (Ossifraga gigantea) were seen in small numbers up to 56°, after
which none were observed for a week. About 49° they again appeared, and a few
were seen almost daily up to the date of arrival at Gough Island.

Dr Bruce tells me that the Giant Petrels observed near Gough Island were dis-
tinctly darker in plumage than those seen in the far south.

A white example was seen on April 1st.

Wilson's Petrel (Oceanites oceanicus) was seen almost daily all the way, but not in
large numbers.*

On the 4th April a " Storm Petrel differing from Wilson's in having a white abdomen,
and this white extending on to the lower surface of the secondaries, except at their
tips," was seen. This very probably refers to Cymodroma grallaria, specimens of
which were obtained further north at Gough Island. It was observed almost daily
thereafter till Gough Island was reached.

* In general, it may be said that Wilson's Petrel, so far as plumage is concerned, could only have been confused
with Oceanodroma leucorrhoa (or allied forms), or with ProcMaria pelagica; the latter, however, has white under the
wings — a character which was at once " spotted " in Oymodroma grallaria.

VOL. IV. 10

276 ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

Several Diving Petrels (Pelecanoides sp.) were observed on April 8th and 9th
(52° 30' S. to 51° S.)-

Priofinus cinereus, logged as the "light brownish-grey petrel," was first noticed
on April 10th, in 49°25'S. On the 13th several were again seen, and thereafter the
species was observed daily till Gough Island was reached (where several examples were
obtained — Section IX.).

(Estrelata brevirostris, a specimen of which had been shot when these birds were
first met with on March '21st, continued to be logged at intervals (to be more exact,
on eight different dates) up to April llth (49° S.), but only in small numbers.

A larger species, otherwise similar to the last, was seen along with it on March 28th,
and again, though alone, on the next three days. Again on April 14th, 16th, and 17th
similar but larger birds were seen. These may have been Majaqueus seqidnoctialis.
Captain Robertson named them " Cape Hens" on seeing them.

(Estrelata mollis was first met with on April 4th in 55° S., and on several other
dates up to the 15th, after which it was seen daily (except on the 19th) till Gough
Island was reached. Off Gough Island it was found in large numbers, and specimens
were obtained.

Dr Brace's description on the 5th April (as well as the Gough Island specimens)
leaves no doubt as to the species, viz. : " About the size of a Cape Pigeon, more slender
in build, general colour pale grey, darker on the upper surface of the wings and top
of the head, white muzzle, dark eye, blackish beak, brownish ring round the neck,
most of the under surface white." It was usually logged as " Black-backed Petrel."

Several other forms of petrel were logged during the voyage to Gough Island, which
it is not possible to identify, as no specimens were obtained. One in particular,
logged as the " Grey-bodied Petrel," was seen almost every day from 5th to 19th April,
and on the 7th and 12th in large numbers. The following description in Dr Bruce's
log for April 5th apparently refers to the "Grey-bodied Petrel" : "One about the size
of a Blue Petrel, but brown instead of bluish grey, and paler brown on the under surface,
with one diffused white patch on its breast."

Skuas (Megalestris antarctica) were met with on the 18th April (two), when the
Scotia was still about 180 nautical miles south of Gough Island. Two were again seen
next day, and one on the 20th (around the island they were numerous).

Terns were logged on the 7th (three), 9th and 20th (one), which were probably
Sterna vittata.

Of penguins, some were observed on April 8th, 9th, and llth, in mid-ocean.
On the two latter dates they were identified as Spheniscus magellanicus and
Catarrhactes sp.

Section IX. — On the Birds of Gough Island, South Atlantic Ocean.*

By WM. EAGLE CLARKE, F.R.S.E., F.L.S., Keeper of the Department of
Natural History of the Royal Scottish Museum.

WHEN homeward bound from the farthest southern point reached — the newly discovered
Coats Land — the Scottish National Antarctic Expedition paid a flying visit to Gough
Island, and made the collection of birds here described. This contains forty -five specimens,
and, with the Antarctic birds, has been placed in my hands for identification and record.

Gough Island, or, to speak more correctly, Diego Alvarez (for such it was named by
its Portuguese discoverers), is situated in lat. 40° 19' S. and long. 9° 44' W. It lies far
out in the South Atlantic Ocean, being some 1500 miles W. by S. of the Cape of Good
Hope and about 2000 miles N. by E. of Cape Horn ; and, with the Tristan da Cunha
group, which lie over 200 miles to the northwards, is among the most remote of all
oceanic islands. It is small and uninhabited ; of volcanic origin ; from seven to eight
miles long, and from three to four wide ; and is lofty, rising to a height of 4380 feet.

The island has been but little visited, except by sealers, who, in days gone by,
found it worthy of their attention ; and the Scotia's party were the first naturalists who
have ever set foot upon its fastnesses.

Dr Harvey Pirie and Dr R. N. Rudmose Brown give the following description of
the island, and Mr D. W. Wilton gives me other valuable information :—

" Gough Island rises on every side abruptly from the ocean in sheer precipices
several hundred feet high. The general aspect of the island, as seen from ship-board,
is very beautiful, with its green slopes and moss- and lichen -covered cliffs, over which
numbers of rushing waterfalls shoot out into the sea with a drop of several hundred
feet. The only apparent landing-place is on the eastern side, where the party from the
Scotia landed. Here a ravine runs down from the interior to the coast, and along it
flows a small stream. Near the seaward end of this ravine are a few acres of level
ground covered with grass or, in the moister parts, with ferns and rankly growing
celery and docks. Here, too, is a narrow beach, perhaps a hundred yards long, strewn
with many large boulders and numerous fern-rhizomes of considerable size. At the
S. W. end of the island there appears to be a plateau of about half a square mile in
extent at an elevation of some 300 feet, but everywhere else the island rises into steep
ridges separated by narrow valleys, which must render its exploration a matter of
extreme difficulty. On the lower ground and up to a height of over 1 000 feet the island
is thickly covered with tussock-grass (Spartina ai'undinacea) and buckthorn-trees

* Reprinted, with slight verbal alterations, from The Ibis, Series VIII., v., 1905, pp. 247-268.

277

278 ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

(Phylica nitida) : the former spread profusely over the steeper slopes, and the latter
gnarled and stunted, yet growing vigorously even on the most exposed ridges. These
trees appear hardly to rise beyond twenty feet in height, and generally bear a thick
growth of lichens on their stems. Under the waterfalls and the sheltered banks of the
streams ferns and mosses grow in luxuriance. More than the general aspect of the
vegetation on the higher ground could not be determined, but the very summit of the
island seemed by its green appearance to be clothed with mosses and lichens."

To this description may be added a few items culled from Mr Comer's notes, to
which reference will be made more particularly anon. He tells us that there are two
kinds of trees on the island, one of which is quite plentiful, but the other is scarce.
These, though stunted, are dense in some places, and retain their leaves all the year
round. The thick bushes, he says, extend up to an elevation of about 2000 feet, while
the grass and brakes grow very rankly and make walking extremely difficult.

According to the Report on the Fisheries and Fishing Industries of the United States,
vol. ii. p. 415 (1817), Gough " Island at one time abounded in fur-seals and sea-elephants,
but is now almost deserted by those animals. About 1825 a party of American sealers
lived there, but met with such indifferent success that the station was abandoned."

The few accounts agree that, owing to the weather conditions usually prevailing,
landing on Gough Island is a matter of great difficulty. In the case of the Scotia,
three days were spent off the island, during which her staff, after landing under the
most trying conditions, were only able to remain ashore about four hours. This was
on the 22nd of April 1904.*

Previous to this visit only twelve species of birds had been identified as forming the
avifauna of the island. The naturalists of the Scotia observed or secured examples of
nineteen species, and added no less than twelve to the ornis of the island, of which two, or
possibly three, are new to science. The total avifauna now stands at twenty-three species.

Only three terrestrial forms are known, all of which are peculiar species — namely,
two buntings of the genus Nesospiza and a flightless Gallinule, Porphyriornis cotneri.

The two species of Nesospiza are the most interesting, not, however, because they
are novelties, nor because we owe our knowledge of them to the researches of the Scottish
Expedition, but because they differ very considerably from their single congener,
N. acunha}, peculiar to Tristan da Cunha, where it is now confined to Inaccessible
Island, though it was formerly also found on the main island of the group.

The Gough Island birds of this genus differ remarkably among themselves, and
whether they represent two species or only one in various stages of plumage is a matter
on which opinions differ. At first 1 was under the impression that they were repre-
sentatives, sexual or otherwise, of a single species ; but an examination of the material
revealed characters which it was difficult to reconcile with such an opinion, and which
led me to describe them as two species. My reasons for doing so will be explained in
the systematic portion of this contribution.

* For a fuller account of Gough Island, see Scot. Geog. Mag., xxi., 1905, pp. 430-440.

ON THE BIRDS OF GOTJGH ISLAND, SOUTH ATLANTIC OCEAN. 279

The Gallinule or " Island Hen " (Porphyriomis comeri) appears to differ only
slightly from the species (P. nesiotis) found on Tristan Island ; though, on the other
hand, both of them would seem, if report is to be relied upon, to be widely different
from the representative of the family found on Inaccessible Island (see Moseley, Natural-
ist on the Challenger, p. 122).

These three terrestrial endemic birds are, moreover, representatives of genera
entirely unknown elsewhere except in Tristan da Cunha ; and thus Gough Island,
though lying over 200 miles to the south, must be looked upon ornithologically as
an outlier of the Tristan group. The relationship is, however, somewhat remote,
for the buntings found on Inaccessible and Gough Islands are specifically very
distinct ; the Tristan possesses a peculiar genus of thrush, Nesocichla, with a single
species (N. eremita), which is not at present known to have any representative on
Gough Island.

The other birds forming the ornis of Gough Island are mostly Tubinares. The
number of species of this order observed during the short visit of the Scotia is quite
remarkable, and it would seem probable that a thorough investigation of the island-
as yet almost untrodden by the foot of man — in the summer season would reveal the
fact that it is a perfect paradise as a breeding-station for " petrels." Indeed, one species
of albatros (Thalassogeron eximius) has not been obtained elsewhere.

The only previous contribution to our knowledge of the avifauna of the island is, I
believe, contained in a communication by Mr G. E. Verrill to the Transactions of the
Connecticut Academy, based upon the collections and experiences of Mr George Comer.
Mr Comer resided on Gough Island from the 22nd of August 1888 until the 23rd of
January 1889, and was engaged, along with others, in the capture of seals. During
the period he got together a very creditable ornithological collection, and among his
specimens were two birds which proved to be new to science, namely, the flightless
Gallinule (Porphyriorms comeri) and an albatros (Thalassogeron eximius). Mr
Comer's notes are of considerable interest, and I have not hesitated to make use of them
where desirable. I have also included the names of the very few species observed by
him of which examples were not obtained or noted by Dr Bruce and his colleagues, in
order to render my account of the avifauna of this most interesting and little-known
isle as complete as possible : these are given within square brackets.

The date of the visit of the Scotia to the island was not the best for ornithological
work, April being mid-autumn in the southern hemisphere. Unfortunately, too, the
state of the weather did not permit of more than a few hours being spent ashore, and
even then the party could not proceed far inland, owing to the uncertainty of the pre-
vailinw meteorological conditions. Under these circumstances the members of the

O O

Expedition are to be congratulated upon having accomplished so much good work.

The colours of the bills and feet, when given, are taken from a series of carefully
prepared drawings made at the time of capture of the various birds by Mr Cuthbertson,
the artist to the Expedition.

280 ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

The few species in the following list the identification of which I consider doubtful
are not numbered.

I have to thank Mr W. P. Pycraft for having most obligingly examined certain
material submitted to him, and for giving me his valuable opinion thereon.

I shall have occasion to make several references to the following works, in addition
to other literature :—

CARMICHAKL. — " Some Account of the Island of Tristan da Cunha and of its Natural Productions." By
Captain DUGALD CAKMICHAKL, F.L.S. Trans. Linn. Soc., xii. pp. 483-513 (1817).

THOMSON. — Voyage of the " Challenger " : The Atlantic. By Sir C. WYVILLB THOMSON. Vol. ii. (1877).

VEBBILL. — " On some Birds and Eggs collected by Mr George Comer at Gough Island, Kerguelen Island,
and the Island of South Georgia, with Extracts from his Notes, etc." By G. E. VERKILL. Trans. Connecti-
cut Acad., ix. pp. 430-478 (1895).

SALVIN. — Catalogue of Birds in the British Museum. Vol. xxv. (Tubinares) (1896).

1. Nesospiza goughensis.

Nesospiza goughensis Eagle Clarke, Bull. Brit. Orn. Club, xv. p. 18.

This species possesses the essential characters of the genus Nesospiza as propounded
by Cabanis ( Journ. fur Orn., 1873, p. 154). It is a larger bird than the type (N.
acunkse) ; it has a longer and more slender bill, in which the culmen is more arched and
the keel of the mandible straighter ; and the third primary is slightly the longest.

Adult Male. — General colour bright olive-green, slightly washed with silvery grey ;
centre of abdomen, under tail-coverts, a stripe from the forehead over and behind the
eye, and a malar stripe olive-yellow ; chin, throat, and lores black ; primaries and
secondaries slate-grey, with silvery-grey tips and bright olive-green margins, and their
under surfaces broadly margined with silvery grey on the inner webs ; under wing-
coverts grey washed with yellowish green ; central pair of rectrices olive-green, the
rest grey edged and slightly washed with green. Bill and feet, in life, clove-brown.
Wing 4'15 inches, tail 3'68, tarsus T18, culmen 71.

This is probably the summer plumage.

Adult Female. — Resembles the male, but is not so brightly coloured, being dull
green. Chin, throat, and lores dusky, and, with the buff stripes over the eye and on
the malar region, inconspicuous ; feathers of the back and outer margins of secondaries
slightly fringed with reddish brown ; secondaries and tertials tipped with greyish buff ;
flanks faintly washed with brown; middle of abdomen and under tail-coverts buff;
central rectrices dull green, the next edged with buff and the outermost with green.
Wing 4'0 inches, tail 3'5, tarsus 1'15, culmen '68.

? Adult in Winter Plumage. — Pileum and hind-neck olive-grey ; back tawny
olive, broadly striped with black on the interscapulary region ; upper surface, wing-
coverts, and tail washed with sage-green ; primaries narrowly edged outwardly with
brown ; secondaries broadly margined with sage-green and tawny olive ; under parts
greyish green, washed with tawny olive on the breast and flanks and "passing into buff
on the lower abdomen and tail-coverts.

ON THE BIRDS OF GOUGH ISLAND, SOUTH ATLANTIC OCEAN. 281

This specimen is unfortunately unsexed, but, since the yellow stripes on the
head and the black on the throat and lores are brighter than in the female, it is
probably a male.

I have ventured to suggest that this bird is an adult in winter plumage, taking my
cue from the fact that a somewhat analogous seasonal change is to be found in the male
of Phrygilus melanodei~us of the Falklands. I may mention that the adult male of
N. goughensis bears a strong resemblance * in colour and markings to the adult male of
P. xanthogrammus of the Falklands, in which apparently the various stages of plumage
are as yet incompletely known, as are also, unfortunately, those of Nesospiza acunhse.

These buntings, with those next to be described, were captured on the low ground
bordering the landing-place, where they were mostly observed hopping about on the
boulders, stranded wood, and seaweed, a little above high-water mark. They were not
so tame as to permit themselves to be caught, but readily allowed an approach to within
two or three yards, and then flitted off to a short distance. They were not observed
in the ravine running inland, nor on its steep sides.

Mr Comer (Verrill, t. c., p. 463), alluding to these birds, says that small birds, like
sparrows, are very common at Gough Island, and are of two kinds — one slatish above,
yellowish beneath, and with a round black spot on the breast : the other much like the
first, but lacking the black spot ; possibly it is the female. Of the habits he merely
remarks that they are very tame and sing.

2. Nesospiza jessise.

Nesospiza jessix Eagle Clarke, Bull. Brit. Orn. Club, xv. p. 18.

Male and Female. — General colour orange-buff (brighter on the lower back), streaked
with black on the head, back, scapulars, lesser wing-coverts, breast and flanks ; primaries
blackish, narrowly edged with dull yellow ; secondaries and wing-coverts blackish,
broadly edged externally with buff ; rectrices with dusky centres and broad buff margins.
Bill and feet blackish. Wing 4'05 inches, tail 3'45, tarsus 1'2, culmen 0'65.

In this form the culmen is nearly straight, the gonys is ascending, the first primary
is equal to the sixth, and the tail-feathers are lanceolate. In all these respects it
differs from N. goughensis.

I at first thought that we had. in these buff specimens the female and young of the
green .V. goughensis, but on dissecting specimens it was found that the green birds
were of both sexes. There then remained the possibility of the buff birds being the
young of N. goughensis, though differing remarkably in plumage from either parent.
An examination of the material, however, disclosed the fact that, although identical
in plumage, some specimens were unmistakably immature, while others possessed
characters which I interpreted as indications of maturity. In these examples the
tendons of the feet were ossified or partially ossified, the fibula was fused to the tibia,

* See Gould's figure in Darwin's Voyage of the Beagle : Birds, plate 33.

282 ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

and the claws were well-worn. In this connection it is important to remember that,
if all young, these birds could only vary a few weeks in their respective ages, for they
were obtained in the autumn, and hence must be " birds of the year." This evidence,
and the other peculiarities alluded to wherein N. jessix differs from N. goughensis, led
me to depart from my first impressions regarding the relationship of these forms to
each other and to consider them distinct species. I may have been mistaken in so
doing, but I think it best to leave it an open question — one to be decided by further
investigations based upon a more complete set of specimens.

Judging from Mr Comer's rough descriptions of the Passerines, he does not seem
to have observed this form, and yet he remained on the island down to the very end
of summer, when young birds must have been in evidence. We must remember,
however, that his observations were not published till some six years after he had
left Gough Island, and therefore much that he has told us may have probably been
from memory.

3. Porphyriornis comeri Allen.

The flightless Gallinule peculiar to this island was abundant in the dense under-
growth along the sides of the stream. Among this it endeavoured to hide, but its
brilliantly coloured bill and feet easily betrayed it. The specimens captured were
secured whilst running amongst this thick mass of vegetation. One ran into a burrow
in its endeavours to escape, and it is probable that others resorted to similar tactics.

Five specimens were secured, one of which is an immature bird, though full-grown
so far as size is concerned. In this undescribed phase the bill and the frontal shield,
which is small, are dull olive-green, and the tarsus and toes dull oil-green. It also
varies in plumage from the adult, having the upper surface a warm vandyke-brown,
a little darker on the head and primaries ; the under surface strongly washed with
sepia-brown ; the flanks brown, two or three of the feathers on each side having short
inconspicuous cinnamon stripes ; the outer margin of the first primary cinnamon-buff ;
the edge of the wing streaked with greyish white ; the feathers of the lower part of the
abdomen edged with greyish white.

In the adult specimens the frontal shield and the basal two-thirds of the bill are
deep coral-red, the distal third of the bill bright yellow. The exposed portion of the
tibia is deep coral-red. Tarsus and toes bright yellow, the former irregularly blotched
with coral-red, while a narrow line of the same colour runs along the sides of the latter.
The amount of yellow and red varies much in different specimens, and in some the red
predominates. The iris is crimson. The wing-measurement of the adults varies from
5 '3 to 5 '95 inches ; in the young specimens it is 5*5 inches.

In form and in the colour of its plumage this bird very closely resembles our
familiar waterhen (Gallinula cMoropus). The bill and frontal shield are larger and
the feet more robust.

Regarding the habits of this bird, which he calls the " Mountain Cock," Mr Comer

ON THE BIRDS OF GOUGH ISLAND, SOUTH ATLANTIC OCEAN. 283

(Verrill, i.e., pp. 434-435) states that they "cannot fly and only use their wings to
help them in running. . . . They are quite plentiful and can be caught by hand. . . .
The bushes grow on the island up to about 2000 feet, and these birds are found as far
as the bushes grow. I do not know how many eggs they lay. . . . When alarmed,
their note is a shrill whistle. They eat the eggs of other birds and also follow the tide
down to feed on small animals left by the receding waves, but are careful never to go
into salt water . . . but are very fond of getting into fresh water and splashing it
over them. . . . They run swiftly when pursued, helping themselves with their wings,
and endeavour to get under a tussock, or some such place, to hide. . . . They have a
very peculiar habit of always hopping over every obstacle, such as a branch or dead
stick, instead of going under it as might be expected."

This species differs from P. nesiotis, of Tristan da Cunha, in having scarcely any
white on the outer edge of the first primary, and in the narrower and smaller streaks of
white on the flanks.

4. Sterna vittata Gmelin.

There are five specimens of this tern in the collection. One of these is an adult,
and the rest are in the pretty first plumage, in which the back is strongly barred with
buff and black, and the buff fore-neck finely dusted with grey. In the young birds in
life the bill is brownish black and the feet pinkish brown. Many of these terns were
seen in the vicinity of the island.

Mr Comer describes this bird as not uncommon at Gough Island, but makes no
mention of its breeding there. It has been recorded from Tristan da Cunha.

5. [Anous stolidus (Linn.).

This noddy was apparently not observed by the members of the Expedition.
Mr Comer's collection contained a specimen from Gough Island, and he remarks : " Quite
a number here " (Verrill, i.e., p. 452). It occurs and nests at Tristan da Cunha.]

[Larus dominicanus Licht.

This gull is included in Yen-ill's lists (t.c. pp. 450 and 464) for Gough Island, but no
specimens appear to have been obtained there. Mr Comer remarks that there were "a
very few sea-gulls ; do not think there were more than six around the island." These
remarks are supposed by Verrill to refer to this species. It did not come under
the notice of the Scottish naturalists, nor does it seem to have been observed at
Tristan da Cunha.]

6. Megalestris antarctica (Lesson).

Many skuas were observed off the island, and two specimens were added to the
collection — probably birds of the year, since they show rufous markings on the mantle,

neck, and lesser wing-coverts.

VOL. IV. 11

284 ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

Under the name of "sea-hen," Mr Comer (Verrill, t.c., pp. 450 and 464) describes
them as being "quite plentiful," and commencing to lay in the middle of September.
" When the penguins lay, the sea-hens come ashore in large numbers, and get their
living by robbing the nests and catching the young penguins. They also kill the
young albatroses, and all the small birds they can catch." Mr Comer killed about 300
of these birds at Gough Island for the sake of their feathers.

This is also a common bird at Tristan da Cunha.

7. Oceanites oceanicus (Kuhl).

Wilson's Petrel, together with Cymodroma grallaria, was observed in some
numbers in the vicinity of the island. One specimen was obtained, but another which
had been shot was carried off by a skua ere it could be retrieved. Although this
species has a remarkably wide range and occurs in the Atlantic as far north as the
European and North American coasts, it has not yet been recorded for Tristan da
Cunha, and did not come under the notice of, or was not identified by, Mr Comer at
Gough Island.

8. Cymodroma grallaria (Vieillot).

This species was observed off the island along with Wilson's Petrel, and was the
more numerous of the two. Several examples were obtained and are in the collection.

Though widely distributed in the southern oceans, it does not seem to have been
hitherto observed at the Tristan da Cunha group or at Gough Island.

9. Puffinus assimilis Gould.

The collection contains a pair of these birds which Dr Pirie captured in a deep
nesting-hole on the steep grass-covered bank flanking the side of the ravine, his atten-
tion being drawn to them by their loud croakings. Similar burrows were extremely
numerous, in fact the bank was honeycombed with them ; and numbers of the birds
were seen off the island. Basal half of the bill pale blue, distal part and nasal tubes
black. Front of tarsus and toes greyish blue, posterior portion of tarsus and sides of
toes black, webs yellowish.

This species does not appear to have been previously recorded from Gough Island,
or from any of the islands of the Tristan group or their neighbourhood.

10. Priofinus cinereus (Gmelin).

There are three specimens of this bird in the collection, all of which have the
feathers of the back, upper tail-coverts, and wing-coverts edged with paler grey than
the rest of their exposed portions. In life the maxilla is dull pea-green, the nostrils,
culmen, and unguis are black ; the mandible with apical plate and cutting-edge is black,
the lower plate is pea-green. The tarsus and toes are pinkish grey, darker or blackish
at the joints, the webs yellowish. In two of the skins the feet are yellow.

ON THE BIRDS OF GOUGH ISLAND, SOUTH ATLANTIC OCEAN. 285

A considerable number of these " light brownish-grey petrels, skin No. 9," were seen
in the immediate vicinity of the island.

This species has not a place in Verrill's list for Gough Island, nor did it come under
the notice of the CJiallenger naturalists at Tristan da Cunha, but Carmichael (t.c.,
p. 497) records it under the name of Procellaria cinerea for the group.

11. Majaqueus sequinoctialis (Linn.).

One specimen was obtained, skin " No. 20, Gough Island," and similar " Black
Petrels " are recorded as having been observed. In the example obtained the bill was
yellowish, with the basal part of the culminicorn, its margins contiguous to the latericorn.
and its tip black ; the distal plate and the narrow median plate of the mandible black.

This species does not appear to have been recorded hitherto from any of the islands
of the Tristan group.

12. CEstrelata mollis (Gould).

A great number of "Black-backed Petrels (skin No. 14)" were observed off the
island, and three examples which were secured are in the collection. In these specimens
the bill was black, the tarsus and basal half of the inner toe and of both webs were
pinkish white, the rest of the foot was deep brown.

This species was not represented among the birds collected at Gough Island by
Mr Comer, and is not included in Verrill's list; but I think that it is probably the un-
known species there alluded to under the name of " Paddy unker," a " dark bird with
white breast and white feet," of which there is one egg, measuring 2'37 x T64 inches,
in Mr Comer's collection (Verrill, t.c., p. 449).

<E. mollis has been recorded from Nightingale Island of the Tristan da Cunha group
(Salvin, Voy. " Challenger" : ZooL, ii. pt. viii. p. 144).

13. \GEstrelata lessoni (Garnot).

Mr Comer's collection contained an example from Kerguelen, and he states (Verrill,
t.c., pp. 448-464) that this "Mutton-bird" was also common at Gough Island. No
specimens were, however, obtained there, nor has the bird been recorded from Tristan
da Cunha. ]

14. Ossifraga gigantea (Gmelin).

Many " Nellies " were observed off the island, and one example, a female, was
obtained — a remarkably dark specimen, without a trace of brown, its plumage being
throughout deep slaty grey, each feather (except the remiges, the greater wing-coverts,
and the tail) terminally or subterminally margined with a slightly paler tint.

The Giant Petrel breeds at Gough Island, where Mr Comer (Verrill, t.c., pp. 447-8
and 464) says that it is not numerous, and commences to lay at the middle of September.
He tells us that it enters the penguin " rookeries. " and carries off the young to eat, and

286 ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

also pulls birds [petrels] out of holes in the ground. This destructive species also
breeds at Tristan da Cunha.

15. Prion vittatus (Gmelin).

There is only one specimen in the collection, and this is seemingly a large example,
for the wing measures 87 inches. The lower plate of the mandible was pale blue, and
the remainder of the bill black. Tarsus and toes cobalt-blue, webs black.

Great numbers of " Blue Petrels or Whale-birds " were also seen, and other specimens
which were shot were carried off by skuas ere they could be picked up from the water.

This Prion has not a place in Verrill's list for Gough Island ; but as " Procellaria
vittata" it is included in the ornis of Tristan by Carmichael (t.c., p. 497), and Wyville
Thomson (t.c., p. 177) mentions it as a breeding species on Inaccessible Island.

16. [Prion desolatus (Gmelin).

Mr Comer obtained skins of this bird at Kerguelen, and remarks ( Verrill, t.c., p. 449)
that it is also found at Gough Island, but he does not appear to have brought back
specimens procured there. Salvin (Cat. B., xxv. p. 435) says that it is found between
35° and 60° S. ; but it has not, so far as I am aware, been recorded from Tristan da
Cunha.]

17. Pelecanoides urinatrix (Gm.).

The collection contains a single example which was captured from the Scotia while
her party of explorers was ashore on the 22nd of April 1904. According to a coloured
drawing, made at the time of capture, the tarsus and toes in life are cobalt-blue and
the webs and claws black.

No Pelecanoides has hitherto been recorded, 1 believe, for any of the islands of the
Tristan group ; and if we follow Salvin and regard P. urinatrix and P. exsul as distinct,
then a very considerable extension eastwards of the range of this species is indicated
by its occurrence at Gough Island, for it does not appear to have been previously
obtained in the South Atlantic elsewhere than at the Falkland Islands. It is even
possible that it breeds at Gough Island, for Verrill (t.c., p. 449) mentions under this
species that there is an egg obtained there in Mr Comer's collection which is " supposed "
to be that of a " Diver." This egg measures 1'57 x T6 inch as against 1'54 x T26 and
1-52x1 '23 for eggs of Pelecanoides obtained by Mr Comer, together with skins, at
Kerguelen, and hence the P. exsul of Salvin (t.c., p. 438).

18. Diomedea exulans Linn.

Many albatroses of this species were observed around the Scotia as she lay off the
island from the 21st to 23rd of April, but no specimens were obtained.

Mr Comer (Verrill, t.c., p. 437) says that this species is common at Gough Island
and breeds there. It commenced to lay at the end of December ; he obtained the first

ON THE BIRDS OF GOUGH ISLAND, SOUTH ATLANTIC OCEAN. 287

eggs on the 26th of that month, and found them quite plentiful by the third of January.
The young, he says, must be ten months old before they can fly, and not more than five
per cent, live to leave their nests ; they are killed by the skuas and Giant Petrels.
Mr Comer considers that the Gough Island race is smaller than those from the other
islands (South Georgia and Kerguelen), and his statements are borne out, according to
Verrill, by both specimens and eggs.*

[Diomedea melanophrys Temm.t

The occurrence of this species must be regarded as doubtful. Mr Wilton records
it in his Log as being abundant off the island on the 21st of April. No specimens of
this albatros are, however, in the collection, nor are they mentioned as having been
obtained. The species is not included in Verrill's list, based upon Mr Comer's material
and observations ; nor does it appear to have been recorded from Tristan da Cunha.]

19. [Thalassogeron eximius Verrill.

This was described as a new species by Verrill (t.c., pp. 440-445), being founded on
a skin and skeleton which were obtained, together with seventy-five eggs, at Gough
Island by Mr Comer.

It is described as similar in plumage to T. chlororhynchus, but the mandible lacks
the transverse yellow bar at its base and is entirely black (except at the extreme outer
end, where it is tipped with light horn-colour). The bright yellow of the culmen
deepens and brightens into orange in the middle, and finally into dull red on the
margins, growing paler towards the tip.

Mr Comer says there is but one kind of "Molly-Mawk" on Gough Island, which
is known as the Blue-head. The birds lay their eggs separately, keeping well apart,
and are scattered about the island among the tussocks and brakes. They commence to
lay on the 20th of September, and their nests are like those of D. exulans, only smaller.

It is possible that some of the birds recorded by Mr Wilton as Diomedea melano-
phrys (but of which no specimens appear to have been obtained), and as being numerous
off the island during the stay of the Scotia, were of this species.]

20. Thalassogeron sp. inc. +

There is in the collection a single specimen of a " Molliemauk " of great interest,
since it does not entirely agree with the description of any known species.

I was inclined to think that it might be an immature example of T. chlororhynchus
in a phase of plumage which, if known, is not described by Salvin (t.c., pp. 451-2). It
differs, however, from the mature bird of that species in having the bill shorter and
entirely black, and the toes much shorter. Its dimensions are: wing 18*2 inches,
tarsus 3'0, middle toe 3 "5, outer toe 3 '2, inner toe 2'3, and culmen 4'75.

* See this volume, p. 292. t Ibid., p. 272. J Ibid., p. 274.

288 ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

This bird was captured under somewhat singular circumstances. Dr Harvey Pirie
was proceeding up the ravine when he came upon it on the top of a mass of tussock-
grass on which it had alighted, but from which it could not rise on the wing again.
It was apparently quite uninjured, and could only have been there a few minutes,
for other members of the party had passed the spot only a little in advance, and
could not have failed to see the bird had it been there, as the ravine is particularly
narrow at the point where it was found. This curious incident might be explained
on the supposition that it was a young bird essaying to reach the sea from its inland
nursery which had halted by the way. Mr Comer tells us that the young of Diomedea
exulans do not fly until they are ten months old ; and if this holds good even for a lesser
period in other species it would help to explain the date and the peculiar circumstances
under which this specimen was found away from the sea.

Mr Rothschild and Dr Hartert, who have examined the specimen, tell me that it
resembles T. carteri, recently described by Mr Rothschild (Bull. Brit. Orn. Club, xiv.
p. 6) from a single example which came ashore with an injured wing at Point Cloates,
N.W. Australia, and that the Gough Island bird only differs in having the toes nearly
•75 inch shorter, and in having the hind-neck washed with grey instead of being white.
As regards my suggestion that the specimen under consideration may possibly be
immature, Dr Hartert remarks that he finds nothing to show that other albatroses
have the bill black in the young, and, moreover, that such an example would hardly
have a face with pure white sides.

21. Phocbetria fuliginosa (Gmelin).

A typical specimen of the Sooty Albatros is in the collection, and many others were
seen, with lesser numbers of Hutton's Albatros (P. cornicoides), around the Scotia as
she lay off the island.

Mr Comer (Verrill, t.c., pp. 445 and 464) describes the species breeding at Gough
Island as having the beak dark with " a yellow stripe on each side." It is common but
does not breed in " rookeries " ; it places its nests separately on cliffs or projecting rocks,
where it is most difficult to get at them. The bird commences to lay by the middle of
September, and, while sitting, keeps up a continual cry similar to that of a young goat.

This is probably the form which also nests at Tristan da Cunha, for Captain
Carmichael (t.c., p. 489) alludes to the bird as the "Black" Albatros (Diomedea
fuliginosa) and describes its breeding-habits.

22. Phosbetria cornicoides Hutton.

I think that there can be little doubt of this being a distinct species. It was only
imperfectly described by Captain Hutton (The Ibis, 1867, p. 192), from specimens
observed at sea, and as a variety of P. fuliginosa.

In a letter dated January 4th, 1905, Captain Hutton tells me that this form has the

ON THE BIRDS OF GOUGH ISLAND, SOUTH ATLANTIC OCEAN. 289

" back and breast grey, head and neck sooty brown," and that his statement about the
mark on the mandible being white was an error. As we now know, this mandibular
stripe is pale blue and much smaller than in P. fuliginosa.

Salvin (i.e., p. 454) says, "If these birds can be traced to a definite breeding-place
where they alone are found, it would be well to assign them specific rank." This Captain
Hutton has done, for he tells me that it is the common form in New Zealand, and
" breeds at the Auckland Islands and Antipodes at the end of October," but that he
does not recollect ever having seen the sooty form in New Zealand. That P. fuliginosa
alone, so far as our present information goes, breeds at Gough Island and Tristan da
Cunha has already been mentioned under that species, and this affords additional evi-
dence in favour of the specific difference of the two forms.

This species was, however, observed off Gough Island by the Scottish Expedition in
somewhat less numbers than the last, but no specimens were obtained there.

23. Catarrhactes chrysocome (Forster).

Several Rock-hopper Penguins were seen ashore, but at places where they could not
be reached. Two skeletons, with some feathers still adhering to them, were found on
the beach where the landing was effected, and were secured.

At Gough Island, Mr Comer says (Verrill, t.c., p. 462), there is no other kind of
penguin. They "number millions" and commence to lay by the 15th of September.
This is the only species of penguin found at Tristan da Cunha, where it breeds on all
three islands.

In addition to the three endemic land-birds, to which special allusion has already
been made, the following marine species have been recorded for the Tristan da Cunha
Isles which have not been observed or obtained at Gough Island, namely : Anous
melanogenys, Pelagodroma marina, and Daption capensis, which Moseley (op. cit.,
p. 134) says breeds there !

Section X. — South Atlantic — Voyage from G-ough Island to Cape Town,

thence to St Helena.

By L. N. G. RAMSAY, M.A., B.Sc.

IN this section the oceanic observations of the voyages from Gough Island to the
Cape of Good Hope (April 24th to May 5th, 1904), and from Saldanha Bay to St
Helena (May 21st to 29th), will be dealt with.

The rich petrel-fauna of the southern ocean was gradually left behind on the voyage
to St Helena, and as the tropical seas were entered, the almost complete absence of
bird-life recorded in the log forms a striking change.

Leaving Gough Island, the Scotia followed an irregular course towards the Cape,
keeping about 38°-40° S. till within a couple of hundred miles or so of the latter.

Sooty Albatroses of both species (Plwebetria fuliginosa and P. comicotdes) were
met with up to the day of arrival at Table Bay. Only the yellow-billed species
(P. fuliginosa) was seen till the second day after leaving the island ; for the next few
days both were common, but after the 27th their numbers decreased greatly. None
were seen after Cape Town was left.

Diomedea melanophrys was plentiful nearly every day up to the arrival at the
Cape, but was not subsequently seen.

Besides the above three species, the Wandering Albatros, D. exulans, was the only
albatros met with ; it was numerous up to the day of arrival at Table Bay, also on the
22nd and 25th May many were about; next day (24° S.) a few were seen, and
those were the last albatroses that the Scotia encountered.

It is here necessary to add a note as to the identity of the large albatroses
observed by the Scotia naturalists, which have all been recorded in these pages as
D. exulans. Two other closely allied species are at present recognised, inhabiting
the southern oceans, to wit, D. regia and D. chionoptera. Of these, Godman *
says : —

"The geographical distribution of D. regia and D. chionoptera appears to be
restricted to certain districts of the southern oceans, whereas D. exulans, the Wandering
Albatros, seems to be dispersed over the whole of the seas inhabited by D. regia and
D. chionoptera. Thus, while D. regia is an inhabitant of New Zealand and the adjacent
islands, D. chionoptera is found only in the South Atlantic."

It further appears that immature stages of D. chionoptera are practically indis-
tinguishable from stages up to the adult of D. exulans, and in fact the identity of

* Monograph of the Petrels, p. 322.
VOL. IV. 291 12

292 ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

the various species remains in a somewhat hazy condition until more material is
available.

All this must be borne in mind in connection with the records of D. exulans
throughout the voyage of the Scotia. Godman adds on p. 323 that he thinks it
highly probable that the albatros of Gough Island (this in reference to Mr Eagle
Clarke's paper, Section IX.) will be found to be D. chionoptera rather than D.
exulans.

It seems improbable that the fully adult white- winged examples of D. chionoptera,
if present in numbers, could have been overlooked by such competent observers as
several of the staff of the Scotia.

The day must be looked forward to when the numerous problems connected with
the albatroses and petrels of the southern oceans may be solved by the despatch of an
expedition having this as one of its main objects.

Three skins of D. exulans were brought back by the Scotia. Data of two of these
are unfortunately lacking at the present time (1912). The description and dimensions
of all three may be given : —

(a) Immature skin. "No. 100." Wing 23 ins. ; culmen 5'6 ins. ; tarsus 4'2 ins. ;
mid-toe 6'25 ins. Bill pale yellow ; forehead, sides of head, and throat white ;
wings above, and scapulars, entirely blackish brown ; crown and rest of upper
parts, fore-neck and breast, and flanks, lighter brown ; axillaries and whole of
under wing-coverts white ; middle of abdomen nearly white, fine brownish
vermiculations on the thighs.

(b) Adult, "7th May 1904."* Wing 24 ins.; culmen 6'5 ins.; tarsus 4'6 ins.;
mid-toe 6 '5 ins. Bill pale yellow. Longer scapulars, and all upper surface of
wing, blackish brown, except for some white mottlings on the coverts forming an
irregular patch near the elbow-joint. Under wing-coverts and axillaries white ;
head, fore-neck, and belly white, but a mottled brown and white patch on the
crown ; back of the head, and all the rest of the upper parts, chest, flanks,
thighs, and under tail-coverts white, closely vermiculated with fine transverse
brown lines ; tail grey-brown.

(c) Adult. Unlabelled. Wing 2375 ins. ; culmen 5'9 ins. ; tarsus 47 ins. ; mid-
toe 67 ins. Colouring as in the last, except that the vermiculated parts are
much whiter in appearance, owing to the brown lines being much further apart.

After Gough Island was left behind, a few examples of Priqfinus cinereus were seen
daily up to May 1st (39° 30' S., 10° 30' E.). These were probably the last met with,
although two of the specimens of this species collected by the Scotia, at present in the
Royal Scottish Museum, are labelled "near Ascension." The original labels of these skins
are lost, and it is highly probable that this locality is due to a mistake, as the Zoological
.Log makes it clear that no petrels of any sort (except a few Storm-Petrels) were obtained

* At this date the Scotia was lying at Cape Town. This is probably one of three examples which were shot on
April 29th.

SOUTH ATLANTIC — VOYAGE FROM GOUGH ISLAND TO CAPE TOWN, ETC. 293

or seen anywhere near Ascension. This record has unfortunately been quoted by
Mr God man in his monograph.

Majaqueus xquinoctialis was seen in small numbers on May 1st to 5th. It was
again observed between Cape Town and Saldauha Bay. and also in numbers on May
22nd and 25th (about 26° S.), and a single one (the last seen) by one of the sailors
on the 27th (about 22° 30' S.).

CEstrelata mollis continued to be met with all the way to South Africa, except on
the last day, and was also met with on the voyage to St Helena (but only at a long
distance from land) in numbers on the 25th and 26th, and a single bird, the last seen,
on the 28th, in about 20° S.

On April 29th a petrel of a species not hitherto met with was shot, and several
others seen. The specimen proves to be CEstrelata macroptera, A.Sm. Others of the
same were seen on- the next three days, and also on May 4th and 5th, when the Scotia
was approaching land. The description and measurements of the specimen obtained
are appended :—

No. 291, killed April 29th, 1904. Wing 11'4 ins.; bill (gape to tip) 2'1 ins.;

tarsus 1'75 ins. ; mid-toe 2'4 ins. ; inner toe 2'1 ins. Bill black, with a whitish

mark on the hook and near the tip of the mandible. Legs and feet black.

Plumage of the upper parts entirely dark blackish brown, with faint paler

margins to the feathers of mantle and hind-neck ; on scapulars and some of

lesser wing-coverts a more reddish-brown tinge. The feathers have light greyish

bases. Forehead and lores greyer than crown. Throat cinereous. Fore-neck and

chest like the back, rest of under parts rather paler, and with a more brownish

tinge. Under tail-coverts dark like the upper ones ; longest ones almost as long

as central tail feathers. Under wing-coverts and axillaries dark like the hack.

Whale-birds (Prion sp.) were seen in small numbers (many on April 26th) up to

May 2nd (38° S., 14° 30' E.). The only subsequent record is a doubtful one, of a

single bird on May 22nd in 22° 30' S.

Single examples of Daption capensis were logged on May 22nd, 25th, and 26th,
(24° S.).

Wilson's Petrel (Ooeanites oceanicus) and White-bellied Storm-Petrels (Cymodroma
i/rallariii.) were seen daily on the voyage to the Cape, but not in large numbers. Very
few Storm-Petrels were observed between the Cape and St Helena, only one or two
being noted ou three different days.

On the last two days before reaching St Helena only a single bird (an CEstrelata
mollis) was seen.

Section XI. — South Africa.
By L. N. G. EAMSAY, M.A., B.Sc.

AFTER spending twelve days at Cape Town, the Scotia left Table Bay on May 17th.
Her course lay northward along the coast, and next forenoon she lay to off Dassen
Island, one of the islands where the Jackass Penguins (Spheniscus demersus) nest
under protection. A permit had been obtained for collecting here, but the heavy
surf prevented a landing being made. Saldanha Bay was reached the same evening,
and here the vessel anchored for two or three days. A small collection of birds,
comprising eight species, was made here, and about midday on the 21st the anchor
was weighed and the Scotia left for St Helena.

The Saldanha Bay collection includes the following : —

Phalacrocorax capensis (Sparrm.). — Two immature skins, shot off Marcus Island,
May 21st, 1904.

This is one of four species of Phalacrocorax which frequent the coasts of South
Africa. It is said to be extraordinarily abundant along the west coast.*

Numerous other shags or cormorants were seen off the coast at Saldanha Bay, and
on Meeuw Island, a tiny islet in the bay, their nesting-places were found. Dr
Rudmose Brown describes these as follows t : — " On low acacia-bushes the shags one
year build a platform of twigs and grass with perhaps a dozen nests on the same level,
next year a second platform with some new nests is built above the former one, and
so on year by year, each successive series of nests covering the previous one, until one
finds a stack higher than a man, of ten to fifteen years' growth. Often, on pulling
such a stack to pieces, deserted eggs of years ago may be discovered."

Larus dominicanus Licht. — Two skins are in the collection, an adult and an
immature (probably second year's bird), shot on May 19th on the shore at Saldanha
Bay, where these gulls were numerous. This species is very common all round the
coasts of South Africa, breeding on islands off the north-west and south coasts.

Lai-us hartlaubi (Bruch.).— Two adults and a young bird are in the Saldanha Bay
collection. This gull is very common along the west coast of the colony.

The young bird mentioned above has the quills and tail-feathers about half-grown
(the wing measures 6'8 ins., as against 1075 for the adult). It is remarkable for the
absence of any trace of a terminal dark band on the tail, which is pure white. A dark

* Fauna of S. Africa: Birds, by A. C. Stark and W. L. Sclater, vol. iv. p. 7 (the information regarding the status
in South Africa of the following species from Saldanha Bay is also taken from this source).
t The Voyage of the "Scotia," p. 277.

295

296 ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

tail-band is, I believe, almost universal in the young and immature stages of other
members of this genus. No mention of this peculiarity is to be found in the descriptions
of the species in the British Museum Catalogue of Birds, nor in the Fauna of South
Africa referred to above. The plumage of this specimen differs from that of the adult
in the presence of brown marks on the tips of the scapulars, median wing-coverts,
crown and ear-coverts, lead -coloured centres to the secondaries, and the smaller
amount of white on the three outer primaries (no white mirrors on Nos. 2 and 3, and
just a trace on No. 1).

Hasmatopus moquini Bp. One specimen is in the collection, shot at Saldanha Bay
on May 19th. It is an adult. The bill was noted as bright crimson, the legs darker
crimson.

The Black Oyster-catcher is common and resident along the shores of Cape Colony.

JEgialitis marginata (Vieil.). — Two specimens of the White-fronted Sand-plover
were shot on the shore at Saldanha Bay. This species is common and resident around
the South African coasts. It is stated to breed at Walvisch Bay in April or May.
These specimens (obtained on May 19th) have the sandy tint on the breast which
characterises the summer plumage.

Although a shore-bird, and apparently similar in habits to its black-and-white
banded congeners, this species has a plumage of a general fawn coloration, with black
reduced to a minimum on the head, 'so that from its appearance it might be taken
for an isabelline desert-form. (Sclater mentions that it has been stated by Layard
to occur on inland waters as well, but that he has never heard of its occurrence except
near the sea.)

Erythropygia coryphfeus (Less.). — A single example of this species, known as the
Cape Ground-robin, was obtained on the 19th. It is a common bird in many parts of
western Cape Colony.

Passer arcuatus Gray. — A male example of the Cape Sparrow was shot on the
19th. This sparrow has assumed similar habits to those of our European House-sparrow
in the towns, but is also found remote from human habitation in the Karroo and
elsewhere.

Serinus imberbis Cab. — A single specimen of this finch was also obtained.

Fringillaria impetuani (Smith). — This buuting, common locally throughout the
colony, is represented likewise by one specimen.

Algernon niv'osa (Swains.). — One specimen was obtained. This lark is found in
South Africa up to the Orange River, and is abundant in the Karroo.

In addition to the above, the following species were observed in South Africa :—

Ibis iethiopica (Lath.). — Numbers of which were seen at Reits Bay, Saldanha Bay.

Megalestris antarctica (Less.). — Skuas seen at sea on the day of arrival at Cape
Town, and also many on the 22nd May after leaving Saldanha Bay, were probably of this
species, which visits South Africa during the southern winter. The Arctic Skua (Ster-
corarius crepidatus) is only found on the South African coasts from October to March.

SOUTH AFRICA. 297

Sula capensis Layard. — These gannets were seen in plenty along the west coast,
and some on the 23rd May, after the Scotia had left for St Helena.

A single specimen of the Lesser Double-collared Sunbird, Cinnyiis chalybeus (L.),
is also in the collection, labelled " Cape Town." It is a species which is common over
the greater part of Cape Colony.

Two papers on Saldanha Bay and its Bird- Islands, by W. L. Sclater, have appeared
in Tlie Ibis (1896, p. 519, and 1904, p. 79).

Section XII. — St Helena, Ascension, and between.
By L. N. G. RAMSAY, M.A., B.Sc.

THERE is not mucli of importance to be recorded of this section of the Scotia's voyage.

A few days were spent at each of these two oceanic islands, and a few specimens of
their scanty avifaunas were obtained. During the voyage between, the tropical seas
were devoid of bird-life, except for a few gannets seen as Ascension was neared (the
h'rst were observed between one and two hundred miles from the island).

St Helena (lat. 16° S., long. 6° W.).— The Scotia touched here from May 30th to
June 2nd (1904). Specimens of three species of birds were obtained, viz. : Anous
stolidus (L.) (several adults), Gygi^ Candida (Gin.) (three), and Caccabis saxatilis,
subsp. cJivknr (Gray) (one). The last is a species of Red-legged Partridge introduced
into the island.

Ascension (lat. 8° S., long. 14° W.). — At this island the Scotia lay from 7th to
10th June.

One of the sights of the island is the " Wideawake Fair," or breeding-ground of the
" Wideawakes " (Sterna fuliginosa), millions of which are said to nest there. The birds
were absent at this time (although some were seen during the first day's sail after the
Scotia left the island), but great numbers of their corpses strewed the ground. A dozen
eggs of these terns were kindly presented by Mr Chalmers of the Eastern Telegraph Co.

On June 10th, while the Scotia was trawling off Pyramid Point before leaving,
Mr Wilton landed from a small boat on a rock near the shore and found gannets and
noddies busy in bringing up their young. The eggs were laid on the bare rock, no
nests being built. Some of the gannets had fresh-laid eggs, others had chicks, some of
which were big birds almost ready for flight. No eggs of the noddy were seen, but
some young birds unable to fly were caught. They were much infested with lice.

The specimens from Ascension include :—

Sula sula (L.).— Three skins, immature.

S. cyanops (Sund.). — Three skins (two adults and an immature bird in dark brown
plumage).

S. piscator (L.). — Two skins (one adult, one nearly so but with traces of brown in
the mantle).

Anous stohdus (L.). — Five specimens, including a naif-grown youngster.

Fregata aquila (L.). — Two skins (adults).

Sterna fuhginosa Gm. — Two carcases in spirit, much damaged.

YOJ,. IV. 299 13

Section XIII.— The North Atlantic.
By L. N. G. RAMSAY, M.A., B.Sc.

THIS, the final section, includes the observations of the voyage from Scotland
southward to St Paul's Rocks in November-December 1902, and the return
journey from Ascension via the Cape Verde Islands and Azores in June and
July 1904.

On the southward trip, very few birds were seen till Madeira was reached on
November 20th. A lark flew aboard about noon on the second day out, in 50° 45' N.,
7° 35' W. (about half-way between Land's End and the Irish coast), in an exhausted
state, and succumbed the same night.

Gulls were logged on the 16th, about 200 miles off the coast of Spain, and two
again on the 18th, still further from land.

After Madeira, petrels were logged almost every day till St Paul's Rocks were
reached, lying near the equator, but none were identified. Porto Grande, in the Cape
Verde Islands, was touched at on December 1st.

On the northward voyage in 1904, the Scotia, after passing through the Cape Verde
Islands on June 21st, steered much further to the westward, and was beyond the
34th meridian of west longitude for two days.

At this point the floating gulf-weed of the Sargasso Sea was passed through from
June 28th to July 2nd (chiefly in lat 29°-34° N.).

Between Ascension and Cape Verde Islands few birds except some Stormy Petrels
were seen. Two Sooty Terns (Sterna fuliginosa) were seen on June 12th, some
300 miles W.N.W. from Ascension. In the neighbourhood of the Cape Verde Islands
gannets were seen, and also several examples of what was probably the Tropic-bird
(Phaethon lepturus), and some boobies (?).

Between the 19th and 35th parallels of latitude only one bird (a small petrel) was
seen. After this the Azores were drawing near, and Stormy Petrels were seen daily till
the English Channel was reached. On 4th July the Scotia stopped to trawl on the
Princesse Alice bank, south of the Azores, and here nine Stormy Petrels (eight of these
which are now in existence are all Oceanites oceanicus) and five large shearwaters
were shot.

Four of the shearwaters I have examined. One of these is a Puffinm grams,
O'Reil., the other three are P. kithli (Boie). They are the Atlantic dark- winged form of
this species, in which the white on the inner webs of the primaries does not extend

301

302 ORNITHOLOGY OP THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

beyond the under wing-coverts (separated by Dr Hartert as P. kuhli flavirostris).
The dimensions of two of the latter are as follows :—

(a) Wing 13'4 ins. ; culmen 2'1 ins. ; tarsus 2'2 ins. ; mid-toe 27 ins.
(I) Wing 14-0 ins. ; culmen 21 ins. ; tarsus 2'2 ins. ; mid-toe 27 ins.

Shearwaters similar to these species were seen on July 7th to 10th, from the Azores
north-westwards to 43° N., 19° 30' W. A Black-backed Gull resembling Larus marinus
was seen on July 3rd and 4th.

Section XIV. — Bibliography of Antarctic Birds.

Only the main works dealing with the birds of the Antarctic regions, properly speaking, are included.
Preliminary notices are omitted in most cases.

(1) ANDBBSSON, K. A., "Das hohere Tierleben im antarktischen Gebiete," Wissen. Erg. Schwed. Sudpolar-

Exped. (1901-03), Bd. v., 2, Stockholm, 1905.

(2) BABIN, R., "Etude de la collection d'oiseaux de 1'antarctique rapportee par la mission Charcot (lre et

2" expeditions, 1903-05 et 1908-10)," Rev. franchise d'ornithologie, ii., 1911, pp. 113-117.

(3) BRUCE, W. S., "Antarctic Birds," KnowMge, xvii., 1894, pp. 208-210.

(4) - - " Animal Life observed during a Voyage to Antarctic Seas," Proc. Roy. Phys. Soc. Edin., xii.,

1894, pp. 350-354.

(5) "Some Ornithological Results of the Scot. Nat. Ant. Exp.," Ornis, xiv., 1905, pp. 271-274.

(6) CLARKE, W. E., " On the Birds of the South Orkney Islands," The Ibis, Series VIII., 6, 1906, pp. 145-187.

(7) "On the Birds of the Weddell and Adjacent Seas," The, Ibis, Series IX., 1, 1907, pp. 325-349.

(8) DONALD, C. W., " On the Penguins of the Antarctic Ocean," Report Brit. Ass. Adv. Set., 63, 1894, p. 808.
(9) " On the Penguins observed during the Sealing Voyage of the s.s. Active, in the neighbourhood

of Erebus and Terror Gulf," Proc. Roy. Soc. Edin., xx., 1895, pp. 170-176.

(10) GAIN, L., "Note sur les oiseaux des regions antarctiques," Bull. Mus. Paris, 1911, pp. 46-48.

(11) LONNBERG, EINAR, "Die A^bgel der Schwedischen Siidpolar-Expedition," Wissen. Erg. Schwed. Sud-

polar-Exped. (1901-03), Bd. v., Stockholm, 1905.

(12) MOSEI.EY, H. N., Notes of a Naturalist on H.M.S. " Challenger," London, 1872.

(13) MURRAY, JAS., Appendix to The Heart of the Antarctic, E. H. Shackleton, London, 1909.

(14) PEALB, TITUS R., Mammalia and Ornithology of the U.S. Exploring Expedition (1838-42),

Philadelphia, 1848.

(15) RACOVITZA, E., Resultats du voyage du s.y. " Belgica" (1897-99), Anvers, 1903.

(16) REICHENOW, ANT., "Vbgel des Weltmeeres," Deutsche Siidpolar-Ezped. (1901-03), Bd. ix., Zoologie,

I., Heft vi., Berlin, 1908.

(17) RICHARDSON, J., and J. E. GRAY, Zoology of Voyage of H.M.SS. "Erebus," and "Terror" (1839-43),

London, 1844-45.

(18) SAUNDERS, H., "Birds," in The Antarctic Manual, Royal Geogr. Soc., London, 1901.

(19) SCHALOW, H., "Ueber die Vogelfauna des Siidpolargebietes," Vortrag. Journ. fiir Ornithol., Leipzig,

1897, pp. 524-533.

(20) SCLATKR, P. L., Scientific Results of the Voyage of H.M.S. "Challenger," Zoology, II., part viii.

(Birds), 1881.

(21) - - " Birds of Antarctica," The Ibis, 1894, pp. 494-501.

(22) SHARPE, R. B., " Birds," Report on the " Soutliern Cross" Collections, London, 1902.

(23) TURQUET, E., "La vie animale an Pole Sud," in Le Fran$ais au Pole Sud, J. Charcot, Paris, 1906.

(24) VALBTTE, V. H., "Viaje 4 las Islas Orcadas Australes," Anales del Min. de Agric. Sect, de Zoologica,

iii. 2, Buenos Aires, 1906.

(25) VANHOFFEN, E., " Bericht iiber die bei der deutschen Siidpolarexpeditiou beobachteten Vbgel," Journ.

fur Ornith., July 1905, pp. 500-515.

(26) WILSON, E. A., (British) National Antarctic Exped., Natural History, ii., Birds, London, 1907.

(27) "On some Antarctic Birds," Ornis, xiv., 1905, pp. 231-234.

(28) WILTON, D. W., and others, " Zoological Log of the Scotia," Report on the Scientific Results of the

Voyage of S.Y. "Scotia" (1902-04), iv., 1, Edinburgh, 1908.

303

Index to Ornithology.

jEgialitis falklandica, 213.

jE. marginata, 296.

Alxmon nirosa, 296.

Anous dolidus, 207, 283, 299.

Anthus correndera, 213.

Aptenodytes forsteri, 223, 233, 256, 257, 258.

A. patagonica, 223.

Butt'o erythronotus, 2 1 2.

Caccaiis saxatilis, 299.
Catarrhades, 219, 276.
C. chrysoconie, 214.

C. chrysolophus, 221, 223, 231.
Cathartes, 212.

C/IIOJIM alia, 216, 217, 220, 221, 223, 244, 253,

256, 270.

Cltloephaga liylrida, 211.
Cinnyris chalyleus, 297.

(Jymodroma grallaria, 208, 275, 284, 293.

Daption capensis, 216, 217, 220, 221, 223, 238, 256,

257, 263, 275, 293.
Dio/nedea chionoptera, 291, 292.

D. exulans, 208, 215, 217, 223, 267, 275, 286, 291,

292.

D. melanophrys, 216, 272, 273, 274, 287, 291.
D. regia, 291.

Erythropyyia coryphaws, 296.
Eudromios modesta, 213.
Eudyptcs clirysocome, 289.

Fregata aquila, 299.

Fregeita melanogaster, 221, 223, 235.

Fringillaria impetuani, 296.

Gallinago paraguayix, 213.

Gannet, 301.

Gulls, 301.

Gygis Candida, 299.

Hxmatopus ater, 213.

H. leucopus, 213.

H. moquini, 296.

Halobsma cwrulea, 216, 217, 256, 257, 264, 275.

Ibis xthiopica, 296.

Lark, 301.

Lartw dominicunus, 213, 216, 220, 221, 223, 241,

253, 283, 295.
L. glaucodes, 212.
L. fiartlaubi, 295.
L. marinus, 302.
L. scoreslryi, 269.

Maj«queus xquinoctialis, 216, 223, 257, 276, 285,

293.
Megalestris antarctica, 213, 216, 220, 221, 223, 242,

251, 253, 256, 276, 283, 296.
M. maccormicki, 223, 244, 256, 257, 270.
Micranous leucocapillus, 207.
Milvago australis, 212.
" Molliemauk," 208, 215, 256, 267.
Muscisaxicola macloviana, 2 1 3.

Nesospiza jessiae, 278, 280.
N. goughensis, 278, 281.
Nycticorax tayazu-guira, 211.

Oceanites oceanicus, 208, 216, 217, 221, 223, 233,

256, 257, 260, 275, 284, 293, 301.
(Eslrelata brevirodris, 257, 262, 276.
(E. incerta, 208.
(E. lessani, 285.
(E. macroptera, 293.
(E. mollis, 276, 285, 293.
Ossifraga gigantea, 208, 214, 216, 217, 220, 221,

223, 237, 253, 256, 257, 263, 275, 285.

Pagodroma nivea, 216, 217, 221, 223, 236, 256, 257,

262.

Passer arcuattis, 296.
Pelecanoides, 276.
P. exsul, 223.
P. urinatrix, 286.
Penguin, 214, 217, 221, 249-253.
Petrel, 207, 209, 257, 276, 301.
Phaethon lepturus, 301.
Phalacrocorax albiventer, 212, 223.
P. atrieeps, 220, 221, 223, 246.
P. capensis, 295.

305

306 ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

Phalacrocorax magellanicus, 212.

Phcebetria, 208.

P. cornicoide*, 216, 256, 257, 266, 27.1-272, 288,

291.

P. fuliginosa, 216, 223, 266, 271-272, 288, 291.
Phrygilus melanoderus, 214.
Porphyriornis coineri, 278, 279, 282.
Priocella glacialoides, 216, 221, 236, 256, 257, 261,

275.

Priqfinus cinereus, 276, 285, 292.
Prion, 208, 216, 275, 293.
P. banksi, 216, 240, 255, 256, 257, 265.
P. desolatus, 223, 265, 286.
P. vittatus, 275, 286.
Pufflnus assimilis, 284.
P. gravis, 301.
P. kuhli, 301.
Pygoscelis antarctica, 216, 219, 220, 221, 223, 250,

252, 256, 259.
P. adelix, 216, 221, 223, 227, 250, 251, 252, 256,

257, 279.
P. papua, 221, 223, 230, 250, 251, 252.

Sennits imberlis, 296.

Spheniscus demersus, 295.

S. magellanieus, 214, 252, 276.

Sterna fuliginosa, 299, 301.

S. hintndinacea, 213, 216, 221, 223, 241, 268.

S. macrura, 223, 257, 258, 268.

S. vittata, 223, 276, 283.

Sula capensis, 297.

S. cyanops, 207, 299.

S. piscator, 207, 299.

S. sula, 207, 299.

Tachyeres cinereus, 211.

Tern, 217, 256.

Thalasaoeca antarctica, 223, 235, 256, 260.

Thalassogeron carferi, 274, 287.

T. cldororhynchus, 215, 273, 275.

T. culminatus, 215, 273, 275.

T. eximius, 279, 287.

Trimja fuscicollis, 213.

T-rupialis militarin, 213.

Turdus faUdandicus, 213.

PRINTRD BY NKILL AND CO., LTD., EDINBUROH.

ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

EXPLANATION OF PLATES.

PLATE I.

The young of the Ringed Penguin (Pygoscelis antardica), showing the two stages in down. Painted by
H. Goodchild.

PLATE II.

Fig. 1. The young of the Snowy Petrel (Patjodroma nivea).
Fig. 2. The young of the Sheathbill (Chionis alba).

Painted by H. Goodchild.

PLATE III.

Fig. 1. Wilson Petrel (Oceanites oceanicus) on nest, with egg.
Fig. 2. Cape Pigeons (Daption capensis) preparing- to nest.
Fig. 3. Young of the Antarctic Skua (Megalestris antardica).
Fig. 4. Nest and eggs of the Southern Black-backed Gull (Larus dominicanus).

Photographs by J. H. H. Pirie.

PLATE IV.

Fig. 1. Tracks of Black-throated Penguin (Pygoscelis adelise). Photo by J. H. H. Pirie.
Fig. 2. Black-throated Penguin building its nest (P. adeiite).
Fig. 3. Gentoo Penguin (P. papua) sitting tight on its eggs after a blizzard.
Fig. 4. Ringed Penguins (P. antardica) at th'eir rookery on Saddle Island.

Photographs by R. N. R. Brown.

PLATE V.

Two new species of Buntings taken at Gough Island.
Fig. 1. Nesospizajessix.
Fig. 2. N. goughenti*.

Painted by H. Goodchild.

PLATE VI.
Eggs of Antarctic birds in natural colour, and reduced to four-fifths their natural size.

Fig. 1. 3fegalestri.< antarctita, Antarctic Skua, taken at the South Orkneys on ] 1th December 1903:
Scotia collections. Egg number 4 : length = 76 mm., weight = 1720 grains. Vide original Log, p. 35.

Fig. 2. M. antardica, taken at the South Orkneys on 2nd December 1903 ; Scotia collections. Egg
number la: length = 74 mm., weight = 1524'7 grains.

Fig. 3. M. maccormitlii. McCormick's Skua, taken in Victoria Land by Shackletou Expedition,
1907-09, per Dr Forbes Mackay. Length = 78 mm.

Fig. 4. Larus dominicanus, Dominican Gull or Southern Black-backed Gull, taken at South Orkneys:
Scotia collections. Length = 70 mm., average weight = 1519'3 grains.

Fig. 5. L. dominicanus, taken at the South Orkneys; Scotia collections. Actual length =71 in in.,
average weight = 1519'3 grains.

Fig. 6. M. maccormicki, taken in Victoria Land by Shackleton Expedition, 1907—09 ; per Dr A.
Forbes Mackay.

Fig. 7. Chionis alba, the Sheathbill or Paddy, taken at the South Orkneys; Scotia collections.
Length = 56 mm.

Fig. 8. C. alba, taken at South Orkneys on 18th December 1903 ; Scotia collections. Length = 55 mm.,
weight = 615 grain.?. Vide Log, p. 36.

Fig. 9. Sterna hirundinacea, the White-rumped Tern, taken at South Orkneys on 12th December 1903;
Scotia collections.

The numbers 1 to 9 read from left to right in each row.

ORNITHOLOGY OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

PLATE VII.

Eggs of Antarctic birds which are either white, pale blue, or, as in the case of Wilson's Petrel, only
faintly spotted. The plate shows the relative size or shape of these eggs, reduced to four-
fifths their natural size.

Fig. 1. Pytjoseelis antarctica, the Ringed Penguin, or Antarctic Penguin, taken at South Orkneys, 1903 ;
Scotia collections. Egg number M. 2 : length = 68 mm., weight =1880 grains. Vide original Log, p. 53.

Fig. 2. P. antarctica, taken at South Orkneys, 1903; Scotia collections. Egg number 11 : length =
68 mm., weight = 1530 grains.

Fig. 3. P. adelix, Black-throated Penguins, taken at South Orkneys, 1903 ; Scotia collections. Length =
67 '8 mm., weight = 1530 grains.

Fig. 4. Phalacrocorax atriceps, the Blue-eyed Shag, taken at Shag Rocks, Brown's Bay, on 9th November
1903 ; Scotia collections. Length = 67 mm., average weight = S64'5 grains.

Fig. 5. Ossifraga gigantea, the Giant Petrel, Nelly, or Stinker, taken at Cape Geddes, South Orkneys ;
Scotia collections. Length = 105 mm., average weight = 3784 grains. Vide Camp Log — W. S. B.

Fig. 6. Pygoscelis papua, the Gentoo Penguin, taken at Scotia Hay, South Orkneys ; Scotia collections.
Length = 66 mm., average weight = 4'41 ounces avoirdupois.

Fig. 7. Daption capensis, the Cape Penguin or Petrel, taken west of Jessde Bay, South Orkneys, on
4th December 1903 ; Scotia collections. Egg number 14 : length 63 mm., weight 876 grains. Vide original
Log, p. 31.

Fig. 8. Oceanites oceanicus, Wilson's Petrel, taken at Churchill, Jessie Bay, South Orkneys, on
4th January 1904 ; Scotia collections. Egg number 3 : length = 33 mm., weight = 155 grains. Vide original
Log, p. 47.

Fig. 9. Pagodroma nivea, the Snowy Petrel, taken on west side of Jessie Bay, South Orkneys, on
5th December 1903. Egg number 26 : length = 55 mm., weight = 685 grains. Vide original Log, p. 29.

The numbers 1 to 9 read from left to right in each row.

MAP I.

Scotland to Coats Land : Track Chart of the Scotia, 1902-04, by William S. Bruce, LL.D.

MAP II.

Weddell Sea: Track Chart, including the trawling stations of the Scotia, 1902-04, by William S.
Bruce, LL.D.

SCOT. NAT. ANT. EXP.

Ornithology of the " Scotia." Plate I.

VOL. IV.

H. Goodchilil, >J.!.]

The young of the Ringed Penguin

{Olirer ,t Logd, Ed'tH., imp.
is iinlarcttfa). showing the two stages in down.

SCOT. NAT. ANT. EXP.

Ornithology of the " Scotia." Plate II.

VOL. IV.

i.

2.

//. GooilchiM, del.] [Olirtr <t Bojrf, £cfin., imp.

1. The young of the Snowy Petrel (Pai/odroma nitta). 2. The young of the Sheathbill (Chionix iillia).

SCOT. NAT. ANT. EXP.

Ornithology of the " Scotia." Plate III.

VOL. IV.

[Photo by J. H. H. Pirie.
1. Wilson Petrel (Oenanttias oeeanicus) on Nest with Egg.

[Photo by J. H. H. Pirie.

2. Cape Pigeons (Daption captnsis) preparing to nest.

[Photo by J. H. H. Pirie.
3. Young of Antarctic Skua (Megalestris anlarctira).

[Photo by J. H. H. Pint.
4. Nest and Eggs of Southern Black-backed Gull (Larvs dominicanus).

SCOT. NAT. ANT. EXP.

Ornithology of the " Scotia." Plate IV.

VOL. IV.

[Pkvtv by J. H. H. Pine.
1. Tracks of Black-throated Penguin (Pygoscelis adtlia).

by R. X. R

2. Black-throated Penguin building its nest.

3. Gentoo Penguin (Pygoscelis papua) sitting tight on its eggs
after a blizzard.

4. Ringed Penguias (Pyqoscelis aiUarctica) at their rookery
(on 'Saddle Island).

SCOT. NAT. ANT. EXP.

Ornithology of the " Scotia." Plate V.

VOL. IV.

H. Gooddiihl, del.]

[Olittr <C Doyd, imp.

Two new species of Buntings, taken at Gough Island.
1. Nesospiza jessue. 2. JV. goiuflienttis.

SCOT. NAT. ANT. EXP. Ornithology of the "Scotia." Plate VI. VOL. IV.

*

'I » , .

v

- ^

1

i . Megalestris ant arc tic a.
4. Larus dom'uricanus.
7. C hion is alba.

Reduced + natural size.

2. M. an tare tic a.

5. Larus dominkanus.

8. Chionis alba.

[Oliver £ B«yd, Bdin., imp.

3. M. maccormicki.
6. M. maccormicki.
9. Sterna hirundinacea.

SCOT. NAT. ANT. EXP. Ornithology of the "Scotia." Plate VII. VOL. IV.

i. Pygoscelis antarctica.
4. Phalacrocorax atriceps.
7. Daption capensis.

Reduced i natural size.

2. P. antarctica.

5. Ossifraga gigantea.

8. Oceanites oceanicus.

[Oliver it Bvyd, Edln., imji.

3. P. adelitz.
6. P. papua.
9. Pagodroma nrvea.

SCOT. NAT. ANT. EXP.

MAP I

Scotland to Coats £and

TRACK CHART OF THE "SCOTIA," 1902-1904, BY WILLIAM S. BRUCE, LL.D.

Jftn-of O Grvenwick 15

MAP II.

PART XV.
FISHES.

XV. -ANTARCTIC FISHES OF SCOTTISH NATIONAL

ANTARCTIC EXPEDITION.

BY C. TATE REGAN, M.A.,

Assistant Keeper in Zoology in the British Museum.

(WITH ELEVEN PLATES.)

The Antarctic Pishes of the Scottish National Antarctic Expedition. By
C. Tate Regan, M.A., Assistant in the British Museum (Natural History). Com-
municated by Dr W. S. BRUCE. (With Eleven Plates. and Six Text-figs.)

(MS. received June 18, 1912. Read December 16, 1912. Issued separately May 23, 1913.)

Our knowledge of the Antarctic fish-fauna has greatly increased during the last
ten years. The Belgian expedition to Graham Land (1897-1899) was followed by
that of the Southern Cross to Arictoria Land (1898-1900), fitted out by Sir GEORGE
NEWNES. Next were the British expedition of the Discovery to Victoria Land and
Edward Land (1901-1904), the German voyage of the Gauss to Kerguelen and Wilhelm
Land (1901-1903), and NORDENSKJOLD'S Swedish expedition to South Georgia, the South
Shetlands, and Graham Land. Then came the voyage of the Scotia to the South
Orkneys and Coats Land (1902-1904), and CHARCOT'S expeditions to the Palmer
Archipelago and Graham Land in the Franccds (1904-1905) and the Pourquoi Pas?
(1908-1910), and finally SHACKLE-TON'S expedition (1908-1909).

The fishes collected during these expeditions have been described in a series of
reports, which may be enumerated in chronological order : —

1902. BOULENGER, Pisces in "Southern Cross" Collections, pp. 174-189, pis. xi.-xviii.

1904. DOLLO, Res. Voy. " Belgica" : Poissons, 240 pp., 12 pis.

1905. LONNBERG, "The Fishes of the Swedish South Polar Expedition," Wissensch. Ergebn.

Schwedisch. Sudpolar-Exped., v. 6, 69 pp., 5 pis.

1906. VAILLAXT, Exped. Antarct. Fran$aise: Poissons, 51 pp.

1907. BOULEXGER, National Antarctic Expedition, Nat. Hist.: II., Fishes, 5 pp., 2 pis.

1911. WAITE, " Antarctic Fishes," in British Antarctic Expedition, 1907-9: Bi ology, pp. 11-16,

pi. ii.

1912. PAPPENHKIM, "Die Fische der Antarktis und Subantarktis," in DeutseJie Siidpolar-Exped.,

1901-1903 : XIIL, Zool., v. pp. 163-182, pis. ix.-x.

Dr DOLLO presented several preliminary notes in the Proceedings of the Royal Society
of Edinburgh * on the fishes of the Scottish National Antarctic Expedition.

The fishes of the second Charcot expedition have been worked out by Professor
ROULE, who has published two preliminary notes (C.R. Acad. Sci. Paris, cliii., 1911, pp.
80-81, &ndBull. Mus.Paris, 1911, pp. 276-28 l),but the final report has not yet appeared.

The important collection of fishes here reported on was made at the Falkland Islands,
the South Orkneys, Coats Land, and Gough Island, and in the Weddell Sea and South
Atlantic Ocean between these localities. As will be seen from the systematic list that
follows, it includes examples of forty-eight species, ten of which are now described as
new to science, whilst three others, known before but wrongly identified, are diagnosed
and given new specific names ; in addition, four species have already been described by

* Proc. Roy. Soc. Edin , xxvi., 1906, p. 172 ; xsviii., 1908, p. 58 ; xxix., 1909, p. 316.
(REPRINTED FROM THE TRANSACTIONS OF THE ROYAL SOCIET? OF EDINBURGH, VOL. xi.ix., pp. 229-292.)

312

MR C. TATE REGAN ON THE

Dr. DOLLO, in whose hands the greater part of the collection has been from 1905 until
March 1912.

The identification of the Notothenioids and Zoarcids has proved a difficult matter in
the present state of our knowledge of these groups, and I have supplemented my report
by a monograph of the former and a revision of the southern genera of the latter ;
further, I have added some notes on the Galaxiidae and Haplochitonidse, as their distri-
bution has given rise to some discussion.

My work on the Notothenioids and Zoarcids is mainly based on the specimens in the
British Museum, including the Erebus and Terror, Challenger, Southern Cross and
Discovery collections, but I have been greatly helped by the loan of specimens from
the Museums at Paris, Berlin, and Stockholm. Thus I have been able to examine all
the species of Notothenia recorded by VAILLANT from Graham Land, two of the three
species of Zoarcids recently described by PAPPENHKIM from Wilhelm Land, and
co-types of some of the Notothenioids described by LONNBERG. For their kindness
in sending me these fishes, and in giving me information about others that could not
be sent, I heartily thank Dr PELLEGKIN, Dr PAPPENHEIM, and Dr LONNBERG.

It need hardly be said that the fishes lend no support to the theory of bipolarity.
Most of the littoral fishes belong to the Nototheniidse and related families, which are
characteristic of and peculiar to the Antarctic seas and the region immediately to the
north of them ; there are also several species of Zoarcidse, generically distinct from
the northern members of the family. Some of the pelagic and abyssal fishes are
Notothenioids peculiar to the Antarctic region ; others also, such as Notolepis, Cynoma-
crurus, and Eugnathosaurus, may not be found elsewhere ; but the rest belong to
widely distributed genera (Synaphobranchus, Bathylagus, Myctophum, etc.) or even
species (e.g. Cyclothone microdon).

In the whole paper the following seven new genera and twenty-one new species
are described : —

Grossolycus, p. 329.
Payetopsisf, p. 368.
Ghxnocepliahts, p. 369.

Bovichthys angustifrons, p. 337.

,, chilensis, p. 338.

,, decipiens, p. 339.
Trematomus loennberyii, p. 345.
Notothenia trif/ramma, p. 348.

„ ramsayi, p. 349.

,, wiltoni, p. 350.

,, vaillanti, p. 354..
ChxnichtJiys rugosus, p. 369
Cryodraco pappeitheimi, p. 370.

XLIX., 230.)

GENKRA.

Eugnathosaurus, p. 316.
Ophthalmolycus, p. 325.
Ausfrolycichthys, p. 326.
Austrolycus, p. 327.

NEW SPECIES.

Bathylagus glacialts, p. 313.
Eugnathosaurus vorax, p. 316.
Synaphobranchus australis, p. 317.
ChaJinura ferrieri, p. 318.
,, whitsoni, p. 318.
Csesioperca coafsii, p. 319.
Neojjhrynichthys marmoratua, p. 323.
Lycenchelys antardicus, p. 324.
Austrolycus depressiceps, p. 327.
Crossolyeun chilensis, p. 329.
Ooitoperca macrophthalma, p. 335.

(ROY. soc. EDIN. TRANS., VOL.

ANTARCTIC FISHES OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 313

I. ANTARCTIC AND SUBANTARCTIC FISHES COLLECTED BY THE " SCOTIA."*

SELACHII.

RAID.E.

I. Raia magellanica, Steind. (PI. I.)
Zool. Jahrb. Suppl., vi., 1905, p. 212.

One specimen from Station 346, Burdwood Bank, depth 56 fathoms; taken on
1st December 1905. Ut, 54° 25' S., long. 57° 32' W. ; temperature 41'8° F.

This is a female of exactly the same size as STEINDACHNER'S type, and apparently in
every way similar, except that there is only a single scapulary spine, instead of a series
of three on each side.

This species is related to R. murrayi, Giinth., from Kerguelen, but has a blunter
snout, a shorter tail, and somewhat different spination.

ISOSPONDYLI.

CLUPEIDJS.

'2. Clupea fuegensis, Jenyns.

Zool. "Beayl*," Pith., p. 133(1842); Smitt, Bihang, Svensk. Vet.-Akad., xxiv., 1898, iv., No. 5,
p. 59, pi. v. fig. 41

Depth of body 4 to 5 in the length. Lower jaw very prominent ; minute teeth in
a single series on the palatines and in an elongate patch on the tongue. Dorsal 17-18 ;
origin equidistant from anterior edge of eye and base of caudal fin. Anal 17-20.
Origin of pelvics vertically below that of dorsal. About 50 scales in a longitudinal
series ; ventral scutes not prominent.

Several specimens, up to 170 mm. in total length, taken at Station 118, Port
Stanley, Falkland Islands, in February 1904, when extraordinary shoals of this herring
visited Port Stanley Harbour.

ARGENTINID^;.
3. Bathylayus glacialis, sp. n. (PI. IX. fig. 2.)

-

Depth of body 6 to 61 in the length, length of head 4| to 4i. Diameter of eye
2i to 2| in the length of head, interocular width 3, interorbital width 6. Dorsal 10 ;
origin nearer to end of snout than to base of caudal. Anal 18. Pelvics 8-rayed,
inserted below middle of dorsal. About 35 scales in a longitudinal series.

* A series of nine water-colour drawings made by Mr COTHBERTSON for the most part represent fishes from Scotia
Bay, South Orkneys, viz. Notolepis coatsii, Harpagifer bispinis, Trematomus newnesii, yotothenia coriiceps, N. nvdifrons,
and N. gibberifrons : there is also a sketch of Lycenchelys antarcticus. In one or two cases I have referred to these
in the text.

(BOY. soc. EDIN. TRANS., VOL. xux., 231.)

314 MR C. TATE REGAN ON THE

There are five examples of this new species, 70 to 100 mm. in total length :—

1. Station 398, 68° 25' S., 27° 10' W., 1 to 1000 fathoms; surface temperature

30° F. ; vertical net ; 29th February 1904.

2. Station 422; 68° 32' S., 12° 49' W., 0 to 800 fathoms; surface temperature

31-1° F. ; temperature at 800 fathoms 32'4° F. ; vertical net ; 23rd March
1904.

3. Station 414, 71° 50' S., 23° 30' W., 0 to 1000 fathoms; surface temperature

29'1° F. ; vertical net; 15th March 1904.

4. Station 417, 71° 22' S., 16° 34' W., 1410 fathoms; temperature at 1410

fathoms 3T90 F. ; trawl; 18th March 1904.

5. Station 418, 71° 32' S., 17° 15' W., 1221 fathoms; temperature at 1221

fathoms 31'9° F. ; trawl ; 19th March 1904.

Bathylagus antarcticus, Giinth., is distinguished by the less graceful form (depth
5 in the length) and the longer anal fin with 22 rays. Bathylagus gracilis, another
Antarctic species recently described by LONNBERG, has the interorbital space very
narrow and deeply concave, and about 41 scales in a longitudinal series.

Other species of Bathylagus have been described from the South Atlantic (GtJNTHER,
LONNBERG), the North Atlantic (GooDE and BEAN), and the North Pacific (GILBERT).

GALAXIID^E.

4. Galaxias attenuatus, Jenyns.

Two examples from Port Stanley and Port Harriet, Falkland Islands, Station 118.

5. Galaxias maculatus, Jenyns.
Several from Port Harriet, Station 118.

HAPLOCHITONID^;.
6. Haplochiton zebra, Jenyns.
One specimen from Port Stanley, Falkland Islands, in fresh water, Station 118.

STOMIATID.E.
7. Stomias boa, Bisso.

One from Station 451, 48° 06' S., 10° 5' W., 1742 fathoms; trawl; 13th April
1904.

8. Cyclothone microdon, Giinth.

Small examples of this widely distributed species were taken at three stations, viz.—
Six at Station 450, 48° 00' S., 9° 50' W., 1332 fathoms; surface temperature

40-0° F. ; trawl; 13th April 1904.
One at Station 422, 68° 32' S., 12° 49' W., 0-800 fathoms ; temperature at 800

fathoms 32 '4° F. ; vertical net; 23rd March 1904.

Four at Station 414, 71° 50' S., 23° 30' W., 0-1000 fathoms; surface tempera-
ture 29-1° F. ; vertical net; 15th March 1904.
(ROY. soc. EDIN. TRANS., VOL. XLIX., 232.)

ANTARCTIC FISHES OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 315

INIOMI.

StJDIDJS.

9. Notolepis coatsii, Dollo.

Proc. Roy. Soe. Edin., xxviii., 1908, p. 58.

Prymnothonus (part.), Giinth., "Challenger" Pelagic Fish, p. 39, pi. v. fig. D (1889).

„ Hookeri (non Richards.), Dollo, Proc. Roy. Soc. Edin., xxvii., 1907, p. 35.

Depth of body 6^ in the length, length of head 5 ; snout half the length of head ;
diameter of eye 6^ in the length of head. Teeth rather small, pointed, uniserial, in
jaws and on palatines. Dorsal 8 ; origin nearly equidistant from head and base of
caudal ; adipose fin rather long and low. Anal 28. Caudal with numerous procurrent rays.
Pectorals narrow, about ^ length of head. Vent below anterior part of dorsal. Scales
deciduous. Myotomes 82, 34 in advance of dorsal fin. Silvery white ; back bluish.

It is with some difficulty that I have put together the above description of the type of
the species, 105 mm. in total length, taken at the surface in Scotia Bay, South Orkneys.
The specimen is in very bad condition,* and everything that one touches falls off;
hence it is not surprising that I cannot see the small pelvic fins described by DOLLO.

In a paper on the classification of the Iniomi (Ann. Mag. Nat. Hist. (8), vii., 1911,
pp. 120-133) I have already called attention to the fact that DOLLO'S family Paralepidae
is not a natural group, and that Notolepis differs from Parolepis apparently only in
the greater length of the adipose fin, a character of very slight importance to anyone
familiar with the species of Siluroids.

Larval and post-larval examples of this species that I have examined are :—
1.— 44 mm. ; 62° 26' S., 95° 44' E. Challenger collection.
2.— 50 mm. ; at Station 422, 68° 32' S., 12° 49' W., 10-800 fathoms ; tempera-
ture at 800 fathoms 32'4°F.; 23rd March 1904. Scotia collection.
3-5.— 38 to 56 mm. ; at Station 414, 71° 50' S., 33° 30' W., 0-1000 fathoms;

surface temperature 29 '1° F. ; 15th March 1904. Scotia collection.
Except that the teeth are relatively stronger and the eye larger, specimens 1 and 2
are extremely similar to the type, and agree with it in the number of fin-rays and
of myotomes ; I cannot find any pelvic fins, nor ascertain the position of the vent, but
the eight-rayed dorsal fin is distinct in both.

Specimens 3 to 5 are the ones described by DOLLO as Prymnothonus hookeri ; these
evidently belong to the same species as the other examples, with which the larger one
agrees in the head, dentition, and approximate number of myotomes. In the smaller
ones the head is relatively smaller and the snout shorter. I am unable to make out
the fins, or position of vent, and I am very doubtful as to whether the so-called
embryonic anal fringe is an actual structure present in the living fish.

Dr DOLLO named this species in honour of the late Mr JAMES COATS, junr., of Paisley,
whose generosity was the chief means of assuring the dispatch of the Scottish National
Antarctic Expedition.

"• This is regrettable, as this specimen was originally so perfectly preserved and was brought home in perfect
condition, and was acknowledged to have been received by Dr DOLLO "en bon elat." — W. S. B., Editor.

(ROY. 800. KDIN. TRANS., VOL. XLIX., 233.)

316 MR C. TATE REGAN ON THE

MYCTOPHID.E.

10. Myctophum antarcticum, Giinth.
Specimens were taken at :—

Station 309, 63° 51' S., 41° 50' W., 2300 fathoms; temperature 31-05° F. ;

trawl; 16th March 1903.
Station 414, 71° 50' S., 23° 30' W., 0-1000 fathoms; surface temperature

29-1° F. ; vertical net; 15th March 1904.
Station 422, 68° 32' S., 12° 49' W., 0-800 fathoms; surface temperature

327° F. ; temperature at 800 fathoms 32'4°; vertical net ; 23rd March 1904.

1 1 . Lampanyctus braueri, Lcinnberg.

One specimen was taken at Station 420, lat. 69° 33' S., 15° 19' W., 2620 fathoms,
by the trawl, on 21st March 1904 ; temperature 31'5° F. The species was previously
known only from the type.

ALEPIDOSAURID^:.

Eugnathosaurus, gen. nov.

Skull very elongate and strongly compressed, with the upper surface somewhat
convex, bearing a fairly prominent median ridge. Snout and lower jaw much produced,
each ending in a fleshy appendage ; lower jaw projecting beyond upper ; suspensorium
directed obliquely forward. Teeth pointed, uniserial ; praemaxillary teeth minute ;
mandibulary teeth sub-conical, erect or somewhat retrorse, strongest in the middle of
the length of the jaw, more spaced posteriorly ; palatine teeth strong, compressed,
curved somewhat forward.

12. Eugnathosaurus vorax, sp. n.

The type of this remarkable new genus and species is a head, measuring 150 mm.
in length from tip of snout to end of operculum, taken in the trawl on 18th March 1904,

FIG. 1. — Ewgnathosaunts vorax.

at Station 417, in lat. 71° 22' S., long. 16° 34' W., off Coats Land, at a depth of 1410
fathoms, by the trawl ; temp. 31 '9° F. That it is related to Alepidosaurits is evident,

(ROY. soc. EDIN. TRANS., VOL. XLIX., 234.)

ANTARCTIC FISHES OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 317

but the form of the skull, the produced jaws, and the different mandibulary and palatine
dentition distinguish it from that genus ; the antrorse palatine teeth are especially
peculiar.

The dentaries of a second specimen were taken at the same locality.

APODES.

SYNAPHOBRANCHID.E.

13. Synaphobranchus australis, sp. n. (PI. VIII. fig. 5.)
Synaphobranchus bathybius (part.), Giinth., "Challenger" Deep-Sta Fish, p. 254 (1887).

The Challenger specimen, 350 mm. in total length, was taken midway between the
Cape of Good Hope and Kerguelen, at a depth of 1375 fathoms, The Scotia example
was obtained on 13th April 1904, at Station 451, in 48° 06' S., 10° 5' W., at a depth
of 1742 fathoms, and measures a total length of 470 mm. The species belongs to
the sub-genus Histiobranchns, Gill, which includes also S. bathybius, Giinth., and
S. mfernalis, Gill. All three are closely related, differing as follows :—

Eye nearer to end of snout than to angle of mouth ; origin of dorsal above base of
pectoral, its distance from end of snout rather less than ^ that from end of
snout to vent . . . . . . . . . • bathybius.

Eye about equidistant from snout and angle of mouth ; origin of dorsal above
posterior part of pectoral, its distance from end of snout somewhat more than
^ that from end of snout to vent ...... infemalis.

Eye about equidistant from snout and angle of mouth ; origin of dorsal a little
behind end of pectoral, its distance from end of snout about 2j in that from
end of snout to vent australis.

ANACANTHINI.

MACRURID.E.

Four species of this family were obtained by the Scotia in Antarctic seas, all belong-
incc to the sub-family Macrurinfe (cf. Ann. Mag. Nat. Hist. (7), xi., 1903, pp. 45S-466),
and to genera with the teeth in the lower jaw uniserial.

14. Nematonurus lecointei, Dollo.
Res. Voy. " Belgica," Poiss., p. 44, pi. vii. (1904); Proc. Roy. Soc. Edin., xxix., 1909, p. 488.

The type was taken in 70° 40' S., 102° 15' W., depth 1526 fathoms. The Scotia
examples are from : (1) Station 313, 62° 10' S., 41° 20' W., 1775 fathoms; temperature
31-0° F. ; trawl; 18th March 1903. (2) Station 451, 48° 06' S., 10° 05' W., 1742
fathoms; 13th April 1904.

(ROY. soc. EDIN. TRASS., VOL. XLIX., 235.)

318 MR C. TATE REGAN ON THE

The prsemaxilLary teeth are biserial, except posteriorly, where the inner series is
replaced by three, forming a narrow band.

15. Chalinura ferrieri, sp. n. (PL II. fig. 1.)

Snout rather strongly produced (for a Chalinura) ; mouth wide, the maxillary
nearly reaching the vertical from posterior edge of eye ; infraorbital ridge fairly
prominent. Diameter of eye less than length of snout, 4f in length of head ; inter-
orbital width 4. Dorsal II 9 ; distance from second dorsal a little more than f the
length of head. Origin of anal at distance from head equal to length of head without
snout. Pectoral 18 or 19-rayed, f the length of head, extending to above origin of
anal. Pelvics 11 -rayed, the outermost ray filamentous, reaching anal. Scales mostly
with 3 parallel series of spinules, but the lateral series sometimes reduced to a single
spine, or absent ; 8 scales between dorsal fin and lateral line.

A. single specimen, 230 mm. in total length, from Station 417, 71° 22' S., 16° 34' W.,
1410 fathoms, off Coats Land; temperature at 1400 fathoms 31'9°F. ; trawl; 18th
March 1904.

This species is named after JAMES G. FERRIER, Esq., F.R.S.G.S., Hon. Secretary of
the Scotia Committee.

16. Chalinura whitsoni, sp. n. (PL II. fig. 2.)

Snout rather produced (for a Chalinura) ; maxillary extending to below posterior
margin of pupil ; infraorbital ridge prominent. Diameter of eye more than length of
snout, 2| to 3^ iii length of head; interorbital width 4 to 4^. Dorsal II 9-10; dis-
tance from second dorsal 5 the length of head. Origin of anal at distance from head
equal to length of head without snout. Pectoral 18-19-rayed. Pelvic 9-rayed, the
outermost ray filamentous, not reaching anal. Scales with 1 series of spinules, but
some on sides of head with 3 series converging anteriorly ; 7 scales between dorsal fin
and lateral line.

Two specimens :—

1. 420 mm.; Station 451, 48° 6' S., 10° 5' W., 1742 fathoms; trawl; 13th

April 1904.

2. 270 mm.; Station 417, 71° 22' S., 16° 34' W., 1410 fathoms, off Coats Land ;

temperature at 1400 fathoms 3r9° F. ; trawl; L8th March 1904.

This species is named after T. B. WHITSON, Esq., C.A., Hon. Accountant of the
Scotia Committee.

17. Cynomacrurus piriei, Dollo. (PL III. fig. 1.)

Proc. Roy. Soc. Edin., xxix., 1909, p. 316.

The type of the genus and species, a specimen of 300 mm., was obtained by the
Scotia at Station 414, 7 1° 50' S., 23° 30' W., in a depth 0-1000 fathoms, surface tempera-
ture 31 '5° F., vertical net, on 15th March 1904. The dentition is very characteristic;
in the prsemaxillaries a narrow band of unequal teeth separated by an interspace from a

(ROY. soc. EDIN. TRANS., VOL. XLIX., 236.)

ANTARCTIC FISHES OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 319

marginal series of small teeth with a strong pair of antero-lateral canines ; in the lower
jaw teeth strong, spaced, unequal,- uniserial.

Other important characters are the large terminal mouth with lateral cleft, the
absence of a barbel, the small eye, and the slender, smooth dorsal spine. The pelvic fins
are 7-rayed, and in counting 12 Dr DOLLO must have reckoned divided rays as two.

This species was named by Dr DOLLO after Dr J. H. HARVEY PIRIE, bacteriologist,
geologist, and surgeon of the Scotia.

PERCOMORPHI.

SERB AN ID JR.
18. C&sioperca coatsii, sp. n. (PL VI. fig. 1.)

Depth of body 2f to 3j in the length, length of head 2f to 3. Diameter of eye 2^
to 2| in length of head, interorbital width 4^ to 5. Interorbital region flat ; maxillary
extending to below middle of eye ; 21 to 24 gill-rakers on lower part of anterior arch.
Dorsal X, 15-18 ; third or fourth spine longest, nearly twice as long as last, f to J
length of head. Anal III 8 ; second spine longest, as long as or longer than longest
dorsal spine. Pectoral shorter than head, asymmetrical, the rays increasing to the
tenth, counting from above, or seventh, from below. Caudal truncate. About 40 scales
in a lateral longitudinal series and 45 in the lateral line, which forms an angle on the
caudal peduncle. Pale reddish brown, with traces of alternating darker and paler longi-
tudinal bands ; upper half of spinous dorsal blackish, or with a series of blackish spots.

Gough Island. Several specimens, up to 135 mm. in total length, taken at Station
461, 40° 20' S., 9° 56' W., off Gough Island, at a depth of 100 fathoms; surface
temperature 54'5° F.; trawl; 23rd April 1904.

This species is of considerable interest, as its three congeners are found on the coasts
of South Australia, Tasmania, and New Zealand. These are distinguished by their
longer pectoral and emarginate caudal fins, and by the convex interorbital region ; but
I am unable to find any characters which would justify the establishment of a new
genus for the new species. The pectoral fin of C. rasor is almost as asymmetrical, and
1 find that the flatness of the interorbital region of C. coatsii is not associated with any
difference in the essential structure of the frontal bones, which are, as in C. lepidofjtera,
smooth and convex posteriorly, and anteriorly consist of a pair of supraorbital flanges
and of a median depression bordered by muciferous canals.

I have pleasure in naming this species after Major ANDREW COATS, D.S.O., a
member of the Scotia Committee, a most generous donor to the funds of the Scottish
National Antarctic Expedition, and himself a polar explorer.

ATHERINID.E.

19. Basilichthys laticlavia, Cuv. and Val.

Several small specimens from Station 118, Port Stanley Harbour, Falkland Islands,
51° 41' S., 57° 51' W. ; shore.

(ROY. soc. EDIN. TBANS., VOL. XLIX., 237.)

320 Mil C. TATE REGAN ON THE

ZOARCID.E.

20. Iluoccctes fimbriatus, Jenyns.
Station 118, Port Stanley, Falkland Islands, 51° 41' S., 57° 51' W. ; shore.

21. Austrolycus depressiceps, Regan.

Several small specimens from Station 118, Port Stanley, Falkland Islands, 51° 41' S.,
57° 51' W. ; shore. This species is described on p. 327.

22. Phucoccctes latitans, Jenyns.

Four small specimens from Station 118, Port Stanley, Falkland Islands, 51° 41' S.,
57° 51' W. ; shore.

23. Lycenchelys antarcticus, Regan.

This new species is described on p. 324, from a single specimen from Station 313,
62° 10' S., 41° 20' W.; depth 1775 fathoms; temperature 31'0° F. ; trawl; 18th
March 1903.

BROTULID^E.

24. Neobytliites brucei, Dollo. (PI. III. fig. 2.)
Proc. Roy. Soe. Edin., xxvi., 1906, p. 172.

Depth of body 6^ in the length, length of head 5, or If in its distance from origin
of anal. Diameter of eye 13 in length of head, equal to width of posterior nostril;
maxillary extending well behind eye ; palatine bands of teeth broad ; no praeopercular
spines ; gill-membranes united for a short distance to each other and to isthmus ;
15 gill-rakers on lower part of anterior arch. About 125 scales in a longitudinal series.
Dorsal 108 ; origin behind base of pectoral. Anal 86. Pectoral nearly as long as
head ; pelvics f- the length of head, 2-rayed, each ray simple, expanded distally into an
ovate blade.

The type, 350 mm. in total length, was taken at Station 291, 67° 33' S., 36° 35' W. ;
depth 2500 fathoms; trawl; 7th March 1903.

From most species of Neobythites this species differs in the gill-membranes attached
to the isthmus and the oar-shaped pelvic rays, and I should be inclined to recognise
GARMAN'S genus Holcomycteronus for this species and N. digittatus, had not GARMAN
stated that the form of the pelvic rays is variable in the latter.

This species was named by Dr DOLLO after Dr W. S. BRUCE, leader of the Scottish
National Antarctic Expedition.

BOVICHTHYID^E.

25. Cottoperca gobio, Giinth.

Station 349, 51° 41' S., 57° 51' W., Port William, Falkland Islands; shore.

(ROY. soc. EDIN. TRANS., VOL. XLIX., 238.)

ANTARCTIC FISHES OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 321

26. Cottoperca macrophthalma, Regan.

At Station 346, 54° 25' S., 57° 32' W., Burdwood Bank; 56 fathoms; surface
temperature 41'8° F. ; otter trawl; 1st December 1903. This new species is described
on p. 335.

27. Boviclithys diacanthus, Carmich.

A specimen of 120 mm. from Gough Island. On comparison with Chilian examples
of the species usually known as B. diacanthus, I find that they are distinct (cf. p. 338).

NOTOTHENIID^E.

28. Harpagifer bispinis, Forst.

Several examples from Station 118, Port Stanley, Falkland Islands, and Station 325,
Scotia Bay, South Orkneys — the latter a new record of locality for this species.

29. Trematomus neivnesii, Bouleng.
Station 325, Scotia Bay, South Orkneys.

30. Trematomus borchgrevinkii, Bouleng.
Station 325, Scotia Bay, South Orkneys.

31. Trematomus bernacchii, Bouleng.
Station 325, South Orkneys.

32. Trematomus hansoni, Bouleng.
Station 411, Coats Land, 161 fathoms.

33. Notothenia trigramma, Regan.

This new species, from Station 118, at the Falkland Islands, is described on p. 348.

34. Notothenia ramsayi, Regan.
This new species, from Station 346, the Burdwood Bank, is described on p. 349.

35. Notothenia tesselata, Richards.
Station 118, Port Stanley, Falkland Islands.

36. Notothenia wiltoni, Regan.

Examples of this new species, described on p. 350, were taken by the Scotia at
Station 118, Port Stanley, Falklands, and at Station 346, the Burdwood Bank.

37. Notothenia brevicauda, Lonnb.
Station 118, Port Stanley, Falkland Islands.

(ROY. soc. EDIN. TRANS., VOL. XLIX., 239.)

322 MR C. TATE REGAN ON THE

38. Notothenia sima, Richards.
Station 118, Port Stanley, Falkland Islands.

39. Notothenia gibberifrons, Lonnb.
Station 325, Scotia Bay, South Orkneys.

40. Notothenia nudifrons, Lonnb.
Station 325, Scotia Bay, South Orkneys.

41. Notothenia coriiceps, Richards.
South Orkneys ; common at Station 325, Scotia Bay.

42. Notothenia cornucola, Richards.
Station 118, Port Stanley, Falkland Islands.

43. Notothenia rossi, Richards.
Station 325, Scotia Bay, South Orkneys.

44. Eleginops maclovinus, Cuv. and Val.
Station 118, Port Stanley, Falkland Islands.

BATHYDRACONID^E.
45. Bathydraco scotise, Dollo.

Station 417, 71° 22' S., 16° 34' W., off Coats Land, at a depth of 1410 fathoms.
This species is described on p. 364.

SCLEROPAREI.

SCORP^NIDAE.

46. Sebastes maculatus, Cuv. and Val.

Specimens from Station 461, Gough Island, at 25 fathoms and 100 fathoms, the
latter with Csesioperca coatsi.

47. Sebastes capensis, Gmel.

A small specimen taken at Station 461, Gough Island, with the preceding; both
these species are found at the Cape of Good Hope.

(ROT. 8OC. EDIN. TRANS., VOL. XLIX., 240.)

ANTARCTIC FISHES OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 323

PSYCHROLUTIDiE.

48. Neophrynichthys marmoratus, sp. n.
Neophrynichthys latus (non Button), Giinth., Proe. Zool. Soc., 1881, p. 20, pi. i.

In this species the dermal appendages on the head and anterior part of the body are
much larger and set further apart than in N. latus. Another striking difference is the
narrower interorbital region, its width measuring only £ of the length of the head in
N. marmoratus, but f in its congener from New Zealand. The dorsal rays number
IX-X, 15-16, the anal 11 or 12 ; the caudal is more rounded than in N. latus.

The irregular marbling gives this fish a very different appearance from the New
Zealand form, with its definite pale spots separated by a brown network.

Three specimens, two in the British Museum collection, from the Straits of
Magellan, 320 and 390 mm. in total length, and one of 160 mm. obtained by the Scotia
at Station 346, 54° 25' S., 57° 32' W., Burdwood Bank, 56 fathoms; surface
temperature 41 '8° F. ; otter trawl; 1st December 1903.

II. A REVISION OF THE ZOARCID.E OF SOUTHERN AMERICA AND THE ANTARCTIC.

The Zoarcidse are principally a northern family, and so far as I am aware none is
known from South Africa, Australia, or New Zealand. Two northern deep-water genera,
Lycenchelys and Melanostigma, are represented in the Antarctic Regions, but the littoral
species, with those of South America and the adjacent islands, all belong to genera
distinct from the northern ones.* There has hitherto been much confusion as to the
characters of these genera and species, which it is the object of this revision to clear up.

Synopsis of the Genera.

I. Pelvic fins present ; mouth subterminal.

A. Snout and lower jaw without fringes.

1. Origin of dorsal fin well behind base of pectoral ; gill-opening cleft

downward nearly or quite to lower end of base of pectoral.
Teeth uniserial or biserial in jaws, uniserial on palatines ; tail long and

slender 1. Lycenchelys.

Teeth in jaws triserial ; two teeth near anterior end of each palatine ; tail

moderately elongate 2. Ophthalmolycus.

2. Origin of dorsal fin above base or anterior part of pectoral ; gill-

opening cleft downward at least to middle of base of pectoral.
a. Mouth large, with wide lateral cleft ; gill-opening cleft down-
wards almost or quite to lower end of base of pectoral ;
teeth in jaws uniserial, with anterior canines in the upper
and lateral canines in the lower.

* The habitat of Gymnelis pictus, Giinth., is unknown, and there is no justification whatever for the statement
that it comes from Magellan Straits.

(BOY. soc. EUIN. TRANS., VOL. XLIX., 241.)

322 MR C. TATE REGAN ON THE

38. Notothenia sima, Richards.
Station 118, Port Stanley, Falkland Islands.

39. Notothenia gibberifrons, Lonnb.
Station 325, Scotia Bay, South Orkneys.

40. Notothenia nudifrons, Lonnb.
Station 325, Scotia Bay, South Orkneys.

41. Notothenia coriiceps, Richards.
South Orkneys ; common at Station 325, Scotia Bay.

42. Notothenia cornucola, Richards.
Station 118, Port Stanley, Falkland Islands.

43. Notothenia rossi, Richards.
Station 325, Scotia Bay, South Orkneys.

44. Eleginops maclovinus, Cuv. and Val.
Station 118, Port Stanley, Falkland Islands.

BATHYDRACONID^E.
45. Bathydraco scotise, Dollo.

Station 417, 71° 22' S., 16° 34' W., off Coats Land, at a depth of 1410 fathoms.
This species is described on p. 364.

SCLEROPAREI.

SCORP^ENIDAE.

46. Sebastes maculatus, Cuv. and Val.

Specimens from Station 461, Gough Island, at 25 fathoms and 100 fathoms, the
latter with Csesioperca coatsi.

47. Sebastes capensis, Gmel.

A small specimen taken at Station 461, Gough Island, with the preceding; both
these species are found at the Cape of Good Hope.

(ROY. HOC. EDIN. TRANS., VOL. XLIX., 240.)

ANTARCTIC FISHES OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 323

PSYCHROLUTIDiE.

48. Neophi~ynichthys mwrnoratus, sp. n.

Neophrynichthys lotus (non Button), Giinth., Proe. Zool. Soc., 1881, p. 20, pi. \.

In this species the dermal appendages on the head and anterior part of the body are
much larger and set further apart than in N. latus. Another striking difference is the
narrower interorbital region, its width measuring only £ of the length of the head in
N. marmoratus, but f in its congener from New Zealand. The dorsal rays number
IX-X, 15-16, the anal 11 or 12 ; the caudal is more rounded than in N. latus.

The irregular marbling gives this fish a very different appearance from the New
Zealand form, with its definite pale spots separated by a brown network.

Three specimens, two in the British Museum collection, from the Straits of
Magellan, 320 and 390 mm. in total length, and one of 160 mm. obtained by the Scotia
at Station 346, 54° 25' S., 57° 32' W., Burdwood Bank, 56 fathoms; surface
temperature 41 '8° F. ; otter trawl ; 1st December 1903.

II. A REVISION OP THE ZOARCHXE OF SOUTHERN AMERICA AND THE ANTARCTIC.

The Zoarcidse are principally a northern family, and so far as I am aware none is
known from South Africa, Australia, or New Zealand. Two northern deep-water genera,
Lycenchelys and Melanostigma, are represented in the Antarctic Regions, but the littoral
species, with those of South America and the adjacent islands, all belong to genera
distinct from the northern ones.* There has hitherto been much confusion as to the
characters of these genera and species, which it is the object of this revision to clear up.

Synopsis of the Genera.

I. Pelvic fins present ; mouth subtenninal.

A. Snout and lower jaw without fringes.

1. Origin of dorsal fin well behind base of pectoral ; gill-opening cleft

downward nearly or quite to lower end of base of pectoral.
Teeth uniserial or biserial in jaws, uniserial on palatines ; tail long and

slender . 1. Lycenchelys.

Teeth in jaws triserial ; two teeth near anterior end of each palatine ; tail

moderately elongate 2. Ophthalmolycus.

2. Origin of dorsal fin above base or anterior part of pectoral ; gill-

opening cleft downward at least to middle of base of pectoral.
a. Mouth large, with wide lateral cleft ; gill-opening cleft down-
wards almost or quite to lower end of base of pectoral ;
teeth in jaws uniserial, with anterior canines in the upper
and lateral canines in the lower.

* The habitat of Gymnelis pictus, Giinth., is unknown, and there is no justification whatever for the statement
that it comes from Magellan Straits.

(ROY. soc. EUIN. TRANS., VOL. XLIX., 241.)

324 MR C. TATE REGAN ON THE

Teeth on vomer and palatines . 3. Iluoccetes.

Palate toothless .... 4. Lycodichthys.

b. Mouth moderate, with short lateral cleft; teeth in jaws

uniserial laterally, usually bi- or tri-serial anteriorly ; no

well marked canines ; teeth on palate.

Head not depressed ; gill-opening cleft downward nearly to lower end of

base of pectoral 5. Austrolycichthys.

Head depressed ; gill-opening cleft downward only to middle of base of

pectoral 6. Austrolycus.

3. Origin of dorsal fin above base of pectoral ; gill-opening small,
above the pectoral ; teeth in upper jaw uniserial, in lower bi-
or tri-serial ; teeth on palate . . . 7 . Phucoccetes.
B. Snout and lower jaw with dermal fringes ; palate toothless.

Teeth conical, bi- or tri-serial ; gill-opening almost entirely above the

pectoral .8. Crossolycus.

Teeth incisor-like, uniserial ; gill-opening cleft downward to middle of
base of pectoral . . . . . . . .9. Platea.

II. No pelvic fins ; mouth terminal ; origin of dorsal just behind head ; teeth

uniserial, in jaws and on vomer and palatines.

Gill-opening cleft downward to middle of base of pectoral . 1 0. Maynea.
Gill opening above base of pectoral . . . .11. Melanostigma.

1. Lycenchelys, Gill, 1884.

Proc. Acad. Philad., p. 110.

Form elongate, with the tail long and slender ; mouth subterminal ; teeth in jaws
slender, uni- or bi-serial ; teeth on vomer ; palatine teeth uniserial. -Gill-opening rather
wide, cleft downwards to lower end of base of pectoral. Dorsal origin well behind head ;
pelvic fins present.

Lycenchelys antarcticus, sp. n. (PI. IX. fig. 3.)

Depth of body 16 in the length, length of head 6 and equal to its distance from
origin of anal fin. Head as broad as deep, its breadth a little more than ^ its length.
Snout twice as long as diameter of eye, which is 6 in length of head ; interorbital
width about 16. Muciferous channels of sides of head and lower jaw with large pores.
Lower jaw included ; teeth in jaws rather slender and obtuse, uniserial, biserial near
symphysis of lower jaw ; teeth on palate acute, wide-set. About 110 rays in dorsal
fin, 9 in caudal, 103 in anal ; origin of anal ^ as distant from vertical through origin
of dorsal as from that through base of pectoral, which fin is a little more than ^ as long
as head. Bluish grey ; head darker ; fins brownish grey.

A single specimen, 128 mm. in total length, from near the South Orkneys, Station

(ROY. soc. KDIN. TRANS., VOL. XLIX., 242.)

ANTARCTIC FISHES OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 325

313, 62° 10' S., 41° 20' W., depth 1775 fathoms; bottom temperature 31'0° F. ;
trawl; 18th March 1903.

The few species of this genus hitherto described are from deep water north of the
Equator.

2. Ophthalmolycus, gen. nov.

Form elongate, compressed. Mouth subtenninal ; teeth rather slender and acute,
in about 3 series in both jaws ; no canines ; 3 teeth on vomer and 2 near anterior end
of each palatine. Gill-opening rather wide, cleft downwards nearly to lower end of
base of pectoral Dorsal origin well behind head ; pelvic fins present.

Ophthalmolycus macrops.

Lycodes macrops, Giinth., " Challenger" Shore Fish., p. 21, pL xi. fig. B (1880).

Depth of body 11| in the length, length of head 5 5. Diameter of eye 3^ in length
of head and 7 times interorbital width. Maxillary nearly reaching vertical from
posterior margin of eye. About 90 rays in the dorsal fin, 80 in the anal, and 10 in
the caudal. Origin of dorsal above posterior £ of pectoral ; origin of anal a head-length
behind the head. Pectoral less than ^ the length of head. Yellowish ; 9 broad dark-
brown cross-bars on back, extending on to dorsal fin ; a series of brown spots on the
side, alternating with the bars ; a brown band from eye to operculum.

Straits of Magellan, 40 to 140 fathoms.

Here described from the type, 135 mm. in total length.

Li/codes concolor, Roule (Bull. Mus. Paris, 1911, p. 280) may belong to this genus.
D. 73 ; A. 68. Coloration uniform.

3. fluoccetes, Jenyns, 1842.

Zool. "Beagle," Fish., p. 165.

Head about as broad as deep ; body compressed ; mouth subterminal, with wide
lateral cleft ; teeth conical, uniserial in jaws, in a patch on the vomer and a single
series on the palatines ; 1 or 2 pairs of canines at the symphysis of the upper jaw ;
1 or 2 teeth on each side of lower jaw enlarged, canine-like. Gill-opening cleft
downward to lower end of base of pectoral. Dorsal origin just behind head ; pelvic
fins present.

Iluoccetes fimbriatus.

Jenyns, i.e., p. 166, pi. xxix. fig. 2.

Lycodes variegatus, Giinth., Cat. Fish., iv. p. 322 (1868).

Phucocaetes variegatus e/usus, Smitt, Bihang Svensk. Vet.-Akad., xxiv., 1898, iv., No. 5, p. 43,

pi. v. fig. 32.
PJtucocietes variegatus micropus, Smitt, I.e., pi. v. fig. 33.

Depth of body 8 to ll£ in the length, length of head 4^ to 5^. Diameter of eye
4 to 5^ in length of head, 3 or 4 times the interorbital width. Maxillary extending to
below posterior margin of eye. Dorsal with 80 to 85 rays, anal with 65 to 70, caudal

(ROY. soc. KDIN. TRANS., VOL. XLIX., 243.)

326 MR C. TATE REGAN ON THE

with about 10. Origin of dorsal above base of pectoral, of anal about a head-length
behind head. Pectoral -| to -f the length of head. Head, body, and fins spotted and
marbled ; sometimes cross-bars on the body ; a more or less distinct band from snout to
eye and eye to operculum ; a series of blackish spots at margin of dorsal and anal.

Falkland Islands ; Magellan Straits ; Chile.

Here described from specimens from the Falkland Islands, 80 to 130 mm. in total
length, including the types of Ly codes variegatus and two obtained by the Scotia at
Station 118, Port Stanley.

I am indebted to Mr L. DONCASTER for the loan of JENYNS' type ; the appearance
of some of the mucous canals as free fringing tubes is due to the bad state of preserva-
tion of the specimen.

SMITT'S Phucoccetes variegatus elongatus (t.c., p. 44, pi. v. fig. 34) seems to be a
distinct species, with the head § of the distance from operculum to origin of anal.

4. Lycodichthys, Pappenheim, 1911.
Sitzungsb. Gesdlsch. Naturf. Freunde, 1911, p. 382.

Closely related to Iluoccetes, differing especially in the toothless palate ; teeth
uniserial ; anterior pair in upper jaw enlarged ; lateral teeth of lower jaw spaced,

canine-like.

Lycodichthys antarcticus.

Pappenheim, t.c., p. 383, and Deutsche Sudpolar-Exped., xiii., Zool., v. p. 180, pis. ix. fig. 6 and x. fig. 4.

Depth of body 8 or 9 in the length ; length of head 5 to 5|. Diameter of eye
5 to 6 in length of head. Maxillary extending to below posterior margin of eye or a
little beyond. Dorsal with 85 to 90 rays, anal with about 65, caudal with about 10.
Origin of dorsal a little behind base of pectoral, of anal 1 to 1^ head-lengths behind
head. Pectoral ^ the length of head. Head, body, and fins spotted or marbled.

Wilhelm Land.

Here described from two co-types, 160 and 200 mm. in total length.

5. Austrolycichthys, gen. nov.

Closely related to Austrolycus, differing in the more compressed form, the head
being at least as deep as broad, and in the more inferiorly placed and somewhat larger
gill-openings, cleft downward nearly to the lower ends of the bases of the pectorals.

(l) Austrolycichthys brachycephalus.

Lycodes brachycephalus, Pappenheim, Deutsche Sudpolar-Exped. , xiii., Zool., v., p. 179, pi. x. tig. 3.

Depth of body 8 to 10| in the length, length of head 5f to 6|. Tail from less than

1^ to If as long as rest of fish. Diameter of eye 5 in the length of head. Maxillary

extending to below anterior part of eye. About 90 rays in the dorsal fin, 70 in the

anal, 10 in the caudal. Origin of dorsal above anterior part of pectoral, of anal

(ROY. soc. KDIN. TRANS., VOL. XLIX., 244.)

ANTARCTIC FISHES OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 327

If to 1^ head-lengths behind head. Pectoral f or f the length of head. Grayish or
brownish.

Wilhelm Land, 380 metres.

Here described from two co-types, 155 and 150 mm. in total length. These differ
greatly in form and proportions, as is shown by the accompanying figures, but I cannot
doubt that they belong to the same species.

FIG. 2. — Austrolyeichthys brachycephaliu.

(2) Austrolyeichthys bothriocephalus.

Ly axles bothriocephalus, Pappenheim, Deutsclte Sudpolar-Exped., xiii., Zool., v. p. 178, pi. x. fig. 2.

Apparently related to the preceding species, the more slender of the two examples
of A. brachycephalus described above showing considerable resemblance to the photo-
graph of the type. But this species is said to have more numerous fin-rays, about 110
in the dorsal and 90 in the anal.

Wilhelm Land, 380 metres.

Total length of the unique type, 181 mm.

6. Austrolycus, gen. nov.

Head depressed ; body compressed posteriorly. Mouth subterminal, with short
lateral cleft ; teeth conical, uniserial on sides of jaws, bi- or tri-serial anteriorly ; teeth
on vomer in a group, on palatines in a single series. Gill-opening cleft downwards to
middle of base of pectoral. Dorsal origin just behind head ; pelvic fins present.

(1) Austrolycus depressiceps, sp. n. (PL V. fig. 1.)

Phucocoetes latitans (non Jenyns) Giinth., Cat. Fish., iv. p. 321 (1862); Smith, Bihang Svensk.
Vet.-Akad., xxiv., 1898, iv., Xo. 5, p. 51, pi. v. figs. 37-39; Carman, Mem. Mus. Comp.
Zool., xxiv., 1899, p. 138.

Depth of body about 10 in its length, length of head 5^ to 6J. Diameter of eye
6| to 9 in the length of head, much less than the interocular, but nearly equal to the

(BOY. soc. EDIN. TRANS., VOL. XLIX., 245.)

328 MR C. TATE REGAN ON THE

interorbital width. Maxillary about reaching vertical from posterior margin of eye,
100 to 110 rays in the dorsal fin, 70 to 80 in the anal, 8 to 10 in the caudal; origin
of dorsal above base of pectoral, of anal If to 2 head-lengths behind head. Pectoral f
the length of head. Brownish ; abdomen pale ; on side of head a sharp line between
the dark brown above and pale yellow below, with the brown projecting downwards on
the cheek as a bar ; a pale transverse band across nape, another above end of pectoral
extending on to dorsal fin, which may be followed by similar bands or spots.

Chile ; Patagonia ; Falkland Islands.

Here described from a large series of specimens measuring up to 250 mm. in total
length, including several obtained by the Scotia at Station 118, Port Stanley,
Falkland Islands.

(2) Austrolycus platei.

Lycodes (Phucoccetes) platei, Steind., Zool. Jahrb., Suppl. iv. p. 320, pi. xix. fig. 8 (1897-98).

Evidently closely related to the preceding species, differing in that the length of
the head is f its distance from the vent, the tail is considerably longer than the head
and trunk (only a little longer in A. depressiceps), and the coloration is different, the
body being marked with broad cross bands, the interspaces between which correspond
to the pale bands or spots on the back and dorsal fin of A. depressiceps.

Chile, Cape Espiritu Santo.

Total length 234 mm.

This may be the Phucoccetes variegatus macropus of SMITT (Bihang Svensk. Vet.-
Akad., xxiv., 1898, iv., No. 5, p. 44, pi. v. fig. 35).

7. Phucoccetes, Jenyns, 1842.*
Zool. "Beagle," Fish., p. 168 (1842).

Head and body compressed. Mouth subterminal ; teeth conical, uniserial in upper
jaw and on palatines, bi- or tri-serial in lower jaw ; anterior pair of teeth in upper jaw,
middle vomerine tooth, and 1 or 2 pairs in lower jaw more or less enlarged and canine-
like. Gill-opening small, above base of pectoral. Dorsal origin just behind head ; pelvic
fins present.

Phucoccetes latitans, Jenyns, I.e., pi. xxix. fig. 3.
Lycodes flavus, Bouleng., Ann. Mag. Nat. Hist. (7), vi., 1900, p. 53.

Depth of body 8 to 10 in the length, length of head 6j to 7. Snout l£ as long as
diameter of eye, which is 6 to 7 in length of head, greater than interorbital width.
Lower jaw included ; maxillary extending to below posterior part of eye. Dorsal with

* GARMAN (Mem. Mus. Cornp. Zool, xxiv , 1899, p. 137) has described a fish from 16° N., 99° W., 660 fathoms, and
has named it Phucoccetes mspectus. It is not a Phucoccetes, nor does it seem to be congeneric with any of the
southern littoral forms.

(HOY. soc. EDIN. TRANS., VOL. XLIX., 246.)

ANTARCTIC FISHES OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 329

about 100 rays, anal with about 80, caudal with 5 or 6. Origin of dorsal above base of
pectoral ; pectoral f , pelvics % as long as head. Brownish ; upper half of head dark
brown, with a pale yellow band from eye to shoulder ; lower part of head pale yellowish.

Falkland Islands.

Here described from two specimens, 65 and 110 mm. in total length, the latter the
type of L. flaviis. Four small examples were obtained by the Scotia at Station 118,
Port Stanley, Falkland Islands.

8. Crossolycus, gen. nov.

Form elongate, compressed. Snout and lower jaw with fringes. Mouth sub-
terminal ; teeth in jaws conical, bi- or tri-serial ; lower jaw with a posterior canine ;
palate toothless. Gill-opening almost entirely above base of pectoral. Dorsal origin
above or a little in advance of base of pectoral ; pelvic fins present.

(1) Crossolycus chilensis, sp. n.

Lycodes (Iluocoetes) fimbriatus (non Jenyns) Steind., Zool. JaJirb., Suppl. iv., 1898, p. 322, pi. xx. fig. 10.

Depth of body equal to length of head, 6f in the length of the fish. Diameter of
eye 7 in length of head and equal to interorbital width. Lips thick. Dorsal 80.
Anal 60. Distance from head to origin of anal 1| the length of head. Pectoral f as
long as head. Head, body, and dorsal fin marbled with brown.

Chile, Cape Espiritu Santo.

STEINDACHNER'S specimen measured 252 mm.

( 2 ) Crossolycm fasciatus.

Ilitoccetesjimbriatus sub-sp. fasciatus, Lonnberg, Swedish S. Polar Exped., Fish., p. 20 (1905).

Depth of body 7| in the length, length of head 5. Diameter of eye 5f in the length
of head and equal to interorbital width. Distance from head to origin of anal 1^ the
length of head. Pectoral a little more than J the length of head. Dark brown, with
5 or 6 whitish transverse bars.

Falkland Islands.

Total length 74 mm.

A specimen of 60 mm. recorded by LONNBERG from Tierra del Fuego, uniform yellow
in colour and differing somewhat in proportions, may belong to another species.

9. Platea, Steind., 1897.

Zool. Jafirb., Suppl. iv. p. 323.

Teeth in jaws uriiserial, incisor-like ; palate toothless. Snout and lower jaw with
fringes. Gill-opening cleft downwards to middle of base of pectoral. Dorsal origin
above anterior part of pectoral. Pelvic fins present

(ROY. soc. KDIN. TRANS., VOL. XLIX., 247.)

330 MR C. TATE REGAN ON THE

Platea insignis.
Steind., I.e., pi. xx. fig. 12.

Depth of body 14£ in the length, length of head 7f. Dorsal with about 100 rays,
anal with about 90. Body with dark spots and bars.
Chile, Cape Espiritu Santo.
Total length 265 mm.

10. Maynea, Cunningham, 1870.

Trans. Linn. Soc., xxvii. p. 471.

Gymnelicldliys, Fischer, Jahrb. Hamburg Wiss. Anst., ii., 1885, p. 60.

Elongate, compressed. Mouth terminal ; teeth conical, uniserial, in jaws and on
vomer and palatines. Gill-opening cleft downwards to middle of base of pectoral.
Dorsal origin just behind head. No pelvic fins.

(1) Maynea patagonica.

Cunningham, I.e., p. 472; Giinth., Proc. Zool. Soc., 1881, p. 881, pi. ii. figs. C and D.

Depth of body 10 or 11 in the length, length of head 6f to 7^. Diameter of eye
5 to 6 in length of head ; interorbital region quite narrow. Maxillary extending to
below anterior ^ or middle of eye. About 120 rays in dorsal fin, 95 in anal, 8 in
caudal. Origin of dorsal above base of pectoral, of anal If to If head-lengths behind
head. Pectoral less than ^ as long as head. Yellowish, with broad brown cross-
bars separated by narrower interspaces.

Patagonia ; Falkland Islands.

Here described from two specimens, the type from the Otter Islands, 150 mm.
in total length, and an example of 90 mm. from the Magellan Straits.

(2) Maynea antarctica.

Gymnelichthys antarcticus, Fischer, Jahrb. Hamburg Wiss. Anst., ii., 1885, p. 61, pi. ii. fig. 9.

Maxillary extending to below posterior maigin of eye. About 97 rays in the
dorsal, 74 in the anal ; origin of latter only \\ head-lengths behind the head. No
cross-bars.

South Georgia.

Total length 220 mm.

11. Melanostigma, Giinth., 1881.
Proc. Zool. Soc., p. 21.

Compressed, elongate ; skin loose, smooth, naked. Mouth terminal, oblique ; teeth
uniserial, in jaws and on vomer and palatines. Gill-opening small, above base of
pectoral. Dorsal origin just behind head ; no pelvic fins.

In addition to the species described below, this genus includes a few from deep
waters of the North Atlantic and Pacific.

(ROT. soc. EDIN. TRANS., VOL. XLIX., 248.)

ANTARCTIC FISHES OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 331

Melanostigma gelatinosum.

Giinth., I.e.

Depth of body about 1 0 in the length, length of head 6. Diameter of eye 3^ in
length of head, interorbital width about 12. Maxillary extending to below middle of
eye. Distance from head to origin of anal equal to length of head ; pectoral nearly
\ as long as head. Sides spotted and marbled with purplish grey ; end of tail blackish ;
inside of mouth, gill-opening, and vent black.

Magellan Straits, 24 fathoms.

Here described from the type, a specimen of 140 mm.

III. A MONOGRAPH OF THE NOTOTHENHFORMES.

The division Nototheniiformes includes Percoids without pungent fin-spines, with
the spinous dorsal, when developed, shorter than the long soft dorsal and anal, the

FIG. 3.— Pectoral fin-skeleton of 1, Cotioperca gobio ; 2, Trematamus nemusii ; and 3, Notothenia coriiceps.
cl, cleithrnm ; sc, hypercoracoid (scapula) ; /, foramen ; cor, hypoeoracoid ; m, metacoracoid process ; 1, 2, 3, radials.

principal caudal rays reduced in number (usually 14), the pectorals typically broad-
based and the pelvics jugular, separated by an interspace, and each formed of a spine
and 5 branched rays. There is a single nostril on each side. The structure and position
of the pectoral radials is highly characteristic ; they are 3 in number, rather large flat
plates ; all or 2 are inserted on the hypoeoracoid, and the lowest is the narrowest and
has its lower edge in contact with the metacoracoid process. In other osteological
characters the more generalised types are very similar to the Perciformes.

(ROY. soc. EDIK. TBANB., VOL. XLIX., 249.)

332

MB C. TATE REGAN ON THE

The group corresponds to the Nototheniidae of BOULENGER and DOLLO, with the
addition of Pleuragramma, which does not at all resemble Leptoscopus, and after the
exclusion of Centropercis, evidently related to Champsodon, and of Acanthaphritis
(Pteropsaron), which is related to Hemeroccetes. Draconetta is allied to the
Callionymidse, and its resemblances to Harpagifer are not due to affinity. As now
restricted the Nototheniiformes are characteristic of and peculiar to the Antarctic seas

Antipodes /s« ' Auckland Is.
Chatham, o //?>'* * 2ea/apd

FIG. 4. — Map showing the localities where Nototheniiform Fishes have been collected.

and the region immediately to the north, ranging to S.E. Australia, New Zealand,
Chile, Argentina, Tristan da Cunha, and St Paul Island.

There is every reason to suppose that the group has always been an Antarctic one,
and seeing that it has become differentiated into four quite distinct families and into
several genera, we may perhaps infer that there has been a large cold southern ocean
throughout the greater part of the Tertiary period.

The group throws no light on the question of former extensions northward of the
Antarctic Continent ; at the present day there are littoral species common to Australia

(BOY. soc. EDIN. TRANS., VOL. XLIX., 250.)

ANTARCTIC FISHES OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 333

and New Zealand (Bovichthys variegatus), to New Zealand and South America
(Notothenia macrocephala, N. cornucola), or to the Antarctic Continent and Kerguelen
(Xotothenia coriiceps, Harpagifer bisjnnis) ; and if under the existing conditions
species may have this wide distribution, the fact that some are more restricted and are
separated from the most nearly related forms by wide expanses of ocean can be
explained without the theory of land-bridges.

Many of the more southern types appear to be circumpolar ; for example,
Trematomus neivnesii, T. borchgrevinkii, T. bernacchii, T. hansoni, T. loennbergii,
Pleuragramma antarcticum, Notothenia coriiceps.

With the exception of Pseudaphritis urvillii from the rivers of Tasmania and S.E.
Australia the Nototheniiform fishes are marine, and the great majority of them are
littoral ; several have been described as frequenting the rocks and weeds, but others
prefer deeper water, the species varying in this respect like the Cottids and Gobies of
our northern seas. Fishes pertaining to four genera (Bathydraco, Gerlachea,
Racovitzaia, Cryodraco) live in the open sea, and probably at some distance below
the surface.

Most of the fishes of this group feed on crustaceans, molluscs, etc. (cf. LONNBERG,
Fish. Swedish South Polar Exped., p. 55), but Gymnodraco and the Chaenichthyidae
are no doubt piscivorous. According to LONNBERG (I.e., p. 52) the breeding season varies,
some species probably spawning in the spring, others in the summer, others in the
autumn. The eggs are smaller in Notothenia and Trematomus than in Artedidraco
and Champsocephalus ; they are probably demersal in all, but certainly in the last two

genera.

Synopsis of the Families.

I. One radial on hypercoracoid, two on hypocoracoid ; gill-membranes separate,

free from isthmus ; teeth on vomer and palatines ; mouth protractile ; snout
not produced ; a spinous dorsal fin . . . . 1. Bovichthyidse.

II. All three radials on hypocoracoid ; gill-membranes united, free or attached to

isthmus, usually forming a fold across it ; palate toothless.

A. Palatine and pterygoids normally developed.

Mouth protractile ; snout not produced ; a spinous dorsal fin 2. Nototheniidse.
Mouth not protractile ; snout produced ; no spinous dorsal fin.

3. Bathydraeonidse.

B. Palatine in great part ligamentous ; no mesopterygoid ; mouth not protractile ;

snout produced and depressed .... 4. Chsenichthyidse.

Family 1. BOVICHTHYIDSE.

This family includes Nototheniiformes more generalised than the rest in the presence
of bands of cardiform or villiform teeth not only in the jaws, but on the vomer and
palatines, and in the separate free gill-membranes. All other members of the group

(ROT. SOC. EDIJf. TRANS., VOL. XLIX., 251.)

334 MR C. TATE REGAN ON THE

have the palate toothless and the gill-membranes united, or joined to the isthmus.
The snout is not produced, the mouth is protractile, the lateral line is complete and
continuous, and a spinous dorsal fin is present. The skeleton is well ossified ; there are
2 radials on the hypocoracoid and 1 on the hypercoracoid (fig. 3, 1) ; the palatine and
pterygoids are normally developed. The vertebrae number 38 to 42 (13-16 + 22-29) ;
praecaudals with parapophyses from the fifth or sixth ; ribs and epipleurals on parapo-
physes, when these are developed.

Littoral fishes, with one species in fresh water.

Three genera.

1. Pseudaphritis, Casteln., 1872.
Proc. Zool. Soc. Victoria, i. p. 72 ; Ogilby, Proc. Linn. Soc. N.S. Wales, xxii., 1897, p. 559.

Body subcylindrical, fully scaled. Head small, scaly, somewhat depressed, narrowed
forward ; interorbital region flat. Teeth in bands in jaws and on vorner and palatines ;
lower jaw projecting. Operculum normal, with a weak spine ; gill-membranes not
united, free from isthmus. Origin of spinous dorsal at some distance behind head ; rays
of all the fins branched.

S.E. Australia and Tasmania ; fresh-water.

Pseudaphritis urvillii.

Aphritis urvillii, Guv. and VaL, Hist. Nat. Poiss., viii. p. 484, pi. 243 (1831) : Giinth., Cat.

Fish., ii. p. 242 (1860).
Pseudaphritis bassii, Casteln., I.e.

Depth of body 5 to 6 in the length, length of head 3| to 4. Diameter of eye
5 J to 7 \ in the length of head, interorbital width 8 to 12. Maxillary extending to
below eye ; about 10 short gill-rakers on lower part of anterior arch. Dorsal VII-VIII,
19-20. Anal 23-25. 60 to 65 scales in the lateral line. Body marbled ; dorsal and
caudal spotted.

Rivers from New South Wales to South Australia and Tasmania.

Here described from eight specimens, 100 to 240 mm. in total length, from South
Australia, Victoria, and Tasmania.

2. Cottoperca, Steind., 1876.
Sitzungsb. Akad. Wien, Ixxii. p. 66.

Head and body compressed, fully scaled. Head large ; snout broad ; interorbital
region concave. Teeth in bands in jaws and on vomer and palatines ; lower jaw some-
what the shorter. Operculum normal, with a weak spine ; gill-membranes not united,
free from isthmus. Spinous dorsal originating above operculum ; rays of soft dorsal and

(BOY. soc. EDIN. TRANS., VOL. XLIX., 252.)

ANTARCTIC FISHES OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 335

anal unbranched ; lower pectoral rays simple, more or less thickened and partly
free distally.

Patagonia ; Magellan Straits ; Falkland Islands..

(1) Cottoperca gobio. (PL IV. fig. 3.)

Aphriti* gobio, Giinth., Ann. May. Nat. Hist. (3), vii., 1861, p. 88; and "Challenger" Shore

Fish., p. 21, pi. ix. (1880).
Cottoperca rosenbergii, Steind., Sitzungsb. Akad. Wien, Ixxii., 1876, p. 67, pi. v. fig. 1.

Depth of body about 4 in the length, length of head about 2£. Diameter of eye
4 to 8 in the length of head, interorbital width 13 to 16. Maxillary extending to
below posterior part or posterior edge of eye ; 5 or 6 short gill-rakers on lower part of
anterior arch. Dorsal (VI) VII, 22-23. Anal 20-23. Dorsal spines and rays increasing
in length with age, the longest varying from 5 to f the length of head. Pectoral
about Y the length of head ; six lowest rays simple, somewhat thickened. Caudal
subtruncate. Least depth of caudal peduncle greater than the diameter of eye, except
in quite young specimens. About 60 scales in a lateral longitudinal series, or 65 in the
lateral line, which is complete and continuous. Orange- yellow, with three broad
brownish cross-bars on upper part of body ; head and sides of body spotted and marbled
with brown.

Patagonia ; Tierra del Fuego ; Falkland Islands.

Here described from nine specimens, 130 to 480 mm. in total length, from Magellan
and the Falklands, at depths varying from 6 to 147 fathoms, including the types of the
species and a specimen from Station 349, Port William, Falkland Islands, taken by the
Scotia in January 1903.

(2) Cottoperca macrophthalma, sp. n. (Pis. IV. fig. 2, and V. fig. 2.)
Depth of body 4 to 5 in the length, length of head (to opercular spine) 2f to 2|.
Diameter of eye 3^ to 5 in the length of head, interorbital width 13 to 16. Maxillary
extending to below posterior part or margin of eye, or a little beyond. 5 to 7 short
gill-rakers on lower part of anterior arch. Dorsal VII (VIII), 21-24. Anal 20-22.
In the young, first dorsal spine longest, j the length of head and as long as soft rays ;
in the adult, fourth or fifth spine longest, sometimes \ the length of head ; longest soft
rays sometimes \ the length of head. Other fins, scales, coloration, etc., as in C. gobio.
Least depth of caudal peduncle not more than diameter of eye.

Ten specimens from Station 346, the Burdwood Bank, south of the Falkland Islands,
54° 25' S., 57° 32' W., taken by the Scotia in 56 fathoms on 1st December 1903, and
three from Magellan Straits, 100 to 450 mm. in total length.

(3) Cottoperca macrocephala.
Roule, Bull. Mus. Paris, 1911, p. 277 (1912).

Eye large. Head longer and fins lower than in C. macrophthalma. Seven simple
pectoral rays. Patagonia.

(BOY. 8OC. BDIN. TBANS., VOL. XLIX., 253.)

336

MR C. TATE REGAN ON THE

3. Bovichthys, Guv. and Val., 1831.
Hist. Nat. Poies., viii. p. 486.

Differs from Gottoperca in the naked head and body, the abnormal operculum,
which has a very strong spine and a superior process which articulates with the post-
temporal, and in the enlarged and partly free posterior anal rays.

FIG. 5. — Distribution of Bovichthys and Trematomus, respectively the northernmost and southernmost Kototheniiform genera.

B = £ovi?hthys. 1. B. variegaius; 2. B. angustifrons ; 3, B. diacanthus; 4. B. chileiais ; 4a. B. sp. ; 5. li. veneris ;

6. B. decipiens ; 7. B. psychrolutes ; 8. B. roseopictus.

T = Trematomus. 1. T. neivnesii ; 2. T.nicolai; 3. T.borchgrevinkii; 4. T. brachysoma ; 5. T.vicarius; 6. T.bernacchii;

7. T.hansoni; 8. T. loemibergii,

Note that in Bovichlhys none, in Trematomus most, of the species appear to be circuni polar.

Chile ; Argentina ; Tristan da Cunha and Gough Island ; St Paul ; S.E. Australia
and Tasmania ; New Zealand and neighbouring islands.

(ROY. soc. EDIN. TRANS., VOL. XLIX., 254.)

ANTARCTIC FISHES OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 337

Synopsis of the Species.
[. Interorbital region distinctly concave.

Interorbital width -J$, pectoral fin f the length of head . 1. variegatus.
Interorbital width ^ or ^ pectoral fin f the length of head . 2. angustifrons.
Interorbital width Tir, pectoral fin £ the length of head . 3. diacanthus.
II. Interorbital region nearly flat.

Interorbital width ^ the length of head, pectoral fin f . . .4. chilensis.

Interorbital width more than J the diameter of eye, which is £ the length of

head ; pectoral extending to vent ...... 5. veneris.

Interorbital width 6 J in length of head ; pectoral extending well beyond origin
of anal ...... . . 6. decipiens.

Interorbital width 5 in length of head . . 7. psychrolutes.

Interorbital width 3^ in length of head . 8. roseopictus.

(l) Bovichthys variegatus.

Richards., "Erebus" and "Terror" Fish., p. 56, pi. xxxiv. figs. 1-4 (1846); Giinth., Cat. Fish.,
ii. p. 250 (1860).

Depth of body 4 to 5 in the length, length of head 2f to 3. Diameter of eye 4 to
4j in the length of head, interorbital width 10. Interorbital region moderately
concave ; maxillary extending to below anterior ^ of eye ; opercular spine equal to or
less than diameter of eye ; 8 gill-rakers on lower part of anterior arch. Dorsal VII-
VIII, 18-20. Anal 14-15. Pectoral f the length of head. Caudal subtruncate.
Caudal peduncle longer than deep. Body with irregular dark cross-bars and usually
with pale spots and vermiculations ; spinous dorsal marbled, sometimes with a blackish
blotch posteriorly ; soft dorsal, caudal, and pectoral with series of dark spots ; anal
with a dark longitudinal band.

S.E. Australia ; New Zealand and neighbouring islands.

Here described from six specimens, 75 to 200 mm. in total length, including the
types of the species, from New South Wales (Haslar), New Zealand (Otago Mus.,
Hutton), and Campbell Island (Southern Cross).

(2) Bovichthys angustifrons, sp. n. (PL IV. fig. 1.)

Depth of body 4| to 5 in the length, length of head 2f to 2|. Diameter of eye
4 in the length of head, interorbital width 12 or 13. Interorbital region moderately
concave ; maxillary extending to below anterior ^ of eye ; opercular spine nearly as
long as diameter of eye ; 8 gill-rakers on lower part of anterior arch. Dorsal VIII, 19.
Anal 14. Pectoral f the length of head. Caudal subtruncate. Caudal peduncle longer
than deep. Dark spots on the head and blotches or bars on the body ; soft dorsal and
caudal with series of spots on the rays.

Here described from two specimens, 160 and 145 mm. in total length, the former
from Tasmania (Allport), the latter without locality (Chatham Museum).

(ROY. 80C. KDIN. TRANS., VOL. XLIX., 255.)

338 MR C. TATE REGAN ON THE

(3) Bovichthys diacanthus. (PI. IX. fig. 5.)

CalUonymus diacanthus, Carmich., Trans. Linn. Soe., xii., 1818, p. 501, pi. xxvi.
Bovichthys diacanthus, Giinth., Cat. Fish., ii. p. 249 (1860).

Depth of body 5 in the length, length of head 3. Diameter of eye 4^ in the
length of head, interorbital width 11. Interorbital region concave ; maxillary extend-
ing to below anterior ^ of eye ; opercular spine f the diameter of eye ; 9 gill-rakers on
lower part of anterior arch. Dorsal VIII, 20. Anal 15. Pectoral | the length of
head. Caudal subtruncate. Caudal peduncle longer than deep.

Tristan da Cunha ; Gough Island.

Here described from a specimen of 120 mm. obtained by the Scotia at Gough
Island, shore.

The species was originally described from Tristan da Cunha, where it is said to be
very common among the rocks and to attain a length of 7 inches. CARMICHAEL
describes the colour as olive, with green blotches and white dots.

(4) Bovichthys chilensis, sp. n.

BovicJithys diacanthus (non Carmichael), Cuv. and Val., Hist. Nat. Poiss., viii. p. 487, pi. 244
(1831); Steind., Zool. Jahrb., Suppl. iv., 1897-8, p. 300, pi. xx. fig. 1.

Depth of body 5 in the length, length of head 3 to 3^. Diameter of eye 4^ to 4|
in the length of head, interorbital width 9. Interorbital space nearly flat ; maxillary
extending to below anterior ^ of eye ; opercular spine as long as or shorter than eye ;
8 gill-rakers on lower part of anterior arch. Dorsal VIII, 21. Anal 14-16. Pectoral
f the length of head. Caudal truncate. Caudal peduncle longer than deep. Body
marbled ; spinous dorsal dusky, with a dark blotch posteriorly ; soft dorsal with 2 or 3
series of dark spots ; caudal dusky with orange posterior margin ; lower fins orange,
more or less spotted.

Chile ; Juan Fernandez.

Here described from two specimens from Chile (Del/in), 92 and 96 mm. in total
length.

BERG (Ann. Mus. Buenos Aires, ii., 1897, p. 298) has recorded a species of
Bovichthys from Argentina as Bovichthys diacanthus. This may prove to be B.
chilensis ; more probably it is a new species, as yet undescribed.

(5) Bovichthys veneris.

Bovichthys psychrolut.es (non Giinth.), Kner, Novara Fische, p. 128, pi. vi. (1869).
,, veneris, Sauvage, Arch. Zool. Exp., viii., 1879, p. 25.

Depth of body 5 in the length, length of head about 3|. Diameter of eye 4 in the
length of head. Interorbital region only slightly concave, its width rather more than
| the diameter of eye. Maxillary extending to below anterior ^ of eye ; opercular

(ROT. SOC. EDIN. TRANS., VOL. XLIX., 256.)

ANTARCTIC FISHES OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 339

spine | the diameter of eye. Dorsal VIII, 19-20. Anal 14-15. Pectoral f or £ the
length of head. Caudal rounded or subtruncate. Caudal peduncle longer than deep.

Island of St Paul.

KNER'S and SAUVAGE'S descriptions are based on specimens of 9 or 10 inches.

(6) Bovichthys decipiens, sp. n. (PL IX. fig. 1.)

Depth of body 4 in the length, length of head 2f. Diameter of eye 3| in the
length of head, interorbital width 6|. Interorbital space nearly flat ; maxillary
extending to below anterior ^ of eye ; opercular spine a little shorter than eye ; 8 gill-
rakers on lower part of anterior arch. Dorsal VIII, 19. Anal 14. Pectoral \ the
length of head. Caudal peduncle longer than deep. Body with cross-bars ; spinous
dorsal with a blotch posteriorly ; dorsal and pectoral with series of spots ; caudal barred.

A specimen of 41 mm. from Cook's Straits (Hector) is very similar in appearance to
B. variegatus.

(7) Bovichthys psychrolutes.
Giinth., Cat. Fish., ii. p. 250 (1860).

Depth of body 4 in the length, length of head 3. Diameter of eye 3i in the length
of head, interorbital width 5. Interorbital region nearly flat ; maxillary extending to
below anterior ^ of eye ; opercular spine as long as eye ; 8 gill-rakers on lower part of
anterior arch. Dorsal VIII, 20. Anal 14. Pectoral f the length of head. Caudal
subtruncate. Caudal peduncle longer than deep. Bluish-olive ; fins pale.

Here described from the type, a specimen 38 mm. in total length, from S.VV. of
the Antipodes Islands (50° 8., 170° W.).

(8) Bovichthys roseopictus.

Hutton, Tram. N. Zeal. Ingt., xxxvi., 1903, art. ix.

Depth of body 5|, length of head 4| in total length. Diameter of eye 3 in length
of head, interorbital width 3^. Top of head smooth, with two small ridges. Dorsal
VIII, 18 (?). Anal 13 (?). Pectoral as long as head. Caudal apparently truncated.
Back dark olivaceous brown, sides and abdomen silvery ; a pink spot at base of operculum
and 5 bright rose-pink bands on each side.

New Zealand ; known from a single specimen of 46 mm. picked up on the beach at
Sumner, Canterbury.

Family 2. NOTOTHENIID^E.

Differ from the Bovichthyidae in the toothless palate, the united gill-membranes,
and in having all 3 radials on the hypocoracoid (fig. 3, 2 and 3). Vertebrae 45-56
(16-20 + 25-35).

In the typical genera the skeleton is well ossified, and the rather strong ribs and
epipleurals are inserted on well-developed parapophyses, or only the first one or two
are sessile. In Pleuragramma the skeleton is weak, with the bones thin and papery,

(ROT. soc. KDIN. TRA.SS., VOL. XLIX., 257.),

340 MR C. TATE REGAN ON THE

the vertebral centra are thin cylinders of bone, parapophyses are developed on the
posterior prsecaudals only, and the ribs and epipleurals are feeble.

Synopsis of the Genera.

I. Body scaly ; gill-membranes forming a fold across the isthmus ; opercles normal.

A. Hypercoracoid enclosing its foramen.

Lateral line scales with tubules or pits . . . 1. Trematomus.
Lateral line scales merely notched. . . 2. Pleuragramma.

B. Foramen partly bordered by hypocoracoid.

1. Two or three lateral lines ; maxillary usually extending to below eye ;

pectoral rounded or vertically truncated.
Teeth in bands ...... . . 3. Notothenia.

Teeth uniserial 4. Dissostichus.

2. One lateral line ; maxillary not reaching eye in the adult fish ; pectoral

very obliquely truncated, the upper rays longest . 5. Eleginops.

II. Body naked ; gill-membranes broadly united to isthmus ; operculum hooked

upwards posteriorly, its upper edge deeply concave ; foramen partly bordered

by hypocoracoid.

A mental barbel ; opercles not spinate .... 6. Artedidraco.

No barbel ; operculum and suboperculum each forming a strong spine.

7. Harpagifer.

1. Trematomus, Bouleng., 1902.
"Southern Gross" Pisces, p. 177.

Body scaly ; 2 lateral lines with tubular or pitted scales. Moutli moderate or
rather large ; jaws with bands of villiform teeth. Gill-membranes united, free or
forming a free fold across isthmus. Skeleton well ossified; vertebrae 52-56 (17-21
+ 32-35) ; most of the prsecaudals with parapophyses to which the ribs and epipleurals
are attached ; hypercoracoid enclosing its foramen (fig. 3,2). A spinous dorsal fin ;
pectoral rounded or sub-vertically truncated.

Coasts of the Antarctic Continent ; South Orkneys and South Georgia (fig. 5, p. 336).

The difference between Notothenia, with the hypocoracoid bordering the foramen,
and Trematomus, with the foramen enclosed in the hypercoracoid, may not be very
important, and PAPPENHEIM believed that he found both conditions in one species
(Deutsche Siidpolar-Exped., xiii., Zool., v. p. 166, figs.); but this seems to have been
an error, the specimen with perforate hypercoracoid being Trematomus hansoni and not
Notothenia lepidorhinus. PAPPENHKIM (t.c., p. 170) states that T. bernacchii is a
Notothenia in the structure of its pectoral arch ; I have examined a large series of
specimens, and find that the hypercoracoid encloses its foramen in all.

(HOT. soc. EDIN. TRANS., VOL. XLIX., 258.)

ANTARCTIC FISHES OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 341

Synopsis of the Species.

I. Upper surface of head naked.

A. Cheeks and opercles fully scaled.

Interorbital width 3^ to 5 in length of head. D VI-VIII, 32-38.
A 32-36 ........ 1. newnesii.

Interorbital width 8 or 9 in length of head. D IV, 37. A 32-33.

2. nicolai.

B. Cheeks and opercles scaly above, naked below ; interorbital width 3 to 5|

in length of head.

D V-VI, 34-37. A 31-33 ... 3. borchgrevinkii.

D IV-V, 30-33. A 29-30 .... 4. brachysoma.

II. Occiput scaly.

A Interorbital region naked, or not fully scaled. D I V-VI, 33-38. A 31-35.

Diameter of eye 4f in length of head, interorbital width about 5
(in a specimen of about 270 mm.) ; 55 to 59 scales in a longi-
tudinal series ........ 5. vicarius.

Diameter of eye 3 to 4^ in length of head, interorbital width 5 to 9
(in specimens up to 340 mm.); 60 to 75 scales in a longitudinal
series ........ 6. bernacchii.

B. Interorbital region fully scaled; width 5 to 10 in length of head.

D V-VII, 36-41. A 33-36 . 7. hansoni.

D V-VI, 33-34. A 31-33 . 8. loennbergii.

(l) Trematomus newnesii.

Bouleng., "Southern Cross" Pisces, p. 177, pL xi. (1902).

Notothenia eyaneobrancha, Vaill., Exped. Atttarct. Franfaise, Poiss., p. 26 (1906).

„ microlepidota, Vaill., i.e., p. 35.

„ hodgsoni, Bouleng., Nat. Antarctic Ezped., Nat. Hist., ii., Fish., p. 2, pi. i. fig. 2 (1907).

Depth of body 4 to 5 J in the length, length of head 3£ to 4£. Diameter of eye 3
to 4j in the length of head, interorbital width 3^ to 5. Maxillary extending to below
anterior part or middle of eye (young) or beyond (adult) ; upper surface of head naked,
cheeks and opercles scaly ; 1 5 to 20 gill-rakers on lower part of anterior arch. Dorsal
VI-VIII, 32-38. Anal 32-36. Pectoral f the length of head or more, longer than
pelvics, which reach the vent in the young, but not in the adult. Caudal truncate.
Caudal peduncle about as long as deep. 68 to 86 scales in a longitudinal series from
above base of pectoral fin to caudal, 40 to 52 in upper lateral line, which ends below
posterior rays of dorsal, 3 to 19 in lower lateral line. Brownish, usually spotted or
marbled or with irregular cross-bars ; spinous dorsal blackish ; other fins dusky, often
with small dark spots.

Here described from a large series of specimens, 50 to 200 mrp. in total length, in-
cluding the types of the species and of N. hodgsoni. The types came from Duke of

(BOY. 80C. KDIN. TBASS., VOL. XLIX., 259.)

342 MR C. TATE REGAN ON THE

York Island, near Cape Adare, 3 to 5 fathoms, and Cape Adare, 4 to 8 fathoms ; those of
N. hodgsoni from the Discovery winter quarters, Ross Island. The Scotia specimens
are all from Station 325, Scotia Bay, South Orkneys ; others in the British Museum
are from the South Shetlands.

(2) Trematomus nicolai.
Notothenia nicolai, Bouleng., '• Southern Cross" Pisces, p. 184, pi. xv. (1902).

Depth of body nearly 4 in the length, length of head 3^ to 3|. Diameter of eye
3 to 3^ in the length of head, interorbital width 8 to 9. Maxillary extending to below
anterior £ or 5 of eye ; upper surface of head naked ; cheeks and opercles scaly ;
11 or 12 gill-rakers on lower part of anterior arch. Dorsal IV, 37. Anal 32-33.
Pectoral f to f length of head, somewhat longer than pelvics, which reach vent or origin
of anal. Caudal rounded. Caudal peduncle nearly as long as deep. 58 to 62 scales
in a longitudinal series from above base of pectoral fin to caudal, 39 to 43 in upper
lateral line, which ends below posterior rays of dorsal, 8 to 18 in lower lateral line.
Brownish, with dark cross-bars and sometimes with small dark spots ; fins dusky.
Victoria Land.

Here described from the types, three specimens 150 to 250 mm. in total length,
from Cape Adare, 5 to 8 fathoms, and Duke of York Island, near Cape Adare, 4 fathoms.

The pectoral arch of this species is exactly similar to that of the closely related
T. newnesii, as figured on p. 331.

(3) Trematomus borchgrevinkii.

Bouleiig., "Southern Cross" Pisces, p. 179, pi. xii. (1902); Pappeuheim, Deutsche Siidpular-
Exped., xiii., Zool., v. p. 171 (1912).

Depth of body 4 to 5 in the length, length of head 3| to 4J. Diameter of eye 4 to 5 in
length of head, interorbital width 3 to 4. Maxillary extending to below anterior ^ of
eye ; upper surface of head naked ; upper parts of cheeks and opercles scaly ; 16 to 19
gill-rakers on lower part of anterior arch. Dorsal V-VI, 34-37. Anal 31-33.
Pectoral f to |- the length of head, longer than pelvics, which rarely reach the vent.
Caudal rounded or subtruncate. Caudal peduncle as long as deep, or deeper than long.
78 to 96 scales in a longitudinal series from above base of pectoral fin to caudal ; lateral
lines vestigial, without or with only a few tubules. Yellowish, with dark spots or
irregular cross-bars ; dorsal and caudal sometimes with series of spots.

Graham Land and neighbouring islands ; Wilhelm Land ; Victoria Land.

Here described from several specimens, 180 to 270 mm. in total length, including
the types of the species, from Cape Adare and Duke of York Island, near Cape Adare
(Southern Gross), and examples from the Discovery winter quarters, Ross Island. A
specimen of 80 mm. was obtained in March 1903 by the Scotia, at Station 325, in
Scotia Bay, South Orkneys; depth 10 to 15 fathoms.

(BOY. soc. EDIN. THANB., VOL. XLIX., 260.)

ANTARCTIC FISHES OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 343

(4) Trematomus brachysoma.
Pappenheim, Deutsche Siidpolar-Exped., xiii., Zool., v. p. 172 (1912).

Depth of body 4^ to 4|- in the length, length of head 3 to 3f . Diameter of eye
3f to 3f in the length of head, interorbital width 4 to 5 J. Maxillary extending to
below anterior part or middle of eye ; upper surface of head naked ; upper parts of
cheeks and opercles scaly ; 16 to 19 gill-rakers on lower part of anterior arch. Dorsal
IV-V, 30-33. Anal 26-30. Caudal rounded. Caudal peduncle a little deeper than
long. 65 to 75 scales in a longitudinal series. Yellowish brown ; head and back dark ;
a series of 6 dark spots from operculum to caudal, and 5 below them at level of base of
pectoral ; a dark spot at tip of spinous dorsal ; soft dorsal with irregular dark cross-bars.

Wilhelm Land.

Total length 93 to 166 mm.

(5) Trematomus vicarius.

Trematomuf berno'-chii subsp. vicarius, Lonnberg, Swedish South Polar Exped., Fish., p. 26 (1905).
INotothenia dubia, Lonnberg, i.e., p. 28, pi. iii. fig. 9.

Depth of body 3| in the length, length of head 3^. Diameter of eye 4f in length
of head, interorbital width about 5. Maxillary extending to below anterior ^ of eye ;

FIG. 6.— Head seen from above of A, Trematomus bcrnaechii, and B, T. viearius.

cheeks, opercles, and occiput scaly ; anterior part of interorbital region scaly in the
middle; about 12 gill-rakers on lower part of anterior arch. Dorsal V, 33. Anal 31.
Pectorals f , pelvics f the length of head. Caudal rounded. Caudal peduncle much
deeper than long. 56 to 59 scales in a longitudinal series, 34 in upper lateral line ;
lower lateral line without tubules.

South Georgia.

LONNBERG'S description is based on a single specimen, 240 mm. in length to base of
caudal fin.

Dr LONNBERG has kindly sent me a sketch of the upper surface of the head of the
type, which is here reproduced, together with a figure of the head of a specimen of

(ROT. SOC. KDIN. TRANS., VOL. XLK-, 261.)

344 MR C. TATE REGAN ON THE

T. bernacchii of the same size. After comparing the type with examples of T. bernacchii
that I sent him, Dr LONNBERG writes that the caudal peduncle is notably deeper and
that the scales are larger.

Dr LONNBERG has also sent me one of the types of N. dubia for examination. The
specimen is 50 mm. in total length, and is a Trematomus, with the head scaled as in
T. bernacchii, but with a smaller eye (3f in the length of head) and broader inter-
orbital region (G| in the length of head) than young examples of that species. I count
Dorsal V, 37. Anal 32. 55 scales in a longitudinal series, 30 in the lateral line. 13
gill-rakers on the lower part of anterior arch. It seems probable that this may be a
young example of T. vicarius, although the interorbital region is scaleless.

(6) Trematomus bernacchii.

Bouleng., "Southern Cross" Pisces, p. 181, pi. xiv. (1902).

Notothenia elegans, Vaill., Exped. Antarct. Franchise, Poiss., p. 28 (1906).

Depth of body 3 to 4 1 in the length, length of head 3^ to 4. Diameter of eye 3 to
4| in the length of head, interorbital width 5 to 9. Maxillary extending to below
anterior part or middle of eye ; occiput, cheeks, and opercles scaly ; interorbital region
naked or with a median series of scales ; 13 to 15 gill-rakers on lower part of anterior
arch. Dorsal IV-VI, 34-38. Anal 31-35. Pectoral about f the length of head;
pelvics just reaching anal in young, but not in adult. Caudal rounded. Caudal
peduncle deeper than long. 60 to 75 scales in a longitudinal series from above base of
pectoral fin to caudal, 30 to 42 in upper lateral line ; lower lateral line usually without
tubules. Large dark spots in 2 or 3 alternating series ; upper \ of spinous
dorsal blackish.

Graham Land and neighbouring islands ; Victoria Land.

Here described from several specimens, 90 to 340 mm. in total length, including
the types of the species from Cape Adare, 5 to 8 fathoms, and Duke of York Island,
near Cape Adare, 3 to 4 fathoms (Southern Cross), examples from the Discovery winter
quarters, Ross Island, and two from Station 325, Scotia Bay, South Orkneys, collected
by the Scotia.

In this species the interorbital region seems to be always scaleless in the young, and
is often so in the adult fish.

(7) Trematomus hansoni.

Bouleng., "Southern Cross" Pisces, p. 180, pi. xiii. (1902).

Tnmatomtu hansoni subsp. georgianus, Lonnberg, Swedish South Polar Exped., Fish., p. 25, pi. v.

fig. 17 (1905).
Notothenia sima, Vaill., Exped. Antarct. Franchise, Poiss., p. 23 (1906).

,, lepidorhinus (part.), Pappenheim, Deutsche Sudpolar- Exped., xiii., Zool., v. p. 169

(1912).

Depth of body 3| to 4| in the length, length of head 3| to 4. Diameter of eye
3f to 5 in the length of head, interorbital width 5 to 6|. Maxillary extending to below

(ROY. soc. EDIN. TRANS., VOL. xiax., 262.)

ANTARCTIC FISHES OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 345

anterior part or middle of eye ; occiput, interorbital region, cheeks, and opercles scaly ;
13 to 16 gill-rakers on lower part of anterior arch. Dorsal V-VII, 36-41. Anal
33-36. Pectoral f to |- the length of head, longer than pelvics, which do not reach
the vent. Caudal subtruncate. Caudal peduncle about as long as deep. 60 to 68
scales in a longitudinal series from above base of pectoral fin to caudal, 38 to 46 in
upper lateral line ; lower lateral line usually without tubules. Brownish, with large
dark spots or cross-bars ; head often spotted ; fins usually barred with series of
dark spots.

South Georgia ; Graham Land ; Coats Land ; Wilhelm Land ; Victoria Land.

Here described from several specimens, 160 to 380 mm. in total length, including
the types of the species, from Cape Adare, 4 to 8 fathoms, and Duke of York Island,
near Cape Adare, 3 to 4 fathoms (Southern Cross), examples from the Discovery
winter quarters, Ross Island, and three from Coats Land, Station 411, 74° 01' S.,
22° 00' W., 161 fathoms ; temperature 28'9° F. ; trap; March 1904.

Dr LOXNBERG'S supposed subspecies from South Georgia is fully identical with the
typical form. He gives "the number of anal rays as (31) 32-33. but the figure shows
36, and in an example that he has kindly sent me I count 35. There is no difference in
the shape of the pectoral.

Dr PAPPEXHEIM has kindly sent me for examination the smallest specimen of his
N. lepidorhinus, 160 mm. in total length, which he has noticed as differing from the
types in the larger number of dorsal rays (38 instead of 32 or 33). It differs also in the
naked snout and praeorbital, shorter pelvic fins, lower lateral line without tubules,
foramen enclosed in the hypercoracoid, etc., and is in every way similar to one of the
types of T. hansoni, with which I have compared it.

(8) Trematomus loennbergii, sp. n. (PL VIII. fig. 4.)

Depth of body 4f to 5 in length, length of head 3 to 3|. Diameter of eye 3 to 4^
in the length of head, interorbital width 7 to 10. Maxillary extending to below
anterior £ of eye ; upper surface of head to nostrils, cheeks, and opercles scaly ; 1 3 gill-
rakers on lower part of anterior arch. Dorsal V-VI, 33-34. Anal 31-33. Pectoral
as long as or a little shorter than head ; pelvics extending to origin of anal or beyond.
Caudal rounded. Caudal peduncle longer than deep. 60 to 70 scales in a longitudinal
series from above pectoral fin to caudal ; 36 to 42 in upper lateral line, which ends below
posterior rays of dorsal ; lower lateral line without tubules.

Victoria Land, Graham Land, and neighbouring islands.

Here described from three specimens, two from the Discovery collection — the larger,
132 mm. in total length, from south-west of the Balleny Islands, 254 fathoms; the
smaller, 65 mm., in total length, from Tent Island, near Ross Island. The third
example, also about 65 rnm., is from Seymour Island, and has been sent to me for
examination by Dr LONNBERG, who has recorded this species as Notothenia nicolai

(ROY. 80C. EDIN. TRANS., VOL. XLIX., 263.)

346 MR C. TATE REGAN ON THE

(Swedish South Polar Exped., Fish., p. 45). I have named the species after Dr
LONNBERG in recognition of his kindness in sending me this and other specimens.

2. Pleuragramma, Bouleng., 1902.
"Southern Cross" Pisces, p. 187 (1902).

Closely related to Trematomus, differing especially in the very thin cycloid scales,
the absence of pitted or tubular lateral line scales, and the feebly ossified skeleton, with
parapophyses developed only on the posterior praecaudal vertebrae.

Coasts of the Antarctic Continent.

BOULENGER has placed this genus in the family Leptoscopidae, but it has no affinity
with Leptoscopus, and, on the other hand, is very near to Trematomus, A comparison
of Pleuragramma antarcticum with Trematomus newnesii shows a very close agree-
ment in external and internal characters, even to the number of fin-rays and vertebrae ;
the pectoral arch is precisely similar. In Pleuragramma the two lateral lines are
marked by scales with notched posterior edges, or, if the scales have been lost, by series
of pores.

Pleuragramma antarcticum.

Bouleng., I.e., pi. xviii. ; Vaillant, Exped. Antarcf. Franfaise, Poiss., p. 48 (1906) ; Pappenheim,
Deutsche Siidpolar Exped., xiii., Zool., v. p. 164.

Depth of body 5 to 6 in the length, length of head 3 \ to 4. Diameter of eye 3| to
3f in the length of head, interorbital width 5 to 6. Lower jaw projecting ; maxillary
extending to below anterior J of eye ; upper surface of head naked ; cheeks and opercles
scaly ; 20 to 25 gill-rakers on lower part of anterior arch. Dorsal VI-VII, 34-37.
Anal 36-38. Pectoral truncated, J to f length of head. Caudal slightly emarginate.
About 55 scales in a lateral longitudinal series. Silvery, back darker ; sides and back
powdered with blackish dots.

Graham Land ; Wilhelm Land ; Victoria Land.

Here described from several specimens, 150 to 200 mm. in total length, including
the types of the species from Victoria Land (Southern Cross) and examples from near-
Cape Armitage, Ross Island, and from south-west of the Balleny Islands (Discovery).
VAILLANT gives D V, 39 ; A 38 ; Sq. 44, for specimens from Graham Land ; and PAPPEN-
HEIM, D V-VIII, 34-38 ; A 36-38 ; Sq. 56-60, for examples from Wilhelm Land.

3. Notothenia, Richards., 1844.

"Erebus" and " Terror" Fish., p. 5; Giinth., Gat. Fish., ii. p. 260 (1860).
Macronotothen, Gill, Proa. Acad. P/iilad., 1861, p. 521.

Differs from Trematomus only in that the hypercoracoid foramen is margined below
by the hypocoracoid (fig. 3, 3, p. 331).

Coasts of the Antarctic Continent and northwards to Patagonia, the Falkland

(ROY. soc. EDIN. TRANS., VOL, XLIX., 264.)

ANTARCTIC FISHES OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 347

Islands, South Sandwich Islands, Marion Islands, Kerguelen, southern New Zealand,
and Chatham Islands.

Although the interorbital region is relatively broader in the larger fish, its width,
compared with the length of the head, is an important specific character. By the
" interorbital width " is here understood the actual width of the osseous interorbital
space ; this is nearly always less, and when narrow proportionately much less, than the
interorbital width as measured by BOULENGER.

Synopsis of the Species.
I. Opercles fully scaled.

A. Upper surface and sides of head scaly, except snout and praeorbital ; inter-

orbital width ^ the length of head, or more.

1. Upper lateral line of 62 to 65 tubular scales ; D VI, 32-34. A. 30-32.

3 lateral lines . . . . . . . . .1. trigramma.

2 lateral lines 2. canina.

2. Upper lateral line of 40 to 55 tubular scales. D V-VII, 32-37. A 31-35.

a. 21 to 25 gill-rakers on lower part of anterior arch. D VII (VIII),

34-36. A. 32-34 3. ramsayi.

b. 14 to 19 gill-rakers on lower part of anterior arch.

a. Upper lateral line not or scarcely extending beyond end of dorsal
fin ; diameter of eye 4^ to 6 in length of head (in specimens up
to 250 mm.). D VI-VII, 32-34. A. 31-34 4. tesselata.
/3. Upper lateral line nearly reaching caudal fin.

D VI-VII, 33-36. A 32-34. 16 to 19 gill-rakers on lower part of anterior
arch. Eye 3| to 4f in length of head (in specimens of 125 to 250 mm.)

5. wiltoni.

D V (VI), 35-37. A 32-35. 16 to 19 gill-rakers on lower part of anterior
arch. Eye 4 to 4J in length of head (in specimens of 90 to 180 mm.)

6. brevicauda.

D V-VI, 34-35. A 32-34. 14 or 15 gill-rakers on lower part of anterior arch.
Eye 3 to 3| in length of head (in specimens of 130 to 180 mm.) 7. longipes.

3. Upper lateral line of 30 to 35 tubular scales; D V-VI, 28-31.

A 27-30 8. sima.

B. Upper surface and sides of head scaly, including praeorbital (and snout also

except in N. scotti).

1. Interorbital width 6 in length of head ; D VI-VII, 32-33. A 35-36.

9. lepidorhinus.

2. Interorbital width 9 to 13 in length of head.

D IV-V, 36-37. A 32 10. squamifrons.

D VI, 37-39. A 38 . . . 11. larseni.

D V, 33. A 31 . . .12. scotti.

(ROY. 80C. EDIN. TRANS., VOL. XLIX., 265.)

348 MR C. TATE REGAN ON THE

C. Upper surface and sides of head scaly (except in N~. nudifrons) ; praeorbital
naked ; interorbital width ^ the length of head, or less.

1. Lower lateral line of 32 to 41 tubular scales; D V1I-V1II, 31-33.
A 3 1-33 13. yibberifrons.

2. Lower lateral line of 15 to 18 tubular scales ; D VI-VII, 28-30. A 28-31.

14. acuta.

3. Lower lateral line without tubular scales.

Upper surface of head scaly. D VII, 32. A 32 . . .15. vaillanti.

Upper surface of head scaly. D IV-V, 35-37. A 33-35 . . 16. mizops.

Upper surface of head naked. D IV-VI, 37-39. A 34-36 . 17. nudifrons.

II. Opercles scaly above, naked below ; upper surface of head scaly.

Interorbital width 10 in length of head. D VII, 29. A 27 18. marionen.ns.
Interorbital width 20 in length of head. D V-VI, 29-30. A 30-31.

19. angustifrons.

III. Opercles scaled only on upper part of operculum ; upper surface of head naked.

A. Anal of 27-33 rays.

a. Interorbital width 12 in length of head. D VI, 33. A 31.

20. elegans.

l>. Interorbital width 4 to 7 in length of head.

Cheek usually scaly behind eye. D IV-VI, 31-34. A 27-31 21. cornucola.
Cheek scaly below and behind eye, its lower \ (young) or \ (adult) naked.

D IV-VI, 33-36. A 30-33 22. cyaneobrancha.

Cheek scaly behind eye. D III-VII, 35-41. A 27-31 . 23. coriiceps.

c. Interorbital width 3} in length of head. D VI-VII, 33-35. A 27-29.

24. rossii.

B. Anal of 22 to 25 rays ; interorbital width 2| to 4| in length of head.
D IV, 29-31. 48 to 56 scales in a lateral longitudinal series.

25. macrocephala.
D VI-VII, 28-29. 52 to 58 scales in a lateral longitudinal series.

26. microlepidota.
D VI-VIII, 26-27. 84 to 92 scales in a lateral longitudinal series 27. colbecki.

C. Anal of 18 to 20 rays ; interorbital width about 4 in length of head.

28. filholi.

(1) Notothenia trigramma, sp. n. (PI. VI. fig. 2.)

Depth of body 5 in the length, length of head 4. Diameter of eye 5 in the length
of head and equal to the interorbital width. Lower jaw projecting ; maxillary
extending to below anterior ^ of eye ; upper surface of head, except snout, cheeks and
opercles scaly ; 15 gill-rakers on lower part of anterior arch. Dorsal VI, 34. Anal 32.
Pectoral longer than pelvics, f as long as head, extending to above anal. Caudal
rounded. About 85 scales in a lateral longitudinal series, 65 in upper lateral line,

(ROY. soc. EDIN. TBANS., VOL. XLIX., 266.)

ANTARCTIC FISHES OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 349

which nearly reaches caudal, 13 in line on middle of tail, and 40 to 45 in a third lower
lateral line, which is separated by 4 or 5 longitudinal series of scales from the base of
the anal fin. Brownish ; fins darker.

Port Stanley, Falklands.

Total length 280 mm.

I was at first inclined to make this species the type of a new genus, but on examining
related species of Notothenia I found a specimen of N. brevicauda with a third lateral
line on one side only, formed of 10 tubular scales and separated from the posterior part
of the anal fin by 3 series of scales.

(2) Notothenia canina.
Smitt, Bih. So. Vet.-Akad. Handl., xxiii., iv., No. 3, p. 32, pi. ii. fig. 22 (1897).

Evidently closely related to N. tesselata, but the outer series of teeth stronger,
spaced, more canine-like, and the upper lateral line with 62 to 65 tubular scales.
Dorsal VI, 32-33. Anal 30-31.

East coast of Patagonia.

Total length 138 mm.

Notothenia acuta, Steind. (non Giinther) (Zool. Jahrb., Suppl. iv., 1897-8, p. 303),
from Chile, is probably closely related to N. canina.

(3) Notothenia ramsayi, sp. n. (PI. VII. fig. 1.)

Depth of body 4 to 5-J- in the length of the fish, length of head about 3£.
Diameter of eye 4 to 4^ in the length of head, interorbital width 4^ to 7. Jaws
equal anteriorly ; maxillary extending to below anterior £ of eye ; cheeks, opercles, and
upper surface of head, to between the nostrils, scaly; 21 to 25 gill-rakers on lower
part of anterior arch. Dorsal VII (VIII), 34-36. Anal 32-34. Pectoral from less
than | to f length of head ; pelvics as long, extending to vent or to anal fin. Caudal
rounded or subtruncate. Caudal peduncle as long as deep, or deeper than long, its
least depth ^ to f the length of head. 60 to 72 scales in a longitudinal series, from
above base of pectoral to caudal fin ; 46 to 54 in upper lateral line, which almost
reaches the caudal ; 8 to 17 in lower lateral line. A lateral series of 5 to 7 dark blotches
or vertical bars.

Several specimens, 200 to 300 mm. in total length, taken on 1st December 1903
from the Burdwood Bank, Scotia Station 346, 54° 24' S., 50° 32' W. ; depth 56
fathoms; surface temperature 41 '8° F. ; one from Isthmus Bay, Magellan Straits,
14 fathoms (CoPPlNGER).

This species is named in memory of ALLAN GEORGE RAMSAY, chief engineer of the
Scotia, who died at Scotia Bay, South Orkneys, on 6th August 1903.

(ROY. soc. EDIN. TRASS., VOL. XLIX., 267.)

350 MR C. TATE REGAN ON THE

(4) Notothenia tesselata.

Richards., "Erebus" and "Terror" Fish., p. 19, pi. xii. figs. 3, 4 (1845) ; Giinth., Cat. Fish.,

ii. p. 260 (1860).
Notothenia veitchii, Giinth., Ann. Mag. Nat. Hist. (4), xiv., 1874, p. 370.

„ brevipes, Lonnberg, Swedish South Polar Exped., Fish., p. 15 (1905).

Depth of body 4^ to G in the length, length of head 3J to 3|. Diameter of eye
4^ to 6 in the length of head, interorbital width 5-£- to 6. Lower jaw rather prominent ;
maxillary extending to below anterior part or middle of eye ; cheeks, opercles, and
upper surface of head, except snout, scaly; 14 to 16 gill-rakers on lower part of
anterior arch. Dorsal VI- VII, 32-34. Anal 31-34. Pectoral from less than
| to more than f the length of head, usually longer than pel vies, which seldom
reach the anal. Caudal rounded. Caudal peduncle deeper than long. 62 to
78 scales in a longitudinal series from above base of pectoral to caudal fin, 41 to 48
in upper lateral line, which ends below or a little behind end of dorsal fin, 6 to 11
in lower lateral line. Body marbled ; spinous dorsal dusky, pale at the base ; soft
dorsal, caudal, and sometimes anal, with series of dark spots.

Chile ; Magellan Straits ; Falkland Islands.

Here described from several examples, 140 to 250 mm. in total length, from the
Falkland Islands, Magellan Straits, and Chile, including the types of the species and
specimens from Station 118, Port Stanley, Falkland Islands, collected by the Scotia.
In young specimens (N. veitchii, N. brevipes) the interorbital width is ] or -J- the length
of head.

(5) Notothenia wiltoni, sp. n. (PI. VII. fig. 2.)

Depth of body 4| to 5J in the length of the fish, length of head 3^ to 3|.
Diameter of eye 3^ to 4| in the length of head, interorbital width 6 to 7. Jaws equal
anteriorly ; maxillary extending to below anterior ^ of eye or beyond ; cheeks, opercles,
and upper surface of head, except snout, scaly; 16 to 19 gill -rakers on lower part of
anterior arch. Dorsal VI-VII, 33-36. Anal 32-34. Pectoral £ or ?{ the length of
head ; pelvics as long or somewhat longer, extending to vent or to anal fin. Caudal
rounded or subtruncate. Caudal peduncle deeper than long, its least depth 2 to -?,- the
length of head. 62 to 70 scales in a longitudinal series from above base of pectoral
to caudal fin, 47 to 54 in upper lateral line, which almost reaches caudal, 7 to 14 in
lower lateral line. Body with irregular dark cross-bars ; spinous dorsal dusky, pale
at base.

Ten specimens, 125 to 250 mm. in total length — one from Orange Bay (Paris Mus.) ;
another from the Straits of Magellan (COPPINGER) ; the others taken by the Scotia at
Port Stanley (Station 118) and Port William (Station 349), Falkland Islands (shore,
51° 41' S., 57° 51' W.), and on the Burdwood Bank (Station 346, 54° 25' S., 57° 32' W.,
56 fathoms ; surface temperature 41'8° F. ; otter trawl; 1st December 1903).

This species is named after Mr D. W. WILTON, zoologist of the Scotia.

(KOY. soc. EDIN. TEANS., VOL. XLIX , 268.)

ANTARCTIC FISHES OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 351

(6) Notothenia brevicauda.

Lonnberg, Swedish South Polar Exped., Fish., p. 6, pi. v. fig. 16 (1905).

Depth of body 4| to 5 in the length of the fish, length of head 3^ to 4. Diameter
of eye 4 to 4^ in the length of head, interorbital width 7 or 8. Maxillary extending
to below anterior \ of eye ; cheeks, opercles, and upper surface of head, except snout,
scaly; 16 to 19 gill-rakers on lower part of anterior arch. Dorsal V (VI), 35-37.
Anal 32-35. Pectoral f to f the length of head ; pelvics as long or a little longer,
extending to the anal. Caudal rounded. Caudal peduncle i to f as long as deep, its
least depth about ^ the length of head. 60 to 70 scales in a longitudinal series from
above base of pectoral to caudal fin, 44 to 50 in upper lateral line, which ends 2 to 4
scales in front of the caudal, 4 to 12 in lower lateral line. Body with irregular dark
cross-bars ; pectorals yellow ; pelvics, anal, spinous dorsal, and base of soft dorsal dusky.

.Magellan Straits ; Falkland Islands.

Twelve specimens, 90 to 1 80 mm. in total length, including examples from Port
Stanley (June 1903, 9-10 fathoms) and from Port William, Falkland Islands (January
1903, 6 fathoms). LONNBERG'S type, a specimen of 120 mm., came from Tierra del
Fuego, and there are examples from that locality in the British Museum.

(7) Notothenia longipes.

Steind., Sitzungsb. Akad. Wien, Ixxii., 1876, p. 70, pi. vi. fig. 7; Gtinth., " Challenger" Shore
Fish., p. 21 (1880).

Depth of body 5| to 6£ in the length, length of head 3j to 3|. Diameter of eye

3 to 3^ in the length of head, interorbital width 7 or 8. Maxillary extending to below
anterior | of eye ; upper surface and sides of head, except snout and pra3orbital, scaly ;
14 or 15 gill-rakers on lower part of anterior arch. Dorsal V-VI, 34-35. Anal 32-34.
Pectoral f to f the length of head, somewhat shorter than pelvics, which reach the anal.
Caudal rounded. Caudal peduncle f to * as long as deep, its least depth from £ to f
the length of head. 62 to 70 scales in a longitudinal series, 46 to 55 in upper lateral
line, which almost reaches caudal, 6 to 1 3 in lower lateral line. Body with irregular
brownish cross-bars.

Patagonia and Magellan Straits.

Here described from four examples, 130 to 180 mm. in total length.

(8) Notothenia sima.

Richards., "Erebvs" and " Terror" Fish., p. 19, pi. si. fig. 1 (1845) ; Giinth., Cat. Fish., ii., p. 262 (1860).
Notothenia squamiceps, Peters, Monatsb. Akad. Berlin, 1876, p. 837.

„ karlandrex, Lonnberg, Swedish South Polar Ejrped., Fish., p. 14, pi. iv. fig. 13 (1905).

Depth of body 4 to 5 in the length, length of head 3^ to 3f. Diameter of eye

4 to 5 in the length of head, interorbital width 6 to 8. Maxillary extending to below
anterior part or middle of eye ; occiput, interorbital region, cheeks, and opercles scaly ;

Y. SOC. EDIX. TRANS., VUL. XUZ., 269.)

352 MR 0. TATE REGAN ON THE

10 to 12 gill-rakers on lower part of anterior arch. Dorsal V-VI, 28-3 L. Anal 27-30.
Pectoral f to f the length of head, about as long as pelvics, which reach the vent.
Caudal rounded. Caudal peduncle much deeper than long. 40 to 46 scales in a
longitudinal series from above base of pectoral fin to caudal, 30 to 35 in upper lateral
line, which ends below posterior rays of dorsal, 2 to 12 in lower lateral line, when
developed. Body with irregular dark cross-bars ; vertical fins more or less dusky, the
caudal often barred and with 2 or 3 dark spots at the base.

Magellan Straits ; Falkland Islands.

Here described from several specimens, 60 to 120 mm. in total length, including
the type of the species, from the Falkland Islands, and a co-type of N. karlandreas.

The Scotia examples are from Station 118, Port Stanley, Falklands, 51° 41' S.,
57° 51' W., and there are others in the British Museum collection from Magellan.

(9) Notothenia lepidorhinus.

Notothenia lepidorhinus (part.), Pappenheim, Deutsche Siidpolar-Exped., xiii., Zool., v. p. 169,
pi. ix. fig. 1 and pi. x. fig. 1 (1912).

Depth of body 4 to 4^ in the length, length of head 3|- to 3f. Diameter of eye
3 to 3£ in the length of head, interorbital width about 6. Maxillary extending to
below anterior margin of pupil ; upper surface and sides of head, including snout and
prseorbital, scaly ; 16 gill-rakers on lower part of anterior arch. Dorsal VI- VI I, 32-33.
Anal 35-36. Pectoral f the length of head ; pelvics extending beyond origin of anal.
Caudal rounded. Caudal peduncle somewhat deeper than long. 72 to 82 scales in a
longitudinal series, 45 to 56 in upper lateral line, 32 to 38 in lower lateral line. Body
with irregular dark cross-bars ; spinous dorsal dark anteriorly ; soft dorsal with dark
oblique stripes.

Wilhelm Land, 385 metres.

The types measure 186 to 240 mm. in total length.

(10) Notothenia squamifrons.

Giinth., " Challenger" Shore Fish., p. 16, pi. viii. fig. C (1880).

Depth of body 4| in the length, length of head 3|. Diameter of eye 3 to 3^ in the
length of head, interorbital width 9 to 12. Maxillary extending to below anterior £ of
eye; upper surface and sides of head, including snout and. prseorbital, scaly ; 14 to 16
gill-rakers on lower part of anterior arch. Dorsal IV-V, 36-37. Anal 32. Pectoral
f the length of head, rather shorter than pelvics, which reach the anal. Caudal
peduncle deeper than long. 55 scales in a longitudinal series from above base of
pectoral to caudal, 44 or 45 in upper lateral line, which ends below end of dorsal, or
just behind it, 15 to 18 in lower lateral line. Body with broad irregular cross-bars;
cheek with two oblique stripes ; spinous dorsal blackish.
Kerguelen.

Here described from the types, two specimens, 110 and 150 mm. in total length.

(BOY. soc. EDIN. TRA.NS., VOL. XLIX., 270.)

ANTARCTIC FISHES OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 353

(11) Notothenia larseni.
Lonnberg, Swedish South Polar Exped., Fish., p. 31, pi. i. fig. 3 (1905).

Depth of body 4^ to 5 in the length, length of head 3|. Diameter of eye 3 in
length of head, interorbital width 11 to 13. Maxillary extending a little beyond
vertical from anterior margin of eye ; upper surface and sides of head entirely scaly.
Dorsal VI, 37-39. Anal 38. Pectoral a little shorter than head, longer than pelvics,
which just reach anal. Caudal rounded. Caudal peduncle as long as deep. 69 to 76
scales in a longitudinal series above upper lateral line, which has 55 or 56 tubes and
nearly reaches caudal ; lower lateral line without tubes. Body with irregular cross-bars ;
dorsal with oblique series of spots.

South Georgia ; length 178 mm.

(12) Notothenia scotti.
Bouleng., Nat. Antarct. Exped. Nat. Hut., ii. Fish., p. 2, pi. i. fig. 1 (1907).

Depth of body 5£ in the length, length of head 3^. Diameter of eye 2| in the
length of head, interorbital width 12. Maxillary extending to below anterior £ of eye ;
upper surface of head, except snout, and sides of head, including prseorbital, scaly ;
12 gill-rakers on lower part of anterior arch. Dorsal V, 33. Anal 31. Pectoral f the
length of head, somewhat shorter than pelvics, which reach anal. Caudal peduncle as
long as deep. 50 scales in a longitudinal series from above base of pectoral to caudal,
probably about 40 in upper lateral line. Body with irregular cross-bars ; spinous
dorsal blackish ; soft dorsal and anal blackish posteriorly.

Near Edward Land.

Here described from the type, a specimen of 110 mm. taken at a depth of 300
fathoms off the Ross Barrier, 27th January 1902. In the original description and figure
the fin-rays are miscounted.

(13) Notothenia gibberifrons.

Lonnberg, Swedish South Polar Exped., Fish., p. 33, pi. iii. fig. 10 (1905); Vaillant, Exped.
Antarct. Franfaise, Poiss., p. 33 (1906).

Depth of body 5 to 5J in the length, length of head 3^ to 3£. Diameter of eye
4 to 4f in the length of head, interorbital width 12 to 16. Jaws equal anteriorly;
maxillary not or barely reaching vertical from anterior margin of eye ; cheeks, opercles,
and upper surface of head to nostrils scaly ; 1 0 gill-rakers on lower part of anterior arch.
Dorsal VII-VIII, 31-33. Anal 31-33. Pectoral f the length of head ; pelvics f to f
length of head, not reaching vent. Caudal truncate. Caudal peduncle nearly as long
as deep ; 55 to 66 scales in a longitudinal series from above base of pectoral to caudal, 36
to 44 (to 5\Jide LONNBERG) in upper lateral line, which ends below posterior part of dorsal,
32 to 41 in lower lateral line. Upper part of body irregularly spotted ; dorsal, caudal,
and pectoral fins with series of dark spots. A water-colour drawing shows the ground
colour yellow, the fins greenish, the spots brown.

(ROY. 8OC. EDIN. TRANS., VOL. XLIX., 271.)

354 MR C. TATE REGAN ON THE

Graham Land ; South Georgia ; South Orkneys ; South Shetlands.

Here described from six specimens, 280 to 340 mm. in total length, taken in July
1903 at Station 325, Scotia Bay, 27 fathoms, and Station 326, Jessie Bay, 10 fathoms,
South Orkneys ; there are also two quite small specimens from the same locality.
LONNBEKG'S types come from South Georgia, and there is a specimen in the British
Museum from the South Shetlands.

(14) Notothenia acuta. (PI. VIII. fig. 3.)

Giinth., " Challenger" Shore Fish., p. 17(1880); Pappenheim, Deutsche Siidpolar-Exped., xiii.,
Zool, v. p. 171, pi. ix. fig. 3 (1912).

Depth of body 6 in the length, length of head 3^. Diameter of eye 3f in the
length of head, interorbital width 16. Maxillary extending to below anterior \ of eye ;
sides and upper surface of head scaly, except snout and praeorbital ; 12 gill-rakers on
lower part of anterior arch. Dorsal VI (VII), (28-29) 30. Anal (28-30) 31. Pectoral
nearly as long as the head, longer than pelvics, which reach the vent. Caudal peduncle
somewhat deeper than long. 60 scales in a longitudinal series from above base of
pectoral to caudal, 38 in upper lateral line, which ends below posterior part of dorsal,
16 to 18 in lower lateral line. Body marbled ; dorsal rays with series of small spots;
caudal barred.

Kerguelen.

Here described from the type, about 62 mm. in total length, from Kerguelen.

(15) Notothenia vaillanti, n. sp.
Notothenia acuta (non Giinth.), Vaillant, Exped. Antarct. Franpaise, Poiss., p. 31 (1906).

Depth of body 5| in the length, length of head 3|. Diameter of eye 3 in the length
of head, interorbital width 14. Maxillary extending to below anterior £ of eye ;
sides and upper surface of head scaly, except snout and praeorbital; 10 gill-rakers on
lower part of anterior arch. Dorsal VII, 32. Anal 32. Pectoral a little shorter
than head, as long as pelvics, which reach the anal. Caudal peduncle a little longer
than deep. 55 scales in a longitudinal series from above pectoral fin to caudal, 34 in
upper lateral line, which ends below posterior part of dorsal ; lower lateral line without
tubular scales. Body with irregular cross-bars, broken up into 3 or 4 series of
alternating spots ; dorsal with small spots ; caudal barred.

Graham Land ; Booth, Wandel, and Wiencke Islands.

Here described from a specimen of 56 (46 + 10) mm. Measurements of this example
are given by VAILLANT (t.c., p. 32), and also those of a much larger fish, 410 mm. in
length to base of caudal, with the eye £ and the interorbital width -,L of the length
of the head.

(ROY. soc. EDIN. TRANS., VOL. xux., 272.)

ANTARCTIC FISHES OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 355

(16) Notothenia mizops.

Gtinth., " Challenger" Shore Fish., p. 16, pi. viii. fig. D (1880).

Depth of body 4^ to 4f in the length, length of head 3f to 4. Diameter of eye
3 to 3| in the length of head, interorbital width about 15. Maxillary extending to
below anterior j or ^ of eye ; cheeks, opercles, occiput, and interorbital region scaly ;
9 to 13 gill-rakers on lower part of anterior arch. Dorsal IV-V, 35-37. Anal 33-35.
Pectoral f to % the length of head ; pelvics longer, reaching the anal. Caudal rounded.
Caudal peduncle deeper than long. 48 to 55 scales in a longitudinal series from above
base of pectoral to caudal, 33 to 38 in upper lateral line, which ends below posterior
part of dorsal ; lower lateral line without tubular scales. Body with 2 series of
large, partly confluent, irregular blackish spots ; cheek with 2 oblique stripes ; a
blackish spot on spinous dorsal ; vertical fins with or without series of dark spots.

Kerguelen.

Here described from the types, five specimens, 70 to 170 mm. in total length.

(17) Notothenia nudifrons.

Notothenia mizops var. nudifrons, Lbnnberg, Swedish South Polar Exped., Fish., p. 30, pi. i.

fig. 2 (1905).
Notothenia mizops, Yaillant, Exped. Antard. Franfaise, Poiss., p. 30 (1906).

Closely related to N. mizops, but occiput and interorbital region naked, fin-
rays usually more numerous (Dorsal IV- VI, 37-39. Anal 34-36), and scales smaller,
55 to 65 in a longitudinal series ; 11 or 12 gill-rakers on lower part of anterior arch.
Coloration of N. mizops. A water-colour drawing shows the fish reddish, the spots
dark brown.

South Georgia ; South Orkneys ; Graham Land.

Here described from nine specimens, 70 to 150 mm. in total length, from Station
325, Scotia Bay, South Orkneys, depth 9 to 10 fathoms (June 1903), from South
Georgia (Swedish Expedition), and from Graham Land (Paris Mus.).

18. Notothenia marionensis. (PI. VIII., fig. 2.)
Giinth., " Challenger" Shore Fish., p. 17 (1880).

Depth of body 5 in the length, length of head 3f . Diameter of eye 4 in the length
of head, interorbital width 10. Jaws equal anteriorly ; maxillary extending to below
anterior ^ of eye ; scales on upper half of cheek and opercles, on interorbital region and
occiput ; a transverse naked strip separating last from scales of nape ; 1 1 gill-rakers on
lower part of anterior arch. Dorsal VII, 29. Anal 27. Pectoral fin f , pelvic f the
length of head. Caudal rounded or subtruncate. Caudal peduncle deeper than long.
48 scales in a longitudinal series from above base of pectoral to caudal fin ; 35 in upper
lateral line, which ends below posterior part of dorsal ; 16 in lower lateral line. Body

(ROT. soc. KDIN. TRANS., VOL. XLIX., 273.)

356 MR C. TATE REGAN ON THE

with irregular dark spots ; a blackish spot on upper part of base of pectoral ; dorsal
and caudal with series of small spots.

Marion Island.

Here described from the type, 82 mm. in total length, from Marion Island,
50 to 75 fathoms.

(19) Notothenia angustifrons. (PI. VIII. fig. 1.)
Fischer, Jahrb. Hamburg Wiss. Atist., ii., 1885, p. 55.

Depth of body 5 in the length, length of head 3|. Diameter of eye 4 in the length
of head, interorbital width about 20. Maxillary extending to below anterior margin
or anterior £ of eye ; upper surface of head scaly to between nostrils ; cheeks and
opercles in great part scaly, but naked below ; 10 or 11 gill-rakers on lower part of
anterior arch. Dorsal V-VI, 29-30. Anal 30-31. Pectoral nearly as long as head,
longer than pelvics. Caudal rounded. Caudal peduncle about as long as deep.
46 to 52 scales in a longitudinal series from above base of pectoral to caudal, 26 to 33
in upper lateral line, which ends below middle or posterior part of soft dorsal, 16 to
23 in lower lateral line. Dark bars across the back, which break up into spots on the
sides of the body ; often a bar through spinous dorsal connecting the bases of the
pectorals ; dorsal, caudal, and pectoral fins with series of small dark spots on the rays ;
pelvics and anal pale, sometimes with a few spots.

South Georgia ; South Sandwich Islands.

Here described from six specimens, 70 to 116 mm. in total length, one from
South Georgia (LoNNBBRG), the rest from the South Sandwich group (ALLAKDYCE).

(20) Notothenia elegans.
Giinth., " Challenger" Shore Fish., p. 21, pi. xi. fig. C (1880).

Depth of body 6 to 7 in the length, length of head 4j. Diameter of eye 3| in
the length of head, interorbital width 12. Maxillary extending to below anterior
^ of eye ; a few scales behind eye and on upper part of operculum, rest of head
probably scaleless ; 10 gill-rakers on lower part of anterior arch. Dorsal VI, 33.
Anal 31. Pectoral f the length of head, rather shorter than the pelvics, which reach
the anal. Caudal rounded. Caudal peduncle somewhat deeper than long. 46 to 48
scales in a longitudinal series from above base of pectoral to caudal, 40 or 41 in upper
lateral line, which ends below last rays of dorsal, 4 to 9 in lower lateral line. Large
dark spots or vertical bars on sides of body ; tip of spinous dorsal pink ; soft dorsal
with 3 or 4 series of small dark spots.

Magellan Straits.

Here described from the types, two specimens 95 mm. in total length, from off
Cape Virgins, Patagonia, 55 fathoms.

(ROY. soc. EDIN. TRANS., VOL. XLIX., 274.)

ANTARCTIC FISHES OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 357

(21) Notothenia cornucola.

Richards., "Erebus" and " Terror" Fish., pp. 8, 18, pis. viii. 6gs. 4, 5, and xi. figs. 3, 4 (1845) ;

GiLnth., Cat. Fish., ii. p. 261 (1860).

Notothenia viryata, Richards., i.e., p. 18, pi. xi. figs. 5, 6 ; Giinth., i.e., p. 262.
„ maryinata, Richards., i.e., p. 18, pi. xii. figs. 3, 4.
„ modesta, Steind., Zool. Jahrb. Suppl., iv., 1898, p. 302, pi. xx. tig. 3.

Depth of body 3| to 4| in the length, length of head 3 to 3|. Diameter of eye
4 \ to 5 in the length of head, interorbital width 5| to 7. Jaws equal anteriorly ;
maxillary extending to below middle of eye ; usually a few scales behind eye and on
upper part of operculum ; 11 or 1 2 gill-rakers on the lower part of anterior arch.
Dorsal IV-VI, 31-34. Anal 27-31. Pectoral about f the length of head, extending
to above origin of anal or a little beyond : pelvics about as long. Caudal rounded.
Caudal peduncle much deeper than long. 47 to 55 scales in a longitudinal series from
above base of pectoral to caudal, 36 to 42 in upper lateral line, which euds below last
2 or 3 rays of dorsal ; 6 to 12 in lower lateral line, when it is developed ; only 2 or 3
scales between lateral line and posterior rays of dorsal. Body usually spotted or
marbled, sometimes with a pale lateral band ; vertical fins dusky.

Patagonia ; Magellan Straits ; Falkland Islands ; New Zealand ; Chatham Islands.

Here described from numerous specimens, 90 to 140 mm. in total length, including
the types of the species, of N. virgata and of N. marginata, mostly from the Falkland
Islands and Magellan Straits ; one small specimen from New Zealand. Some examples
were taken at Station 118, Port Stanley, Falkland Islands, shore, by the Scotia.

(22) Notothenia cynaneobrancha.

Richards., "Erebus" and " Terror" Fish., p. 7, pi. iv. (1844); Giinth., Cat. Fish., ii. p. 261 (1860).
Notothenia purpuriwps, Richards., I.e., pi. ii. figs. 3, 4; Giinth., i.e., p. 262.

Depth of body 4 to 5 in the length, length of head 3 to 4. Diameter of eye
4 to 6 in the length of head, interorbital width 5 to 6. Jaws equal anteriorly ;
maxillary extending to below middle or posterior part of eye ; upper surface of head
naked except for a few temporal and post-temporal scales, which may be absent in the
young ; cheek scaly behind and below eye, the lower \ (young) or £ (adult) naked ;
upper part of operculum scaly ; 10 to 13 gill-rakers on lower part of anterior arch.
Dorsal IV-VI, 33-36. Anal 30-33. Pectoral f the length of head, extending to
above vent or origin of anal (adult) or a little beyond (young) ; pelvics about as long.
Caudal rounded. Caudal peduncle much deeper than long. 60 to 70 scales in a
longitudinal series from above base of pectoral to caudal, 32 to 39 in upper lateral
line, which ends below posterior part of dorsal ; lower lateral line, when developed,
with 6 to 15 tubular scales. A dark oblique stripe from eye to angle of praeoperculum,
another below it.

(BOY. soc. EDIN. TRANS., VOL. XLIX., 275.)

358 MR C. TATE REGAN ON THE

Kerguelen.

Here described from several specimens, 120 to 260 mm. in total length, including
the type of the species.

(23) Notothenia coriiceps.

Richards., "Erebus" and "Terror" Fish., p. 5, pi. iii. figs. 1, 2 (1884); Giinth., Cat. Fish., ii.
p. 261 (1860); Vaill., Exped. Antarct. Franfaise, Poiss., p. 24 (1906).

Depth of body 3| to 4^ in the length, length of head 3 to 3£. Diameter of eye
4| to 7 in the length of head, interorbital widtli 4 to 5. Jaws equal anteriorly ;
maxillary extending to below middle (young) or posterior margin (adult) of eye ; head
naked except for a few scales behind eye, on upper part of operculum, and on post-
temporal region; 10 to 14 gill-rakers on lower part of anterior arch. Dorsal III-VII,
35-40. Anal 27-31. Pectoral from less than f (in large specimens) to £ the length
of head, extending to above origin or anterior rays of anal ; pelvics shorter, not or
barely reaching vent. Caudal subtruncate. Caudal peduncle nearly as long as deep.
54 to 68 scales in a longitudinal series from above base of pectoral to caudal, 34 to 49
in upper lateral line, which ends below posterior part of dorsal, 8 to 17 in lower lateral
line ; 4 or 5 scales between lateral line and posterior dorsal rays. Colour varying from
dark greenish black to a pale orange, with or without spots or marking; usually one
or two oblique dark bars across cheek, sometimes broken up into spots ; head sometimes
with pale spots enclosed in dark rings ; spots on body and dorsal fin sometimes large
and tesselated, more often smaller and scattered, rarely uniting to form longitudinal
stripes ; soft dorsal and anal usually with a pale edge.

Graham Land and neighbouring islands ; Kerguelen ; Victoria Land.

Here described from a large series of specimens obtained by the Scotia at Station 325,
Scotia Bay, South Orkneys, 160 to 450 mm. in total length, in addition to the type
of the species from Kerguelen, examples from Duke of York Island and Cape Adare,
Victoria Land (Southern Cross), and from Graham Land (Franqais).

(24) Notothenia rossii.

Richards., "Erebus" and "Terror" Fish., p. 9, pi. v. figs. 1, 2 (1844); Giinth., Cat. Fish., ii.

p. 263 (1860).

Macronotothen rossii, Gill, Proc. Acad. Philad., 1861, p. 521.
Notothenia marmorata, Fischer, Jahrl. Hamb. Wiss. Anst., ii., 1885, p. 53 ; Lonnberg, Swedish

South Polar Exped., Fish., p. 34.

Depth of body 4^ to 4£ in the length, length of head 3| to 3f. Diameter of eye
5 to 5^ in the length of head, interorbital width 3|. Jaws equal anteriorly ; maxillary
extending to below anterior margin of pupil ; scales on upper part of cheek and operculum
and on temporal region ; upper surface of head papillose ; 1 2 gill-rakers on lower part
of anterior arch. Dorsal V-VII, 33-35. Anal 27-29. Pectoral f the length of head,
longer than pelvics. Caudal truncate. Caudal peduncle as long as or a little longer
than deep. 58 to 62 scales in a longitudinal series from above base of pectoral fin to

(HOY. 8OC. EDIN. TRANS., VOL. XLIX., 276.)

ANTARCTIC FISHES OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 359

caudal, 46 to 52 in upper lateral line, which ends below posterior rays of dorsal, 10
to 18 in lower lateral line. Body marbled; dorsal with 2 or 3 series of dark spots ;
anal and caudal with a dark band.

South Georgia ; South Orkneys.

Here described from two specimens, 250 mm. in total length, from South Georgia ;
a little fish, 62 mm. in total length, obtained by the Scotia at Station 325, Scotia
Bay, South Orkneys, seems to belong to this species.

Notothenia rossii was based on a large stuffed specimen, 850 mm. in total length,
with the dorsal spines short and blunt as they often are in large Nototheniids.

(25) Notothenia macrocephala.

Giinth., Cat. Fish., ii. p. 263 (1860).

Notothenia maoriensis, Haast, Trails. N.Z. Inst., v., 1873, p. 276, pi. xvi. fig.

,, anguttata, Hutton, Ann. Mag. Nat. Hist. (4), xvi., 1875, p. 315.

,, hassleriana, Steind., Sitzungsb. Akad. Wien, Ixxii., 1876, p. 69, pi. vi. fig.

., antaretica, Peters, Monatsb. Akad. Berlin, 1876, p. 837.

„ arguta, Huttou, Trans. N.Z. Inst., xi., 1879, p. 339.

porteri, Delfin, Rev. Chilen. Hist. Nat., iii., 1899, p. 117.

Depth of body 3 to 4 in the length, length of head 3£ to 3f . Diameter of eye
4 to 6 in the length of head, interorbital width 2£ to 3£. Jaws equal anteriorly ;
maxillary extending to below anterior ^ of eye ; imbricate scales behind eye and on
upper part of operculum ; upper surface and sides of head otherwise naked, papillose ;
10 to 12 gill-rakers on lower part of anterior arch. Dorsal IV (III- VI), 29-31. Anal
22-25. Pectoral f to % the length of head, considerably longer than pelvics. Caudal
truncate or slightly emarginate. Caudal peduncle usually somewhat longer than deep,
48 to 56 scales in a longitudinal series from above base of pectoral fin to caudal ; 36
to 44 in upper lateral line, which ends below posterior rays of dorsal, 6 to 12 in lower
lateral line. More or less distinct longitudinal stripes or series of spots on the sides ;
dorsal dusky, sometimes reticulated ; caudal, anal, and pelvics sometimes similarly
coloured.

Patagonia ; Magellan Straits ; Falkland Islands ; New Zealand ; Auckland Island ;
Campbell Island.

Here described from several specimens, 130 to 280 mm. in total length, from New
Zealand, Campbell Island, Magellan Straits, and the Falkland Islands. In addition
to the type of the species, types of N. arguta and N. angustata have been examined.

(26) Notothenia microlepidota.

Hutton, Trans. N.Z. Inst., viii., 1876, p. 213; Waite, Subantaretic Id. N. Zealand, Pisces,

p. 590, fig. (1909).
Notothenia parva, Hutton, Trans. N.Z. Inst., xi., 1879, p. 339.

Depth of body 4 to 5 in the length, length of head 3^ to 3£. Diameter of eye
4^ to 6£ in the length of head, interorbital width 3£ to 4^. Upper surface of head

(ROY. soc. EDIN. TRASS., VOL. XLIX., 277.)

3GO MR C. TATE REGAN ON THE

naked, papillose ; sides mostly naked, scaly behind the eye and on upper part of
operculum ; 11 or 12 gill-rakers on lower part of anterior arch. Dorsal VI-VII, 28-29.
Anal 23-25. Pectoral ^ (adult) to f (young) the length of head ; pelvics nearly as
long, not reaching vent. Caudal rounded or subtruncate. Caudal peduncle deeper
than long. 52 to 58 scales in a longitudinal series from above base of pectoral fin to
caudal, 52 to 60 in upper lateral line, which ends near end of dorsal fin, 10 to 15 in
lower lateral line. Head reticulated ; body and fins spotted.

New Zealand ; Auckland Island ; Campbell Island.

Here described from five specimens, 90 to 500 mm. in total length, from Auckland
and Campbell Islands, the smallest the type of N. parva.

(27) Notothenia colbecki.

Bouleng., "Southern Cross" Pisces, p. 185, pi. xvi. (1902) ; Waite, Subantardic hi. N. Zealand,
Pisces, p. 594 (1909).

Depth of body 4 to 5 in the length, length of head 3^ to 3|. Diameter of eye
5 to 7 in the length of head, interorbital width 3 to 4. Maxillary extending to below
anterior part or middle of eye ; head mostly naked, with granular papillae, scaly behind
the eye and on upper part of operculum ; 15 to 18 gill-rakers on lower part of anterior
arch. Dorsal VI-VII I, 26-27. Anal 23-24. Pectoral £ to f the length of head, some-
what longer than pelvics, which do not reach the vent. Caudal emarginate. Caudal
peduncle longer than deep, 84 to 92 scales in a longitudinal series from above base of
pectoral to caudal, 62 to 69 in upper lateral line, which ends near end of dorsal fin,
24 to 35 in lower lateral line. Brownish above, yellowish below ; a pair of oblique
stripes across the cheek ; dorsal and caudal dusky.

Auckland Island ; Campbell Island.

Here described from specimens 125 to 550 mm. in total length, including the types
from Campbell Island and a large stuffed specimen from Auckland.

(28) Notothenia filholi.

Sauvage, Bull. Soc. philom. (7), iv., 1880, p. 228; Passage de Venus, iii. p. 345 (1885); Vaillant,
Exped. Antarct. Franpaise, Poisa., p. 22 (1906).

Depth of body about 6 in the length, length of head 3|- to 4. Diameter of eye
4f to 5 in length of head, interorbital width about 4. Head mostly naked, with
granular papillae ; scaly behind the eye and on upper part of operculum. Dorsal
VI-VII, 24-27. Anal 18-20. Caudal emarginate. Caudal peduncle longer than deep.
Scales in a longitudinal series 100 to 110 (SAUVAGE) or 78 (VAILLANT), the discrepancy
probably due to different methods of counting. Lower lateral line extending forward
to above middle of anal, its anterior 15 scales overlapped by the upper. Brownish.

Campbell Island.

Total length 150 mm.

(ROY. BOC. EDIN. TRANS., VOL. XLIX., 278.)

ANTARCTIC FISHES OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 361

4. Dissostichus, Smitt, 1898.
Bih. Svensk. Vet.-Akad. Handl., xxiv., iv., No. 5, p. 1.

Differs from Notothenia in that the teeth are uniserial, spaced, canine-like.
Patagonia to Graham Land.

Dissostichus eleginoides.

Smittj i.e., p. 2, pi. i. figs. 1-1 1 ; Vaillanfc, Exped. Antarct. Fran^aise, Poiss., p. 36.

Depth of body about 6 in the length, length of head 3. Diameter of eye about
5 in the length of head, interorbital width about 4£ (? 5^). Maxillary extending to
below middle of eye ; upper surface of head to nostrils, cheeks and opercles scaly.
Dorsal IX-X: 27-28. Anal 28-30. Pectoral 5 the length of head. Caudal truncate
or slightly emarginate. Caudal peduncle much longer than deep. About 124 scales
in a longitudinal series ; upper lateral line extending back beyond dorsal, lower extend-
ing forward nearly to the pectoral.

Total length of the type, 228 mm.

Magellan Straits ; Graham Land.

5. Eleginops, Gill, 1861.

Proc. Acafl. Philad., 1861, p. 522.

•Eleginus (non Fischer), Cuv. and Val., Hist. Nat. Poiss., v. p. 158 (1830); Giinth., Cat. Fifh.,
ii. p. 247 (1860).

This genus differs from Notothenia in the rather small mouth, in the complete
absence of the lower lateral line, and in the shape of the pectoral fin.
Chile ; Patagonia ; Falkland Islands.

Eleginops maclorinus.

Eleginus maclovinus, Cuv. and Val., I.e. ; Giinth., I.e.

„ chilensis, Cuv. and Val., o.c., ix. p. 480 (1833) ; Giinth., l.c.
Aphritis undulatus, Jenyns, Zool. "Beagle" Fish., p. 160, pi. xxix. fig. 1 ; Giinth., t.c., p. 243.

„ porosus, Jenyns, t.c., p. 162; Giinth., I.e.
Klef/inus falklandicus, Richards., "Erebus" and "Terror" Fish., p. 30, pi. xx. figs. 1-3 (1845).

Depth of body 4£ to 5| in the length, length of head 3| to 4. Diameter of eye
5 to 8 in the length of head, interorbital width 3 to 5. Maxillary just reaching
vertical from anterior margin of eye in the young, but not in the adult ; occiput, inter-
orbital region, cheeks, and opercles scaly ; 14 or 15 gill-rakers on lower part of anterior
arch. Dorsal VIII-1X, 24-26. Anal 22-24. Pectoral obliquely truncated, with the
upper rays longest, nearly as long as head. Caudal truncate or emarginate. About
60 scales in a lateral longitudinal series, and 65 in the lateral line, which nearly reaches
the caudal fin. Body often spotted or marbled.

Chile ; Patagonia ; Falkland Islands.

Here described from several specimens, 120 to 450 mm. in total length.

(ROT. 800. EDIN. TRASS., VOL. XilX., 279.)

362 MR C. TATE REGAN ON THE

6. Artedidraco, Lonnberg, 1905.
Swedish South Polar Exped., Fish., p. 39.

Differs from Notothenia in the naked body, the absence of the lower lateral line,
the presence of a mental barbel, the broad union of the gill-membranes to the isthmus,
and the hooked operculuni.

Coasts of the Antarctic Continent and South Georgia.

(1) Artedidraco mirus.
Lonnberg, i.e., p. 40, pi. i. fig. 4 and pi. iv. fig. 14.

Barbel club-shaped in the male. Depth of body 4 in the length, length of head
2f to 2|. Diameter of eye 3^ to 4 in the length of head ; iriterorbital width 1\ to 8f .
Dorsal III, 23-24. Anal 17.
Length of types 40 to 92 mm.

South Georgia.

(2) Artedidraco skottsbergi.

Lonnberg, i.e., p. 48, pi. ii. fig. 7, pi. iv. fig. 15; Vaillant, Exped. Antarct. Francaise, Poiss.,

p. 46 (1906).

Depth of body 4£ to 5 in the length, length of head 3. Diameter of eye 3 to 3f
in the length of head, interorbital width 9 or 10. Dorsal III, 24-25. Anal 18-19.
Graham Land, 125 m.
Length of type 57 mm.

(3) Artedidraco shackletoni.

Waite, Brit. Antarctic Exped., Fish., \>. 15, pi. ii. (1911).

Depth of body 4 in the length, length of head about 3. Diameter of eye 3| in the
length of head, interorbital width 10. Dorsal V, 27. Anal 20.
Length of type 146 mm.
Victoria Land ; off Cape Royds, Ross Island, 30 to 80 fathoms.

7. Harpagifer, Richards., 1844.
"Erebus" and " Terror" Fish., p. 11.

Differs from Artedidraco in the absence of the mental barbel and the development
of the operculum and suboperculum as strong spines.
Patagonia to Graham Land and Kerguelen.

Harpagifer bispinis.

Batrachus bispinis (Callionymus bispinus, Forster), Schneid., Block's Syst. Ichth., p. 45 (1801).
Harpagifer bispinis, Richards., i.e., pp. 11, 19, pis. vii. figs. 1-3, xii. figs. 8-9 ; Gtinth., Cat. Fish.,

p. 263 (1860); Smitt, Bihang Svensk. Vet.-Akad. Handl., xxiv., 1898, iv., No. 5, p. 17;

Vaillant, Exped. Antarct. Francaise, Poiss., p. 44 (1906); Pappenheim, Deutsche Siidpolar-

Exped., xiii., ZooL, v. p. 177 (1912).
Harpagifer palliolatus, Richards., i.e., p 20, pi. xii. figs. 5-7.

Dorsal I1I-V, 21-25. Anal 16-20. Coloration variable, usually with bars or
blotches.

(ROY. soc. EDIN. TRANS., VOL. XLIX., 280.)

ANTARCTIC FISHES OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 363

Total length 100 mm.

Patagonia ; Magellan Straits ; Graham Land ; Falkland Islands ; South Georgia ;
South Orkneys ; Marion Islands ; Kerguelen.

The Scotia examples are from Station 118, Port Stanley, Falkland Islands, and
Station 325, Scotia Bay, South Orkneys.

Family 3. BATHYDRACONID^E.

The depressed head, produced snout, non-protractile mouth, and absence of the
spinous dorsal fin distinguish this family externally from the Xototheniidae ; the genera
with spatulate snout may be distinguished from the Chsenichthyids without a spinous
dorsal fin by their scaly body.

The skeleton of Gymnodraco differs from that of Notothenia in the more depressed
skull and more produced rostrum, the elongation of the palatine, which loses its lateral
ethmoid attachment, and the separation of the mesopterygoid and the metapterygoid,
so that the upper margin of the quadrate is free. The pectoral arch is as in Notothenia.
There are 49 vertebrae (20 + 29), the praecaudals with parapophyses behind which
the long slender epipleurals are inserted, and the feeble ribs attached to the epipleurals
at some distance from the centra.

I have ascertained that Bathydraco agrees with Gymnodraco in the structure of
the palatine and of the pectoral arch, and in the presence of ribs.

Synopsis of the Genera.

I. Body scaly ; snout spatulate ; teeth villiform or cardifonn, in bands, without
canines.

A single lateral line running to or towards middle of base of caudal fin ; body
completely scaled . . . . . . . 1. Bathydraco.

Lateral line running near base of dorsal fin ; body completely scaled.

2. Gerlachea.
Lateral line running near base of dorsal fin ; scales in scattered groups.

3. Racovitzaia.

II. Body naked ; snout pointed ; teeth curved, compressed, close-set in a single

series, with strong anterior canines . . . . .4. Gymnodraco.

1. Bathydraco, Giinth., 1878.

Ann. Mag. Nat. Hist. (5), ii p. 18.

Body scaly ; a single lateral line, running to or towards middle of base of caudal.
Snout spatulate ; jaws with small villiform teeth in bands.

Antarctic, in deep water.

GUNTHER has stated 10 branchiostegals for B. antarcticus, but I find only 7.
DOLLO gives 6 for B. scotite, but I count 7 in that species also.

(ROT. soc. EDIN. TRANS., VOL. XLIX., 281.)

364 MR C. TATE REGAN ON THE

(l) Bathydraco antarcticus.
Giinth., I.e., and " Challenger" Deep-Sea Fish., p. 47, pi. viii. fig. A (1887).

Elongate, subcylindrical, the depth 9 in the length, length of head 3. Snout 1| as
long as diameter of eye, which is 4 in length of head, interorbital width 20. Lower
jaw projecting ; maxillary reaching vertical from anterior margin of eye ; cheek com-
pletely scaled; 16 gill-rakers on lower part of anterior arch. Dorsal 36. Anal 31.
Caudal subtruncate. Pectoral truncated, as long as head without snout, reaching origin
of anal. About 1 40 scales in a lateral longitudinal series, about 60 in the lateral line,
which is complete. Brownish ; fins dusky.

South-east of Heard Island, 1260 fathoms.

Here described from the type, 260 mm. in total length.

(2) Bathy draco macrolepis.
Bouleng., Nat. Antarctic Exped. Nat. Hist., ii., Fish., p. 4, pi. i. fig. 3 (1907).

Depth of body 9 in the length, length of head 3. Snout If as long as diameter of
eye, which is 4^ in the length of head, interorbital width 14. Lower jaw projecting;
maxillary reaching vertical from anterior margin of eye ; cheeks naked below the sub-
orbitals ; 11 gill-rakers on lower part of anterior arch. Dorsal 34. Anal 29. Caudal
subtruncate. Pectoral as long as head behind middle of eye, reaching origin of anal.
About 90 scales in a lateral longitudinal series, about 55 in the lateral line, which is
complete. Brownish ; fins dusky.

South-west of Balleny Islands, 252 fathoms.

Here described from the type, 210 mm. in total length. In the original description
the number of gill-rakers was erroneously given as 6, and of dorsal rays as 39 ; the
latter number is also shown in the figure.

(3) Bathydraco scotise. (PI. IX. fig. 4.)
Dollo, Proc. Roy. Soc. Edin., xxvi., 1906, p. 65.

Depth of body 9 to 10 in the length, length of head 3^. Snout If as long as eye,
the diameter of which is 5 in the length of head, interorbital width 12 or 13. Lower
jaw projecting ; maxillary not reaching the vertical from anterior margin of eye ; cheek
naked below the suborbitals ; 19 to 22 gill-rakers on lower part of anterior arch.
Dorsal 38. Anal 31. Caudal subtruncate. Pectoral as long as head without snout,
extending a little beyond origin of anal. About 100 scales in a lateral longitudinal
series, 36 to 40 in the lateral line, which ends at a distance from the caudal equal to
^ its own length.

Two specimens, 133 and 145 mm. in total length, taken by the Scotia at Station 417,
71° 22' S., 16° 34' W., off Coats Land: depth 1410 fathoms; temperature 3T9° F. ;
trawl; 18th March 1904.

(ROY. EOC. EDIN. TRANS., VOL. XLIX., 282.)

ANTARCTIC FISHES OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 365

2. Gerlachea, Dollo, 1900.
Bull. Acad. Roy. Belg. Sciences, p. 195.

Differs from Bathydraco in that the lateral line runs near the base of the dorsal
fin ; a second short lateral line above base of anal.
Deep water off Graham Land.

Gerlachea australis.

Dollo, i.e., p. 196, and Res. Voy. "Belyica," Poiss., p. 25, pi. iL fig. 1 and pi. v. fig. 2 (1904).
Depth of body 8f in the length, length of head 3j. Snout twice as long as diameter
of eye, which is 5 in the length of head, interorbital width 11. Maxillary not reaching
vertical from anterior edge of eye ; cheek fully scaled. Dorsal 47. Anal 35.
Pectoral f the length of head. Caudal emarginate.
71° 14' S., 89° 14' W., 246 fathoms.
Total length 180 mm.

3. Racovitzaia, Dollo, 1900.

Bull. Acad. Roy. Belg. Science*, p. 317.

Body with scattered groups of scales ; a single lateral line running near base of
dorsal fin ; an incubatory pouch between pelvic fins and vent ; in other characters
similar to Gerlachea.

Deep water off Graham Land.

Racovitzaia glacialis.

Dollo, i.e., p. 318, and Res. Voy. " Belgica," Poiss., p. 29, pi. ii. figs. 2, 3, pi. v. fig. 3 (1904).

Depth of body 12 in the length, length of head 3£. Diameter of eye 4 in the length
of head, interorbital width 25. Maxillary not reaching vertical from anterior margin
of eye. Dorsal 30. Anal 27.

71° 19' S., 87° 37' W., 237 fathoms.

Total length 82 mm.

o

4. Gymnodraco, Bouleng., 1902.

"Southern Cross" Pisces, p. 186.

Body naked, depressed anteriorly, compressed posteriorly. Head depressed ; snout
produced, pointed ; jaws with curved compressed teeth, close-set in a single series and
with large anterior canines, those of the mandible exposed in front of the snout.
Operculum with a strong spine with a hooked branch ; suboperculum with a short
spine ; 6 branchiostegals ; gill-membranes forming a fold across isthmus. Two
lateral lines.

Coasts of the Antarctic Continent.

(ROY. soc. EDIN. TRANS., VOL. XLIX., 283.)

366 MR C. TATE REGAN ON THE

Gymnodraco acuticeps.
Bouleng., I.e., pi. xvii. ; Pappenheim, Deutsche Sudpolar-Exped., xiii., Zool, v. p. 176, pi. ix. tig. 4 (1912).

Depth of body 8 in the length, length of head about 3. Snout as long as post-
orbital part of head. Diameter of eye 5 to 6 in length of head, interorbital width 6 to
7. Lower jaw strongly projecting ; maxillary extending to below anterior margin of
eye ; gill-rakers short, sometimes almost vestigial except near the angle. Dorsal 28-30.
Anal 24-26. Pectoral truncated, | as long as head. Caudal truncate. Large dark
spots on head and body ; fins dusky.

Victoria Land ; Wilhelm Land.

Here described from the types, 200 to 300 mm. in total length, from Cape Adare,
4 to 8 fathoms.

Family 4. CH.ENICHTHYID.E.

This family differs externally from the Nototheniidse in the naked body, produced
spatulate snout, and non-protractile mouth. The skeleton of Champsocephalus esox
shows several peculiarities. The skull is depressed, with the long rostral lamina formed
by the frontals and the ethmo-vomer ; the parasphenoid meets the frontals between the
small lateral ethmoids at the anterior margin of the orbit. The palatine is represented
by the maxillary process, attached to the lateral edge of the rostral lamina near its
anterior end, and by a posterior portion articulating with the lateral ethmoid, these
being connected by a long and slender ligament ; the pterygoid is slender, and there
is no mesopterygoid. The prseorbital is large, but the suborbitals are unossified. The
pectoral arch is as in Notothenia. There are 57 vertebrae (28 + 29) ; the epipleurals
are sessile, but the ribs are not ossified.

I have ascertained that Chtenichthys and Pagetopsis are essentially similar in the
structure of the rostrum, the palato-pterygoids and the pectoral arch, and in the
absence of ribs. In all the genera the mouth is very distensible and the dentaries
are freely movable on the articulars ; this is the case, to a certain extent, in the
Bathydraconidse also.

Synopsis of the Genera.

I. Two lateral lines ; pelvic fin-rays all branched or bifid, the middle ones the longest.

A. Lateral line without bony plates ; a spinous dorsal fin, subcontinuous

with the soft dorsal.
No spine on snout ; spinous dorsal of 9 or 10 spines, not more than ^ the length

of soft dorsal . . . . . . .1. Champsocephalus.

A median spine near end of snout ; spinous dorsal of 12 to 15 spines, more than

\ as long as soft dorsal ...... 2. Pagetopsis.

B. Lateral line with bony plates.

A spinous dorsal fin ........ 3. Chienichthys.

No spinous dorsal fin 4. Parachxnichthys.

(ROY. soc. EDIN. TKANS., VOL. XLIX., 284.)

ANTARCTIC FISHES OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 367

II. Two lateral lines ; outer rays of pelvic fins longest ; dorsal fins separated by
an interspace ; lateral line without bony plates . 5. Chsenocephalus.
III. Three lateral lines.

Pelvic fins of moderate length ...... 6. Chionodraco.

Pelvic fins elongate, the rays simple ... 7. Cryodraco.

1. Champsocephalus, Gill, 1861.
Proc. Acad. Philad., p. 509.

Body naked, elongate ; 2 lateral lines, without bony plates. Eye nearly in
middle of length of head ; no spine on snout. Jaws with rather narrow bands of small
sharp teeth, forming only 2 series laterally ; lower jaw not projecting. Gill-rakers
short, but well developed on all the branchial arches, dentigerous, about 20 on lower
part of anterior arch. Spinous dorsal fin well developed, its base less than ^ that of
the soft dorsal, with which it is almost continuous ; pelvics comparatively short, with
the rays normally branched, the middle ones the longest

Patagonia ; Magellan Straits ; South Georgia.

(1) Champsocephalus esox. (PI. X. fig. 1.)

Ohsenichthys esox, Giinth., Ann. Mag. Nat. Hist. (3), vii., 1861, p. 89.

Depth of body 7 to 8 in the length, length of head 3 to 3^. Snout a little longer
than postorbital part of head. Diameter of eye 7 in the length of head, interorbital
width 4 to 5. Supraorbital edges not raised. Maxillary extending to below anterior
part or middle of eye. Uppermost opercular spine shorter than and quite distinct from
the middle one. Dorsal (IX) X, 33-36. Anal 32-35.

Body with dark cross-bars.

Patagonia ; Magellan Straits.

Here described from five specimens, 200 to 300 mm. in total length, including the
type of the species.

(2) Champsocephalus gunnari. (PI. X. fig. 2.)
Lbnnberg, Swedish South Polar Exped., Fish., p. 37 (1905).

Depth of body 6| in the length, length of head 3 J-. Snout as long as postorbital
part of head. Diameter of eye 5 in the length of head, interorbital width 3J.
Supraorbital edges not raised. Maxillary extending to below anterior ^ of eye.
Uppermost and middle opercular spines only free distally, appearing as a single bifid
spine. Dorsal IX (X), 37-40. Anal 36-38. Plumbeous, with some broad darker
cross-bars.

South Georgia.

Here described from a specimen of 420 mm.

(BOY. 8OC. EDIN. TRANS., VOL. XLIX., 285.)

368 MR C. TATE REGAN ON THE

2. Pagetopsis, gen. nov.

Body naked, moderately elongate; two lateral lines, without bony plates. Eye
behind middle of length of head ; an antrorse curved spine near end of snout. Teeth
in jaws small, sharp, biserial ; lower jaw slightly projecting ; gill-rakers vestigial or
absent. Spinous dorsal fin well developed, its base more than \ that of the soft
dorsal ; pelvics rather long, the rays bifid or slightly branched.
• Coasts of the Antarctic Continent.

Pagetopsis macropterus.

Champsocephalus macropterus, Bouleng., Nat. Antarctic Exped. Nat. Hist., ii., Fish., p. 3, pi. i. ;
Pappenheim, Deutsche Sudpolar- Exped , xiii., Zool., v. p. 174 (1912).

Depth of body 4 to 5 in the length, length of head 2f to 2f . Snout nearly | the
length of head. Diameter of eye 5 in the length of head, interorbital width 4.
Maxillary extending to below anterior ^ of eye. 3 or 4 opercular spines, the upper-
most with an antrorse hook. Dorsal XII-XV, 28-31. Anal 25-27. Pectoral f,
pelvics f the length of head. Dark spots and vermiculations on head ; irregular
double cross-bars on body.

Victoria Land ; Wilhelm Land.

Here described from the types, 160 to 250 mm. in total length, from the stomach
of a seal near Cape Armitage, Ross Island.

3. Chsenichihys, Richards., 1844.

"Erebus" and "Terror" Fish., p. 12; Gunth., Cat. Fish., ii. p. 249 (1860).

Differs from Champsocephalus in having a spine on the snout, the teeth in
broader bands, and in the bony plates of the lateral line. Dorsal fins separated by an
interspace. Gill-rakers short, dentigerous.
Kerguelen.

(l) Clisenichihys rhinoceratus.

Richards., i.e., p. 13, pi. vi. ; Gunth., Cat. Fish., ii. p. 249 (1860); Pappenheim, Deutsche
Siidpolar-Exped., xiii., Zool., v. p. 193.

Depth of body 6 in the length, length of head 2f. Snout nearly \ the length
of head. Diameter of eye 5J to 6^ in the length of head, interorbital width 5 to 5§.
Maxillary extending beyond middle of eye (adult). Head moderately rugose ; supra-
orbital edges slightly raised. Dorsal VII, 33-34. Anal 30-33. Second and third
rays of spinous dorsal longest, thence decreasing rapidly. 79 to 84 plates in upper
lateral line ; a few plates on middle of side. Brownish, with darker spots and
reticulations.

Description from the type, a specimen of 450 mm. and a second example of
175 mm., from Kerguelen.

(BOY. soc. EDIN. TRANS., VOL. XLIX., 286.)

ANTARCTIC FISHES OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 369

(2) Chxnichthys rugosus, sp. n.

Eye smaller than in C. rhinoceratus, diameter 8 in head. Head rougher and
supraorbital edges more elevated. Maxillary shorter, not quite reaching middle of
eye. Dorsal VIII, 30 ; third and fourth spines longest, fifth as long as first. Anal 29.
62 plates in upper lateral line ; a nearly continuous series of plates on middle of side.

A specimen of 400 mm. from Kerguelen.

A stuffed example with VIII, 34 dorsal and 30 anal rays, and 72 plates in the
lateral line, appears to belong to this species.

4. Parach&nichthys, Bouleng., 1902.
"Southern Cross" Pisces, p. 176.

Differs from Chtenichthys in the absence of the spinous dorsal fin.
South Georgia ; Graham Land.

Parach&nichthys georgianus.

Chsenichthys georgianus, Fischer, Jahrl. Hamburg Wiss. Aiustalt, ii., 1885, p. 50, pi. i. figs. 1, 2.
? „ charcoti, Vaillant, Fatped. Anlarct. Fran^aise, Pois*., p. 39, fig.

Maxillary not nearly reaching the vertical from anterior margin of eye. Inter-
orbital region narrow, its width less than \ the diameter of eye. Dorsal 44.
Anal 32.

South Georgia ; Graham Land.

Total length 490 mm.

It seems probable that the imperfect fish described by VAILLANT, from Graham
Land, belongs to this species. The figure of the upper surface of the head is at first
sight rather different from FISCHER'S, but the differences may be due to the expansion
of the jaws and opercles and the smaller size of the specimen (head 136 as against
173 mm.).

5. Clixnocephalus, gen. nov.

Body naked, elongate ; two lateral lines without distinct bony plates. Eye some-
what behind middle of head ; a small prominence at anterior end of ethmoid ; jaws with
small sharp teeth forming rather broad bands, there being several series even at the
sides ; lower jaw not projecting ; gill-rakers absent except for 3 or 4 very short
ones below the angle of the first arch. Spinous dorsal fin well developed, its base
about 5 that of the soft dorsal, from which it is separated by an interspace ; pelvics
comparatively short, with the two outer rays the longest, enveloped in thick skin,
but bifid, the others normally branched.

South Georgia.

(ROY. soc. EDIN. TRANS., VOL. SLIS., 287.)

370 MR C. TATE REGAN ON THE

Chaenocephalus aceratus. (PL XL)

Chxnichthys aceratus, Ldnnberg, Kungl. Svensk. Vet.-Akad. Handl., xl., 1906, No. 5, p. 97.

Depth of body 5 to 6 in the length, length of head 2£ to 2|, Snout a little less
than | the length of head. Diameter of eye 5 to 6 in the length of head, interorbital
width about 5. Supraorbital edges raised ; operculum with 3 radiating ridges ending
in spines, the uppermost bifid. Maxillary extending to below middle of eye or beyond.
Dorsal VII-VIII, 39-40 ; third spine longest, J to more than f the length of head.
Anal 37-38. Pectoral and pelvic fins subequal in length, nearly | the length of head.
Greyish, with 4 or 5 dark cross bands, the first from spinous dorsal through base of
pectoral, the second downwards' from origin of soft dorsal, the others less regular and
sometimes with narrower bars developed between them.

South Georgia.

Four specimens, 480 to 530 mm. in total length, collected by Mr DAVID FERGUSON,
and presented to the Scottish Oceanographical Laboratory by Messrs SALVESEN.

6. Chionodraco, Lonnberg, 1906.
Kungl. Svensk. Vet.-Akad. Handl., xl., No. 5, p. 99.

Apparently intermediate between Chssnocephalus and Cryodraco, resembling the
former in fin-structure, the latter in the three lateral lines and the well-developed
rostral spine.

Graham Land.

Chionodraco hamatus.

Cheenichthys rhinoceratus subsp. hamatus, Lonnberg, Swedish South Polar Exped., Fish., p. 47 (1905).
Chionodraco hamatus, Lonnberg, Kungl. Svensk. Vet.-Akad. Handl., xl., 1906, No. 5, p. 99.

Head 3 in total length (with caudal). Snout nearly ^ length of head, nearly twice
diameter of eye, and 1^ interorbital width. Dorsal VIL, 37. Anal 33.
Snow Hill.
Total length 330 mm.

7. Cryodraco, Dollo, 1900.*
Bull. Acad. Roy. Belg. Sciences, p. 129.

Differs from Chsenocephcdus especially in the structure of the pelvic fins, with the
rays simple, the two outer enlarged and prolonged, and in the presence of an additional
lateral line at the base of the anal fin.
Graham Land ; Wilhelm Land.

(1) Cryodraco antarcticus.

Dollo, i.e., p. 130, and Res. Voy. " Belgica," Poiss., p. 20, pi. i. pi. v. fig. 7 (1904).
Depth of body 8 in the length, length of head 3^. Snout 2, eye 4, interorbital width
5 in the length of head. Dorsal III, 44. Anal 43. Pelvic fin more than £ the length

* A fish from Wilhelm Land, 69 mm. long, is recorded by PAFPENHEIM under the name Pagetodes antarcticus. The
number of fin-rays (D IV, 31. A 31) scarcely justifies this determination, and the fish may well belong to an undescribed
species. But as it is so juvenile and even its generic position uncertain, I refrain from giving it a specific name,

(ROY. soc. EDIN. TRANS., VOL. XLIX., 288.)

ANTARCTIC FISHES OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 371

of the fish, extending nearly to end of dorsal fin. Body with 7 dark transverse
bands.

71° 18' S., 88° 2' W., 450 metres.

Total length 200 mm.

o

Length to Base
of Caudal.

Head to End of
Opercular Bony
Flap. Opercufum.

Snout.

Eye.

Interorbital
Width,

. 173

?56 53

26-5

13-25

10-6 f

168

68 ?64

32

13

16

(2) Cryodraco pappenheimi, sp. n.

Pagetodes * antarcticus (non Dollo), Pappenheim, Deutsche Siidpolar-Exped., xiii., ZboZ., v. p. 175.

Length of head 2f in the length of the fish. Snout 2 in the length of head, diameter
of eye 5, interorbital width 4. Dorsal V, 45. Anal 39. Pelvics only reaching four-
teenth ray of dorsal (the prolonged rays perhaps not entire).

Wilhelm Land.

Length of the type, 168 mm. to base of caudal. This species is known to me only
from Dr PAPPENHEIM'S description and from some notes and measurements that he has
kindly sent me. Some of these may be given for comparison with those of the type
of C. antarcticus. The measurements are in millimetres.

0. antarcticus .
C. pappenheimi .

IV. THE SYSTEMATIC POSITION AND GEOGRAPHICAL DISTRIBUTION OF THE
GALAXIID^E AND HAPLOCHITONID.E.

The Galaxiidse and Haplochitonidae are Teleostean fishes of the order Isospondyli,
that is to say, they are malacopterous physostomes with a truly homocercal caudal fin,
with abdominal pelvic fins, and with ribs inserted on autogenous parapophyses. In
this order the name Salmonoid may be given to a group of fishes with an adipose fin
usually present, with one supramaxillary or none, with parietals well developed, and
with oviducts absent or incomplete. The relations of the Salmonoid families are
indicated in the following synopsis :—

I. An orbitosphenoid ; an opisthotic ; a mesocoracoid ; vertebrae upturned at base
of caudal fin ........ 1. Salmonidae.

II. An orbitosphenoid; no opisthotic; no upturned vertebrae; meso-pterygoids

toothless.
A mesocoracoid ; parapophyses inferior .... 2. Argentinidse.

No mesocoracoid ; parapophyses lateral . . . S. Microstomidse.

* The fish named Pagetodes by RICHARDSON (" Erebus " and " Terror" Fish., p. 15, pi. viii. fig. 3) may have belonged
to the genus Cryodraco, but in the form of the body, the length of the pelvic fins, and the continuous dorsals it shows
more resemblance to Pagetopsis. Until RICHARDSON'S species is rediscovered, the name Pagetodes cannot be used.

t DOLLO'S figure of the upper surface of the head is enlarged, the length of the head, to the end of the bony
operculum, to 80 mm. and the interorbital width to 16 mm.

(ROY. soc. EDFN. TRANS., VOL. XLIX., 289.)

372 MR C. TATE REGAN ON THE

III. No orbitosphenoid ; no opisthotic ; no upturned vertebrae ; mesopterygoids
toothed (absent in Salangidse).

A. A mesocoracoid ; maxillaries dentigerous, entering gape . 4. Osmeridee.

B. No mesocoracoid ; maxillaries dentigerous, entering gape.

Head compressed ; mesopterygoid well developed, dentigerous ; ribs ossified.

5. RetropinnatidfB.

Head strongly depressed ; no mesopterygoid ; ribs not ossified 6. Salangidx. '
0. No mesocoracoid ; maxillaries toothless, behind prsemaxillaries.

Prsemaxillaries not extending whole length of maxillaries ; roof of myodome
unossified ; no adipose fin . . . . . . 7. Galaxiidse.

Prsemaxillaries nearly reaching extremities of maxillaries ; roof of myodome
ossified ; an adipose fin . . . . . 8. Haplochitonidse.

The Argentinidse and Microstomidse are inhabitants of rather deep water, but the
rest are littoral fishes, many of them entp.ring fresh water and often forming colonies,
races, or species confined to fresh water.

It is of some interest to note that the Galaxiidse and Haplochitonidse are related to,
but more specialised than, the Osmericlse, or Smelt family, of northern seas. Retropinna,
from the coasts and rivers of Australia and New Zealand, is still nearer to the Galaxiidse
and Haplochitonidse ; both these families occur in Australia, Tasmania, New Zealand,
South America, and the Falkland Islands, and there are even two species of Galaxias
at the Cape of Good Hope. All the species enter fresh water, and the majority seem
to be strictly fluviatile or lacustrine, but in a few cases species of Galaxias have been
observed in the sea.

In 1906 (Proc. Zool. Soc., 1905, ii. pp. 36.3-384, pis. x.-xiii.) I published a
revision of the Galaxiidse, and then wrote :—

" The occurrence of Galaxias maculatus in the sea has been recorded by VALEN-
CIENNES and by PHILIPPI, off the Falkland Islands and off the coast of Chile respectively.
The observations of JOHNSTON in Tasmania and of HUTTON and CLARKE in New Zealand
are to the effect that Galaxias attenuatus descends to the sea periodically to spawn.
Mr RUPERT VALLENTIN has seen shoals of little fishes, which I identify with the
Galaxias gracillimus of CANESTRINI, in the sea at the Falkland Islands. Recently
Galaxias brevipinnis also has been found to be marine, G. bollansi, described by
HUTTON from the Auckland Islands, proving to be identical with this species."

WAITE (Subantarctic Islands of New Zealand, p. 586) has recently shown that
BUTTON'S conclusion as to the marine habit of G. brevipinnis was probably incorrect.
EIGENMANN (Rep. Princeton Exped. Patagonia, iii., Zool., 1909, p. 274) says of G.
gracillimus : " This is undoubtedly the young of attenuatus " ; and if this opinion, which
does not appear to be the result of an examination of specimens, be accepted, the known
marine species of Galaxias would be reduced to two only.

In my revision I distinguished G. gracillimus from G. attenuatus by the more

(ROY. soc. EDIN. TRANS., VOL. XLIX., 290.)

ANTARCTIC FISHES OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 373

slender form, the smaller head, etc. My specimens, 53-55 mm. in total length, were of
the same size as the smallest examples of G. attenuatus, but, bearing in mind the extra-
ordinary larval history of Anguilla, Albula, etc., I wrote : " Possibly this species may
be based on a larval form of G. attenuatus, but if so it is remarkable that it has been
recorded only from South America, and that larval forms of other species have not been
described." A series of Galaxias attenuatus from the Falkland Islands, since received
from Mr VALLENTIN, includes specimens of 20 to 30 mm. which agree with those of 55 to
60 mm. in form, size of head, etc., and show pretty conclusively that G. gracillimus does
not represent a stage in the life-history of this species. Mr VALLENTIN'S collection also
includes some young examples of G. smithii, hitherto known only from the type from
Sir ANDREW SMITH'S collection; these are yellowish, with numerous brownish irregular
vertical stripes.

The South American species of Galaxias are seven in number, viz. :—

1. Galaxias attenuatus, Jenyns, 1842.

S.E. Australia ; Tasmania ; New Zealand and neighbouring islands ; Chile ; Pata-
gonia ; Tierra del Fuego ; Falkland Islands.

2. Galaxias gracillimus, Canestrini, 1864.
Chile ; Falkland Islands.

3. Galaxias maculatus, Jenyns, 1842.
Chile ; Patagonia ; Tierra del Fuego ; Falkland Islands.

4. Galaxias alpinus, Jenyns, 1842.
Alpine lakes of Tierra del Fuego.

5. Galaxias bullocki, Regan, 1908.

Ann. Mag. Nat. Hist. (8), i. p. 372.
Temuco, Chile.

6. Galaxias platei, Steind., 1897.

Galaxias titcambi, Everm. and Kendall, Proc. U.S. Nat. Mus., xxxi., 1907, p. 92, fig.
Patagonia ; Argentina.

7. Galaxias smithii, Regan, 1906.
Falkland Islands.

It should be noted that only the marine species occur both at the Falkland Islands
and on the continent of South America, and there can be little doubt that Haplochiton
zebra, with this distribution, will prove to be marine.

The conclusion that the Galaxiidse are originally marine and are establishing them-
selves in fresh water is strengthened by their relationship to the Osmeridae; their distribu-
tion has little bearing on the question of a former extension of the Antarctic Continent.

The expense of the publication of this Memoir is defrayed from the Government
Publication Grant administered by the Royal Society of London.

(ROY. 80C. EDIN. TRANS., VOL. XLIX., 291.)

374 ANTARCTIC FISHES OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

LIST OF THE PLATES.

PLATE I.
Raia magellanica x J.

PLATE II.

Fig. 1. Chalinura ferrieri.
Fig. 2. ,, whitsani.

PLATE III.

Fig. 1 . Gynomacrurus piriei.
Fig. 2. Neobythiies brucei.

PLATE IV.

Fig. 1. Bovichthys augustifrons.
Fig. 2. Cottoperca macrophthalma.
Fig. 3. „ gobio.

PLATE V.

Fig. 1. Austrolycus depressiceps.
Fig. 2. Cottoperca macrophthalma x

PLATE VI.

Fig. 1. Geesioperca coatsii.

Fig. 2. Notothenia trigramma x f-.

PLATE VII.

Fig. 1. Notothenia ramsayi.
Fig. 2. „ wiltoni.

PLATE VIII.

Fig. 1. Notothenia angustifrons.
Fig. 2. „ marionensis.

Fig. 3. „ acuta x 1|.

Fig. 4. Trematomus loennbergii.

Fig. 5. Synaphobranchus australis.

PLATE IX.

Fig. 1. Bovichthys dect'piensx 1|.
Fig. 2. Bathylagus glacialin.
Fig. 3. Lycenchelys antardicus.
Fig. 4. Bathydraco scotiee.
Fig. 5. Bovichthys diacanthua.

PLATE X.

Fig. 1. Champsocephalus esox.
Fig. 2. ,, gumiari x

PLATE XI.
Cheenocepl/alus aceratus.

(ROY. SOC. EDIN. TRANS., VOL. XUX., 292.)

SCOT. NAT. ANT. EXP.

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PART XVI.
FISHES.

XVL-ATLANTIC FISHES OF SCOTTISH NATIONAL

ANTARCTIC EXPEDITION.

BY R. S. CLARK, M.A., B.Sc.,

Scottish Oceanographical Laboratory, Edinburgh.

(379)

Scottish National Antarctic Expedition : " Scotia " Collection of Atlantic Fishes.
By R. S. Clark, M.A., B.Sc., Zoological Assistant, Scottish Oceanographical
Laboratory, Edinburgh. (With Five Text-figures and Map.)

(MS. received January 14, 1913. Issued separately June 30, 1913.)

INTRODUCTION.

The fishes examined and discussed in the following pages were collected by Dr W. S.
Bruce on the Scotia between latitudes 40° N. and 36° S.* The specimens number
about seven hundred and fifty, and have been divided into two classes :

Littoral Fishes (p. 380).
Pelagic Fishes (p. 399).

The line of demarcation, however, is by no means clearly drawn, as several pelagic
forms occur in the littoral, and several littoral forms in the open sea.

And first as to Littoral fishes. These, numbering about six hundred and fifty,
were caught near the seven different points at which the Scotia touched, namely :
Azores. Madeira. Cape Verde Islands, Ascension, St Helena, Cape Colony, and St Paul's
Rocks. 79 genera are represented, comprising 116 species, none of which are new.
Nevertheless, the collection is of importance as considerably increasing our knowledge
of the geographical distribution of several of the species. As all have been previously
described in detail, it is not intended to enter into particulars, except in the case of the
St Helena fishes, where several points of special interest have been noticed. In certain
other cases, a few notes have been appended, dealing with divergent characteristics of
certain of the species.

With regard to Pelagic fishes, the number of specimens taken is considerably fewer
than in the previous case, extending only to about a hundred. These are referred to
11 genera and 14 species. Here again, there are no new species.

Much of the work of identification has been carried out at the British Museum,
where, through the courtesy of Dr Harmer, Keeper of Zoology, comparison was made
available with the type specimens. To Mr C. Tate Regan, my heartiest thanks are
due for the active assistance he has given me in the identification of many of the
species, to Dr Bruce for the opportunity of making these investigations, and to Mr J. R.
Park for help in compiling this report.

The whole collection is deposited in the Scottish Oceanographical Laboratory,
Edinburgh, along with the more important Antarctic collections of the Scottish National
Antarctic Expedition.

* Vide chart of the route of the Scotia, VoL IV.

380 MR R. S. CLARK ON

A. — LITTORAL FISHES.

I. COLLECTION OF FISHES FROM HORTA FAYAL, AZORES.

Eleven species were secured by the Expedition from traps both inside and outside
the breakwater on July 5th and 6th, 1904. One species, Box vulgaris, was taken in
abundance. All have been previously recorded from these islands.*

Mursena helena Linn.

,, unicolor (De la Roche).
Conger vulgaris Guv.

SPHYR^ENID^E.
Sphyr&na vulgaris (Linn.).

GADID.E.
Phycis mediterraneus De la Roche.

Box vulgaris Cuv. and Val.
Pagrus vulgaris (Linn.).
Pagellus bogaraveo (Briinn.).

LABRID^E.

Coris julis (Linn.).

CARANGID.E.
Tracliurus trachurus (Linn.).

Scorp&na scrofa Linn.

II. COLLECTION OF FISHES FROM MADEIRA.

The collection of fishes from Madeira contains examples of twenty-five different
species. Most of these were bought at the fishmarket of Funchal during the three

* Regan : " On a Collection of Fishes from the Azores," Ann. Mag. Nat. Hist. Land. (Ser. 7), 12, 1903.
Collett : R&. Camp. Sci. de S.A.S. le Prince de Monaco, fasc. x., Monaco, 1896.

THE "SCOTIA " COLLECTION OF ATLANTIC FISHES. 381

days, November 20th to 22nd, 1902, that the Scotia remained there. As all of these
species have been recorded either from Madeira* or from localities not far distant, a list
only of those secured is given.

SQUALID^;.
Centrophorus squamosus (Gmelin).

RAIID^E.
Rcda niaderensis Lowe.

ScOMBRESOCIDjE.

Belone gracilis Lowe.

MUGILID.E.
Mugil chelo Cuv.

SPHYR.ENID.E.
Spkyrama vulgaris (Linn.).

Phyds tnediterraneus De la Roche.

Beryx decadactylus Cuv. and VaL

SERRANID.E.

Anthias sacer (Linn.).
Polyprion amemcanus (Bl. Schn.).
Sen*anus atricauda Giinther.
Priacanthus ci*uentatus (Lacep.).

PRISTIPOMATID.E.
Smarts insidiator Cuv. and Val.

Pagrus vulgaris Cuv. and Val.
Pagellus centrodontus (De la Roche).

* Lowe : (a) "Fishes of Madeira," Proc. Zool. Soc. Land., 1833.

(6) "Synopsis of the Fishes of Madeira," Trans. Linn. Soc , vol ii., London, 1841.

382 MR R. S. CLARK ON

Mullus surmuletus Linn.

LABRID^E.

Labrus mixtus Fries och Ekstrom.
Cossyphus scrofa Cuv. and Val.

CARANGID.E.

Trachurus trachurus (Linn.).
Caranx dentex (Bl. Schn.).

TRICHIURID^E.

Aphanopus carbo Lowe.

Thyrsites prometheus (Cuv. and Val.).

ZEIDJE.
Zeus faber Linn.

ECHENEIDID.E.
Echeneis brachyptera Lowe.

Sebastes kuhlii (Bowd.).
Scorpdena dacti/loptera I)e la Roche.
Sebastes dactylopterus.

III. FISHES COLLECTED AT ST VINCENT, CAPE VERDE ISLANDS.*

Twenty species were secured during the short stay on 24th December 1902.
Interesting occurrences at these islands are the West Indian species Belone caribbsea
and Mugil nigrostrigatus, while the record of Merluccius vulgaris may prove the
southern limit of this species.

SCOMBRESOCID^:.

Belone caribbsea Lesueur.
Hemirhamphus brasiliensis (Linn.).
Exocoetus obtusirostris Giinther.

* Vide Giinther, Challenger Report, "Shore Fishes."

THE " SCOTIA " COLLECTION OF ATLANTIC FISHES. 383

MUGILID.E.

Mugil nigrostrigatus Giinther.

The original locality of this species seems doubtful,* though it has been considered
as belonging to the West Indian fish fauna. It is closely allied to M. chelo, a
Mediterranean form, and the possibility is that the two are not distinct species. The
Scotia specimen is identical with the types of Jf. nigrostrigatus in the British Museum,
though much larger. Total length of specimen 400 mm.

PoLYNEMIDiE.

Galeoides polydactylus (Vahl.).

SPHYR^ENHXE.
Sphyrtena vulgaris (L. Gmel.).

GADID^E.
Merluccius vulgaris (Linn.).

SERRANID.E.
Epinephelus alexandrinus (Cuv. and Val.).

SPARID^E.
Sargus rondeletii Cuv. and Val.

MULLIP^E.
Upeneus prayensis Cuv. and Val.

CARANGID,E.

Caranx sanctse-helenaB Cuv. and Val.
,, crumenophtholmus Lacep.
,, dentex (Bl. Schn.).
Tracliynotus glaucus Linn.t

Lichia glauca.
Argyreiosus setipinnis (Mitchill).

Thynnus pelamys (Linn.).

* Brit. Mus. Cat., iii. p. 457.

t Regan : " On the genus Lichia of Cuvier," Ann. Mag. Nat. Hist. Lotid. (Ser. 7X 12, 1903.

384 MR E. S. CLARK ON

PLEURONECTID.E.
Hemirliombus soleieformis (Agass.).

Gobius nigri Giinther.

DACTYLOPTERID^E.
Dactylopterus volitans (L. Gmel.).

BLENNIID.E.
Blennius sanguinolentus Pall.

IV. COLLECTION OP FISHES FROM ASCENSION.*

The trawling in 40 fathoms off Pyramid Point, Ascension, on June 9th, 1904,
revealed the presence of a large number of flat-fish. These were chiefly Rhomboid-
ichthys podas, though Hemirhombus soleieformis and Arnoglossus capensis were
tolerably abundant.

An interesting occurrence at this locality was Chtetodon dichrous, which Cunning-
ham t considered peculiar to the St Helena fauna.

In the list of twenty-six species, about 50 per cent, occur at the West Indies, a
third at the Cape of Good Hope, and a fifth at Cape Verde and Madeira islands.

LAMNID.E.

Lamna glauca Mull, and Henle.
Caught in trammel net on June 9th, 1904. Total length 79 inches; weight, 110 Ibs.j

Mursena moringa Cuv.

One large specimen, 1240 mm. in total length. Stomach contents: one complete
specimen of Holocentrum longipinne.

SCOPELID^;.

Trachinocephalus my ops (Forster).§
Saurus myops.

* Vide Giinther, Challenger Report, " Shore Fishes."

t "Fishes of St Helena," P.Z.S., 1910, part 1.

j Extract from Fish and Seal Log, kept by Dr Bruce and Mr 1). W. Wilton.

§ Vide Jordan and Evermann, Fishes of North and Middle America, 1896.

THE "SCOTIA" COLLECTION OF ATLANTIC FISHES. 385

SCOMBRESOCIDJE.

Exoccetus obtusirostris Gthr.

BERYCID.E.

Holocentrum longipinne Cuv. and Val.
sp. juv.

SERRANID^E.
Epinephehis ascensionis (Osbeck).

CAPROID.E.
Antigonia capros Lowe.

CH^TODONTIDJS.
Ch&todon dichrous Gthr.

Several specimens were taken in the trawl off Pyramid Point, Ascension. They
were kept alive in a tank on board the Scotia and presented to the Marine Station at
Millport, where they survived only for a short time.* These are wrongly named
Chelmon in the Zoological Log.

CARANGID/K.

Caranx ascensionis (Osbeck).
,, dentex Bl. Schn.
„ hippos (Linn.).
., crumenophthalmus (BL).
„ sanctse-helenss Temm. and Schleg.
Trachynotus glaucus (Linn.).
Lichia glauca.

SCOMBRID.E.
Scomber colias Gmelin.
TJiynnus albacora Lowe.

Three specimens were caught by line on June 9th, 1904, in Clarence Bay. The
largest weighed 105 Ibs., and measured 63 inches from tip of snout to tip of tail. t

PLEURONECTID^E.

Rhomboidichthys podas (De la Roche).
Hemirhombits soleseformis (Agass.).
Amoglossus capensis Blgr.

* Vide Zoological Log of Scotia.

t Extract from the Fish and Seal Log, kept by Dr Bruce and Mr D. W. Wilton.

386 THE "SCOTIA" COLLECTION OF ATLANTIC FISHES.

ECHENEIDID^E.

Echeneis remora Linn.

Scorp&na scrofina Cuv. and Val.

Batistes vetula Linn.
Monacanthus scriptus (Osbeck).

OSTRACIONTID^;.

Ostracion quadricornis Linn.

var. notacanthus Bleek.

DIODONTTD.E.
Diodon maculatus Lacep.

"SCOTIA" COLLECTION OF ATLANTIC FISHES. 387

V. — FISHES COLLECTED AT ST HELENA.

(MS. received 13th January 1913. Read 24th February 1913. Issued separately, 30W /«n*1913.)

THE 23 species here recorded were taken by the "Scotia" during a short
stay at St Helena on her return voyage from the Antarctic. Though all
have been previously described, yet, as a result of this capture, four now fall
to be included in the fish fauna of the island.

These are —

Gonorhynchus greyi.

Exoccetus speculiger.
Thynnus pelamys.
Sargus rondeletii.

Little importance can be attached to the occurrence of Exoccetus speculiger
and Thynnus pelamys, as these are typical open Atlantic forms and widely
distributed, though their appearance at this island seems rare.

As Mr J. T. Cunningham (1) has already stated, the fish fauna of the
island is extremely interesting owing to its wide relationships. Its members
show affinities with the fishes from Ascension, Cape Verde Islands, Madeira,
Azores, and the Mediterranean ; and also with those from Brazil and the
West Indies, and from the Cape of Good Hope.

To Dr Harmer, Keeper of Zoology, British Museum, I wish to express
my indebtedness for granting me all facilities to compare my material with
the type specimens ; and also my cordial thanks to Mr C. Tate Eegan, whose
method of description I have adopted throughout, for kindly assisting me
in the identification of several of the species, and for personal supervision
of my work.

The following are the "Scotia" species, with a detailed description of
two — Gonorhynchus greyi and Synodus synodus.

Gonorhynchus greyi (Richardson).

Depth of body 7| to 10 in the length ; length of head 4f to 5^. Snout
If to 24 as long as diameter of eye, which is 4^ to 5^ in length of head ;
interorbital width 4 to 5. Lips fringed and papillose ; barbel reaching
margin of upper lip. Gill rakers, 12 to 15 in lower anterior arch.

Dorsal 11 to 12 ; first ray three-fifths of second, which is about two-fifths
length of fourth ; fourth and fifth rays the longest, 1| to 2 in length of head.

(KEPKINTED FROM THE PROCEEDINGS OF THE ROYAL PHYSICAL SOCIETY, VOL. XIX., PP. 47-53).

388

MR R. S. CLARK ON THE

Origin of dorsal almost above end of base of pelvics. Anal 9 to 10. Origin
of anal equidistant from base of pelvic and base of caudal.

Pectorals 1 g to 1^ in length of head ; second and third rays the longest.
Pelvics 1^ to 2| in length of head. Caudal emarginate. Caudal peduncle
If to 2^ as long as broad.

FIG. l.
Side view of Gonorhynchus greyi.

170 to 200 scales in a longitudinal series ; 20 to 21 between origin of
dorsal and lateral line. Body yellowish brown — darker above than below.
Scales, especially above the lateral line, with minute dark spots. Fins,
except caudal, with terminal black spots, margined with white. Caudal with
central black blotch.

Description based on 6 specimens, 289 mm. to 437 mm. in total length.

These include specimens from New Zealand, Tasmania, West Australia,
Cape of Good Hope and St Helena. The " Scotia " collection is represented
by 1 specimen, 338 mm. in total length, from St Helena.

A list of measurements in millimetres is added to show individual
variations.

a

6

c

d

«

/

9

Length to base of Caudal .

403

371

338

286

279

260

268

Length of Head

80

79

69

55

52

54

63

Length of Snout

34

32

28

22

21

21

26

Length of Eye ....

15

16

13

12

11

12

13

Interorbital width .

16

17

15

13

13

13

16

Length of Pectoral .

55

59

48

44

39

42

50

Length of Pelvic

38

37

36

30

28

29

26

Length of longest Dorsal Ray .

40

41

40

31

29

30

33

Length to origin of Dorsal

277

244

230

197

196

183

194

(ROY. PHY. soc. PROC., VOL. xix., 48.)

"SCOTIA" COLLECTION OF ATLANTIC FISHES. 389

1 a Specimen from False Bay — Cape of Good Hope.

b „ „ East London, Cape Colony.

c „ „ St Helena—" Scotia."

d „ „ New Zealand.

e „ „ West Australia.

/ „ „ Tasmania.

g Gonorhynchus abbreviatus Schlegel (Fauna Japonica).

The measurements of Gonorhynchus abbreviatus Schlegel, are placed along-
side the others. There seems very little of a distinctive character except
the longer pelvics. The number of scales in a longitudinal series is 165.
The dark extremities of the fins seem to be deeper in colour, while the caudal
has a black spot on each lobe.

Synodus synodus (Linnaeus).
Saurus synodus Cuv. <fe Val.

Depth of body 4| to of in the length ; length of head 3£ to 34,. Snout
li to If as long as diameter of eye, which is 54 to 6f in length of head ;
interorbital width 7-J- to 8^. Upper jaw projects slightly. Praemaxillary
If to 1^ in length of head. Teeth in both jaws in a double row, lanceolate,
and with a tendency to arrow-shape. Teeth alternately long and short,
especially in upper jaw. Palatine teeth much more slender and in several
rows; inner rows longer, anterior teeth largest. Tongue covered with
retrorse teeth.

Top of head striated ; the striations few and very pronounced. Supra-
orbital bones prominent and slightly striated.

Gill rakers reduced to rows of minute teeth.

Dorsal 14-15. Origin of dorsal just behind base of pelvic.

Dorsal fiu equidistant from tip of snout and root of caudal ; fourth ray
the longest, 21 in length of head. First dorsal ray two-thirds length of the
fourth ; subsequent rays diminish in length. Adipose very small. Distance
from adipose to root of caudal three-fifths distance from adipose to base of
dorsal. Anal 9-10. Length of base half the length of dorsal base.
Pectorals 2+ in length of head. Pelvics eight-rayed ; sixth ray the longest,
which is seven-tenths length of head. Caudal forked.

60 to 62 scales in a longitudinal series ; 5 between origin of dorsal and
lateral line.

1 a, b, d, e, /, g are British Museum specimens.
(ROY. PHY. soc. PKOC., VOL. xix., 49.)

390

MR R. S. CLARK ON THE

Body and fins speckled dark grey on a silvery grey background ; black spot
on shoulder and on tip of snout ; several indistinct and irregular cross bars on
the body. Description based on six examples, 244 mm. to 274 mm. in total
length. The Indo-Pacific form (Saurus varius Glinther) is closely allied to
this species, but differs in having longer pectorals and pelvics, fewer dorsal

FIG. 2. Synodus synodus.
Reproduction of coloured drawing by Mr Cuthbertson— from fresh specimen.

FIGS. 3 and 4.
Side view of two specimens of Synodus synodus.

and anal rays, body more elongate, and coloration lighter. Saurus
meleagrides Val., from Buenos Aires, is also nearly related. The pectorals
are longer, and the dorsal and anal rays are different. It is figured in
D'Orbigny's Voyage dans I'Amtrique mtridionale.

There seems sufficient proof that the " Scotia " specimens are identical
with those described from St Helena and from Bahia by Cuvier and

Valenciennes.

(ROY. PHY. soc. PROC., VOL. xix., 50.)

"SCOTIA" COLLECTION OF ATLANTIC FISHES.

391

The following are a few of the measurements of the specimens in milli-
metres, to show individual variations : —

a

6

c

d

1

/

Length to base of Caudal .

254

253

250

250

245

226

Length of Head

72

64

70

69

67

59

Length of Snout

19

18

18

17

16

15

Length of Eye ....

11

11

12

11

10

10

Interorbital width .

8

9

8

7

7

7

Tip of Snout to origin of Dorsal

101

103

105

100

100

89

Length of Pectoral .

28

29

28

26

25

24

Length of Pelvic

53

56

55

53

46

45

Length of Praeiuaxillary .

48

43

47

46

43

38

I

Aulostoma coloratum Mull. & Trosch.

2 specimens, young aiid adult, taken off St Helena, 171 mm. and
566 mm. in total length.

Exocoetus obtusirostris Giinther.

Dorsal 13 ; anal 13. Depth of head equal to distance from extremity
of snout to hind margin of prjeoperculum. Snout f diameter of eye, which
is 3| in length of head. Origin of dorsal opposite origin of anal.

1 specimen, total length 137 mm.

Exocoetus speculiger Guv. <k Val.

Depth of body 6 in the length ; length of head 4|. Snout equal to
diameter of eye, which is 3J in length of head ; interorbital width slightly
more than diameter of eye.

Dorsal 10 ; first pectoral ray over half the total length of the pectoral fin,
which reaches just beyond the extremity of the dorsal. Anal 11. Origin of
anal slightly in advance of the dorsal. Pectoral on upper side blackish, with
an oblique white band and white margin. 1 specimen, total length
240 mm.

Holocentrum longipinne Bl. Schn.

Several specimens averaging 230 mm.

(KOY. PHY. soc. PROC., VOL. xix., 51.)

392 MR E. S. CLARK ON THE

Epinephelus ascensionis (Osbeck).

1 specimen, total length 203 mm.

Rhypticus saponaceus (Bl. Schn.).

Dorsal III 23. Anal 16.
. The maxillary in this specimen reaches to two-thirds diameter of eye.

Priacanthus cruentatus (Lacep).

First few dorsal spines rather low. Last spine twice as long as the second.
All are strongly spinulose. liays not much larger than longest spines.

2 specimens, total length 210 mm.

Sargus capensis Smith.
Several specimens, total lengths 232 mm. and 280 mm.

Sargus rondeletii Guv. & Val.
2 specimens, total lengths 176 mm. and 205 mm.

Chastodon sanctee helenae Giinther.

A painting of this specimen by Mr Cuthbertson, the "Scotia"
artist, represents the colour of the body as a light lavender, which becomes
darker towards the vertical fins. A narrow yellow band runs from the
supra-orbital along the dorsal fin, across the caudal, and ends at the origin of
the anal. The soft dorsal and anal fins have a thin black margin.

Cossyphus pectoralis (Gill).
1 specimen, total length 255 mm.

Scarus strigatus Giinther.
Several specimens averaging 350 mm.

Caranx dentex (Bl. Schn.).
1 specimen, total length 594 mm.

Caranx sanctse-helenee Guv. <fe Val.
Numerous young specimens.

Trachynotus glaucus (Linn.).
Lichia glauca (Linn.).

1 specimen taken off St Helena, total length 333 mm.
(ROY. PHY. soc. PROC., VOL. xix., 52.)

"SCOTIA" COLLECTION OF ATLANTIC FISHES. 393

Scomber colias Gmelin.
Scomber pneumatophorus De la Roche.

Several specimens, young and adult.

Thynnus pelamys (Linn.).
1 specimen, total length 424 mm.

The Belted Bonito has a wide tropical and sub-tropical distribution. It
has been recorded from Luce Bay in the South of Scotland.

Thyrsites prometheus (Cuv. <fe Val.).
1 specimen, total length 285 mm.

Scorpsena mellissii Giinther.
1 specimen, total length 217 mm.

Scorpaena scroflna Cuv. <fe Val.
Several specimens, 249 mm. to 305 mm. in total length.

Tetrodon sp.

Probably T. cutaneus, but too mangled for identification. This was
found in the stomach of Thynnus pelamys.

BIBLIOGRAPHY OF ST HELENA FISHES.

1. CUNNINGHAM, J. T. On Marine Fishes and Invertebrates of St Helena, Proc.

Zool. Soc. London, 1910, part 1.

2. CUVIER ET VALENCIENNES. Histoire naturelle des Poissons, Paris, 1828-1849.

3. GUNTHER, A. Report on a Collection of Fishes made at St Helena, by J. C.

Melliss, Esq., Proc. Zool. Soc. London, 1868.

4. MELLISS, J. C. St Helena: a Physical, Historical and Topographical Descrip-

tion of the Island, including its Geology, Fauna, Flora and Meteorology,
London, 1875.

(ROY. 1'HY. SOC. 1'ROC., VOL. XIX., 53.)

394

MR R. S. CLARK ON THE

VI. — COLLECTION OF FISHES FROM CAPE COLONY.

Thirty-seven species were secured in May 1!)04 chiefly from trawlings made at
Table Bay, False Bay, oil' Dassen Island and at Saldauha Bay.

The collection contains a large number of specimens of sharks of the families
Carchariidat, Scyliorhinidce and Squalidce.

Two species have not been previously recorded from the Cape, — Mustelus
maculatus and Scatharus grcecus — while a third, Caranx dentex, has been con-
sidered a doubtful record.

The occurrence of Mustelus maculatus at Station 483 is extremely interesting,
as the locality of this species was hitherto unknown.1

All the others have been recorded either from the Cape, by Dr Gilchrist,2 or
from Natal, by Mr C. Tate Regan.3 Three species are recorded by Dr Giinther in
the Challenger Report on Shore Fishes.

The following details are extracted from the Station Log of the Scotia, and
are inserted here for greater convenience.

Station

Time
May 1904

Locality

Depth in
Fathoms

Method of
Capture

Remarks

478

May 5 to 17

Table Bay

...

...

...

479

,, 6 to 16

False Bay

480

„ 18

Off Dassen Island

•••

...

481

,, 18 to 21

N.W. of Ijzer
Fontein Point

35

Otter Trawl

Sand

482 («)
fr)
(c)

„ 18 to 21

Houtjes Bay and
off Salamander Pt.

8
5
2

Trawl
Trap
Trammel

Shells (a)
Sand (6)
Sand (c)

483

„ 21

Entrance to
Saldanha Bay

25

Otter Trawl

Sand and
Kelp

BDELLOSTOMATIU^;.
Bdellostoma cirrkatum Forst.
Station 482 (b).

SCYLIORHINID^E.

Scyliorhinus africanus (Gmel.).
Station 483.

Scyliorhinus edwardsii (Cuv.).
Stations 482 (a) and 483.

1 Brit. Mus. Cat., vol. viii., p. 388.

2 Marine Investigations in South Africa, vol. i. Annals of the South African Museum, vol. vi., pts. 2 and 3.

3 Annals Natal Government Museum, vol. i., pt. 1, 1906 ; pt. 3, 1908.

COLLECTION OF FISHES FROM CAPE COLONY. 395

CARCHARIID^;.
Mn.<tt'li(* la'rifi Risso.
Station 483.

Mit*t<-1n.< iiKU-nlatHf Kner. & Stein.

Description similar to that given in the British Museum Catalogue, vol. viii.,
p. 387.

Black spots fairly large.
Station 483.

SQUALID^;.

SqiKtlii* acutipiniii* Regan.
Stations 482 (a) and 483.

RAIID.E.

Raia batis Linn.

This young specimen is similar to the adult, now in the British Museum, which
was recorded by Dr Gilchrist from the Buffalo River locality.
Station 482.

Raia sp. juv.

Probably Raid rhizacanthm Regan.
Station 482.

TQBPEDIHIDJL

Torpedo marmorata Risso.
Station 483.

CHDCJtRmJE.

CaOorhynckiu a>itarcti<-><.< Lacep.
Station 483.

SlLURID^E.

Galeichthys feliceps Cuv. & Val.
Station 483.

SYXGXATHID.E.
Syiifitxithiix in-Hit Linn.
Station 482 (b).

396 MR R. S. CLARK ON THE

At/ierina bremceps Cuv. & Val.
1 young specimen with monstrosity in lower jaw.
Station 482 (c).

MUGILID.E.
Mugil capita Cuv.
Station 478.

GADID^E
Merlueeius capensis Regan.

Stations 481 and 483.

Genypterus capensis Smith.

Stations 481 and 483.

Scicena aquila Risso.
Station 483.

SPARIDJE.

Box salpa Linn.
Station 478.

Scatharus grcecns1 Cuv. & Val.
Station 482.
This Mediterranean species has not been previously recorded from the Cape.

Sarffus cermnus Lowe.
Station 481.

Sargus nigrofasciatus Regan.
Previously recorded only from Natal.
Station 482 (b).

Pagrus laniarim Cuv. & Val.
Station 478.

CARANGID.E.

Caranx dentex (Bl. Schn.).
One specimen was secured at Station 478.

1 It is possible, though scarcely probable, that a confusion of labels may have here occurred, for the
species is only previously recorded from the Mediterranean.

COLLECTION OF FISHES FROM CAPE COLONY. 397

SCOMBRID^.

Scomber colidf Gmelin.
Station 479.

TlUCHIURIDjE.

Lepidopw cat/<l<(t".< (Euphrasen).
Station 481.

Thyrsites atun (Euphrasen).
Station 481.

PLEURONECTID^E.

Cynoglofsus capetms Kaup.
Stations 478, 481, 482, and 483.

Synaptura microlepis Blkr.
Stations 478 and 483.

GOBIHXE.

Gobi/i* iimlici'ps Cuv. & Val.
Station 479.

TRIGLID.E.

Tnglaperonii Cuv. >S: \*al.
Stations 482 and 483.

GOBIESUC1D-K.

Chvrisochimiw tlctitex Pall.
Station 482 (r).

BLENNID.K.

Clintf* atipi'n-ili >v/.< Linn.
Stations 478 and 482 (c).

Clitntit acuiitiit'itiis Cuv. & Val
Station 482 (c).

Clinus dorsalis Bleeker
Station 482 (c).

398

MR R. S. CLARK ON THE

OSTRACIONTID.E.
Ostracion diaphanus Bl.
Station 483.

TETRODONTID.E.
Tetrodon honckenii Bl.
Station 479.

DlODONTID^E.

Diodon maculatus Gthr.
Station 482 (a).

VII. — SHARKS FROM ST PAUL'S ROCKS.

At St Paul's Rocks, Scotia Station 53, eight specimens of Cnrcharias menisorrah,
Mull, and Henle, were captured on 10th December 1902. These sharks have been
reported from this locality by various expeditions, but their identification has not
been definitely recorded. Carcharias menisorrah, though an Indian species, seems
therefore to be cosmopolitan. The dimensions of these specimens are given just as
they were tabulated by Dr Bruce and Mr D. W. Wilton in the Scotia Fish and Seal
Log. For permission to reproduce the photograph of St Paul's sharks from "Three
Voyages of a Naturalist," I am indebted to the author, Mr M. J. Nicoll, while I
wish also to thank Messrs Witherby for the use of the half-tone block.

Several small specimens of Echeneis remora were found attached to these sharks.

SHARKS FROM ST PAUL'S ROCKS.

399

Measurements in inches of 8 specimens, and weights in Ibs. of 5 specimens of
Carcharias menisorrah Miill. and Henle.

A?

«

c,

D?

E?

F?

G?

H

a

78

73

65i

54

59

65

62J

52i

b

57

53i

46}

391

43

47

45

38J

c

i

7

i

f

i

7

f

f

d

10

91

81

H

*\

H

7

6|

e

8

?!

6

5

5i

5

5

*i

f

H

2^

21

u

If

If

4

If

9

16

13}

13

8

1U

10J

10

7f

h

n

H

H

4

5i

5

5

H

1

j

2i

2

if

If

4

4

B-

If

i

3

2T

H

e

21

if

if

i

4

k

...

...

2 38

45

43

21

A, B, C, etc. Xuraber of specimens.

a = Tip of snout to tip of tail.

b = Tip of snout to root of tail.

c = Diameter of eye.

d= Interorbital space.

e = Breadth of angle of lower jaw.

/= Length of 1st gill slit.

g = Length of pectoral fin.

A = Length of base of 1st dorsal fin.

t = Length of base of 2nd dorsal fin.

j = Length of base of anal fin

k = Weight in Ibs.

B. — PELAGIC FISHES.

This list contains only a small percentage of the surface fishes, as most of
the larval forms obtained in the tow-nettings could not be identified with any

degree of accuracy.

SCOPELID^;.

Myctophum punctatum (Rafinesque).
29° 54' N. 34° 10' W. Station 537.

CESTRISCIIXS:.
Centriscus scolopax L.
34° 2' S. 49° 7' W. Station 98.

Syngnathus pelagicus L.
33° 53' N. 32° 27' W. Station 539.

400 MR R. S. CLARK ON THE

SCOMBRKSOCIDTE.

Hemirliamphm sp. juv.
15° 15' N. 25° 9' W. Station 25.
T 20' S. 34° 38' W. „ (51.

Exocmtm nigricans Benn.
18° 28' N. 24° 28' W. Station 21.
This specimen was picked up on the deck of the Sfotia.

Exoccptus obtusirostris Gthr.
Station 21. 18° 28' N. 24° 28' W.
Several secured on deck of the Scotia.

Exoccetus evolans L.

18° 28' N. 24° 28' W. Station 21.
5° 25' N. 26° 7' W. Station 42.

Exoccetus speculiger Guv. and Val.
1° 22' N. 28° 10' W. Station 52.

CARANGIDJE.

Nauc rates ductor juv. L.
Station 521. 5° 47' N. 20° 28' W.

ECHENEIDIDJE.

Echeneis remora juv. Linn.
Station 70. 9° 6' S. 34° 38' W.

DACTYLOPTERIDjE.

Dactylopterus volitam juv. (L. Gm.).
Station 82. 20° 40' S. 38° 20' W.
87. 24° 42' S. 40° 34' W.

ANTENNARIID^E.

Antennanus marmoratns juv. (Shaw).
Station 539. 33° 53' N. 32° 27' W.

TETRODONTIDJE.

Tetrodon sp. juv.

Station 67. 7° 20' S. 34° 38' W.
Station 78. 15° 24' S. 37° 12' W.

PELAGIC FISHES. 401

-

Linn.
Station 106. 39' 01' S. 53 40' W.

Orth<i{i<n->#-<i* iitof'i L.1

The following measurements (in inches) are from the Scotio /•'/>•// <ut<l S«/I Log.-

Length (total) . . 80

Depth . . 51

Girth ... 120

Dorsal fin 34

„ base . .16

Anal fin 31

„ base . . 13

Pectoral fin ... 9g

„ base . - 6

Length from eye to pectoral fin 12

Diameter of eye . 3^

Length of gill slits . . 3^

Weight . . . . \ ton

Two specimens of Mii-m-i* remora L. were found attached to the inside of the

mouth.1

1 Vide Zoological Log, pi. iv., p. 5.

• Fish and Stal Log, by Dr W. S. Bruce and Mr D. W. Wilton.

PART XVII.
PTEBOBBANCHIA,

XVIL-THE PTEBOBRANCHIA OF THE SCOTTISH
NATIONAL ANTARCTIC EXPEDITION.

BY S. F. HARMER, Sc.D., F.R.S., Keeper of the Department of Zoology in the British
Museum; and W. G. RIDEWOOD, D.Sc., Lecturer on Biology in the Medical
School of St Mary's Hospital, University of London.

(WITH FIVE TEXT FIGURES AND TWO PLATES.)

The Pterobranchia of the Scottish National Antarctic Expedition (1902 to
1904). By S. F. Harmer, Sc.D., F.R.S., Keeper of the Department of Zoology
in the British Museum ; and W. G. Ridewood, D.Sc., Lecturer on Biology in the
Medical School of St Mary's Hospital, University of London. Communicated by
Dr J. H. ASHWORTH. (With Five Text-Figures and Two Plates.)*

(MS. received February 15, 1913. Read March 17, 1913. Issued separately July 4, 1913.)

INTRODUCTION.

The genus Cephalodiscus was instituted by M'!NTOSH for a species, C. dodecalophus,
which had been obtained by the Challenger Expedition. After the publication of
the full account of this species, by M'!NTOSH and HARMER (87) in the Challenger
Report, succeeding papers for nearly twenty years were all based on the original
Challenger material. The subject was in particular re-investigated by MASTERMAN
in a series of papers (971, 972, 98, 99, 03). The Siboga report, published by
HARMER (05) in 1905, added three Oriental species to the genus. SCHEPOTIEFF (05,
07, 08) devoted several papers to a further description of the Challenger material ;
while, more recently (09), he has described an interesting new species, C. indicus,
from Ceylon. In 1906 RIDEWOOD (06) described C. gilchristi from South Africa;
and in the following year he gave an account (071) of the two species which had been
dredged by the Discovery Expedition ; and (072) of the development of the plumes in
four species of Cephalodiscus. One of the Discovery species, C. nigrescens, had been
described two years earlier by LANKESTER (05), in a preliminary paper ; and RIDEWOOD
( 1 2) has recently brought forward evidence to show that this species had been dredged
by the Erebus and Terror Expedition in 1841 or 1842. In 1907 ANDERSSON (07)
added no less than six species to the genus, in describing the results of the Swedish
South Polar Expedition, though we give some evidence (pp. 435-439) to show that one
of them, C. inxquatus, is synonymous with C. hodgsoni, which had been described
by RIDEWOOD (071) earlier in the same year, from the Discovery collection. The
most recent addition to the list of species has been given by GRAVIER (12), who has
published an account of C. anderssoni, a new species which was obtained by the second
French Antarctic Expedition.

Full references to the literature of the subject, up to the dates of the respective
publications, have been given in the works of HARMER (05), RIDEWOOD (071), and
ANDERSSON (07). A general account of the group has more recently been given by
SPENGEL (12).

The characters of the coenoecium are so well marked that it appears justifiable to

* Published by permission of the Trustees of the British Museum.
(REPRINTED FROM THE TRANSACTIONS OF THE ROYAL SOCIETY OF EDINBURGH, vox,. XLIX., pp. 531-565.)

408 DR S. F. HARMER AND DR W. G. RlDEWOOD ON THE

use them, as a rule, as a ready means of distinguishing one species of Cephalodiscus
from another. It is remarkable that what one would suppose to be the uncorrelated
efforts of the numerous zooids of a colony do in fact produce so uniform a result.
The comparison of larger series of specimens may perhaps show in the future that
the coenoecial characters are less reliable than they appear to be at present ; but the
fact remains that the common house of the colony has, as a rule, so distinct a character
of its own that it is difficult to believe that it cannot be used for systematic purposes.
It has thus been possible to distinguish the following subgenera of Cephalodiscus :—

(1) Demiothecia, Hidewood (071), in which the coenoecial cavity is continuous and the
zooids occur separately or in groups in any part of it, being free to wander about in it.

(2) Idiothecia, Kidewood (071), in which each zooid, with a certain number of its
buds, occupies an independent tube-like cavity in the ccenoecium.

(3) Orthoecus, Audersson (07), in which each zooid has a tube of its own, but
the tubes are free for the greater part of their length, instead of being embedded in
the common ccencecial mass as in Idiothecia.

The descriptions which have been published by the observers who have been
referred to above show that there is a singular uniformity of structure in the zooid
throughout the genus. In such fundamental characters as the three divisions of
the body, and their associated ccelomic cavities, the notochord, the proboscis-canals,
the collar-canals, and the gill-slits, there is practically no variation. The remarkable
character of the male C. sibogse, as described by HARMER (05), indicates that in that
species at least there is a striking sexual dimorphism. In those other species in
which both sexes are known there appears to be no essential difference between
the two sexes except as regards their gonads. An exception must, however, be made
in the case of C. insequatus (probably = C. hodgsoni], where most of the female zooids
have five pairs of arms, while most of the males have six pairs (see pp. 436-438).

Making use of the coencecial characters which have been mentioned above, it
appears to be possible to distribute the known species of Cephalodiscus among the
three subgenera there indicated, and in this way to separate species which it might
be hard to characterise from the structure of their zooids. But within a single sub-
genus it becomes necessary to rely more largely on the characters of the zooid ; and
the discrimination of species on anatomical grounds may offer considerable difficulty.

In the earlier accounts of species of Cephalodiscus some stress was laid on the
proportions of the zooid and of its stalk. This procedure has been criticised by
ANDEUSSON (07), who has had the unique opportunity of examining certain species in
the living state. When account is taken of the highly muscular character of the
zooid, and of the evidences of contraction afforded by wrinkling of the skin of
the stalk and other parts, it is easy to believe that ANDERSSON'S criticism is to a large
extent well founded. It may, nevertheless, be true that some weight may be ascribed
in certain cases, as in discriminating between zooids in a similar degree of contraction,
to features of this kind.

(ROY. soc. EDIN. TRANS., VOL. XLIX., 532.)

PTEROBRANCHIA OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 409

The difficulty of finding reliable specific characters in the structure of the zooid is
increased by the fact that the material has as a rule not been preserved with any
special refinements of technique. The study would be greatly facilitated by being
able to make use of specimens which had been preserved in a fully extended condition.
It would then be comparatively easy to ascertain the number of the arms or plumes
with certainty. Under other circumstances this point, for instance, cannot always be
made out without the possibility of mistake. Even in a well-prepared series of
sections it is sometimes excessively difficult to count the arms, which may be cut in
planes most unfavourable for study. The method of dissection is capable of giving
valuable evidence ; but here, too, mistakes are not out of the question. There is
reason to believe that in some species of the genus the number of arms is variable
(cf. RIDEWOOD, 071). It must also be remembered that in some cases the number of
the arms has been given as the result of the study of sections of a very small number
of individuals. The comparison of series of sections of the same species might
suggest differences which are really due to the degree of maturity or of contraction,
or to the planes in which the sections are cut. Some caution is thus necessary
both in making statements and in accepting those which have been made by other
observers.

It can hardly be doubted, however, that the number of pairs of arms does provide
a character which is of value systematically. Other characters which can be specially
relied on are probably : — the number of buds, whether few or many, which are borne
at any one time by the budding region of the stalk ; the presence of one or three
nerve-tracts in the stalk, as pointed out by HARMER (05) and ANDERSSON (07) ; the
presence or absence of deep pigmentation of the skin ; the shape of the operculum
or postoral lamella ; the size of the free ova and the structure of the embryos ; and
the mode of development of the arms in the buds.

Using some of these characters, the single species dredged by the Scotia may be
thus characterised :—

Cephalodiscus agglutinans, n. sp. Colony massive, branching, somewhat resembling
that of C. nigrescens in the size of its branches, but the material of the ccenoecium
includes large quantities of shells of Foraminifera, small fragments of shells of Mollusca
and spines of Echinoids, and rounded particles of slate. The pieces of colony vary in
size up to 100 or 115 mm. in greatest length, and 45 or 55 mm. in greatest breadth.
Cavity of the cceuoecium in the form of a tubular labyrinth continuous throughout
(except for a few isolated septa), much branched in the interior, but the superficial
parts of the tubes are radially set, and open at a fairly definite angle (about 80°) to
the surface. Diameter of the peripheral tubes 1 mm. Ostia elliptical, size 1'2 by
•8 mm., each with a single thick lip or spine* projecting about 3 mm. beyond the
general surface. No peristomial tubes. Mean distance from the middle of an ostium
to the middle of the ostium nearest to it, 3 or 4 mm. ; at the free extremity of the

* The spines are broken off in the greater part of the material examined.
(ROY. soc. EDIN. TRASS., VOL. XLIX, 533.)

410 DR S. F. HARMER AND DR W. G. RIDEWOOD ON THE

branch, however, the ostia are more crowded. Length of zooids from the free ends of
the arms to the end of a fairly extended body, 4 '5 mm. ; length from base of arms to
end of body, 3'2 mm. ; width of body, '8 mm., but if the body is much contracted its
width is that of the tubes, i.e. 1 mm. Arms usually nine pairs, but often fewer and
exceptionally more ; no end-bulbs with refractive beads. Stomach and the succeeding
U-shaped intestinal loop long. Females not known ; and no buds having more than
five pairs of arms have been found.

MATERIAL.

The specimens of Cephalodiscus submitted to us for study were sent in six bottles,
five of which were of 2^ litres capacity, and the sixth of about half that size. This
material was dredged on December 1, 1903, and was obtained in a single haul on the
Burdwood Bank, to the south of the Falkland Isles (Station 346 ; lat. 54° 25' south ;
long. 57° 32' west; fathoms 56). The contents of the trawl on this occasion
amounted to about half a ton,* the largest and richest catch of the expedition ; and
the members of the staff worked up to two o'clock in the morning sorting and bottling
the specimens. All the Cephalodiscus material was preserved in alcohol, and no
special measures were taken for killing the zooids in an extended condition.

CCENCECIUM.

The distinguishing feature of the present species, the agglutination of foreign
particles in the soft substance of the crenoecium, is apparent at a glance, and the
semitransparent or translucent appearance of the co?noecium of other species of
Cephalodiscus is only to. be detected in certain parts of branches where the
inclusions are more scanty than usual. The general appearance of pieces of colony
is whitish or whitish yellow, with dark spots which mark the ostia or openings
of the tubular cavities inhabited by the zooids (text-fig. 1, A and B). The branches
are massive, resembling those of C. nigrexcens in size and bulk. The substance of
the branch is weak and easily crumbled, the included shelly particles adding to
the weight and not adding to the strength of the coenoecium. A branch taken out of
alcohol and held horizontally in the hand by one end breaks across by its own weight.

The majority of the pieces of colony are about 80 to 90 mm. in length, and
oval in cross section, measuring about 30 mm. in the long diameter and 20 mm.
in the short diameter. Some of the small pieces measure about 70 mm. by 20 mm.
by 15 mm. The largest pieces measure about 100 to 115 mm. long, and have three
or even four branches. The total width of such branching pieces of colony is
45 or 55 mm. On Plate I. are represented five pieces which are the largest in the
whole collection and best suited for showing the characters of the branching. Since
these are drawn of the natural size, a detailed description of them individually is

* Report of the Scientific Results of the Voyage of the S.Y. "Scotia," 1902-4, vol. iv., Zoology, part i., Zoological
Log, 1908, p. 61.

(ROY. soc. EDIN. TKANS., VOL. XLIX., 534.)

PTEROBRANCHIA OF TEE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 411

unnecessary. Most of the pieces of material show no branching, but resemble the
piece of which a photographic reproduction is shown in text-fig. 1, A.

The surface of the colony appears rough, and, if the ostia or openings of the tubes
are not clearly visible, does not differ much from the surface presented when a piece
is broken across (text-fig. 1, D and E). But in sheltered situations, as between the
branches of a piece of colony, such as is shown in PI. I. figs. 1, 4, there are blunt

A

B

D

m

TEXT-FIG. ]. — Photographic reproduction of pieces of colony of C. agglutinans ; of the natural size. A, a piece broader at
the free end (uppermost in the figure) than at the basal end, showing no branches ; B, a piece showing a branching into
two ; C, a longitudinal median section of a piece of colony ; D and E, transverse sections through a piece of colony. The
darker areas are the tubular spaces occupied by the zooids, the white specks are small pieces of shell included in the
coanoecial substance.

brownish processes or "spines" projecting to an extent of 3 mm. beyond the general
surface. Inspection shows that these are really the projecting lips of the inhabited
tubes, one to each tube, similar to the lips that occur on the surface of a piece of
C. nigrescens (07, pi. iv. fig. 11 ), but differing in being less regular in form. These spines
are built up by the superposition of solid caps of coenoecial substance upon the summit
of pre-existing spines, just as are the long spines of species of Cephalodiscus such as
C. dodecalophus, C. hodgsoni (07, pi. iv. fig. 21), and C. gilchristi (06, pi. iii. figs. 9,
10, 11), but they differ in not exceeding a length of 3 mm., and in being restricted in

(ROY. soc. EDIN. TRAXS., VOL. XLIX., 535.)

412 DR S. F. HARMER AND DR W. G. RIDEWOOD ON THE

number, one to each ostium. They further differ in that they include shelly particles
(see text-fig. 3), although these are present to a less extent than in the general
substance of the coenoecium. They are fairly pointed at the extremities, although the
actual shape may be the result of the size and form of the included shelly particles,
and they broaden out at the base and curve round the ostium for about one third of
its margin. In protected situations the inhabited tubes are seen to widen out suddenly
as they reach the surface, and resemble very short funnels, and it is from the rim of
the funnel that the lip projects. This accounts for the distance between the spine and
the tube in the surface-view of the branch shown in text-fig. 2, B.

A careful examination of the outer surface of a piece of colony and a dissection into
the substance of the coenoecium between the tubes reveals the presence of broken
spines in association with the several ostia, and leads to the conclusion that the whole
of the surface of the colony is in the natural state provided with short lips or spines,
but that, owing to the rough treatment which the specimens must have experienced in
coming up in the trawl with a great weight of other specimens, their surface has been
scoured and deprived of the projecting spines, which are now only to be seen in their
full development in protected situations between branches which have not been broken
apart, as in PI. 1. fig. 1.

Of particular interest in this connection is the fact that although in G. agglutinans
the zooids do not live in isolated tubes, but in tubes forming parts of a common tubular
system, the single lip or spine to each ostium is in other known species of Cephalodiscus
only met with in species of the subgenus Idiotkecia (C. gilchristi, G. nigrescens, and
C. levinseni) and Orthoecus (C. solidus), whereas in the known species of Demiothecia
the spines are multiple, four or five being present around each ostium in the case of
C. liodgxoni, for instance.

The tubular cavities which open to the exterior, and in which the zooids occur, are
approximately 1 mm. across, and the average distance from the middle of the orifice
of a tube to the middle of that nearest to it is 3 or 4 mm., except at the free end of a
branch, where the tubes are more crowded. The walls immediately bounding these
cavities are composed of a thin layer of yellowish-brown coenoecial substance, much
tougher than the jelly-like albuminoid that fills in the intervals between the tubes and
makes the whole so compact. The tubes proper do not include in their walls any of
the foreign bodies that give to the coenoecium of this species its characteristic appear-
ance ; but some of the shelly particles of the softer material of the coenoecium are so
tightly fixed to the outer surface of the tubes that in dissecting a branch the tubes are
liable to be torn in the attempt to remove the particles.

The tubes in the outer part of a piece of colony are fairly uniform in diameter (one
millimetre), except at the junction of two tubes, where irregularities in shape and size
may occur. A longitudinal section of a branch shows that the peripheral parts of the
tubes are not set at right angles to the surface, but slope somewhat towards the free
end of the branch (see text-fig. 1, C). In the middle of the piece the tubes are very

(HOY. soc. BDIN. TRANS., VOL. XLIX., 536.)

PTEROBRANCHIA OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 413

irregular iu their disposition, but the cavities may still be termed tubular in spite of
irregularities iu their shape.

The internal tubular system is continuous throughout the whole branch, and if it
were not for probable resistance offered by other zooids one could conceive of a zooid
travelling internally from any one part to any other part of the system. In the middle
portion or core of a branch the tubes not only exhibit branching, but here and there a
complete tubular circuit can be seen, as is shown at b, b, b, b in the diagrammatic text-
tig. 2, A. The existence of such continuous circuits is explained by the part of the
circuit nearest the base of the branch being due to the bifurcation of a tube during an
early stage of the growth of the branch, when this part occupied an apical position ;
whereas the part of the circuit farthest from the base of the branch is due to the bury-
ing of a superficial groove that connected two young ostia at a time when this part
of the branch constituted the apex. There can be no doubt, from a study of the apex
of a branch, that growth is apical. The tubes that open to the surface at the apex are
much shorter and closer together than in other parts of the branch, although of the
same diameter. Evidence of the branching of a tube at the apex is forthcoming, and
also evidence of the enclosure of a superficial groove between two young ostia. such as
produces the continuous circuits above referred to.

The ostia show on the surface of a piece of colony as dark areas, due partly to the
fact that one is looking into a tubular cavity, and partly to the fact that the actual wall
of the tube does not contain the shelly particles that make the softer substance of the
ccenrecium .so white in colour. In rare cases in the material studied the zooid occupy-
ing a peripheral tube could be seen without cutting into the branch, but in most cases
the zooids had all retreated into the middle parts of the tubular system, and had become
entangled and intertwined with one another. The ostia proper are slightly funnel-shaped
and oval, measuring about 1*2 nine, in long diameter and '8 mm. in short diameter,
whereas the tube itself is roughly circular in section, and of about 1 mm. diameter.

So far as can be seen from the study of the surface in protected parts between two
branches that have not broken apart, the tubes do not project beyond the general sur-
face, as "peristomial tubes," in the manner of the tubes of C. levinseni (05, pi. ii. fig. 11)
and the tubes in certain parts of the colony of C. nigrescens (07, pi. iii. fig. 5), and this
conclusion is supported by a study of the short tubes at the free extremity of a branch.
The filling up of the intervals between the new tubes seems to take place pari passu
with the additions made to the mouths of the tubes themselves. (Incidentally it may
here be mentioned that the " habit " or " facies " of a colony of Orthoecus differs from
that of a colony of Idwthecia owing to the peristomial tubes of the former being of
extraordinary length ; indeed, in C. rarus the whole colony consists of peristomial tubes
with only a small amount of softer coenoecial substance binding the blind ends of the
tubes together.)

The distinction between the sub genera Idwthecia and Demiothecia was drawn in
1907 (07 *, p. 7) because the isolation of the zooids in separate tubes in the coanoftcium

(BOY. soc. KDIN. TEAKS., VOL. XLIX., 537.)

414 DR S. F. HARMER AND DR W. G. RIDEWOOD ON THE

of the species grouped in the former subgenus appeared to be of systematic importance.
The study of the present material, however, goes some way to break down the distinction.
In Cephalodiscus agglutinans the internal system of spaces in the coenoecium is con-
tinuous throughout (except for occasional septa, see below), and zooids can move fairly
freely through the system, as may be seen by the manner in which they have retreated
to the internal parts of the branches in the material under consideration. So far as one
can judge from a study of pieces of colony preserved in alcohol, the growth of a branch
in a species of Idiothecia is by the buds (or possibly larvae) settling at the apex of the
branch, and each secreting a tube of its own, independent of those already existing,
whereas in the present species the (presumably young) zooids at the apex secrete tubes
continuous with the existing system of spaces within the branch. But C. agglutinans
differs from all known species of Demiothecia in having long tubes, i.e. spaces of
approximately even calibre extending for considerable distances ; for in the recorded
species of Demiothecia there is a single irregular branching cavity in the interior of the
piece of colony, and this opens abruptly to the exterior here and there on the surface of
the branch, or by means of very short tubular passages (see G. hodgsoni, 07 *, pi. iv.
fig. 22).

The disposition of the tubes within a branch of the colony may be studied by dis-
secting away the soft part of the ccenoecium and the included shelly particles, but the
process is laborious and not altogether satisfactory in other respects. Much better
results can be obtained by decalcifying a branch in a 1 per cent, solution of nitric
acid for ten days or a fortnight, and then bisecting the branch lengthwise and dissect-
ing out such of the tubes as are laid bare. The process of decalcification removes all
the shelly particles, and leaves only the grains of slate (see p. 418), which are not
numerous and can be easily picked out. The piece of colony after decalcification is of
a dirty brown colour, and is translucent, without the opacity and whiteness which is
such a marked feature of the ccenoecium before it is placed in acid. In text-fig. 2, A,
is shown a diagrammatic representation of the cut surface of such a bisected branch.
(Compare this with the diagrammatic longitudinal section of a branch of C. nigrescens,
in 071, pi. iv. fig. 10.)

This diagram (text-fig. 2, A) explains the main features of the tubular passages of
the ccencecium. It shows that in the most superficial parts the tubes are fairly straight,
and of uniform diameter, and that they slope more or less radially outwards and incline
somewhat towards the apex. The tubes in the middle are irregular in their arrange-
ment, but on the whole are set at a uniform distance from one another. What appear
to be blind ends of tubes at d and d' are merely parts of tubes that are cut across
obliquely because they are leaving the plane of section.

In some of the passages, especially in the more basal parts of a branch, as at s and
s', there are thin concavo-convex partitions or septa, sometimes two or three in close
succession. These septa, however, are not common. Their concave faces may be
directed towards the apex of the branch, as at s', or away from it, as at s.

(ROY. soc. EDIN. TRANS., VOL. XLIX., 538.)

PTEROBRANCHIA OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 415

In the apical region of a branch, as shown in the upper part of text-fig. 2, A, the
ostia are closer together than on the sides of the branch, and the tubes that open to
the exterior are much shorter than elsewhere ; but the tubes are of the same calibre as
in the other parts of the branch. The substance of the coenoecium is more delicate
and the lining of the tubes thinner and less brown in colour than in other parts, and
the tubes have no spines. These circumstances lead to the conclusion that the growth
in the length of a branch takes place at the apex.

B

TKXT-FJG. 2.— A, a diagrammatic longitudinal section of a branch of C. agglutinans. a, a short peripheral
tube at the apex of the branch ; 6, b, b, b, a part of the inner tubular system showing a complete open
circuit ; c, comcecial substance, a soft albuminoid with shelly particles included ; d, d', parts of tubular
passages cut obliquely : they are not blind ends ; /, lip or spine set at the opening of a peripheral tube ;
s, s', thin concavo-convex septa set across the tubes. Approximately x 1J. For further explanation
see the text, page 414.

B, a stout branch of C. agghitinans with the ostia and their spines or lips indicated over a portion of
the surface. The tube-openings are shown in black : the oval and crescentic areas represent the broken
bases of spines, a, a', spines situated on the side of the tube towards the apex of the branch ; b, b',
spines situated towards the basal side of the tube ; c, c', spines more or less lateral in position ;
s, s', spines with no tubes alongside them. Approximately x 1J. See text, p. 417.

In the case of species of Idiothecia and Orthoecus it may be surmised from a study
of the characters of the tubes of the coenoecium that youug zooids, on separating from
the stalk of the parent zooids, migrate over the surface, and settle down, in the case
of species of Idiothecia * at the apex of the branch, and in the case of species of
Orthoecus at the margin of the colony-mass. In C. (Demiothecid) insequatus,

* Except in C. indicus (ScHEpoiiEFF, 09), about which we know too little to be able to speculate upon the mode
of growth of the colony.

(ROY. soc. EDIN. TBAJSS., VOL. XLIX., 539.)

416 DR S. F. HARMER AND DR W. G. RIDEWOOD ON THE

ANDERSSON has actually seen living zooids creeping over the surface of the coenoecium
(07, p. 7).

In the case of C. agglutinans the state of preservation of the material does not
enable us to conclude with certainty that each superficial tube opening to the surface
was inhabited by a single zooid and its set of buds, but the probabilities are much in
favour of this. The even calibre of each tube that opens to the surface points to its
gradual growth outwards, as the branch thickens, being the work of a single zooid, as
in the case of species of Idiothecia and Orthoecus. There would, therefore, be in any
one piece of colony approximately as many fully formed zooids as there are ostia.
What occurs in the middle part of the tubular system it is not easy to conjecture ;
these parts can scarcely be inhabited in the ordinary sense of the term, since there is
probably in the deeper tubes no circulation of sea-water such as would suffice for
nutrition and respiration of zooids living there. One must conclude that these internal
tubes are rather in the nature of deep retreats for the zooids in time of danger, and
that when conditions of life are favourable and all the full-grown zooids have their
proboscis and arms, and perhaps the whole of their body except the stalk, projecting
out of the mouth of the tube, the inner part of the tubular system is empty, or serves
only as a safe situation in which the young buds may grow.

When a bud is so far grown that it severs its connection with the parent and
wanders off on. its own account towards the apex of the branch to settle down as an
independent zooid with a peripheral tube of its own, it may possibly reach the apex
by the internal tubes, where, as suggested above, it would meet with but little resistance
from other zooids ; but it is much more likely that the young zooid takes a superficial
route to the end of the branch, as one must suppose a young zooid of C. nigrescens
does. But in the present case the young zooid on arriving at the apex does not com-
mence to form a new short csecal tube for itself, as is the case in C. nigrescens, but,
in conjunction with other young zooids like itself, elaborates the tubular labyrinth
already existing. At any one time one must suppose that every tube that opens upon
the apex has in it a single young zooid which is adding to the mouth of the tube to
increase its length, and is filling up with softer coenoecial material and shelly fragments
the interval between its tube and those around. A newly arrived young zooid would
therefore have to contend with some other zooid for the possession of a tube, and
what seems to happen is that a modus vivendi is arrived at by the two agreeing upon
a branching in an apical direction of what is at present a single tube. This, of course,
is but speculation, but it seems to be the most plausible conclusion from a careful study
of the form and disposition of the tubes at the free extremity of a branch.

In the older parts of a branch it would be interesting to know what happens to a
peripheral tube when the adult zooid dies. The probability is that the vacant tube
would soon be appropriated by some newly liberated bud or young zooid, either from
within or from without. In some cases, as is shown on p. 418, the vacant tube is
sealed up, but this is very exceptional.

(BOY. soc. EDIN. TRANS., VOL. XLIX., 540.)

PTEROBRANCHIA OF THE SCOTTISH NATIONAL ANTARCTIC. EXPEDITION. 417

The life of a colony is apparently not indefinite, for some of the pieces of the Scotia
collection are " dead," i.e. no zooids at all are to be found in them. The tubes are
either empty, or are occupied by a powdery brownish black mass, which, judging from
its colour, may possibly consist of the products of decomposition of zooids. Among
this debris are numerous specimens of a Tanaid Crustacean, of which no adult males
have been found, so that the determination of the species is difficult. The specimens
have been submitted to Mr T. V. HODGSON, who will doubtless refer to them in his
report upon the Tanaidacea obtained on the Scotia Expedition.

The spines represented in text-fig. 3 show the mode of their construction by the
superposition of successive caps of coenoacial substance and the inclusion of foreign
particles in these caps. Each fragment of shell is, of course, embedded in the particular
increment that was being applied at the time that the fragment was picked up, and
consequently it may cause a distortion of the surface of that particular cap, and the

-

TEXT-FIG. 3. — Three spines showing the successive caps of ccencecial substance and the foreign

particles included.

subsequent one, but it does not project into the preceding, more basally placed cap.
In text-fig. 3 it might appear in certain places as though the particle were pro-
jecting into the cap below it, but this effect is produced by the particle and the
optical section of the cap drawn in the figure being in different planes ; the particle
is more or less behind or in front of the median longitudinal section of the spine.

The diagrammatic text-fig. 2, A, shows that, except at the apex of a branch, the
lip or spine at the edge of the ostium may be situated towards the apex or towards the
base of the branch, or more or less laterally, and that it has not the regularly basal
position that it has in C. nigrescent (071, pi. iv. fig. 10). This irregularity is illustrated
further in text-fig. 2, B, which represents the carefully drawn details of a part of the
surface of a stout branch. Owing to the fact that most of the spines on the surface
of a piece of colony are broken off short, it is necessary to dissect into the substance
of the ccencecium to see the relation of the spine to the ostium. In the portion dis-
sected it will be noticed that the spine is situated towards the apical end of the branch
at a and a', but towards the basal side of the ostium at b and b', and it is laterally
placed at c and </. The spines broaden out as one dissects into the branch, and can
be traced along the tubes for about a centimetre, when they cease to be distinguishable

(EOT. soc. EDIN. TRASS., VOL. XLIX., 541.)

418 DR S. F. HARMER AND DR. W. G. RIDEWOOD ON THE

from the soft general material of the crencecium. This is in conformity with what
one sees at the apex of a branch, where the very young tubes have no spines (text-
fig. 2, A).

The part drawn in text-fig. 2, B, shows four spines which have no ostia alongside
(e.g. s). Three of these spines did not reach the surface, but had been buried over
during the increase in the thickness of the branch. Occasionally, but rarely, a buried
spine is found to have a closed-up tube by its side. Such an occluded tube is closed
by a thin, curved partition, such as is seen in some of the inner tubes (as at 5 and s' in
text-fig. 2, A), the convexity in the present case being towards the outer surface ; and
over this and also over the spine the soft material of the coenoecium is deposited
uniformly. One must suppose that in this case the tube was not occupied at the time
that the zooids in the immediate vicinity were lengthening their tubes and filling in
the intervals between them, and that the septum was formed subsequently from
within. The relations of the spine alongside the occluded tube, and the succession of
caps composing the spine, prove that the blind end of the tube is a closed-up ostium,
and is not the initial end of a tube, blind from the commencement, such as one sees in
the middle part of a branch of C. nigrescens (071, pi. iv. fig. 10).

Although the present species is denominated O. ayglutinans, from the manner in
which the zooids have embedded great quantities of shelly fragments in the material of
the coenoecium, the habit is not peculiar to this species, for HARMER has stated in his
account of the new species which were obtained on the Siboga Expedition (05, p. 8),
that the cosnoacium includes foreign particles as a rule, and that they are specially
obvious in C. sibogse (05, pi. ii. fig. 17, 18). ANDERSSON quotes as one of the characters
by which C. solidus is distinguishable from C. rarus, that the tubes are encrusted with
sand, diatoms, etc. (07, pp. 11, 12) ; and he further states that the tubes of C. densus
are covered with sand-grains. GRAVIER, again (12, p. 2), says that in C. anderssoni
(which, if not synonymous with C. densus, is very closely allied to it) sand-grains and
shells of Foraminifera resembling Polystotnella adhere to the outer face of the tubes,
and are even incorporated in the substance of the wall. The basal parts of the pieces
of colony of C. nigrescens are also known to contain fragments of shells (RIDEWOOD,
12, p. 551, Specimen C).

The inclusions in the coenoecium of C. agglutinans are present in such quantity as to
make the branches of the colony much more readily crumbled and broken than are those
of C. nigrescens, which in general massiveness bear some resemblance to the branches
of the present species. The particles included are mostly calcareous, and disappear
when a branch is soaked in a 1 per cent, solution of nitric acid for ten days or a
fortnight. The only foreign particles remaining after this treatment are some rounded,
dark grey, or blackish grains, which Dr J. W. EVANS, Mineralogist of the Imperial
Institute, has been good enough to examine for us. He reports that they are water-
worn grains of slate. They mostly vary in size from '3 to 1 mm. in diameter, but one
was found as large as 6 mm. across. A few of them are paler in colour, and softer,

(HOY. soc. EDIN. TRANS., VOL. XLIX., 542.)

PTEROBRANCHIA OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 419

than the majority, and are nearly white when dry ; these are grains of slate more
decomposed than the others.

While the slate-grains are all rounded and water-worn, the calcareous inclusions
are remarkably free from signs of attrition. Delicate shells of Foraminifera, minute
bivalve Molluscs, extremely thin, and with the two valves still in their natural relation,
pieces of Echinid spines, and portions of Polyzoan colonies occur in a remarkably fresh
and undamaged state, and can be picked out clean from the soft albuminoid material
of the coenoacium. Among the other inclusions one can recognise pieces of fairly large
Lamellibranch shells, fragments of Gastropod shells, and pieces of Serpulid tubes.
These broken pieces of shell show sharp edges in the great majority of cases, but a few
are rounded. The fragments are, many of them, but little larger than the shells of the
Foraminifera, but most are from '5 to 1 mm. in longest diameter. The largest pieces
picked out were 8, 10, or 12 mm. long.

The included particles are all clean and free from mud, and from their even
disposition in the coenoecium one is tempted to conclude that they have not drifted
by accident against newly secreted coenoecial material which has not yet " set " or
hardened, but that the particles are definitely selected by the zooids and built into the
wall of their colonial residence, somewhat in the manner in which the arenaceous
Foraminifera form their shells, and the larvae of the Caddis-flies their tubes; and
similar instances might also be quoted from among the tubicolous Polychaet Worms
and the Rotifers.

The inclusion of foreign particles is not always uniform in the same piece of colony.
It frequently happens that one side of a branch is whiter than the other, and a longi-
tudinal section shows that the whiter half has more calcareous particles embedded than
the other, and fewer tubes.

The pieces of colony brought home by the Scotia are too fragmentary to enable
us to draw any conclusions as to the total dimensions of a colony of C. agglutinans.
None of the pieces show at their basal ends any signs of an area of attachment to a
solid substratum, and while each of the larger pieces, such as are figured in PI. I.,
may be a separate colony, it is just possible that they may be portions of a large
branching colony. This is rather suggested by all the zooids found being males, and
by the fact that all the buds are in about the same stage of development — namely, very
young buds, up to buds with four pairs of arms : only two buds were found having as
many as five pairs of arms. There is, however, no evidence of the existence of stouter
branches than those shown in PI. I., such as would be capable of sustaining the great
weight of the specimens in question, and, further, it must not be overlooked that the
shelly fragments in the ccencecium add to the weight of the branches without adding
to their strength, and this may impose a limit to the size of an individual colony.

There is yet. however, a further possibility that the whole colony may consist of a
broad, plate-like part closely adherent to some other object, and that from this there
may stand up a forest of pieces such as are figured in PI. I. The trawl in passing over

(ROY. soc. EDIX. TRAXS., VOL. XLIX., 543.)

420 DR S. F. HARMER AND DR W. G. RIDEWOOD ON THE

such a colony would snap off several of the upright pieces, but would not dislodge the
basal plate. This view of the constitution of the colony is supported by the fact that
the diminutive colony of C. sibogse is composed of a basal encrustation growing on a
stone, and a series of upstanding pieces (05, p. 13, and pi. i. fig. 2).

EXTERNAL APPEARANCE OF THE ZOOIDS.

The Scotia specimens are, unfortunately, in a rather poor state of preservation.
In many cases the zooids had obviously retreated, on being killed, into the deeper
recesses of the coencecium, where they did not come properly into contact with the
preserving fluid. The result is that they are usually in a highly and often very
irregularly contracted state, and their tissues are a good deal macerated. They are
frequently in closely aggregated and deeply pigmented masses, which have somewhat
the appearance of the zooid with its buds of C. nigrescens shown by RIDEWOOD (071,
pi. iii. fig. 7). They are, in fact, more similar in external appearance to the zooids of
that species than to those of any of the species of Demiothecia previously described.
In well-extended and fairly straight individuals the average length from the end of the
metasome to the free ends of the arms is 4 '5 mm., the length .from the end of the
metasome to the bases of the arms about 3 '2 mm. The thickness of a zooid is
about '8 mm.

The zooids are, however, probably all shorter than they would have been in their
fully extended condition, since the wrinkling of the skin indicates a strong contraction
of the muscles, particularly of the longitudinal muscles of the anterior side of the
metasome and of their continuation into the stalk.

All parts of the epidermis contain a dark pigment, so that the zooids have a deep
brown or black colour in the preserved material. It is obvious that the pigment is
not completely dissolved out by alcohol ; but it is probable that its diffuse arrangement,
as seen in the preserved specimens, does not represent the exact disposition during life.
This is suggested by the comparison of sections of certain individuals of C. nigrescens
(Discovery Expedition) which had been preserved with special care with sections of
zooids which had not been so treated (see also p. 426). The former series show pigment-
granules in sharply restricted areas (cf. EIDEWOOD, 071, pi. v. fig. 28), while the latter
show a diffuse arrangement, suggesting that the pigment had been partially dissolved
out, and then redeposited as a general staining of the tissues.

PROBOSCIS.

The proboscis or buccal shield of the zooids in the material brought back by the
Scotia is in such a poor state of preservation that it is not possible to make a state-
ment as to its average shape and size. By an examination of whole zooids under a
dissecting microscope, sufficient can be seen to show that the proboscis offers no
exceptional features ; there is a main lobe, and a lower lobe, thinner than the main

(ROY. soc. BDIN. TRANS., VOL. XLIX., 544.)

PTEROBRANCHIA OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 421

portion, and possessing the red line which is so curiously constant a feature in the
various species of Cephalodiscus. But the epidermis is hopelessly disintegrated in
most cases, and it is only in the buds that the proboscis is sufficiently well preserved
for its form and proportions to be adequately determined (PI. II. figs. 1-4 ; and
text-fig. 5).

As in other species of Cephalodiscus, the proboscis is a highly mobile organ, and
its parts appear in very varying positions in different individuals. It is shown in
the sagittal section, fig. 15, and in the frontal sections, figs. 6-10, of PI. II.

PL II. fig. 14 represents a section which has cut the proboscis-stalk at its narrowest
region, and it demonstrates the two proboscis-canals (p-p.) in their usual relation to
the pericardium (per.) and the anterior dorsal horns (6.c.2a) of the collar-cavity.

COLLAR-REGION.

The principal parts of the collar are the arms or plumes, dorsally ; and the
operculum or postoral lamella, ventrally.

(i.) Tentacle-bearing Arms or Plumes.

The number of these structures is greater than in any of the species previously
described, since the full number appears to be nine pairs. This number is, however,
not quite constant, as is indicated below. The species which most nearly resemble it
in the number of their arms are the three species of Orthoecus (C. densus, C. rants,
C. solidus), which were described by ANDERSSON (07), in which the number is stated
to be constantly eight pairs ; and C. nigrescens, in which the number is usually
seven pairs, although it may rise to eight pairs (RiDEWOOD, 071). None of the species
of Demiothecia which have been previously described have more than six pairs ; and
this is the commonest number in the subgenus in question.

The sections shown in figs. 6-1 1 of PI. II. are chosen to demonstrate that nine pairs
of arms may really be present. They have been cut at right angles to the long axis
of the zooid, and therefore in a plane which has been described in the Siboga
report (HARMER, 05) as " frontal."

Fig. 6 shows the first seven and the ninth arms of the left side, and the first seven
arms of the right side. The study of the entire series of sections shows that all these
arms except L.9 and R.1 are directed dorsally, in a line prolonging the main axis of the
zooid. The eighth and ninth pairs of arms could probably have assumed the same
general direction during life ; but they are actually lying in a position which makes
their interpretation more difficult than in the case of the first seven pairs.

In fig. 7, which is the eighth section from that represented in fig. 6, the first
seven arms of the left side are connected with one another. The lophophoral region of
the collar has, in this section, not yet split up into the separate arms. Four of the
membranous partitions which separate the arm-cavities for a short distance at their

(ROT. soc. EDIN. TEANS., VOL. XLIX., 545.)

422 DR S. F. HARMER AND DR W. G. RIDEWOOD ON THE

bases, and are formed by the "Grenzmembran," * are seen in this section. On the right
side the first five arms are similarly connected, but R.6 and R.7 are already free. A
comparison of this figure with fig. 8 will show that the first right arm has its axis directed
more or less horizontally (or backwards) at the base, and that the arm then curves
dorsally to assume the position in which it was seen in fig. 6. On the left side of
fig. 7 the lophophore is about to separate from the rest of the collar, and the anterior
dorsal horn of the collar-cavity is cut nearly at its tip. On the right side most of the
lophophore is separated, in the section, from the rest of the collar, but the first arm is
cut in a plane parallel to its food-groove, at its base. As the two sides of the groove
are continuous with one another in an adjacent section, the difference between the
two halves of the lophophore is thus not so great as it appears to be at first sight.
The food-grooves are obvious in all the arms except in L.6, where the groove has
opened out.

Fig. 8, which is eight sections further on, shows the dorsal parts of the collar-cavity
(ft.c.2) separated by a well-marked dorsal mesentery, at the ventral end of which lies the
notochord (nch.), followed by the pericardium (per.) containing the heart. Some of
the arm-grooves have disappeared. The eighth arm (L.8) is visible on the left side, and
the ninth arm (L.9) of the same side shows indications of joining the lophophore. The
relations of the first right arm (R.1) have been explained in describing the previous
section. The sixth right arm has joined the lophophore, but the seventh arm of that
side is still distinct.

Fig. 9 represents the tenth section from that shown in the preceding figure. The
eighth arm of the left side is now separate and is cut longitudinally, while the ninth
arm of the same side is cut in a plane parallel to its long axis, so that its two sides are
completely separated by the food-grove. It will be noticed that this arm, the last of
the series, is closely related at its base to the opercular lobe (o-p.l.) of the same side. On
the right side, R.7 has joined the lophophore ; R.8 is beginning to appear, although still
separated from the lophophore ; and the tentacles of R.9 are commencing to be visible.

Fig. 10, which is four sections further on, shows the last two arms of both sides,
both ninth arms being connected with the corresponding opercular lobe at their base.

In fig. 11, which is seventeen sections nearer the aboral end of the animal, the
eighth left arm has passed across the dorsal side to the right of the metasome, where
it lies close to the eighth right arm ; while the ninth right arm has passed across the
ventral side of the zooid. The section passes through the region of the gill-slits and
collar-canals. Parts of the gill-slits (g.s.) are seen, in the region of their external
openings and of their origin from the pharynx (ph.), on both sides of the section.
The left collar-canal appears as a single cavity, but the right canal is cut twice, the
two parts being connected by the strong muscle of the canal. The reason for this
appearance is that the organ has a concave outline on its dorsal side, where the section

* SPENGEL, J. W., "Die Enteropneusten d. Golfes v. Neapel," Fauna u. Flora d. Golfes v. Neapel, 18 Monogr.,
1893, p. 452.

(BOY. soc. EDIN. TKANS., VOL. XLIX., 546.)

PTEROBRANCHIA OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 423

passes. Part of the ccelomic opening (c.c.i.) is visible in the figure, while ,a line
crossing the epidermis just outside the canal indicates the commencement of the
external orifice. On the dorsal side of the section are visible the two testes (t.L, t.r.),
with the genital mesenteries passing from the median dorsal mesentery (d.m.3) of the
metasome to the gonads. The left testis (t.L} of this individual is large and functional,
but the right testis (t.r.), which is cut close to its external opening, has remained
undeveloped. It appears in only a few sections of the series, and it does not anywhere
reach a size much larger than is shown in the section under consideration. The
asymmetry of the gonads is to be regarded as an individual peculiarity of this zooid
(see also p. 4'29).

The demonstration of the number of arms, beyond the possibility of doubt, being
a matter of some importance, a plasticine reconstruction of this individual was made
according to the method described on p. 21 of the Siboga report (HARMER, 05).
A slight improvement was, however, introduced by drawing the sections on thin
drawing card (" Bristol board "). After cutting out the outlines drawn, the card was
covered with a layer of plasticine and was attached to the part of the model already
made. The cards representing the sections were left in situ in the model. The special
advantage of this mode of procedure is that the cut edges of the cards remain in the
model as a record of the outlines of the sections from which it has been constructed.
This obviates the danger of destroying the outlines of the sections when smoothing out
the intervals between two successive sections. If it should appear, moreover, that two
sections have not been fitted together in correct " registration," it is easy to separate
them at any time and to fit them together more accurately without losing the evidence
on which the model was built up. The study of this reconstructed model has enabled
us to obtain a clear idea of the position of the individual arms, of the parts of the
operculum, and of the arrangement of the gill-slits and collar-canals, with other features
of the zooid.

Making use of this evidence, the correctness of which can be estimated from the
figures (figs. 11-6), the following description of the arms may be given. It will, of
course, be understood that their position in the individual under consideration is to a
large extent fortuitous, and that during life the arms could undoubtedly have assumed
many other positions.

The first seven arms of both sides are directed dorsally, in line with the long axis
of the zooid. On the right side, however, the first arm passes at first posteriorly, at
right angles to. the long axis, and then curves dorsally. On both sides the eighth and
ninth arms are in the main directed ventrally. The ninth right arm passes ventrally
from its origin, but soon curves across the ventral or anterior side of the zooid to assume
the position shown in fig. 11. The eighth arm begins by passing dorsally for a short
distance, and then doubles back sharply in a ventral direction. Fig. 10 represents the
arm at the point where this flexure is taking place. It will be seen that the food-
groove faces posteriorly. The part which is marked R.8 represents the arm after the

(ROY. soc. BDIN. TRANS., VOL. XLIX., 547.)

424

DR S. F. HARMER AND DR W. G. RIDEWOOD ON THE

flexure has taken place; and in the later sections of the series (fig. 11), the arm is
seen in transverse section as a continuation of the part marked R.8 The elongated
part which stretches from this region towards the ninth arm shows the food-groove
(R,8*) opened out and cut obliquely at or near its flexure.

The eighth left arm (figs. 11-8) also has its food-groove directed posteriorly ; but
it lies in such a position that it is cut almost longitudinally as it passes across the
dorsal side of the principal part of the collar. The ninth left arm (figs. 10-6) at first
lies between the eighth arm of the same side and the region of the central nervous
system (figs. 10-8). It runs more or less horizontally near its base, but soon curves
dorsally, so as to appear even in the first section (fig. 6) which has been represented.

Although there are eighteen arms in the zooid which has thus been described in
detail, this number is not constant in the species. A series of selected zooids were
dissected and were found to give the results set forth below. It is possible, however,
that arms may have been lost in some of the zooids examined, either by accident
during life or as the result of violent contraction at the moment of preservation, and
that some margin of error may thus have been introduced into the table. The facts
as observed are as follow : —

12 arms

3 individuals ....

13

1

• I

14

3

. /

15

4

. 1 r

16

11

. \

17

11

1 -i^

18

6

• 1

19

1

'. . '. . I

Total 40 individuals

40

If it may be inferred that in a zooid with thirteen arms, for instance, there were
seven arms on one side and six on the other, it follows that of the forty individuals
dissected there were only three which had less than seven arms on one side at least,
and these appear to have had six pairs. In four cases there were seven arms on one or
both sides ; in fifteen cases there were eight arms on one or both sides ; in seventeen cases
there were nine arms on one or both sides ; and in one case there were ten arms on one
side. Representing these as percentages, it follows that the observations indicate :—

Total Number
of Arms.

Maximum Number
of Arms on
One Side.

Individuals.

Percentages.

17 or 18

9

17

42-5

15 or 16

8

15

37-5

13 or 14

7

4

10-0

12

6

3

7-5

19

10

1

2-5

Total .

40

100-0

(BOY. 8OC. EDIN. TRANS., VOL. XL1X., 548.)

PTEROBRANCHIA OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 425

It results from these figures that 42 "5 per cent, of the zooids had nine arms on one
side at least ; while 80 per cent, had either eight or nine arms on one side at least.
We accordingly draw the conclusion that the typical number of arms, in this species,
is nine pairs, but that more than a third of the cases observed had a pair less ; while
a smaller percentage had seven, six or even ten arms as the maximum number on
one side.

B

TEXT-FIG. 4. — Arms of Cephalodiscus agglutinans. A, an arm well extended ; B, an arm in a
moderate state of contraction ; O-E, terminal portions of arms.

The number of tentacles or pinnules on an arm of a full-grown zooid is usually
between thirty-five and forty-five pairs, but in some instances the number may be
fifty pairs, and in younger zooids the number is less than thirty-five pairs.

It is difficult to say with certainty whether the dorsal epidermis of the axis of the
arm is thick and black in C. agglutinans, as it is in C. nigrescens, because in most
cases the epidermis has disintegrated, and there is more than a suspicion that in those
cases in which the epidermis is still present the pigment-granules, which might have
been present in them, have become bleached. This suspicion is founded upon the fact

(ROY. soc. EDIN. TRASS., VOL. XLIX., 549.)

426 DR S. F. HARMER AND DR W. G. RIDEWOOD ON THE

that in the Discovery material some of the pieces of C. nigrescens were carefully fixed
in Perenyi's fluid and some in picric acid solution, but most of the material was
preserved in 5 per cent, formalin ; and in slides made from the former material (fixed
in Perenyi's fluid), the pigment is strongly marked (RIDEWOOD, 071, pi. v. fig. 28),
whereas in sections prepared from the formalin-preserved material, the thick dorsal
epidermis is not darker than the other parts of the section. The material of
C. agglutinans was preserved in alcohol and not in formalin, it is true, but the
preservation is not good, and the appearance of the large cells on the dorsal surface
of the axis of the arms seen in paraffin-prepared serial sections is remarkably like that
in the sections of the formalin-preserved zooids of C. nigrescens.

The dorsal epidermis of the arms of C. agglutinans is least disintegrated in contracted
arms, and in the arms of fairly young zooids having but twenty-five to thirty-five pairs
of tentacles. Text-fig. 4, A shows the appearance of a well -extended arm, and B an arm
in a moderate state of contraction. The terminal part of the arm has no end-bulb
with highly refractive beads such as distinguish the arms of C. dodecalophus (M'lNTOSH,
87, pi. iv. fig. 1 ; pi. v. fig. 1 ; and RIDEWOOD, 071, text-fig. 1, p. 4) and C. hodgsoni
(071, pi. v. fig. 32). In extended arms the extremity is bluntly pointed ; in contracted
arms it is rounded and even hemispherical. Some terminations better preserved than
usual are shown in text-fig. 4, C-E (cf. C. nigrescens, 071, PI. v. figs. 23-27).

(ii.) Opercitlum or Postered Lamella.

The interpretation of the sections shown in figs. 8-11 is at first sight by no
means easy, so far as the operculum is concerned. The study of the plasticine
reconstruction has, however, enabled us to come to a clear conclusion as to its general
form. It is seen to be deeply emarginate in the middle line, and the median sagittal
sections examined thus show only a feebly developed lower lip (e.g. fig. 15, op.).
On either side of the mouth, however, the operculum is produced into a large lateral
lobe, a great part of which is free. The comparison of this series with other series
of sections, and with what is known of other species of Cephalodiscus, shows that the
operculum is a highly mobile organ, the parts of which can assume very different
positions at the will of the zooid. In the specimen under consideration the right
lobe (op.r.) is directed in the main dorsally (figs. 10-8), while the left lobe is sharply
reflected ventrally, close to its origin from the collar. Figs. 10 and 9 cut this left
lobe at or near the point where the flexure takes place. A consideration of fig. 9
will show that if the lobe of the left operculum (op.l.) nearer to the proboscis be
imagined to have been unbent, so as to pass in a dorsal direction as a prolongation of
the part of the operculum of the same side which is nearer the central nervous
system, it would not have appeared in fig. 9 ; and the two sides of the section would
have been fundamentally similar. If the zooid had died with the left lobe in the
position indicated, both lobes would have appeared in fig. 8 in the position actually

(ROY. SOC. EDIN. TRA.NS., VOL. XLIX., 550.)

PTEROBRANCHIA OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 427

seen on the right side. This section shows the large right lobe of the operculum in a
region where it is free from the rest of the collar. The species appears to be charac-
terised by having these free lateral lobes of the operculum well developed. It may
be supposed that the size of the lateral lobes is to some extent correlated with the
number of the arms ; since it can hardly be doubted that one of the most important
functions of the operculum is to help to convert the ventral grooves of the arms into
channels leading the food to the mouth. A large development of the arms would
accordingly require a corresponding development of the opercular lobes, which, during
the act of feeding, would presumably be thrown into the position seen on the right
side in fig. 8.

Comparison with other species of Cephalodiscus in this respect cannot be
attempted very satisfactorily, but it may be noted that the operculum of C. hodgsoni,
figured by RIDEWOOD (07l, pi. vi. fig. 53), has a close resemblance to that of
C. agglutinans, here described. Except with the aid of solid reconstructions, which
have not been made in many cases, it is often difficult to ascertain the exact form of
this organ ; but it may be remarked that in C. gracilis, of which a plasticine reconstruc-
tion has been figured in pi. iii. fig. 25, of the Siboga report (HARMER, 05), the
operculum is hardly emarginate in the middle line, and has no special development
of its lateral lobes. The arms in that species are comparatively few, only five being
present on each side.

Two further points may be noted with regard to the operculum of C. agglutinans : —

(a) The last arm of each side is continuous with the corresponding opercular
lobe, as in other species of Cephalodiscus (cf. HARMER, 05, pp. 31, 36 ; and RIDEWOOD,
072. p. 229). This is sufficiently illustrated by fig. 10.

(b) The free edge of the operculum is more or less scalloped, as is indicated by
fig. 11, where two of the end-lobes are cut separately. Evidence that the edge has
this form has also been obtained in some of the dissected specimens. The same
character has been figured by RIDEWOOD (071, pi. vi. fig. 53) in C. hodgsoni.

(iii.) Collar-canals.

These structures agree in general form and position with those of other species of
the genus. Seen in side view the ventral outline is very convex, while the dorsal
outline may be concave. In a frontal section of the zooid (fig. 11), a collar-canal
which is cut near its dorsal side may appear as two separate parts. The inner part
(c.c.i.) opens into the collar-cavity, while the two parts of the tube are connected by
the strong collar-muscle. The epithelium of the canal is much thinner on the dorsal
side than it is ventrally ; and it is reasonable to suppose that it is more fiexible there.
The muscle would thus seem to have the function partly of dilating the coelomic
opening and partly of acting in antagonism to the prolongation of the principal
muscular mass of the metasome ; which, as shown in the sagittal section, fig. 16, ms.,

(ROY. soc. EDIN. TRANS., VOL. XLIX., 551.)

4'28 DR S. F. HARMER AND OR W. G. RIDKWOOD ON THE

ends in close relation with the collar-canal (c.c.i.). The modus operandi of the collar-
canals has previously been discussed by both of us (HARMKR, 05, pp. 41-46 ; RIDEWOOD,
07, pp. 41-43).

GILL-SLITS.

The gill-slits (PI. II. figs. 5, 11, g.s.) agree closely with those of other species of
the genus. In the individual represented in figs. 6-1 1 , the external openings of the
gill-slits are seen from the plasticine reconstruction to lie in strongly marked
longitudinal grooves, each of which occurs nearer to the middle line of the anterior
wall of the metasome than the external opening of the corresponding collar- canal.
These grooves are, however, probably the result of contraction at the moment of
death. They can be traced along the metasome for a considerable distance, in an
aboral direction, beyond the collar-pores. The walls of the gill-slits are composed
of vacuolated epithelium (MASTERMAN'S " pleurochords "), as in other species of
Cephalodiscus.

ALIMENTARY CANAL AND NOTOCHORD.

The great length of the intestinal loop, which commences at the stomach and is
situated in the caecal end * of the metasome has been alluded to above in describing
the distinctive features of this species. It is well seen in the dissected specimen,
fig. 12, and in the sagittal section, fig. 16. In both figures the metasome is probably
shortened by muscular contraction ; and in its full state of extension the folds seen in
the figures would have probably disappeared. Owing to the length of this part of
the alimentary canal, the stomach (stom.) is separated by a considerable interval from
the caecal end of the metasome — a character in which the present species resembles
C. levinseni. In one individual which has been examined, the intestinal loop contains
a number of Diatoms and other microscopic organisms, and many Sponge-spicules.

The notochord (nch.) is seen in sagittal section in fig. 15 ; and as shown by this
figure, as well as by figs. 8-10, it has a well-developed lumen.

GONADS.

Every individual in which the gonads have been examined has proved to be of the
male sex. Although the amount of material at our disposal has been large, it was all
dredged at one time, and there is nothing to forbid the assumption that all the pieces
brought back by the Scotia were parts of a single large male colony.

The testes vary a good deal in size in the different individuals examined. In their
highest development (figs. 5, 12) they are elongated organs, which may be simple
tubular structures, or may be marked by several constrictions extending traversely to

* ANDERSKON (07, p. 7) lias pointed out, from observations on living zooids, that the " csecal end " of the metasome
disappears in fully extended zooids, where the stalk appears as a direct continuation of the body. The appearance
of a caecal end is none the less very characteristic of the majority of the contracted zooids.

(BOY. soc. EDIN. TRANS., VOL. XLIX., 552.)

PTEROBRANCHIA OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 429

their main axis. These are probably in part the result of the contraction of the zooid.
The aboral end of the testis may extend at least as far as the origin of the stalk from
the metasome.

In some cases, though not in all, we have noticed the appearance indicated at the
aboral end of the testis in fig. 12. A tube, which seems to end blindly, extends into
the cavity of the organ as an invagination of the aboral end. The most probable inter-
pretation of this structure is that it is merely part of the wall of the testis which has
been invaginated, and that it has no special morphological significance. Its walls are
sometimes more pigmented than those of the rest of the testis, so that it may be a
conspicuous structure in entire preparations.

When both testes are fully developed, as in fig. 5, t.l., t.r., they appear symmetric-
ally disposed on either side of the median dorsal mesentery of the metasome. The
central part of the testis is occupied by a mass of fully developed spermatozoa, while
there is a peripheral zone, of varying width, which contains very numerous nuclei,
indicating stages in spermatogenesis. In other cases, as in fig. 11, the two testes are
very different in size ; and in this particular case the left testis (t.l.) is fully developed,
while the right testis (t.r.) has remained vestigial. The figure shows the two trans-
verse genital mesenteries which run from the dorsal mesentery of the metasome, near
the region where the testes open to the exterior, to the two gonads. These transverse
mesenteries are no doubt the bearers of blood-vessels which pass from the dorsal vessel
to the gonads.

NERVOUS SYSTEM.

We have no new observations of importance with regard to this part of the anatomy,
and it will be sufficient to note that the nervous system of C. agglutinans agrees in
essential respects with that of other species of the genus.

ANDERSSOX has, however, stated (07, pp. 7, 8, 32) that while in the subgenus
Orthoecus the lateral nerves unite ventrally on the metasome to give rise to a nerve-
tract which has a single median thickening, Demiothecia differs from it in having a
subsidiary thickening on each side of this median one. The occurrence of the three
nerve-tracts in the ventral region of the metasome and in the stalk was first pointed
out by MASTERMAN (98, p. 513) in C. dodecalophus ; but it is not certain that the
character in question can be used to discriminate the subgenera of Cephalodiscus.
Thus in C. levinseni (HARMER, 05, p. 51, pi. xi. fig. 132), which belongs to Idiothecia,
a single nerve-tract is present in the stalk, while in C. nigrescens (RiDEWOOD, 071, p. 40,
text-fig. 15), belonging to the same subgenus, there are three nerve-tracts. ANDERSSON
has, moreover, not sufficiently taken into account the fact that in C. gracilis and
C. sibogas (HARMER, 05, pp. 52, 53), which belong to Demiothecia, only a single nerve-
tract is present ; while, on the other hand, C. dodecalophus, C. hodgsoni, and C. aequatus,
belonging to the same subgenus, have three nerve-tracts. C. agglutinans (fig. 13, n.t.)
agrees with the species of Orthoecus described by ANDERSSON, and with C. (Idiothecia)

(ROT. soc. EDIN. IBANS., YOL. sux., 553.)

430 DR S. F. HARMER AND DR W. G. RIDEWOOD ON THE

levinseni, C. (Demiothecia) gracilis, and C. (D.) sibogfe in having a single nerve-
tract in the stalk.

The number of the nerve-tracts in the region in question is no doubt correlated with
the way in which the metasomatic muscles are arranged at the point where they pass
into the stalk. In C. agglutinans, as in C. levinseni, C. gracilis, and C. sibogse, the
layer of metasomatic muscles forms a single fold extending deeply into the cavity of
the stalk, round the single nerve-tract, in the region where the stalk is separating
from the body. In C. dodecalophus, C. hodgsoni, and C. sequatus, in which three nerve-
tracts are present, there is, in addition, a pair of muscular folds similarly related to the
lateral nerve-tracts.

BUDS.

The buds seem to be produced in exceptionally large numbers in the present species,
and an examination of the budding disc at the end of the stalk of full-grown zooids
shows that in some cases as many as seven pair of buds are present at the same time.
As in other species, the free surface of the disc is of the nature of a sucker for adhering
to foreign objects ; the disc is wider than the end of the parental stalk, and the buds
develop in pairs in the groove between the stalk and the disc. The youngest buds
cannot be said to have any stalks ; they are merely clavate or pyriform outgrowths,
which soon show a transverse groove when the proboscis differentiates from the rest of
the bud. The stalks of the other buds are remarkably long ; that of one bud with two
pairs of arms developing, of the stage shown in text-fig. 5, (.!, measured 3'9 mm. ;
another of the same stage of development measured 4 '3 mm.

There is a remarkable uniformity to be noticed in the size of the stalked buds present.
The proboscis of a bud in which the first pair of arms are just about to develop, such
as is represented in text-fig. 5, A, is not very much smaller than that of a bud in
which five pairs of arms can be recognised — and it is the proboscis which practically
determines the size of the bud. Buds intermediate in size and development between
minute stalkless buds just forming from the disc and buds like that shown in text-fig.
5, A, are scarce; and no buds have been found older than that shown in text-fig. 5,
F. Indeed, only two buds of the latter stage were discovered. It would seem, there-
fore, that buds develop very rapidly up to the time when the first pair of arms make
their appearance, and that their subsequent growth is slower. What happens after the
stage with five pairs of arms it is impossible to say. The buds of this age seem to be
too young to separate off as independent zooids and to leave the colony — their alimentary
canal is still csecal, and the ten arms have not yet produced any tentacles (text-fig. 5, F)
—and in some other species of Cephalodiscus (e.g. C. hodgsoni and C, nigrescens,
RIDEWOOD, 072) buds may be found still attached by their stalk to the parent having the
full number of arms -characteristic of the adult zooids. The absence of old buds in the
present material may perhaps be accounted for by some seasonal cause ; or buds may only
begin to be produced when the development of the gonads has reached a certain stage.

(BOY. soc. EDIN. TKANS., VOL. xi.ix., 554.)

PTEROBRANCHIA OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 431

In text-fig. 5 are shown composite figures of buds based upon thirty -five camera lucida
drawings. The proboscis is reproduced in the successive figures as gradually increasing
in size ; this is correct in the main, but buds with the same number of arms may have

TEXT-FIG. 5. — Figures illustrating the development of the arms or plumes in the buds of Cephalodiscus agglvtinans, for
comparison with the figures of C. gilchristi and C. nigrestens in RIDEWOOD, 072, pp. 243 and 236. The lettering is the
same as before, namely : — 1, 2, 3, 4, 5, the first, second, third, fourth, and fifth arms of the buds ; b, the "body" or
metasome of the bud ; b.s., main portion of the proboscis or buccal shield ; m, mouth ; p. I., lower lobe of the proboscis ;
po.l., operculum or postoral lamella ; s, stalk of the bud.

A, aboral view of a bud in which no arms have yet developed ; B, a bud with one pair of arms ; C, a bud with two pairs ;
D, a bud with three pairs ; E, a bud with four pairs ; F, a bud with five pairs of arms ; G, the oral surface of the body of a
bud in which the lower lobe of the proboscis was turned well upward, and has been cut away. The operculum is seen to
be continuous with the last developed (fourth) arm, and is as yet incomplete below the mouth. Approximately x 60.

the proboscis of different sizes, and there are many buds with three pairs of arms which
have a smaller proboscis than others with two pairs or even one pair only. This irregu-
larity may be partly accounted for by disparity in the rates of growth of the proboscis
and the collar-region, but is to a large extent also to be explained by the fact that the

(BOY. soc. EDIN. TRAUS., VOL. XLIX., 555.)

432 DR S. F. HARMER AND DR W. G. RIDEWOOD ON THE

proboscis is not only a very mobile organ, but is evidently also capable of considerable
change in superficial extent ; a large proboscis on a bud with three pairs of arms is
thinner than a smaller proboscis on another bud with the same number of arms.

In buds with three, four, or five pairs of arms the first portion of the gut can be
seen in the interior of the " body " in clarified preparations ; it appears as a caecal tube,
connected with the mouth, and with a slender cord passing from the blind extremity
towards the stalk. The state of preservation of the material is so poor that the study
of the internal structure of the buds by means of serial sections did not seem advisable.
The development of the internal organs in buds of Cephalodiscus is not likely to differ
much among the various species, and any time and energy spent upon a study of these
parts is most profitably expended by restricting oneself to material that is specially
suited to the methods of microscopical technique.

On comparing the buds of C. agglutinans with those of other species of Cephalo-
discus, one is struck by the fact that in the relatively late appearance of the arms this
species more closely resembles C. gilchristi, a species of the subgenus Idiothecia, than
it resembles species of the subgenus Demiothecia, such as C. dodecalophus and C.
hodgsoni (RIDEWOOD, 072, pp. 231 and 225). Incidentally it may be mentioned that the
buds of C. sequatus and C. insequatus, examined by us in material collected on the
Swedish South-Polar Expedition, agree very closely with those of C. dodecalophus and
C. hodgsoni. In C, dodecalophus, C. hodgsoni, C. insequatus (which we regard as
synonymous with C. hodgsoni, see p. 4:»6), and C. sequatus the first pair of arms reach
the end of the proboscis at a stage when three pairs of arms are recognisable (072, p. 231,
text-fig. 4, G and H, and p. 225, text-fig. 2, G) ; whereas in C. agglutinans, as also
in C. nigrescens and C. gilchristi (072, pp. 236 and 243), the arms at the three-pair
stage (text-fig. 5, D) are but insignificant bead-like outgrowths from the sides of the
collar, and are very remote from the edge of the proboscis.

SYSTEMATIC POSITION OF C. AGGLUTINANS.

We have not found it an easy matter to come to a definite conclusion with regard
to the affinities of the Scotia species of Cephalodiscus. In the present state of our
knowledge it is probably best to accept provisionally the subdivision of the genus into
the three subgenera mentioned on p. 408. We may remark, however, in passing that
it is surprising that a crowd of independent zooids should be able to build up a
coanoecium, by their united efforts, which has a definite character of its own. But, on
the other hand, there is sufficient resemblance between the ccenoecia of C. levinseni,
C. nigrescens, and C. gilchristi to justify their inclusion in a single subgenus, Idio-
thecia ; and this becomes the more significant when it is remembered that these species
have been recorded from such widely distant localities as between Japan and Corea,
the Antarctic Ocean, and South Africa respectively. There is, moreover, a considerable
amount of resemblance in anatomical structure between the zooids of species which

(ROY. 8OC. EDIN. TRANS., VOL. XLIX., 556.)

PTEROBRANCHIA OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 433

have been referred to Demiothecia, as well as between those which have been placed
in Orthoecus.

According to the subgeneric diagnoses which have previously been given, C.
agglutinans should come in Demiothecia, since the cavity of the ccenoecium is con-
tinuous throughout (as distinguished from Idiothecia, in which each zooid inhabits a
separate tube of its own), while the tubes are not in the main distinct from one
another and with separate cavities, as in Orthoecus. But in other respects the
ccenoecium of C. agglutinans shows a greater resemblance to Idiothecia, to which we
are inclined to refer it, in spite of the continuity of the ccenoecial cavity.

The ccenoecium of the known species of Demiothecia consists of a more or less
branching system traversed by a continuous cavity, which is of such a kind that a
transverse section of a branch shows a central cavity surrounded by a layer of the
ccenoecial substance, irregularly thickened in various places. A transverse section of
the ccencecium of C. agglutinans does not show this simple arrangement ; but it
exhibits a massive ccencecial substance, crammed with foreign inclusions, and traversed
here and there by the relatively small tubes inhabited by the zooids. The arrangement
of these tubes is shown diagrammatically in text-fig. 2, A, on p. 415. In its massive
character and in the disposition of the outer parts of the tubes this has considerable
resemblance to the coencecium of C. (Idiothecia) nigrescens, as figured by RIDEWOOD
(071, pi. iv. fig. 10). It differs from it in the fact that the tubes which lead to the
external ostia do not end blindly at their lower ends, but are connected together by a
complicated system of branching tubes which occupy the axial part of the branch.
But the fact must be emphasised that the arrangement of the outer parts of the tubes
agrees substantially with that of the entire tubes of C. nigrescens, and that no species
of Demiothecia yet described possesses a system of regularly arranged long tubes lead-
ing from the interior to the ostia. The ostia of Demiothecia are little more than
perforations of the common ccencecial wall, so that the central cavity may be described
as opening directly to the exterior by means of the ostia. We think, therefore, that
there is much to be said for the view that C. agglutinans may be regarded as an
Idiothecia in which the inner parts of the tubes of the zooids are connected by a set
of tubes, branching in the axial region of the ccenoecium and placing all the cavities
in communication with one another.

A further argument in favour of this view may be obtained from a consideration
of the mode of growth which appears to be indicated by a study of the coencecium
(cf. p. 415). We have given reasons for believing that the growth of the branches
is to a large extent apical, as appears to be the case in C. nigrescens and C. levinseni,
both of which belong to Idiothecia. This may be inferred by the regular arrangement
of the outer parts of the tubes, in all parts of the colony, as well . as by the fact that
the apical tubes are shorter than the others, and apparently younger.

The occasional occurrence of septa across the tubes of C. agglutinans (p. 414) may
indicate some approach to the more typical species of Idiothecia in which each tube

(BOY. soc. EDIN. TRANS., VOL. XLIX., 557.)

434 DR S. F. HARMER AND DR W. G. RIDEWOOD ON THE

is completely cut off from its neighbours (compare the septa in C. nigrescens, RIDEWOOD,
071, pi. iv. fig. 10).

The single short spine or lip at the side of each ostium in C. agglutinans finds a
parallel in C. (I.) nigrescens, C. (/.) levinseni, and C. (0.) solidus. In C. (1. ) gilchristi
there are long spines, about as numerous as the ostia, but not very distinctly related
to them ; whereas in C. (I.) indicus, C. (0.) densus, and C. (0.) rarus there are no
lips to the tubes. In C. (D.) dodecalophiis, C. (D.) hodgsoni, and C. (D.) fequatits,
however, the spines are long and numerous, usually four or five to each ostium.

The study of the zooids may be held to give some support to the view that the
nearest ally of C. agglutinans is to be found in the subgenus Idiothecia. There is
considerable resemblance between the zooids of C. agglutinans and C. nigrescens in
their relatively large size, in the general proportions of the body, in the large number
of arms, and in the dense pigmentation of the skin. Too much weight, however, must
not be attached to this last character, since the pigmentation may not really be an
indication of affinity, but may be of a purely physiological nature, as seems to be the
case in certain deep-water pelagic Fishes.* C. sibogze, a species of Demiothecia, has,
moreover, a deeply pigmented epidermis (HARMER, 05, p. 8).

In the absence of end-bulbs with refractive beads at the ends of the arms, C. agglu-
tinans agrees with the species of Idiothecia and Orthoecus, and differs from those of
Demiothecia.

Attention has already been drawn (p. 432) to the fact that, in the relatively late
appearance of the arms in the buds of C. agglutinans, this species more closely resembles
C. gilchristi and C. nigrescens than species of Demiothecia such as C. dodecalophus,
C. hodgsoni, and C. sequatus.

The special characters of the coenoecium of C. agglutinans might be taken to justify
the institution of a new subgenus for the reception of this species. We think it un-
necessary to adopt this course ; but it is obvious that, if the species is rightly referred
to Idiothecia, the original diagnosis of that subgenus (RlDEWOOD, 071, p. 10) must
be amended by adding to the statement, " tubular cavity . . . having no connection
with the other cavities of the tubarium," some statement to the effect that the
tubes may be connected with one another. The following amended diagnosis is
suggested :—

Subgenus Idiothecia. Each ostium leading into a long tube lodged in a common
coanoecial substance ; the tubes definitely arranged at a more or less constant angle to
the surface, usually completely separated from one another, and each containing a single
zooid with its buds, but sometimes connected with one another by an intercommuni-
cating system of tubes.

It may be pointed out in conclusion that the name Idiothecia would not be
less strictly applicable in consequence of the extension of the scope of the subgenus,

* Cf. Sir JOHN MURRAY and J. HJORT, The Depths of the Ocean, London (Macmillan & Co.), 1912, pp. 618, 624,
677, and elsewhere.

(ROY. soc. EDIN. TRANS., VOL. XLIX., 558.)

PTEROBRANCHIA OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 435

for the derivation of the word (idios, one's own, personal, private ; and theke, a case,
box, vault) refers to the zooids, with their buds, having each their own tubes ; and this
appears to be the case in C. agglutinans, in spite of the fact that the tubes communicate
with one another in the axial part of the coenrecium (see p. 416).

ADDENDUM ON THE SYNONYMY OF C. HODGSONI.

Through the kindness of Dr H. THEEL, the British Museum has received from the
Riksmuseum at Stockholm duplicate specimens of all the species of Cephalodiscus
which were described by ANDERSSON in the report (07) on the Pterobranchia of the
Swedish South-Polar Expedition. We have been much interested in comparing this
material with the other specimens which have been available for study.

The species of Cephalodiscus which have previously been described from Southern
localities are as follows: — C. dodecalophtts (Challenger and Swedish Expedition);
C. ivquatus and C. in&quatus (Swedish Expedition) ; C. nigrescens and C. hodgsoni
(Discovery Expedition) ; C. gilchristi (Cape Colony, Dr J. D. F. GILCHRIST) ; C. densus,
C. rarus, and C. solidus (Swedish Expedition) ; and C. anderssoni (second French
Antarctic Expedition). With the exception of the three Oriental species described
in the " Siboga " report, and of C. indicus, from Ceylon, more recently described
by SCHEPOTIEPF (09), this list accounts for all the species of Cephalodiscus hitherto
recorded. ANDERSSON (07, p. 16) has stated that within the Antarctic area
Cephalodiscus is one of the most characteristic members of the marine fauna,
in the neighbourhood of Graham's Land at least ; and that it occurs there, as a
rule, in depths of 100 to 150 metres, wherever the bottom has a firm consistency
and consists of gravel ("Kies") and small stones. We may agree with him, in
view of these facts, in thinking that the headquarters of the genus lie in Antarctic
and Subantarctic waters.

Of the species already mentioned, C. densus, C. rarus, and C. solidus were placed
by ANDERSSOX in his subgenus Orthoecus ; and C. anderssoni, more recently described
by GRAVIER (12), clearly belongs to the same assemblage. None of these species have
at present been found a second time, and we have no criticism to make with regard
to them, except that C. anderssoni may perhaps prove to be a synonym of either
C. densus or C. rarus.

C. nigrescens and C. gilchristi are two very distinct species of the subgenus
Idiothecia,

The rest of the Antarctic or Subantarctic species, namely, C. dodecalophus,
C. aequatus, C. vnxquatus, and C. hodgsoni, belong to the subgenus Demiothecia, in
which the coeno?cial cavity is continuous and is not represented by a number of
distinct tubes, each belonging to one zooid. According to the diagnoses originally
given, all these species are characterised by the possession of six pairs of arms in each
sex, with the exception of C. in&quatus, in which ANDERSSON states that there are five

(ROY. soc. EDIN. TRANS., VOL. XLIX., 559.)

436

DR S. F. HARMER AND DR W. G. RIDEWOOD ON THE

pairs in the female and six pairs in the male. According to this observer (p. 8),
C. sequatus is closely related to C. hodgsoni, but on p. 9 he says that it is nearly
allied to C. dodecalophus.

At the first examination of the coenoecium of the specimens obtained by the
Swedish Expedition it appeared to us that there was an even closer resemblance-
amounting, in fact, to a practical identity of characters- — between C. hodgsoni and
C. insequatus. Since the latter is said to be characterised by a difference between the
two sexes in respect of the number of arms (female five pairs, male six pairs), it became
important to make a new examination of C. hodgsoni to ascertain whether any
difference could be detected between the two sexes in that form. Although the number
of male individuals examined was but small, the result of this inquiry was to confirm
the conclusion, suggested by the appearance of the coenoaciuin, that C. insequatus
is a synonym of C. hodgsoni. It may be noted in passing that although the
memoirs containing the accounts of the two species in question were published
in the same year (1907), the name C. hodgsoni has the priority, as is shown by
the inclusion of the title of BJDEWOOD'S paper in the bibliography given by ANDKRSSON
on p. 115.

In the original description of C. hodgsoni it is stated (RIDEWOOD, 071, p. 55) that
" the normal number of plumes is twelve, but the sixth pair develop late, and a full-
sized polypide, with buds of its own, and with well-developed ovaries, may have only
ten fully-grown plumes." It is further remarked, however, that a vestigial sixth pair
can " usually " be detected in such cases, but that no individual has more than six pairs.
In his later paper (072, p. 230) the same author states that " some polypides of
Cephalodiscus hodgsoni of full size and with mature gonads have five pairs of plumes
only." He does not discuss, in either place, the possibility that the difference to which
he alludes might be correlated with a difference in sex.

We are in a position to confirm the substantial accuracy of ANDERSSON'S state-
ment with regard to the number of arms or plumes in the specimens described by
him as C. insequatus, though we think he has gone rather too far in asserting
(07, p. 6) that the number is invariable, in each sex, in the species examined by
him. The following is the result of an examination of 38 zooids in which the
arms were dissected from one another sufficiently to enable them to be counted
when the zooid was mounted on a slide. There is no doubt some element of
uncertainty in some of these estimations, particularly with regard to the presence or
absence of vestigial sixth arms ; but in the main the results can probably be accepted
as trustworthy.

The material examined consisted principally of female zooids ; since, when the
gonads were sufficiently advanced in development to be recognisable as to their sex,
19 specimens were female and 3 were male. The remaining 16 specimens were
mostly advanced buds or quite young zooids in which the sex was not certainly
distinguishable.

(ROY. soc. EDIN. TRANS., VOL. XLIX., 560.)

PTEROBRANCHIA OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 437

The evidence as to the number of arms may be given in a tabular form, as
follows : —

Number of Individuals.

Sex.

Number
of Arms.

Remarks.

Stat. 5.

Stat. 94. Total.

Female

10

13

2 15

11

3

3

In one of these cases the evidence

for the occurrence of more than

ten arms was not conclusive.

JJ

9

1

...

1

Unknown

10

5

9

14

»

11

1

1

Perhaps not more than ten arms.

>»

12

...

1

1

» »».

Male

12

2

2

»

11

...

1

1

1

38

It is thus clear that the majority (29 out of 38) of the zooids examined have ten
arms ; that of this number 15 were proved to be female, while the remaining 14 may
have belonged to that sex ; and that no individual proved to be male has ten arms.
It should be noted, however, that we are assuming, with ANDERSSON, that the specimens
from Stations 5 and 94 belong to the same species. Of three zooids in which twelve
arms are certainly or doubtfully present, two were ascertained to be male. There is
evidence that the female may occasionally have eleven or nine arms instead of the
normal number (ten), and that the male may have only eleven arms instead of the
number (twelve) stated by ANDERSSON to be characteristic of that sex. The evidence of
the occurrence of eleven arms in two female specimens appears to be quite satisfactory.
In counting the arms, an arm -bud has been reckoned as an arm even though, as in one
of these two, it is very small and but little developed.

Evidence of a similar nature was obtained from sections of the material received
from Stockholm. In six series of sections of female zooids, five arms could be counted
on one or both sides ; while in a single series of sections through a male zooid twelve
arms were ascertained to be present.

Turning to the Discovery specimens described by RIDEWOOD as C. hodgsoni, the
number of the arms has been re-examined in eight series of sections, with the following
result : —

Ten arms are present in 2 female zooids, and in 2 others in which the gonads are
too minute to be certainly distinguishable as ovaries. Ten is probably the number of
the arms in 1 other case in which the sex is not determinable. Three zooids are male ;
and of these 1 has twelve arms, 1 has not less than eleven arms, and 1 has not less than
ten arms, while the possibility of the occurrence of twelve arms is not excluded.

(BOY. SOC. EDIX. TRASS., VOL. XLIX, 561.)

436 DR S. F. HARMER AND DR W. G. HIDEWOOD ON THE

In six individuals of C. hodgsoni which were partially dissected and in which ovaries
could be identified with certainty the results were somewhat less uniform. Two of
these zooids had ten arms each ; 1 had eleven ; and 3 had twelve arms. But even
here there is a sufficient amount of correspondence with the results arrived at by
ANDERSSON to give some support to the conclusion that C. ingequatus is synonymous
with C. hodgsoni, on the assumption that the number of arms is not invariable.

CffiNCECIUM.

It can hardly be doubted, from the published figures, that there is at least a consider-
able resemblance between the coenoecium of C. hodgsoni and that of C. insequatus.
ANDERSSON'S figure (07, pi. ii. fig. 1) of the latter closely resembles RIDEWOOD'S (071, pi. ii.
fig. 1) of the former. It may be noted that both figures represent the natural size of
the object. In both cases the general appearance of the coenoecium and its mode of
branching are substantially the same ; and the same result is arrived at by a comparison
of the actual specimens. RIDEWOOD states (p. 51) that the spines are simple, forked, or
trifid, and that the length of their free part is variable, usually within the limits 5 to
15 mm. ANDERSSON states (p. 10) that the length of the " Ausliiufer," by which we
understand the free parts of the spines, may reach 20 mm. ; and (p. 19) that they may
be divided into two or three branches. Although this latter statement is made in a
paragraph which refers to two other species as well as to C. in&quatus, an inspection
of his figure shows that it really refers to the species under consideration.

We may add that the comparison of the actual specimens shows that C. ingequatus
resembles C. hodgsoni in general size and proportions of the colony, in the colour of the
ccenoecium, and in the length and thickness of the spines. In both forms the ostia are
elliptical and their average long diameter is 3 mm. ; while in both the number of spines
associated with each ostium is about four or five.

MEASUREMENTS OP ZOOIDS.

According to RIDEWOOD (071, p. 53), the zooids of C. hodgsoni measure about 2 mm.
from the caecal end of the body to the dorsal border of the proboscis. ANDERSSON
(07, p. 10) gives the length of a zooid of C. ingequatus which is not too strongly
contracted as about 3 mm. It must be remembered, however, that no special care had
been taken, in preserving the Discovery material, to obtain well-extended zooids ;
and that ANDERSSON had the opportunity of examining the zooids of the form described
by him in the living condition. It is, moreover, not quite clear how his measurement
was taken.

We have ourselves measured several zooids of both forms in more or less median sagit-
tal sections, the measurement being taken in each case from the extremity of the caecal
end of the metasome to the dorsal border of the proboscis. According to these measure-

(EOY. soc. EDIN. TRANS., VOL. XLIX., 562.)

PTEROBRANCHIA OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 439

x

ments, four individuals of C. hodgsoni, of both sexes and in varying degrees of con-
traction, measured from 1'568 to 1'856 mm. Four individuals of C. in&quatus,
similarly measured, fell within the limits 1'760 to 2'240 mm. The longest of these
zooids in particular was obviously well extended. It may accordingly be concluded
that while, on the whole, the zooids of the material collected by the Swedish Expedition
are rather larger than those obtained by the Discovery, there is no essential difference
in size between the zooids of the two forms under consideration.

Similar results were obtained by a study of the length and thickness of the noto-
chord, the size of which ANDERSSON claims to be of some taxonomic value (07, p. 60).
The measurements taken from sections of C. inasquatus are on the whole greater than
those of C. hodgsoni, but the two series of measurements overlap.

The buds of C. in&quatus were examined to see if there is any appreciable difference
between them and those of C. hodgsoni ( RIDE WOOD, 072, p. 225), particularly in respect
of the mode of development of the arms. In size and general proportions, and in the
mode of development of the arms, the two sets of preparations are in very close
agreement.

We are accordingly of opinion that C. insequatus should be regarded as a synonym
of C. hodgsonj,.

REFERENCES.

(07) ANDERSSOX, K. A., "Die Pterobranchier der Schwedischen Siidpolarexpedition, 1901-1903," Wiss.

Eryebn. Schicedischen Siidpolar-Expedition, Bd. v., 1907, pp. 1-122, eight plates.
(12) GRAVIER, CH., "Sur une espeee nouvelle de Cephalodiscus (C. Anderssoni, nov. sp.) provenant de la

seconde Expedition antarctique francaise," Bull. Mus. Hist. not. Paris, 1912, No. 3, pp. 1-5,

two text-figures.
(87) HARMER, S. F., Appendix to Report on Cephalodiscus, " Challenger" Reports: ZooL, vol. xx. pt. Ixii.,

1887, pp. 39-47, four text-figures.

(97) - - ''On the Notochord of Cephalo'litru*.'" Zool. Anzeiger, xx., 1897, pp. 342-346.

(05) " The Pterobranchia of the Siboga Expedition, with an Account of other Species." Resultats des

Explorations entreprises aux Indes N?erlandai«es Orientates en 1899-1900 a bord du " Siboga,"
livr. xxii., Mouogr. xxvi. bis, Leiden, July 1905, pp. 132, fourteen plates and two text-figures.

(05) LANKESTBR, E. R., "On a New Species of Cephalodiscus (C. nigrescens) from the Antarctic Ocean," Proc.
Rny. Soc. London, 1905, vol. Ixxvi. B, pp. 400-402, one plate.

(87) M'lsiosH, W. C., "Report on Cephalodiscus dodecalophut," "Challenger" Reports: ZooL, vol. xx.
pt. Ixii., 1887, with Appendix by S. F. HARMER, pp. 48, seven plates and six text-figures.

(971)MASTERMAX. A. T., "On the Diplochorda: II. The Structure of Cephalodiscus," Quart. Journ. Micr.
Sci., N.S.. xl. pt. ii., 1897, pp. 340-366, four plates.

(97-) " On the ' Notochord ' of Cephalodiscus," Zool. Anzeiger, xx., 1897, pp. 443-450, five text-
figures.

(98) - - " On the further Anatomy and the Budding Processes of Cephalodiscus dodecalophus," Trans.

Roy. Soc. Etlin., xxxix. pt. iii., Dec. 1898, pp. 507-527, five plates.

(99) - - "On the ' Notochord ' of Cephaltdiscus," Zool. Anzeiger, xxii., 1899, pp. 359-363.

(03) "On the Diplochorda: IV. On the Central Complex of Cephalodiscus dodtcalophus" Quart.

Journ. Micr. Sci., N.S., xlvi., 1903, pp. 715-727, two plates.

(ROY. soc. KDIN. TRANS., VOL. XLIX., 563.)

440

DR S. F. HAEMER AND DR W. G. RIDEWOOI) ON THE

(06) RIDEWOOD, W. G., "A New Species of Cepltalodiscus (C. gilchriati) from the Cape Seas," Marine

Investigations, South Africa, iv., Cape Town, 1906, pp. 173-192, three plates and five text-figures.
(071) " Pterobranchia : Cephalodiscus," National Antarctic Expedition: Natural History, vol. ii.,

London [Brit. Mus.], 1907, pp. 1-67, seven plates and seventeen text-figures.
(072^ « On the Development of the Plumes in Buds of Cephalodiscus," Quart. Journ. Mier. Sri.,

N.S., li., 1907, pp. 221-252, eleven text-figures.

(12) "On Specimens of Cephalodiacus nic/rescens supposed to have been dredged in 1841 or 1842,"

Ann. Mag. Nat. Hist. (8), x., Nov. 1912, pp. 550-555, one text-figure.
(05) SOHEPOTIEFF, A., " Zur Organisation von Cephalodiacus," Bergens Museums Aarbog, 1905, No. 8,

pp. 20, two plates.

(07) "Die Pterobranchier : Anatomic von Cephalodiscus," Zool. Jahrb., Abth. filr Anat., xxiv., 1907,

pp. 553-608, eleven plates.

(08) " Die Pterobranchier : Knospungsprozess von Cephalodiscus, und vergleichend-anatomischer Teil,"

Zool. Jahrb., Abth. filr Anat., xxv., 1908, pp. 405-491, four plates.

(09) "Die Pterobranchier des Indischen Ozeans," Zool. Jahrb., Ablh. fur Systematik, xxviii., 4, 1909,

pp. 429-445, two plates.
(12) SPENGEL, J. W., "Pterobranchia," in Handwi'irterbuch der Naturwissensch often (Korschelt and others),

vii., 1912.

LIST OF REFERENCE-LETTERS.

an., anus.

b.c.1, proboscis-cavity.
b.c.z, collar-cavity.
b,c?a., anterior dorsal horn of collar-cavity.

b.c.s, metasomatic cavity.

b.c.2b., part of metasomatic cavity in loop of ali-
mentary canal.
c.c., collar-canal.

c.c.i., internal opening of collar-canal.
c.n.s., central nervous system.
d.m.z, dorsal mesentery of collar.
dm.3, dorsal mesentery of metasome.
g.s., gill-slit.
int. , intestine.
L^-L.9, left arms.
l.n., lateral nerve.
m., mouth.
ms., longitudinal muscles of metasome.

nch., notochord.
n.t., nerve-tract.

op., operculum or postoral lamella.
op.l., left lobe of operculum.
op.r., right lobe of operculum.
op.rec., recess at base of operculum.

p., proboscis.
per., peristome.
ph., pharynx.
p.p., proboscis-pore.

?•., rectum.
R^-R.0, right arms.

st., stalk.
ftom., stomach.

t., testis.
t.L, left testis.
t.r., right testis.
i:.m.~, ventral mesentery of collar.

(ROY. 8OC. EDIN. TKANS., VOL. XLIX., 5(>4.)

PTEROBRANCHIA OF THE SCOTTISH NATIONAL ANTARCTIC EXPEDITION. 441

EXPLANATION OF PLATES.
PLATK I.

Figs. 1—5. Selected pieces of colony of Cephalodiscus agglutinans, n. sp., drawn by Miss G. M.
WOODWARD. Natural size. The specimen drawu in fig. 1 is selected as the type-specimen of the species.

PLATE II.
Cephalodiscus agglutinans, n. sp.

The figures were drawn with the Zeiss objectives severally indicated, and were then reduced to two-fifths
linear. The scale for the two objectives used in drawing the figures is given, in hundredths of a millimetre,
at the bottom of the plate.

Fig. 1. A young bud. Obj. A.

Fig. 2. A slightly older bud. Obj. A.

Fig. 3. An older bud. Obj. A.

Fig 4. A still older bud. Obj. A.

Fig. 5. Obliquely frontal section, showing both testes (t.l., t.r.), parts of both gill-slits (</.?.), and the
left collar-canal (c.c.). Obj. C.

Figs. 6-11. Frontal sections of one zooid.

Fig. 6. Showing eight of the left arms (as numbered) and seven of the right arms. Obj. C.

Fig. 7. Showing the same fifteen arms, the anterior dorsal horns of the collar-cavities (b.c.*a.), the peri-
cardium (per.) enclosing the pericardial sinus or heart, and the internal openings of the proboscis-pores (p.p.).
Obj. C.

Fig. 8. Showing the anterior end of the notochord (nch.) and of the right lobe (op.r.) of the operculum.
Obj. C.

Fig. 9. Through the mouth (m.). The double appearance of the left lobe (op.l.) of the operculum is
due to the fact that the organ in question is folded ventrally. To see the correspondence between the two
sides of the section, the part of the left lobe which is nearer the proboscis (p.) must be imagined to have
been folded dorsally. It would then not have appeared in the section, and the left lobe of the operculum
would have had much the same appearance as the right lobe. Obj. C.

Fig. 10. The mouth (m.) is still visible. The reflected part of the left lobe (op.l.) of the operculum is
almost separate from the rest of the organ. Obj. C.

Fig. 11. Through the region of the collar-canals (c.c.) and gill-slits (g.s.). The right testis (t.r.) is very
small in this zooid. Two of the terminal lobes of the scalloped edge of the right half (op.r.) of the operculum
are visible. Obj. C.

Fig. 12. Stomach (atom,), intestine (int.), rectum (r.), aiid one testis (t.*) of an old zooid. Obj. A.

Fig. 13. Stalk, in transverse section. Obj. C.

Fig. 14. Proboscis-stalk, in transverse section, showing both proboscis-pores (p.p-), the anterior dorsal
horns (b.c.-a.) of the collar-cavities, the anterior end of the central nervous system (c.n.s.), and the peri-
cardium (per.) with the included pericardial sinus. Obj. C.

Fig. 15. Median sagittal section of the anterior end of a zooid, showing the notochord (nch.). Obj. C.

Fig. 16. Nearly median sagittal section of another zooid, showing the alimentary canal. Obj. C.

N.B. — In several of the sections, figured parts of the epithelia were somewhat macerated, as the result
of imperfect preservation. The epithelium of the stomach in fig. 16, part of the anterior epidermis of the
proboscis in figs. 6-9, and part of the epidermis containing the central nervous system in figs. 10 and 11
have accordingly been restored where necessary.

(HOY. soc. EDIN. TRANS., VOL. XLIX., 565).

Scot. Nat. Ant. Exp.

HARMER & RIDEWOOD : PTEROBRANCHIA,

Vol. IV.

I

G.M.Woodward del.etlith.

imp.

Scot. Nat. Ant.Exp. Vol. IV.

HARMEK AND RIDEWOOD : " SCOTIA" PTEROBKANCHIA— PLATE II.

Fig. 16.

Obj. A.

Obj. C.

trFarlanr 4. Enkine. Lilh Edin

PART XVIII.
SEALS.

SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

XYIII.-SYSTEMATIC ANATOMY OF A FCETAL SEA

LEOPARD (Stenorhynclnis leptonyx) AND MICROSCOPICAL

ANATOMY OF SOME OF THE ORGANS.

BY HAROLD AXEL HAIG, M.B., B.S., M.R.C.S.,

Late of University College, Cardiff; M'Robert Research Fellow, University of Aberdeen.
(WITH FOUR PLATES AND THREE TEXT FIGURES.)

XVIII. — Scottish National Antarctic Expedition : A Description of the Systematic
Anatomy of a Foetal Sea- Leopard ( Stenorhynchus leptonyx), with Remarks upon
the Microscopical Anatomy of some of the Organs. By Harold Axel Haig,
M.B., B.S., M.R.C.S., late Lecturer in Histology and Embryology, University
College, Cardiff; M'Robert Research Fellow, University of Aberdeen. Com-
municated by Professor ARTHUR ROBINSON, M.D. (With Four Plates and Three
Text-figures.)

(MS. received January 26, 1914. Read February 16, 1914. Issued separately April 30, 1914.)

During the Scottish Antarctic Expedition of 1892-93 Dr W. S. BRUCE secured
fcetuses of Stenorhynchus leptonyx and Lobodon Carcinophaga, and on his return
passed them over, with other material, to Professor D'ARCY W. THOMPSON for the
Zoological Museum of University College, Dundee. While some of the material has
unfortunately been lost sight of during these twenty-one years, one specimen, viz. that
of a foetus of Stenorhynchus leptonyx, was still in existence, and was returned by
Professor D'ARCY THOMPSON to Dr BRUCE, who in turn asked me to examine and
report upon it. Furthermore, during the voyage of the Scotia embryos of Leptony-
chotes weddelli were obtained by the Scotia naturalists, and these were passed on for
description to Professor WATERSTON, at that time in the University of Edinburgh. It
is on this material that the present monograph is based.

PRELIMINARY CONSIDERATIONS.

The foetus of Stenoryhnchus leptonyx, which is in a good state of preservation,
measures 122 mm. from the tip of the tail to the most prominent part of the mid-
brain ; the greatest breadth is about 43 mm., and the greatest dorsi-ventral measure-
ment is in the mid-dorsal region, measuring 35 mm. The umbilicus is situated 36 mm.
from the cloacal aperture, the umbilical cord being relatively short, owing to the
fact that the umbilical vessels soon undergo division into several large branches ; a
portion of the placenta with fragments of the foetal membranes is still attached
to the cord. The actual mode of placentation and the disposition of the foetal
membranes are points which will be discussed at a later stage (see infra, on the
Placenta).

The skin is in many places thrown into folds, some of these being normal, but
others undoubtedly due to shrinkage consequent upon the action of the preservative
medium. The flippers are fully formed, nail rudiments being present upon the
dorsal aspects of the distal phalanges.

(REPRINTED FROM THE TRANSACTIONS OF THE ROYAL SOCIETY OF EDINBURGH, VOL. L., PP. 225-251.)

448 MR HAROLD AXEL HAIG ON

The head is at this stage not large in proportion to the trunk ; rudiments
of the vibrissae are to be seen at the sides of the snout, and the eyelids are
formed although the palpebral fissure is not as yet open. The tongue is an
elongated organ, with a cleft tip, the two divisions appearing between the lips in
the middle line.

No sign of an external ear is present, nor is there any opening indicating the
position of an auditory meatus.

The whole trunk is curved towards the ventral aspect, but some of the curvature
is probably due to mechanical causes subsequent to removal of the foetus from
the uterus.

Reference to PL I. fig. 1 will render the above points clear.

SECTION I.

GENERAL TOPOGRAPHY AND ANATOMY.
A. Appearance of the Main Viscera in situ, from the Ventral Aspect.

A median ventral incision was made, and flaps of skin and deeper tissues turned
back to expose the structures in the neck, thorax, and abdomen ; the sternum and
ribs being also removed for the purpose of demonstrating the thoracic viscera, whilst
the attachments of the diaphragm to the lower ribs were likewise severed. A
separate flap was raised in the neck region to expose the larynx, trachea, thyroid, and
parathyroids, and finally the parietal pericardium was cut away.

In the region of the umbilicus, care was taken not to sever the connection of the
urinary bladder with the umbilical cord, and the umbilical vein passing from the
cord to the liver was also kept intact ; subsequently, however, these connections were
severed for greater convenience of examination.

Reference to fig. 2, Plate L, will show that the heart is at this stage a large
organ filling the greater part of the thoracic cavity ; from the ventral aspect, the
right ventricle appears larger than the left, and the right auricular appendix wider
than the left ; the right appendix has a deep notch in its lower border, whilst the
left one possesses three such notches.

A portion of the aortic trunk shows above and dorsal to the right appendix.

The thymus is relatively large, and extends from the root of the neck, where it is
bifurcated, towards the left of the middle line, until it reaches a point just anterior
to the left auricular appendix : the main mass of the thymus is subdivided into a
number of lobes and lobules, and there are a few small isolated masses situated at
the sides of the trachea just anterior to the upper bifurcated extremity.*

The lungs lie compressed against the walls of the thorax, and are not very obvious
from the ventral aspect, the right upper lobe being most prominent, and below

* Microscopical examination showed that these isolated masses possessed a typical thymus structure.

(ROY. soc. KDIN. TRANS., VOL. L., 226.)

THE SYSTEMATIC ANATOMY OF A FCETAL SEA-LEOPARD. 449

this a small portion of the middle lobe, whilst none of the lower lobe is visible ;
both lobes of the left lung show, the anterior edges and portions of the lateral
surfaces being seen. In the neck region the larynx and trachea form prominent
features, whilst at the sides of the trachea the lateral lobes of the thyroid gland
with the lower parathyroids are to be seen ; no thyroid isthmus is, however, to be
detected, a point which is noteworthy.

The abdominal viscera from the ventral aspect (PI. I. fig. 2) : — The liver
forms the most prominent organ in the abdomen, its right and left lobes, together
with certain accessory lobes, occupying about one-half the available space ; the right
lobe appears to be larger than the left, but in reality this is not the case, since, when
viewed from the dorsal aspect (see PI. III. fig. 5), the left lobe is seen to be much
the bulkier of the two. A fissure passes obliquely downwards and inwards from the
middle of the lateral aspect of the right lobe, and effects a partial subdivision of this
lobe into two, but the cleft does not extend deeply into its substance ; whilst a
small flap of the upper of the two subdivisions is seen a short distance internal to
the right lateral margin, and partly conceals an aperture in the lobe in which the
fuiidus of the gall-bladder appears (PL I. fig. 215).

Between the right and left lobes there is a fairly wide cleft, in which may be seen the
umbilical vein passing from the umbilicus towards the ductus venosus.

Coils of small intestine are seen lying caudal to the liver, but the stomach is not
visible, being largely hidden by the left hepatic lobe.

The urinary bladder is a very elongated structure, and is attached ventrally to
the umbilicus : it opens caudally into the cloaca. The umbilical arteries pass along
the lateral aspects of the viscus towards their destination in the placenta, and are
well seen in transverse sections. (See PI. III. fig. 4.)

The chief points of interest in connection with the above description of the
ventral aspect of the viscera are : —

(i.) The relative size of the thymus, which, although as a rule large at this stage
of development, appears in the present case to be markedly so ; the left
lateral deviation of its caudal extremity is also a point worthy of note.
The presence of isolated lobules of this gland suggests a possible origin of
these from some of the higher branchial clefts.

(ii.) The deep notching of the auricular appendices in the heart is a feature which
seems very striking upon opening the pericardial cavity. One other point
also, viz. the well-marked interventricular furrow, is a characteristic which
becomes more marked as development proceeds, the external subdivision of
the ventricular portion of the heart giving it in the full-grown seal an
almost bifid appearance.

(iii.) The great longitudinal extent of the liver, an organ occupying at this stage of
development a relatively large proportion of the abdominal cavity.
(ROT. soc. EDIX. TRANS., VOL. L., 227.)

450 MR HAROLD AXEL HAIG ON

(iv.) The peculiar shape of the urinary bladder. According to HEPBURN,* the
bladder in the adult seal is represented by the whole length of the foetal
organ, since no shrinkage takes place in the cephalic portion to form a
definite urachus.

B. General Topographical Anato^ny of the Head.

I. The oral, nasal, and cranial cavities, with their contents (PL II.).
. A median sagittal section of the head was made, so that the oral, nasal, and
cranial cavities were laid bare ; whilst, in order to expose the contents of the nasal
fossae, the nasal septum was subsequently removed.

Rudiments of the incisor milk-teeth were cut through in the upper and lower
jaws, and the surface of the mesial section of the hypophysis cerebri exposed, the
hypophysis lying in the sella turcica of the developing sphenoid bone at the base of
the skull (PI. II. fig. 33).

Other points worthy of note in such a hemisection of the head are :—

(a) The elongated tongue (PI. II. fig. 32).

(b) The falx cerebri, covering the mesial surface of the right cerebral hemisphere

(PI. II. fig. 39) ; about the middle of the concave edge of the falx the
corpus callosum is seen in section (PL II. fig. 39'.)

(c) The optic thalamus (PL II. fig. 38) continued posteriorly into the

mesencephalon and pons ; dorsal to these latter, the Sylvian aqueduct with
its roof, in which the rudiments of the corpora quadrigemina (fig. 37)
are to be made out ; posterior to these are seen the mesial section of the
cerebellar vermis, and, ventral to this, the 4th ventricle and medulla
oblongata (PL II. fig. 33'6'6'4).

(d) The cavity of the 3rd ventricle, with its extension into the infundibulum of

the hypophysis.

After the removal of the falx cerebri and nasal septum, the following additional
structures come into view :—

In the cranial cavity (PL II. fig. 5) :—

(e) The mesial surface of the cerebral hemisphere (h.).

(/) The olfactory lobe (olf. I.), lying between the fore-brain and the anterior
boundary of the cranial cavity : the lobe is not a large one, and does not
appear to give off many nerve-filaments to the ethmoidal region of the
nasal fossa.

In the region of the nasal fossae :—

(g) The rudiment of the ethmo-turbinal bone (eth. t.) lying at the superior angle ;
this rudiment, which at the present stage is cartilaginous, shows three ridges

* Tram. Roy. Soc. Edin., vol. xlviii. part i., No. 3, 1913.
(ROY. soc. EDIN. TRANS., VOL. L., 228.)

THE SYSTEMATIC ANATOMY OF A FCETAL SEA-LEOPARD. 451

with two intervening depressions, and represents the olfactory region of the
nasal fossa.

(h) The large maxillo-turbinal rudiment (mx. t.), presenting upon its mesial
surface a number of narrow lamellae with intervening sulci : it is carti-
laginous and is covered by a mucous membrane lined by ciliated epithe-
lium. Anteriorly the maxillo-turbinal bone is attached to the outer wall
of the fossa, whilst posteriorly it fuses by an elongated pedicle with the
periosteum of the base of the skull.

The buccal cavity is lined by a mucous membrane covered by stratified epithelium
with but few cell-layers. At the present stage of development, the membrane bones
of the cranial vault are partially ossified, but the bones of the base of the skull (basi-
sphenoid, basi-occipital) are still partly cartilaginous ; the sphenoid bone (see text-fig.
3, 6) is in the " irruption "-stage of endochondral ossification, the process being seem-
ingly delayed by the rather late persistence of the epithelial connection between the
hypophysis cerebri and the buccal epithelium.*

The brain. — General superficial anatomy (PI. II. figs. 1 and 2). The brain of
the present specimen shows some very well-defined characters, and on the whole
may be said to conform to the mammalian type : viewed from the dorsal aspect
(PI. II. fig. l), the following features are obvious :—

(i.) The large cerebral hemispheres separated by the longitudinal sulcus in which

the falx cerebri is normally lodged,
(ii.) Behind the hemispheres, the crura cerebri and region of the mesencephalon,

covered by fragments of torn pia mater,
(iii.) The cerebellum, consisting of the mesially situated rudiment of the vermis,

and on either side of this the cerebellar hemisphere : the latter already

present a number of flattened laminae with intervening sulci.
(iv.) The 4th ventricle in its lower half, with the restiform bodies bounding it

on either side ; the floor of the ventricle is seen owing to previous removal

of the roof of pia mater.

The surface of each cerebral hemisphere is quite smooth upon its superior aspect,
and shows no indications as yet of any convolutions.

Viewed from the side, several additional features become apparent, viz. (PI. II.
fig. 2) :-

(v.) The olfactory lobe, and its connection with the ventral part of the fore-brain,
(vi.) A wide shallow depression passing from the lower aspect of the fore-brain
upwards and backwards towards the hind-brain (future occipital lobe) :
this depression indicates the position of the future Sylvian fissure (PI. II.
fig. 22), and ventral to its posterior extremity a forward extension of the
hemisphere represents the rudimentary temporo-sphenoidal lobe.

* See also P. T. HERRING ou " Development of Mammalian Pituitary Bodv," Journ. Exper. Physiol., 1909.

(ROY. 8OC. EDIN. TRANS., VOL. L., 229k)

452 MB HAROLD AXEL HAIG ON

Upon the ventral aspect, at the base of the brain, the infundibulum forms a down-
ward projection from the floor of the 3rd ventricle (PI. II. fig. 2s), whilst half way
between the infundibulum and a projection which marks the position of the ventral
part of the pons the 3rd nerve forms a noticeable feature (PI. II. fig. 24).

Additional features to be made out in a sagittal section taken through a cerebral
hemisphere, the optic thalamus and mid- and hind-brain (PI. II. fig. 4) : — In such
a section the general relations of the various regions of the brain may be studied to
a certain extent, and, moreover, by staining with hsematoxyliu and eosin the main
distribution of grey and white matter at this stage of development may be made out.
The cortex cerebri (PL II. fig. 4*' 5) is made up of two parts, viz. :—

(i.) A superficial layer of white fibres and neuroglia, forming a very narrow zone

(PI. II. fig. 46).
(ii.) A deeper layer of deeply-staining nerve-cells, with neuroglia, of somewhat

wider extent than (i.).

Deep to the cortex comes the white matter of the hemisphere, which is at this
stage not very thick. A section taken so as to pass completely through the cavity of
the prosencephalon shows in addition an inner layer, somewhat deeply stained,
which later on will form the ependymal lining of the lateral ventricle.

Below the main mass of the hemisphere, part of the descending horn of the lateral
ventricle is seen (PL II. fig. 4"'), and this becomes continuous anteriorly with
the rhinencephalon, passing into the olfactory lobe. The anterior boundary of the
descending horn is formed by a layer of grey matter, which above takes the form of
radiating streaks, alternating with white matter.

In the region of the optic thalamus, small isolated patches of grey substance
appear near the dorsal surface, but in the region of the corpora quadrigemina white
matter seems as yet to predominate.

The grey matter of the cerebellar hemisphere appears to lie chiefly on the surface,
but there is a deeper zone of small nerve-cells, the two layers being separated by a
clear zone of white matter : the cells of Purkinje do not as yet appear to have become
differentiated as a distinct line of neuroblasts. In the region of the medulla oblongata,
the nuclei of the 10th and 12th cranial nerves form a series of groups of rather large
nerve-cells (PL II. fig. 41) ; whilst the pyramidal tract appears as a well-defined
longitudinal set of fibres.

The infundibulum, with a small portion of the cavity of the 3rd ventricle, has been
already noted ; the ventral mass of the pons is a very obvious feature lying just
anterior to the medulla (PL II. fig. 4).

A fold of pia mater (PL II. fig. 4s) shows between the overhanging posterior
extremity of the cerebral hemisphere and the mesencephalon, whilst in the deep
fissure anterior to the optic thalamus a small piece of choroid plexus appears.

Points for comparison with the brains of other carnivora are the following :—

(ROY. soo. EDIN. TRANS., VOL. L., 230.)

THE SYSTEMATIC ANATOMY OF A FCETAL SEA-LEOPARD. 453

(a) The relative shortness of the corpus callosum, a structure which, for instance, in
the cat, at a corresponding stage of development has assumed a much longer
antero-posterior measurement ; moreover, it would seem that in this Seal
embryo the corpus callosum is taking a more vertical course than is usual.

(6) Altogether the general appearance and stage of development of the brain under
discussion corresponds very closely with one figured by His * of a three-
months human foetus, with the exceptions that the hypophysis cerebri is
much further advanced (see infra) and the cerebellum has assumed greater
complexity.

C. Anatomical Details of the Remaining Viscera.

The thyroid and parathyroids, the thymus, pancreas, and tongue, will be described
under the histological section of this pamphlet ; in the present instance the following
organs will be considered, and comparisons made with other types, where this is

possible : —

(i.) The lungs.

(ii.) The heart.

(iii.) The liver, stomach, and intestines,
(iv.) The spleen.

(v.) The kidneys and adrenal bodies,
(vi.) The genital glands and ducts : the urinary bladder.

The histology of some of these will be dealt with later.

(i.) The lungs (PI. III. fig. 3). — Both lungs are somewhat compressed against
the thoracic walls, owing to the large size of the heart, and it is only upon removing
the latter organ that a good view can be obtained of the roots of the lungs and their
ventral aspect ; posteriorly the inner margins of the upper and lower lobes of both
lungs are grooved longitudinally by the vertebral column, whilst laterally the surfaces
of all the lobes are grooved by the ribs.

In the right lung, three lobes, upper, middle, and lower, can be distinguished t ;
the middle lobe is peculiar in that at the root it gives off two caudally directed
subdivisions, the largest of which is almost a separate lobe, and presents dorsal,
right lateral, and anterior surfaces (PI. III. fig. 3 a). The cranial and caudal lobes
of both lungs are somewhat similar in shape, the caudal lobes being the larger, being
not unlike the corresponding lobes of a human lung. Marked trabeculse of con-
nective tissue are to be seen upon the surfaces of all lobes, with finer strands passing
off in all directions, and these subdivide the surface into polygonal areas, the bases,
as it were, of the superficial lobules ; on being placed in water the lungs sink.

At the root of the lung the bronchi and pulmonary vessels are seen entering (or

* HERTWIG, Handbuch dtr Enticickelungslehre der Wirbeltiere.

t The terms " upper," etc., are here used with reference to the vertical position of the trunk, and not the natural
position of the adult animal.

(ROY. soc. EDIN. TRANB^ VOL. L., 231.)

454

MR HAROLD AXEL HAIG ON

emerging from) the upper lobes, the bronchi being dorsal to the pulmonary arteries,
although the left bronchus is much closer to the artery than the left.

(ii.) The heart (PI. II. fig. 2 and text-fig, l). — All four cavities of the heart
contain firm clot, which extends into the auricular appendices ; on section in the
coronal plane, the cavities of the auricles appear to be larger than those of the
ventricles, whilst the auricular appendices add considerably to the auricular capacity.
Moreover, although from the ventral aspect the right ventricle appear larger than
the left, there is not the same relative difference between the capacities of the
ventricles, both appearing to possess much the same size in median coronal section ;
the thickness of the myocardium is rather greater in the left than in the right
ventricle, that of the auricles being about equal on both sides. The aortic bulb is

FIG. 1.— Dissection to show the relations of the aortic trunk, pulmonary trunk, and ductus arteriosus.

L.V. Left ventricle in section. R.S. Right subclavian.

R.V. Right ventricle in section. AO. Aorta.

B.AO. Aortic trunk and bulb. P. A. Pulmonary artery.

IN. Innominate trunk. D.A. Ductus arteriosus.
R.C. Right common carotid.

The dotted lines show the connection of the pulmonary trunk with the right ventricle ;
the lumen of the aortic bulb is opened, and two semilunar valves are seen.

full of firm clot, and passes at once into a relatively short but thick aortic arch ; the
pulmonary trunk shows the same relations to the aortic trunk as it does in the case
of the human foetus of a corresponding stage of development, whilst the ductus
arteriosus is relatively wide and is a continuation of the main pulmonary trunk after
the latter has given off the two branches to the lungs (see fig. 2). The ductus joins
the arch of the aorta close to the origin of the latter from the bulbus aortse.

The cardiac valves are well developed, the semilunar valves consisting of thin
plates of fibrous tissue, the free edges of which project into the aortic and pulmonary
trunks, whilst the mitral and tricuspid valves have their free edges projecting into
the ventricular cavities. The coluinnse carnse are not marked features, nor do the
chordae tendinse appear to be strongly developed at this stage.

(ROY. soc. EDIN. TKANS., VOL. L., 232.)

THE SYSTEMATIC ANATOMY OF A FCETAL SEA-LEOPARD. 455

Serial sections taken coronally through the heart show that the foramen ovale is
a feature at this phase, but the aperture is a very narrow one, and will not admit a
glass seeker of more than 1 mm. diameter.

From the above description it will be seen that the heart of Stenorhynchus during
early foetal life corresponds more or less closely with the typical mammalian organ
in its developmental aspects ; the iuterventricular furrow, however, is a marked ex-
ternal feature, and during development, at least in the Weddell Seal, it becomes
deeper, so that the full-grown heart possesses a bifid apex, the tips of the two
ventricles being separated by a deep notch.*

(iii.) The liver, stomach, and intestines (PI. III. figs. 1 and 5). — In some respects
the liver of this specimen shows characters similar to those of the human foetal organ,
but the accessory lobes, and the clefts which produce these, are to a certain extent
atypical ; thus the right lobe shows from the ventral aspect a fissure which passes
obliquely inwards from the lateral surface, but this fissure is not seen from behind,
as it extends for only a short distance into the substance of the lobe.

From the dorsal aspect (PI. III. fig. 5) it appears that the left lobe is the bulkier
of the two, and the inner surface of this lobe shows depressions corresponding to
the ventral surface of the stomach, a small area of the spleen, and anteriorly the
oesophagus and vena cava ; the inner surface of the right lobe shows dorsally de-
pressions corresponding to the numerous subdivisions of the right kidney, whilst the
lateral aspects of both lobes are grooved by five or six of the posterior ribs. In the
middle line ventrally the umbilical vein (PI. III. fig. 5, l.v.) forms a thick cord,
passing towards the ductus venosus.

The gall-bladder is not seen from the dorsal aspect except when the lobes are
widely separated, when it appears as an elongated sac, deeply embedded in the deep
surface of the right lobe, the fundus presenting veutrally in the small opening noted
in the topographical description (see supra).

Certain accessory lobes show up when the two main lobes are separated from one
another : these may possibly be the homologues of the Spigelian and quadrate lobes
of the human organ, but their relations are somewhat different.

The main points for comparison to be noted in this organ are : —

(i.) The relative longitudinal extent of both lobes, this being distinctly greater

than is the case with most other carnivora.
(ii.) The small accessory flap guarding the aperture in which, ventrally, the fundus

of the gall-bladder is to be seen.
(iii.) The large volume of the abdominal cavity, occupied by the whole liver, at a

stage when other abdominal viscera have assumed a relative importance

in size.

The stomach and intestines (PI. III. fig. l) are seen from the ventral aspect, after

* See HEPBURN, Trans. Roy. Soc. Edin., vol. xlviii., 1913.
(ROY. soc. EDIN. TRANS., VOL. L., 233.)

456 MR HAROLD AXEL HAIG ON

removal of the liver, by which organ they are largely concealed ; the stomach is
placed with its long axis nearly parallel to the long axis of the foetus, and possesses
a well-marked cardiac extremity lying with its furidus pressed against the diaphragm,
and a pyloric end which is narrow and passes by a sharply curved portion into the
first part of the small intestine. The lesser curvature of the stomach looks towards
the ventral aspect, the greater curvature and fundus are dorsally situated, whilst
a peritoneal fold, the representative of a great omentum, is attached to the whole
length of the larger curvature ; the lesser curvature and the duodenum have
passing between them a narrow peritoneal sheet, which holds up the duodenum so
that its first part runs parallel with the stomach.

The stomach is an inch and a half in length, and at its cardiac end is joined by
the oesophagus, the latter being a somewhat wide tube, two inches long and quarter
of an inch in diameter.

The first part of the small intestine (duodenum) makes three bends, and possesses
four distinct portions, the first of which is parallel with the smaller curvature of the
stomach, whilst the second, third, and fourth divisions enclose a portion of peritoneum,
between the layers of which the pancreas is held in position. The coils of small
intestine proper are already many in number, and their general arrangement may be
made out by reference to PI. III. fig. 1.

With regard to the large intestine, the position of the caecal pouch is noteworthy :
this pouch is placed opposite the level of the third bend of the duodenum, being con-
nected with the latter by a short fold of peritoneum. No sign of a vermiform ap-
pendix is to be made out ; as a matter of fact, this organ is not seen in the full-grown
animal.*

(iv.) The spleen (PL III. fig. 6) has a situation upon the left side of the abdomen,
parallel to the greater curvature of the stomach and attached to this by a fold of
peritoneum ; in thickness this organ does not measure more than one-eighth of an inch,
but in length exceeds two inches. There are no notches in either its ventral or its
dorsal edge, and the hilus occupies a large proportion of a ridge forming its inner
margin which lies close to the stomach ; the outer surface of the spleen is grooved
by two or three of the posterior ribs.

(v.) The kidneys (PI. III. fig. 4). — These organs are somewhat elongated oval
bodies lying low down at the back of the abdominal cavity and close to the middle
line ; each kidney belongs to the type common to the Pinnipedia, viz. the per-
manently subdivided type, where the organ is made up of a large number of anatomi-
cally distinct renal pyramids, t the secreting tubules of which open into a common
pelvis, from which latter a ureter conducts the secretion to the urinary bladder.

In this specimen there are in each kidney about two hundred and forty small

* HEPBURN (Trans. Roy. Soc. Edin., vol. xlviii. part i., No. 3, 1913) regards the csecal diverticulum as a combined
caecum and vermiform appendix.
t " Renculi " of German authors.

(ROT. soc. EDIN. TRANS., VOL. L., 234.)

THE SYSTEMATIC ANATOMY OF A FCETAL SEA-LEOPARD. 457

raised areas, circular in contour, representing the bases of the renal pyramids : these
are mostly uniform in size, but some, smaller than the majority, lie rather below the
general surface. A mesial coronal section of the kidney passes through many of
the pyramids and also opens up the calyces and pelvis ; according to CHIEVITZ,* a
certain amount of reduction takes place during development, so that whereas in certain
instances about two hundred calyces may be present, in the full-grown animal only
one hundred and forty remain, the reduction apparently commencing in those of the
6th and 7th order of origin.

The ureter emerges from about the middle of the inner and dorsal margin of the
kidney, and passes down parallel to the mid-line, crossing the Miillerian ducts dorsally at
right angles, and finally opens dorsally into the lower segment of the bladder ; the
hilus of the kidney from which the ureter emerges and into which the renal vessels
pass is not a marked feature. The " pelvis " mentioned above is also but little
developed, since the ureter divides almost at once into two main branches, the latter
undergoing further subdivision in the kidney until the final divisions are reached
close to the surface of the organ in the cortical zone of each renal pyramid.
Each kidney measures about one inch by half an inch. The adrenal bodies (PI. III.
fig. 4) are small reniform structures lying one on each side just anterior to the
kidney ; there is a loose connection with the latter organ by means of a peritoneal
band. In actual shape the adrenal is a flattened pyramid with a convex base facing
outwards and ventralwards, whilst the hilus is placed at about the middle of the
inner, or rather dorsal, edge, close to the vertebral column ; the length of each gland
is half an inch, and the breadth one quarter inch. Some points in the histology of
both the kidney and the adrenal gland will be described later.

(vi.) The genital glands with their ducts (PI. III. fig. 4) are in the present
specimen already sufficiently established to be able to determine the sex of the animal
— i.e. they are ovaries, and the Fallopian tubes pass from the outer ends of the glands
to fuse in the middle line dorsal to the bladder and ventral to the rectum, and from
the rudimentary uterus ; the outer ends of the Fallopian tubes are dilated to form
large ampullae for the reception of the ova, and these ampullae lie ventral to the
outer pole of each ovary.

Each gland is an ovoid body lying obliquely from without backwards and inwards
just behind the posterior pole of the kidney ; from its anterior extremity an elongated
narrow muscular band, the diaphragmatic ligament of the mesonephros, passes
towards the diaphragm and becomes attached to the dorsal wall of the abdomen.

The urinary bladder (PI. III. fig. 4s) is an elongated organ attached above by a
patent allantoic duct to the umbilicus ; on transverse section, three apertures are seen
—a median one, the bladder, and two lateral, the lumina of the umbilical arteries.
The latter are passing to the umbilicus from their origin in the aorta.

The bladder opens into the cloaca by a transversely elongated slit-like aperture,

* Archiv Anat. u. EmbryoL, Supplement, 1897.
(ROY. soc. KDIN. TRANS., VOL. L., 235.)

458 MR HAROLD AXEL HAIG ON

and the ureters are to be seen opening into the bladder laterally upon its dorsal
aspect ; it is noteworthy that in the seals the bladder is represented by the entire
intra-abdominal extent of the allantois.*

Summary of the Anatomical Features.

From the foregoing description it will be readily gathered that the Sea-Leopard
Seal, during the middle phase of foetal life, presents fairly typical embryological
features ; the age of the present specimen can hardly be worked out with any
approach to accuracy, but it may be stated that the stage of development of most
of the organs would place the fetus at about the end of the first third of intra-
uterine life. The exact period of gestation of seals is, however, somewhat difficult
to determine owing to the peculiar habits of mating which these animals have, so
that the above estimate should be accepted with some reservation.

In summarising the main anatomical features, it is possible to pick out the
following more obvious characters : —

(a) In the brain : firstly, the relatively advanced stage of the cerebellum, the

hemispheres of this portion showing distinct evidence of folds and sulci
which are not to be made out in a three-months human foetus ; secondly,
the comparatively advanced development of the pituitary body, a feature
which will be more readily appreciated when the histology of the hypo-
physis is considered (see infra).

(b) In the heart : the most prominent feature is the late persistence of the bulbus

aortse, whilst the peculiar shape of the auricular appendices is also worthy
of note.

(c) In the lungs : the possession of an accessory lobe by the right lung, and the

relative size of these organs, which are certainly large, are points of com-
parative value.

(d) Other points which may be emphasised are the large size of the thymus and

the left-sided deviation of this organ, the apparent absence of a thyroid
isthmus, and the large size of the lowest parathyroids ; in connection with
the alimentary tract, the forward position of the csecal diverticulum and
the great length of the lobes o'f the liver. The kidneys are noteworthy on
account of their surface lobulation into numerous renal pyramids, whilst
the adrenal bodies do not lie directly upon the anterior poles of the kidneys.
Moreover, the adrenals are relatively much smaller than is the case with
these bodies in the human foetus at a corresponding stage of development.
Lastly, certain external characters, such as the absence of an external ear and
the protruding bifid tip of the tongue, are features so obvious as to need
no further comment.

* HEPBURN, Trans. Roy. Soc. Edin., vol. xlviii. part i., No. 3, 1913.
(ROY. soc. EDIN. TRANS., VOL. L., 236.)

THE SYSTEMATIC ANATOMY OF A FCETAL SEA-LEOPARD. 459

SECTION II.

AN OUTLINE OF THE MICROSCOPICAL ANATOMY OF SOME OF THE ORGANS.

The histological characters of certain of the viscera were examined in this foetus,
and in a few instances, viz. kidney and pituitary body, were found interesting from
the point of view of the histogenesis of their essential secreting portions. The
following account must, however, be looked upon as purely descriptive in character,
since from the mere observation of features presented by a single specimen at one
stage of development it is hardly possible to formulate a complete account of the
histogenesis of any one organ.

(i.) The thyroid and parathyroids (see PI. III. fig. 2). — The lateral lobes of the
thyroid gland are situated rather far forward in the neck region, reaching the level
of the lower border of the cricoid cartilage ; each lobe measures about half an inch
in length. The largest parathyroid is the posterior one, and is placed upon the
mesial surface of the thyroid at the ventral and posterior angle of the lobe. The
anterior parathyroid is quite small, and is found about the middle of the dorsal
margin of the lateral lobe.

In minute structure the thyroid is seen to be made up of a large number of small
vesicles, some of them showing evidence of recent origin from branched tubular
columns of cells, and in some of them colloid is already to be detected ; the vesicles
are lined by a cubical epithelium, and there is a small amount of interstitial connec-
tive tissue, but no sign of a basement membrane outside the epithelium. The whole
lobe is subdivided into relatively few rather large lobules by coarser trabeculse of
connective tissue which are given off from the inner surface of a rudimentary
capsule which surrounds the lobes. Blood-vessels are fairly numerous, the larger
branches being supported by the coarser trabeculae of connective tissue lying
between the lobules.

The parathyroid (PI. III. fig. 2, p.th.) shows a quite typical structure : it is
enclosed by a capsule of somewhat open connective tissue, which gives off trabeculae
passing into the interior of the gland. The essential secreting cells occur in the form
of folded columns of cuboidal or polyhedral cells, whilst these columns are in close
apposition to the blood-channels, the latter being at this stage lined by a definite
endothelium ; the latter, however, is not so marked a feature as it is in the blood-
channels occurring in the pars anterior of the pituitary body of this foetus.

(ii.) The thymus shows on section and microscopical examination a number of
lobulated masses of lymphoid tissue supported by a framework of open connective
tissue in which run a few rather large blood-vessels ; here and there in the actual
substance of the lymphoid masses there occur a few areas which appear clearer and
which probably represent the remains' of the original lumina of the epithelial tubules
from which the gland arises.

(ROY. SOC. EDIN. TRANS., VOL. L., 237.)

460 MR HAROLD AXEL HAIG ON

The accessory portions of the thymus, noted in the general description of the
viscera as being isolated from the main mass, are in all probability derived from the
2nd branchial cleft, and do not fuse with the portions derived from the 3rd and 4th
clefts.

GROSCHUFF and VERDUN state that the thymus in carnivora arises invariably from
the 3rd and 4th clefts, but in the rabbit, according to VERDUN, additional parts may
arise from the 2nd cleft.* In the present instance, paired accessory portions were
found lying dorsal to the sterno-mastoid muscle on either side of the trachea, and
these on examination showed a typical thymus structure.

The lymphoid nodules of the thymus present a more or less uniform density with
the exception of the occurrence of the above-mentioned clearer areas, and no sub-
division into cortex and medulla is as yet obvious ; nor are any corpuscles of
Hassall to be observed.

(iii.) The lungs (PI. IV. fig. 3). — -Microscopical examination reveals in these
organs a structure entirely comparable to that of a compound tubular gland, the
branching tubules of which lie embedded in connective tissue ; the latter exists in
large amount and is of a fibro-cellular character. The epithelium lining all of the
ramifications of the bronchi is of the high columnar type, with the nuclei lying next
the basement membrane, whilst immediately outside the latter there is seen a fairly
wide zone of tissue, more densely cellular than the true interstitial connective tissue ;
this denser zone is the anlage of the fibro-muscular and elastic coats of the bronchioles.

The epithelium of the branching tubules is ciliated, and it is only during later
stages that the cilia disappear in those portions of the bronchioles where the latter
expand into the infundibula and alveoli ; at the present stage, although in some places
the tubules appear to widen out into sac-like expansions, the epithelium remains of
the ciliated variety, since no true alveoli with air-sacs are as yet developed.
The interstitial connective tissue, which during later stages becomes compressed
and relatively diminished in amount by the preponderating development of the
alveoli, contains some large blood-vessels, but these are as yet relatively few in
number.

(iv.) The pancreas (PL IV. fig. 5). — The histological features presented by this
gland are quite typical : branching tubules supported by a fine meshwork of con-
nective tissue, the whole enclosed by a capsule of somewhat open character, from
which trabeculse pass into the substance of the gland. The tubules are lined by
columnar epithelium, but there is as yet no definite basement membrane ; f the
lumina of the developing alveoli are quite small. The blood-vessels are quite small
and apparently not very numerous at this stage ; no signs of any cell-groups com-
parable to islets of Langerhans are to be detected, but probably it is too early for

* See HERTWIG, Handbuch der Entwickelungslehre rfer WirbeUiere, 1906.

t The connective tissue appeared to have shrunken away from the tubules, leaving a considerable space between the
two (see fig. 5).

(ROY. soc. EDIN. TRANS., VOL. L., 238.)

THE SYSTEMATIC ANATOMY OF A F(ETAL SEA-LEOPARD. 461

these to have been formed. When a section of the pancreas stained with haema-
toxylin is examined, it appears that the cells lining the alveoli are not characterised
by the deeply-staining outer zone which is so marked a feature in the pancreas
of the armadillo ; moreover, the nuclei lie at the outer or attached borders of
the cells.

(v.) The spleen. — In minute structure, the spleen presents the following features :—

(a) Externally, a somewhat dense fibrous capsule, in which also unstriped muscle

cells are undergoing development.

(b) Trabeculae passing from the capsule into the substance of the gland, and

forming a network in the interior : near the surface some large arteries
may be seen passing in, more especially in the vicinity of the hilus, and
corresponding venous branches are emerging.

(c) The bulk of the organ is made up of a mass of erythrocytes, erythroblasts,

and lymphocytes ; here and there cells suggesting the splenic cells of the
adult organ may be seen, but giant-cells are apparently not present. A
section taken through the splenic artery and vein with the blood contents
of these vessels shows quite clearly that the vein contains many more
lymphocytes than the artery, a feature which indicates that the lymphoid
function of the spleen is already established ; whilst the presence of ery-
throblasts in fairly large numbers in the spleen-pulp leads to the inference
that haemogenesis is also a splenic function at this stage — -a point which
is well established in the case of the rabbit and some other mammals.

No Malpighian corpuscles are to be seen in a section of the organ, but in some
places the lymphocytes seem to be more densely aggregated than in others, with an
indication of a small artery in their neighbourhood : these masses are not, however,
well defined.

(vi.) The kidneys and adrenal bodies (PI. IV. figs. 1 and 2). — At this stage the
kidney presents histogenetic features which correspond fairly closely with those seen
in the kidney of a four-months human foetus ; that is to say, each renal pyramid
when sectioned in a plane passing through the cortex and the apex of the papilla is
seen to be made up of the following parts : —

(a) A cortical zone, in the outermost layer of which are to be seen the di-
chotomously branched endings of the tubules derived from the ingrowth
into the metanephros of the diverticulum from the Wolffian duct ; in many
of them the ampullary portions are continued into a coiled tubule — cut
across many times and in various planes — the first or proximal convoluted
tubule. At a somewhat deeper region of the cortex, the first set of
Malpighian capsules are to be seen, these being relatively large as com-
pared with those arising later : no sign of Henle's tube is as yet evident,
(ROT. soc. EDLK. TRANS., VOL. L., 239.)

462 MR HAROLD AXEL HAIG ON

since no downgrowth has occurred from the convoluted tubule to form
the loop.*

(b) A deep zone, in which as yet connective tissue preponderates, and through
which course the branching tubes derived from the Wolffian diverticulum ;
these tubules possess wide lumina, and are lined by a clear cubical epi-
thelium. They represent the rudiments of the straight and collecting
tubules, those nearest the papilla becoming later on the ducts of Bellini ;
in the Phocidse, according to CmKvrrz,t many of the secondary and tertiary
branches of the Wolffian diverticulum disappear during development.

The epithelium lining the convoluted tubules of the cortical zone is clear and
cubical, the nuclei staining but feebly with basic stains : the portion of the tubule,
however, which joins the ampulla is lined by smaller cells, the nuclei of which stain
deeply. The glomerulus in each Malpighian capsule is a well-developed capillary
tuft with already an indication of lobulation.

Between the renal pyramids and supporting them there is a certain amount of
connective tissue (columns of Bertini) in which small blood-vessels are seen cut across
(capsular vessels of later stages).

The adrenal bodies (PL IV. fig. 2). — Relatively speaking, the adrenals are much
smaller than one would expect at this phase of development, but their histogenetic
features are none the less instructive : each gland is enclosed in a capsule of connec-
tive tissue in which course branches of the adrenal artery and vein, whilst smaller
vessels (arterial) pass at right angles to the surface into the gland, being supported
by the fine septa which are given off from the inner aspect of the capsule.

The substance of the adrenal is made up of the following parts :—

(a) An outer zone, the commencing zona glomerulosa, composed of folded columns

of small cuboidal cells.

(b) A wide intermediate zone composed of anastomosing broad columns of large

polyhedral cells : this is the developing zona fasciculata, amongst the
columns of which ramify the small vessels noted above as passing in from
the capsule. The cells of this zone are characterised by their small clear
nuclei, and their deeply-staining cytoplasm, which takes up eosin very
readily.

Throughout this zone are scattered small masses of rounded cells, with deeply-
staining nuclei, the sympathetic ganglion rudiments. These are aggregated
in larger masses in the central region of the gland, where they form the
anlage of the medulla. In the medullary region the blood-channels are
wide, and as yet are lined by a well-defined endothelium.

* CHIEVITZ (Archiv Anat. u. Embryol., Supplement, 1897) found Henle's tubule present in an embryo of Phoca,
145 mm. in length. The present fretus is only 122 mm. in length,
t Op. cit.

(BOY. soc. EDIN. TRANS., VOL. L., 240.)

THE SYSTEMATIC ANATOMY OF A FCETAL SEA-LEOPARD. 463

Compared with mammals, such as the pig, the adrenal of this Seal appears to
be somewhat behindhand in the relative rate of its development ; thus, in a pig
embryo of 119 mm. the medulla is well defined, and most of the sympathetic
derivatives are confined to it alone ; but in a pig embryo of 70 mm. the histology of
the gland is much as has been described above.*

(vii.) The genital glands (PI. IV. fig. 4). — As has been noted, the genital glands
in the present instance are ovaries ; each ovary has a well-defined histological appear-
ance, and it is possible to recognise —

(a) An outer layer of rather high cubical epithelium.

(6) A wide cortical zone composed of masses of rounded cells or primordial ova
(some of these being considerably larger than others, and forming potential
ova which will later become surrounded by a follicle of smaller cells to
form the commencing Graafian follicles). Between the ova a good deal
of fibro-cellular connective tissue is to be seen, and this forms a dense
interlacing network supporting the egg-cells.

(c) Deep to the above cortical zone comes a layer of fibro-cellular stroma, com-

parable to the tunica albuginea of the testicle, but having a relatively
different position in the gland.

(d) A central portion, composed for the most part of dense stroma, with here and

there masses of ova, which are the remains of the so-called medullary
cords of somewhat earlier stages.

o

In the mesovarium, which forms a wide peritoneal band of attachment, there are
to be seen mesonephric tubules and glomeruli, whilst at the point of attachment of
this band to the ovary there are some tubules, supported by stroma, forming the
rete ovarii.

The above histological appearances correspond closely with those to be seen in
the ovary of a cat embryo of 94 mm. ; t no signs of developing Graafian follicles are
to be seen, as the follicular epithelium has not as yet been formed round any of the
larger ova.

(viii.) The placenta (Text-fig. 2). — The details of the placenta and placenta tion
have been acquired in the present instance from the study of a specimen lent to the
author by Professor WATERSTON of King's College, London ; this specimen shows a
foetus of the Weddell Seal, a closely allied species, in situ in the uterus, with the
membranes in their proper relative position with regard to foetus and placenta.J The
relations of the amniotic sac to the foetus and umbilical cord are such that the
former appears to be enclosed in a small complete sac, which is quite closed towards
the ventral aspect of the embryo, the umbilical cord upon reaching the line of
closure dividing up into a number of branches each containing twigs from the

* See HERTWIG, op. at. + Figured by COEBT, Inaug. Dissert, Leiden, 1898.

I Figured in Sir WILLIAM TURNER'S Catalogue of Marine Mammals.

(KOY. SOC. EDIS. TRASS., VOL. L., 241.)

464 MR HAROLD AXEL HATG ON

arteries and vein. The amnion then spreads out on either side of the line of closure,
the resulting folds passing anteriorly, posteriorly, and laterally to the margins of the
placenta, over which it then spreads, covering the fcetal surface of that structure.
The branches of the umbilical arteries and vein are conducted by the above-mentioned
folds to the margins of the placenta, and finally divide up into numerous twigs which
enter the substance of the chorion.

The placenta itself is of the zonary type, occupying a median zone of the uterus :
its histology corresponds very closely with the descriptions of GROSSER * and DUVAL
for the zonary placentae of the cat and bitch, with the exception of certain minor

FIG. 2. — A portion of a transverse section of the uterine wall and placenta of the
Weddell Seal. (Semi-diagrammatic. )

m. Muscle layers of uterine wall.
b. Maternal blood-vessels.
«. Epithelium lining uterine glands.

I. Lumina of glands containing secretion (em-

bryotrophe of Grosser).
tr. Interglandular fastening septum.

v. Villi of placental labyrinth.

The foetal blood-vessels exist in considerable numbers in the villi, but are not represented in the figure ;
the foetus contained in the above uterus had undergone about one-third of its development.

details. Thus we find that a vertical section taken through the placenta and uterine
wall presents the following main features (fig. 2) :—

(a) A sheet of amnion (not shown in the figure) covering the fcetal surface of the

numerous "cotyledons" of the placenta.

(b) The fcetal portion of the placenta, composed of the rather wide laminae of the

so-called " placental labyrinth " t between which mesodermic tissue lies ;
the sheets of this labyrinth are made up of a syncytium (derived from
the trophoblast of earlier stages) which encloses and surrounds large

* Vergleichende Anat. und Entvrickelungsgesch. der Eihaute u. Placenta, 1909.
t See GROSSER, op. cit.

(ROY. soc. EDIN. TRANS., VOL. L., 242.)

THE SYSTEMATIC ANATOMY OF A FCETAL SEA-LEOPARD. 465

numbers of foetal capillaries, whilst maternal vessels of rather wider
calibre lie in the mesodermic septa and become at times surrounded by
portions of the syncytiuni.

(c) A zone in which an invasion of the superficial gland-layer of the uterine

mucosa has taken place, the syncytium of the villi having at an earlier
phase converted the uterine epithelium into what GROSSER terms a " sym-
plasma," and becoming as it were welded with the mucosa at numerous
points ; the partitions between the glands become also fixed to other
villous tufts.

(d) A deeper layer which comprises the bases of the uterine glands, and lies

next the uterine muscle. The muscular coat, which is in the present case
thin, contains large branches of the maternal blood-vessels, and these,
where the interglandular septa pass to become fixed to the syncytium
of the villi, pass into the mesoderm lying between the lamellae of
the placental labyrinth.

It appears that zone c noted above (so-called " umlagerungszone " of STRAHL and
GROSSER) is, during the earlier phases, of the greatest importance in establishing the
connection between the uterine epithelium and the syncytium of the villi ; a further
action of the syncytium is to convert some of the decidual cells lying between the
uterine glands into trophoblastic masses not unlike the invading syncytium itself.

During earlier stages, stress is also to be laid upon the probability that the
secretion of the uterine glands serves as an additional source of nutriment (" embryo-
trophe ") to the foetus.

From the above account it will be seen that there is a considerable histological
similarity between the zonary placentae of the cat and Seal ; one notable difference
is to be seen in the relatively narrow extent of the gland-layer in the uterus of the
Seal, and another in the greater width of the laminae of the placental labyrinth.

(ix.) The internal ear (PI. IV. fig. 6). — The semicircular canals, utricle, ampullae,
and cochlea are well advanced in development, and lie in the cartilaginous rudiment
of the osseous labyrinth ; the membranous labyrinth is represented by a somewhat
thick connective tissue with a certain amount of elastic tissue entering into its
composition.

A transverse section across one of the semicircular canals in situ shows that the
canal is placed very excentrically, lying against one side of the cartilaginous labyrinth,
to which the connective tissue fixes it quite firmly. From the projecting part of the
canal, strands or bundles of fibres pass to the opposite circumference of the cartila-
ginous tube, and, joining here a continuation of the membrane, help to fix the canal,
so that practically no contraction of the lumen is possible. The spaces between the
fixing strands are filled with perilymph, whilst the canal itself is lined by a somewhat
flattened epithelium, which later on secretes endolymph. In the utricle and ampullae

(ROY. 8OC. EDIN. TRANS., VOL. L., 243.)

466 MR HAROLD AXEL HAIG ON

small elevations or " cristae" project into the lumen, and are lined by a much higher
type of epithelium than that found in the canals ; but as yet there is no evidence of
hair-like structures upon the free internal borders of the component cells.

The cochlea is rather more advanced in development than would be the case in a
three-months human foetus ; according to KRAUSE,* the organ of Corti in the human
fetus at birth shows that the membrana tectoria is only commencing to form, whilst
the sensory epithelium lying upon the basilar membrane shows only a larger and a
smaller group of columnar cells. The author's preparations of the cochlea of a three-
months human foetus show the sensory epithelium as a group of columnar cells higher
than the remainder in the tube, but no sign of the membrana tectoria. The scala
tympani is present, but no membrane of Reissner as yet divides the upper cavity into
scalae media and vestibuli, whilst the rudiments of the spiral ganglion and cochlear
nerve are certainly to be made out, but are not at all advanced.

In the Seal embryo under discussion, all three scalae are present, the spiral ganglion
is a marked feature, the membrane of Reissner is well defined, and the epithelium of
the organ of Corti is becoming differentiated, the component cells being higher upon
the outer side, and their free borders showing a well-defined clear zone.

(x.) The pituitary body (see text-fig. 3).- — A specimen of the pituitary gland of
an adult Weddell Seal (Leptonychotes weddelli) was examined histologically by
the author some time since, t and found to possess all three portions, viz. pars anterior,
pars intermedia, and pars nervosa, highly differentiated. The Sea-Leopard Seal in its
earlier phases of development possesses a very interesting hypophysis, which more-
over sheds some light upon the origin of the pars intermedia. Although a complete
account of the histogenesis of this structure is not possible in the present instance,
there are certain points in its development which are worthy of somewhat detailed
description, and for purposes of comparison the developing hypophyses of the rabbit
(at the twelfth, fourteenth, and nineteenth days) and of the three-months human
foetus were submitted to microscopical examination.

A nearly median sagittal section taken through the pituitary of this Seal embryo
(the gland being in situ in the sella turcica of the ossifying sphenoid bone) shows the
following features :—

(i.) The anterior lobe (pars anterior), consisting of irregular columns of rather
large polyhedral cells, separated by wide blood-channels, the latter possess-
ing a well-marked endothelial lining ; at the posterior extremity of the lobe
the blood-channels are observed to open into large venous tributaries, which
ultimately join up and communicate with the cavernous sinus, the latter
being seen in section lying just anterior to the dorsum sellse of the sphenoid
bone (fig. 3, g).

* See HERTWIG, op. cit.

t Trans. Roy. Soc. Kdin., vol. xlviii. part iv., No. 31, 1913.

(ROY. soc. EDIN. TRANS., VOL. L., 244.)

THE SYSTEMATIC ANATOMY OF A FCETAL SEA-LEOPARD.

467

(ii.) An intermediate cleft lying dorsal to the pars anterior, which is the remaining
evidence of Rathke's pouch ; this cleft is towards its lateral aspects (not
shown in the figure) partially filled by proliferating columns of epithelial
cells, some of which are to be seen in the mesial section at the anterior
extremity of the cleft.

(iii.) The pars nervosa, or posterior lobe, lying dorsal to the cleft, and connected
with the floor of the 3rd ventricle by the infundibulum ; part of the cavity
of the ventricle is seen extending into the infundibulum, but this feature
disappears during later stages to a large extent.

FIG. 3. — A vertical section through the base of the skull to show the hypophysis cerebri m situ, lying
in the sella turcica of the sphenoid bone. (Semi-diagrammatic.)

a. Epithelium of the roof of the nasal cavity.

6. Developing basi sphenoid bone ; the " irruption "-stage of ossification iu cartilage is represented.

c. Large anterior lobe of hypophysis showing cell- columns and intervening blood-vessels ; the

dark masses are cells staining more deeply with eosin.
rf. Infundibulum, showing communication with the 3rd ventricle of brain.
e. Posterior lobe (pars nervosa) of hypophysis.
/. Cleft between the anterior and posterior lobes ; at the anterior end of this is seen a mass of

cells, derived by an infolding from the anterior lobe, which will ultimately give rise to the

pars intermedia,
g. Cavernous sinus.
h. Dorsum sellse of sella turcica.

In connection with the pars intermedia, it appears that the columns of cells found
partially filling the above-mentioned cleft are derived by a proliferation of cells at
the upper and anterior angle of the pars anterior. HERRING * describes (in the cat)
the pars intermedia as an epithelial investment of the posterior lobe, but in the Seal,
at least during later stages, the intermediate mass is a well-defined strip of closely

* Journ. of Exper. P/IJWO/., 1909.

80C. KDIN. TRANS., VOL. L, 245.)

468 MR HAROLD AXEL HAIG ON

packed cells, which curves round the ventral aspect of the posterior lobe and appears
to end abruptly in a broad club-shaped extremity at the posterior and ventral margin
of that lobe. The dorsal portion, however, passes over the upper surface of the pars
anterior and seems to fuse with the anterior and dorsal margin of that lobe. No
remnant of Rathke's pouch is to be seen in the fully formed pituitary of the Seal,
whereas in the rabbit, cat, and man the pouch persists as a distinct cleft between the
pars anterior and the pars intermedia.

In the present instance the roof of Rathke's pouch is lined by a columnar
epithelium, which at the anterior extremity merges into the cell-columns which later
on fill up the pouch. Upon the dorsal aspect of the posterior lobe no epithelial
investment is to be seen, whilst the floor of Rathke's pouch is formed by the super-
ficial cells of the pars anterior, which are arranged in the form of an epithelium.

The very close union maintained between the buccal and cerebral portions of the
pituitary from the earliest stages is commented upon by HERRING * and emphasised
as having a direct bearing upon the functional importance of the gland. In the
Seal, at the stage of development now under discussion, the union between
the two portions is very intimate, but becomes even more marked as develop-
ment proceeds.

Compared with the hypophysis of a three-months human foetus, that of this Seal
is a good deal in advance. In the human foetus at three months the connecting
strand passing between the pars anterior and the buccal epithelium is still present,
whilst the posterior lobe is only represented by a very narrow diverticulum from the
floor of the 3rd ventricle.

Again, sections taken sagittally through the pituitary of the rabbit three days
before birth show that the size of the posterior lobe is small as compared with the
anterior, and the cavity in the infundibulum is represented by a mere cleft lined by
ependyma. During earlier phases in this animal (twelfth and fourteenth days) the
relation between the size of the lobes is such that the posterior lobe is proportion-
ately larger than during later stages. In the Seal, at the stage under discussion, the
posterior lobe is at least one-half the size of the pars anterior ; but this relation does
not hold in the full-grown animal, where the anterior lobe is five or six times as large
as the pars nervosa.

More detailed histological examination of the pituitary of the present foetus
•shows certain characters which differ from those observed in similar anatomical
regions during later stages ; in the first place, the blood-channels in the pars anterior
still retain their endothelial lining, a feature which disappears during development
to a large extent, although in some parts of the full-grown anterior lobe endothelial
cells are to be seen forming at least a partial lining to the blood-vessels. The cell-
columns of the anterior lobe are composed of closely packed polyhedral cells, which,
when stained with an acid stain such as eosin, may be differentiated into two

* Op. cit.

(ROT. SOC. EDIN. TRANS., VOL. L., 246.)

THE SYSTEMATIC ANATOMY OF A F(ETAL SEA -LEOPARD. 469

varieties, viz. a majority which stain lightly, and here and there isolated cells
staining intensely ; the cytoplasm of the latter cells is also more granular in
appearance. In the fully formed gland these deeply staining cells are more
numerous, and collected into small groups instead of being isolated cells. The pro-
liferating columns seen in Rathke's pouch are composed of rather cuboidal, or in
some cases columnar, cells, which do not possess any special affinity for eosin, but
the nuclei of which stain deeply with basic dyes ; no evidence of the syncytial
structure seen in the pars intermedia of later stages is to be made out, nor does any
colloid appear to have been formed as yet. The pars nervosa appears to possess
much the same minute structure as does that part of the adult gland ; for the most
part, neuroglia cells and fibres predominate, some of the former being spindle-shaped
and occurring in that portion of the lobe which lies next the epithelium forming
the roof of Rathke's pouch, their long axes being at right angles to the ventral
surface of the lobe. The portion of the cavity of the 3rd ventricle which passes
into the infundibulum is lined by an ependyma of the usual type, viz. rather high
columnar epithelium, the component cells being ciliated, whilst the basal portions
of these cells are continued as neuroglia fibres into the substance of the pars nervosa.
There is no colloid to be detected as yet in this part of the pituitary, although in the
fully formed gland small particles of colloidal material derived from the pars inter-
media are to be seen scattered throughout the posterior lobe. Lying just dorsal to
the infundibulum is a folded portion of the floor of the 3rd ventricle, which encloses
a small fold of pia mater ; this portion later on becomes modified to form a small
ovoid mass lying on the dorsal aspect of the posterior lobe, and contains syncytial
strands of nucleated cytoplasm more fully described by the author in a previous
communication.* When fully formed it is very vascular, but its functional signi-
ficance is not at all obvious. The above somewhat brief description of some of the
developmental aspects of the pituitary of the Sea-Leopard Seal are, of necessity,
incomplete, on account of the fact that only a single specimen was available for
investigation. It would be interesting to follow some of the earlier phases, inasmuch
as the gland appears in this animal to have a high functional significance — quite as
much so, in fact, as in some higher types. t

GENERAL COMPARATIVE CONCLUSIONS IN CONNECTION WITH THE ANATOMICAL AND
HlSTOLOGICAL FEATURES PRESENTED BY THE F(ETUS OF STENORHYNCHUS.

(i.) The foetus, in the light of the above considerations, shows many points in
common with the human foetus at the beginning of the fourth month of intra-uterine
development. If the relative rates of growths during the earlier developmental

* Op. tit.

t For further details of the cytological characters of the various regions of the mammalian pituitary, see
SCHAFER, " Text- Book of Microscopic Anatomy " (Quain's Anatomy, vol. ii. part i.).

(ROT. SOC. BDIN. TRANS., VOL. L., 247.)

470 MR HAROLD AXEL HAIG ON

phases are at all comparable in the two cases, then the present fetus should have
completed about one-third of its development. In many respects, however, this
fetus shows an advance upon the three-months human embryo, notably in connection
with the pituitary gland, the cerebellum, and the internal ear.

(ii.) The other mammals (rabbit, cat, and pig) with which incidentally the fetus
has been compared show on the whole a fairly close agreement, both anatomically
and histologically ; one marked exception, from the anatomical point of view, is in
connection with the kidney, which places the Seals in a small sub-group of the
carnivora, to which the bear also belongs.

(iii.) The developmental features of the pituitary, kidney, and brain are sufficiently
instructive to necessitate, when possible, investigation of the earlier phases of
development of these organs ; the acquisition of early Seal embryos is, however, a
difficult matter, but they might with advantage be sought for, as large numbers of
Seals are killed annually.

In conclusion, the author wishes to thank Dr. W. S. BRUCE and Professor
WATERSTON for their courtesy in lending Scotia specimens of foetal Seals for investi-
gation ; also Professor HEPBURN for many valuable suggestions in connection with
anatomical details, and Mr. T. H. BTJRLEND, of University College, Cardiff, for much
useful criticism concerning the arrangement of the above report.

BIBLIOGRAPHY.

BAILEY and MILLER, Text-book of Embryology, 1909.

BRYCB, T. H., Quain's Anatomy : vol. i., Embryology.

CHIBVITZ, Archiv Anat. u. EmbryoL, Supplement, 1897.

GROSSER, Vergleich. Anat. und Entwickelungsgeschichte der Eihaute u. Placenta, 1909.

HAIG, Trans. Roy. Soc. Edin., ibid.

HEPBURN, Trans. Roy. Soc. Edin., vol. xlviii., 1913.

HERRING, Journ. of Exper. Physiol., 1909.

HERTWIG, Handbuch der Entwicleelungslehre der Wirbeltiere, 1906.

REFUEL and MALL, Text-book of Embryology, 1912.

SCHAFER, E. A., Quain's Anatomy: vol. ii. part, i., Text-book of Microscopic Anatomy.

EXPLANATION OF PLATES.
PLATE I.

Fig. 1. Foetus of the Sea- Leopard Seal : —
a.f. Anterior flipper.
p.f. Posterior flipper.

t. Tail.

cl.a. Cloaeal aperture.
u. Umbilicus.

am. Amnion.
p. Placenta.
b. Tongue : the bifid extremity showing

between lips.
pal.f. Palpebral fissure.

u.c. Umbilical cord.

(ROY. soc. EDIN. TRANS., VOL. L., 248.)

THE SYSTEMATIC ANATOMY OF A FCETAL SEA-LEOPARD.

471

Fig. 2. Dissection of the Sea-Leopard Seal, to show the principal viscera in situ (ventral aspect) : —

1. Thyroid and lowest parathyroid.

2. Trachea.

3. Thymus.

4. Left lung (upper lobe).

5. Left auricular appendix.
5a. Left lung (lower lobe).

6. Ventricular portion of heart.

7. Diaphragm.

8. Left lobe of liver.

9. Umbilical cord.

1 0. Patent allantoic duct.

1 1 . Urinary bladder.

12. Coils of small intestine.

13. Umbilical vein.

14. Right lobe of liver.

15. Gall bladder.

15a. Right lung (middle lobe).

16. Right auricular appendix.

17. Right lung (upper lobe).

18. Arch of aorta.

PLATE II.
Anatomical details of the brain and cranial and nasal fossae.

Fii,r. 1. The brain from the dorsal aspect : —

1. 4th ventricle (floor).

2. Lamellae of cerebellar hemisphere.

Fig. 2. The brain from the side : —

1. Olfactory lobe.

2. Fissure of Sylviu?.

3. Infundibulum.

4. 3rd nerve.

3. Superior surface of cerebral hemisphere.

4. Crura cerebri.

5. Medulla oblongata.

6. Cerebellum.

7 Cerebral hemisphere.

Fig. 3. Mesial surface of a median sagittal section of head : —

1. Lower jaw.

2. Tongue.

21. Lamina terminalis.

3. Hypophysis cerebri.

4. Medulla oblongata.

5. 4th ventricle.

6. Cerebellum.

7. Corpora quadrigemina.

8. Optic thalarnus.

9. Falx cerebri.

9'. Corpus callosum.

10. Nasal fossa (lower).

11. Xasal septum.

12. Buccal cavity.

Fig. 4. Nearly mesial sagittal section of brain to show distribution of grey and white matter, and general

relations of parts : —

1. Nuclei of vagus and hypoglossal nerves.

2. Grey matter of cerebellar hemisphere.

3. Pia mater.

4. Grey matter of cortex cerebri.

5. Superficial white matter of cortex.

6. Grey matter lining cavity of prosen-

cephalon, just showing.

7. Portion of choroid plexus.

8. Portion of 3rd ventricle in infundibulum.

9. Grey matter in optic thalamus.

ii. Part of prosencephalon communicating witli

lateral ventricle.
iv. 4th ventricle.

Fig. 5. Mesial surface of sagittal section of head, the nasal septum and falx having been removed : —

t. Tongue.
hy. Hypophysis.
m.o. Medulla oblongata.
cb. Cerebellum.
p. Pons.

op. Optic thalamus.

h. Cerebral hemisphere.

olf.l. Olfactory lobe.

eth.t. Ethmo-turbinal bone.

mx.t. Maxillo-turbinal bone.

(ROT. soc. EDIN. TRANS., VOL. L., 249.)

472

MR HAROLD AXEL HAIG ON

PLATE III.

Anatomical features of some of the viscera.
Fig. 1. The stomach and intestines from the ventral aspect : —

1. Coils of small intestine.

2. Peritoneal fold attached to greater

curvature of stomach.

3. Pyloric end of stomach.

4. Cardiac end of stomach.

5. (Esophagus.

6. First bend of duodenum.

7. Pancreas.

8. Part of transverse colon.

Fig. 2.
th.
p.th.

v.

Fig. 3.
a.
b,
c.

Fig. 4.
1.
2.
3.
4.
5.
6.

Fig. 5.
r.l.

k.

Fig. 6.
a.

A coronal section of the thyroid and lowest parathyroid : —

Lobules of the lateral lobe.

Parathyroid, showing arrangement of cell-columns with intervening blood-channels and con-
nective tissue.
Vein in capsule, with a small nerve-trunk.

The lungs from the ventral aspect : —
Part of right middle lobe.
Upper and lower lobes of left lung.
Bronchi entering root of lung.

The retroperitoneal and pelvic viscera :-

Cloacal aperture.

Severed allantoic duct.

Urinary bladder.

Ureter (cut across).

Genital duct.

Left kidney.

The liver from dorsal aspect : —
Right lobe.

Surface of right lobe which lies over
kidney.

Deep surface of the spleen : —
Elongated hilus with blood-vessels.

d. Pulmonary vein emerging from right lung.

e. The three lobes of right lung.

7. Adrenal gland.

S. Diaphragmatic ligament of mesonephros.
9. Genital gland.
10. Rectum, with rudimentary uterus just ventral

to it.

l.l. Left lobe

l.ii. Umbilical vein ( = ligamentuni teres of later
stages.)

b. and c. Dorsal and ventral margins.

PLATE IV.
Figures (semi-diagrammatic) illustrating the histological features of some of the viscera.

Fit;. 1. Portion of a longitudinal coronal section of a kidney showing a quadrant of one renal pyramid
and some intervening connective tissue : —

a. Ampullae of terminal branches of

Wolffian diverticulum.
con.t. Convoluted tubule seen cut across

several times.
m.c. Malpighian capsules.

Fig. 2. Part of a longitudinal coronal section of an adrenal body : —

c. Capsule. *. Sympathetic ganglion cell masses.

w. Secondary, tertiary, etc., branches of Wolffian

diverticulum.

i. Intervening connective tissue between neigh-
bouring pyramids.

z.g. Zona glomerulosa.
«./. Zona fasciculate.
m. Medulla.

b. Blood-channels in medulla between large
groups of sympathetic cell-masses.

(ROY. soc. EDIH. TRANS., VOL. L., 250.)

THE SYSTEMATIC ANATOMY OF A FCETAL SEA-LEOPARD. 473

Fig. 3. Part of a section of lung, showing the branching tubules embedded in discrete masses of con-
densed connective tissue with rudiments of muscular and elastic layers lying next the tubules. The in-
terstitial connective tissue is also present in large relative amount.

Fig. 4. Portion of a longitudinal section of the ovary : —

ep. Surface epithelium (germinal epi-
thelium).
e. Cortical" zone with primitive ova and

s. Dense stroma forming a kind of tunica

albuginea.
m. Medulla with a few masses of ova and

some stroma. much stroma separating them.

Fig. 5. Part of a section of the pancreas, showing typical arrangement of branching tubular acini, with
but little interstitial connective tissue.

Fig. 6. Portion of a vertical section through the developing cochlea showing one turn of the spiral
in section : —

s.v. Scala vestibuli.
It. Membrane of Reissner.
c.c. Canal of the cochlea.

B. Basilar membrane, with rudimentary organ

of corti resting upon it.
S.T. Scala tympani.
G. Spiral ganglion.

(ROY. soc. EDIS. TRAXS., VOL. L., 251.)

SCOT. NAT. ANT. EXP. Vol. IV. Plate I,

HAIG: SYSTEMATIC ANATO/AY OF A FOETAL SEA-LEOPARD.

J

SCOT. NAT. ANT. EXP. Vol. IV. Plate II.

HAIG: SYSTEMATIC ANATOMY OF A FOETAL SEA-LEOPARD.

7

n

mx.t

\

olf.l.

eth.t.

SCOT. NAT. ANT. Exp. Vol. IV. Plate III.

HAIG: SYSTEMATIC ANATOMY OF A FOETAL SEA-LEOPARD.

y 5

.th.

r.l.

SCOT NAT. ANT. EXP. Vol. IV. Plate IV.

HAIG: SYSTEMATIC ANATOMY OF A FOETAL SEA-LEOPARD.

-i

I

PART XIX.
WHALE FISHEEIES.

SCOTTISH NATIONAL ANTARCTIC EXPEDITION.

XIX.-THE WHALE FISHERIES OF THE FALKLAND
ISLANDS AND DEPENDENCIES.

BY THEODORE E. SALVESEN, F.R.S.E., Leith.

(WITH TEN PLATES.)

( 479 )

XIX. — The Whale Fisheries of the Falkland Islands and Dependencies.
By Theodore E. Salvesen, F.R.S.E., Leith. (With Ten Plates.)

(MS. received December 9, 1913. Issued separately May 12, 1914.)

Time was, and not long ago. when the only human enemies of the Cachalot or Sperm
whale (Physeter macrocephalus], and of the southern Right whale (Baltena australis)
in the South Atlantic and Antarctic oceans, were the crews of British and American
whaling ships. The whales of the Finner species, such as the Blue whale (Balaenoptera
sibbaldii), the Finner whale (Baltenoptero, musculus or physalus), and the Humpback
whale (Megaptera boops), were left entirely unmolested, the reason being that the
method of capture was unsuitable. Rowing boats were employed, where the harpooner
stationed in the bow threw by hand or shot from a gun a harpoon, with or without
a bomb, and attached to a whaling line. The Cachalot and Right whales generally
remain some time on the surface, thereby enabling a rowing boat to approach
them ; whereas the Finner species are much more active in their movements,
and are only awash for a very short time when breathing. However, there might
now and again be opportunities for even a rowing boat to approach and kill a Finner
whale, but it would not be possible to bring the dead whale to the surface. This is
simply owing to the fact that the Finner whales sink after being killed, whereas the
Cachalot and Right whales float when dead. The weight of the carcase of the dead
Finner whale is very much greater than the buoyancy of an ordinary whale boat, and
if an attempt were made to lift the dead whale to the surface the whale boat would
naturally be drawn under.

These whalers were wooden sailing vessels of about 150 to 450 tons register, and
carried on pelagic whaling, touching only at ports in order to refit. They were
complete in themselves, inasmuch as they carried not only their whale boats and
gear, but also the products of the catch, the blubber being rendered into oil on board.

The industry was commenced in the beginning of the eighteenth century, the first
British whaling fleet of 12 vessels sailing in 1725 for the southern seas. In the first
half of the nineteenth century as many as 500 to 600 vessels were employed in the
southern hemisphere hunting Cachalot and southern Right whales. Since then the
number of vessels has been very greatly reduced owing to the fall in value of the
products, viz. oil and baleen, combined with increased working expenses and smaller
catches. At present this class of whaler is practically unknown in the Antarctic Regions.

The existence of enormous quantities of whales of the Finner tribe in the southern
seas was well known in whaling circles, and by members of Antarctic scientific ex-
peditious. Various attempts had been made to form companies for the purpose of

480 MR THEODORE E. SALVESEN ON THE

prosecuting whaling operations in the south by the same methods as employed by the
Norwegians in the north, but these efforts were unsuccessful until the year 1903.

Captain C. A. Larsen of Sandefjord had noticed the enormous numbers of Finner
whales when, in command of the sealer Jason, he visited the Antarctic in 1 892 and
1893 in search of seals and Right whales, and also about ten years later, as master of
the Antarctic, he accompanied Nordenskiold's scientific expedition to the Antarctic
regions. His efforts to form a company in his native land to start whaling operations
met with no success, but having friends in Buenos Aires, he succeeded in raising
sufficient capital there, and the Compania Argentina de Pesca of Buenos Aires was
founded, with Captain Larsen as manager.

He fitted out his expedition in Sandefjord, Norway, and arrived in Cumberland
Bay, South Georgia, in December 1904, with one modern steam whaler and two small
sailing vessels as transports, and immediately started building a whaling factory in
King Edward's Cove, on the same site as had been used for trying-in plant for seal
blubber in the early part of the preceding century. The satisfactory results obtained
led to the formation of a large number of companies, which now carry on in the depen-
dencies of the Falkland Islands the largest whaling business in the world.

METHODS.

Before enumerating in detail the various companies presently operating, it may be
of interest to have the method of pursuit and capture carried on by the whalers fully
explained, as it differs in every essential from that practised by the men who hunted
whales in the South Atlantic and Antarctic oceans up to the last decade.

The successful pursuit and capture of the Finner whales was inaugurated in 1866 by
a Norwegian, Captain Svend Foyn, who noticed the large numbers of this species
when passing the northern coast of Norway on his way to the Arctic ocean in search of
seals. After three years of arduous labour he at last solved the difficulties, and the
methods introduced by him, with considerable improvements, are those now in use by
all the modern whaling companies the world over.

In order to be able to pursue the active Finner whales and get within striking
distance, Captain Foyn substituted steam propulsion instead of men rowing the whale
boats, and he overcame the buoyancy question by increasing the size of the vessel.
The most approved type of a modern steam whale-catcher (Pis. I. and II. fig. 1) has a
length of from 98 to 115 feet over all, with a beam of 18 to 22 feet, and a moulded
depth of from 11 feet to 12 feet 9 inches, flush-decked, and cut away at both bow and
stern in order to make the vessel answer her helm very quickly. The lines of the hull
are designed for a speed of from 11 to 12 knots, and the engines are from 350 to 650
effective horse-power. Steam steering-gear is provided in order to operate the rudder
to most advantage, and a powerful double whaling-winch of two cylinders each is fitted
on the deck abaft the foremast. In the bow a glycerine recoil muzzle-loading cannon
is fixed on a swivel, and so delicately balanced that, when loaded with powder and

WHALE FISHERIES OF THE FALKLAND ISLANDS AND DEPENDENCES. 481

harpoon, it can be raised and lowered and turned from side to side by the harpooner
without apparent exertion. The bore of the cannon varies from 3 to 3^ inches, and its
length is about 45 inches. The harpoon is made of the finest tempered Swedish steel,
and is about 6 feet in length, weighing slightly over 1 cwt. It has four prongs, which
spring out to an angle of about 45° on tightening the line after the harpoon is lodged in
the whale's body. The point of the harpoon consists of a cast-iron head or shell of
conical shape, about 14 inches in length and weighing from 10 to 12 Ibs. This shell is
charged with gunpowder, which is fired by a time-fuse about three seconds after the
harpoon has been shot from the cannon. A foregoer of 60 fathoms in length and from
3| to 4-^ inches in circumference, made of the finest Italian hemp, is attached to a ring in
the shank of the harpoon. To the other end of the foregoer a 5-inch to 5 J-inch whale
line of 120 fathoms is spliced on. The length may be increased by adding additional
lines as may be required. The bow of the vessel is fitted with a double set of rollers over
which the lines run, and along the keelson, from the collision bulkhead forward to the
stokehold bulkhead, are arranged double rows of powerful spiral steel springs, connected
to two snatch-blocks above the shrouds of the foremast by strong flexible wire ropes.

The usual charge for a whaling gun is about 14 ounces of powder, and the range is
only 25 yards. If a whale be struck in a vulnerable part, death will usually be instan-
taneous. The animal will then immediately sink, drawing with it the whale line. The
steamer is brought to a standstill, and when the rope hangs vertically it is stoppered
in the bow, and slack is given inboard to allow it to be passed over the snatch-block at
the shrouds, from whence it goes to the winch, which raises the whale to the surface.

The snatch-block is, as stated above, attached to powerful steel springs, and the
object of this arrangement is to compensate for the rise and fall of the whaler caused
by wave motion, and to effect this these steel springs should be compressible to an
extent equal to the difference of level between the top and the bottom of the waves.
The elasticity of the whale line itself is not sufficient, and were no accumulators, as
these springs are called, fitted, the strain on the rope when heaving in the whale, caused
by the rise and fall of the steamer, would be greater than the strength of the line.

If the whale be not shot dead on the spot, the gunner has to play it in a very
similar manner to that of the angler who has hooked a salmon. In both cases the line
is much weaker than the weight exercised at its extremity, and the most experienced
skill and careful handling have to be exercised. The whaling steamer plays the whale
by manipulating the steam-winch, and also by moving ahead and astern, but it often
happens that the rope will be snapped, caused by a sudden jerk.

The animal having been raised to the surface, a chain is passed round the tail and
made fast at the bow of the ship, the foregoer is cut off at the harpoon, and the whale
is towed at the side of the vessel tail foremost. In order to make the carcase float and
thus lessen its towing weight, a hole is pierced through the body into the lungs or the
stomach, and the whale is inflated with air by means of a steam air-pipe in the engine-
room of the whaler.

482 MR THEODORE E. SALVESEN ON THE

SPECIES.

The following are the only species of whales which are hunted in the South
Atlantic :—

The southern Right whale (Balsena australis) (PI. II. fig. 2) is a sub-antarctic species,
and has a length of about 45 to 50 feet, the baleen measuring up to 7 feet in length and
weighing from 4 to 5 cwts. A full-grown specimen in good condition will yield from 60
to 70 barrels of oil. The price of baleen being now so low, no special efforts are made
to kill it, and the annual catch varies from ten to fifteen only.

The Blue whale (Balsenoptera sibbaldii) is the largest living animal in the world,
and may attain a length of 100 feet, and yields on an average 70 to 80 barrels of oil.

The Finner whale (Bal&noptera musculus or physalns) (Pis. III. figs. 1, 2, IV.
fig. 1) has a length from 50 to 70 feet, and its yield of oil averages 35 to 50 barrels.

The Fish whale or Seihval (Balsenoptera borealis) is the smallest of the whales killed
in the south, measuring only from 25 to 40 feet, and yielding 10 to 15 barrels. It is
thus of very small value, and is only pursued when there is a scarcity of the larger kinds.

The Humpback whale (Megaptera loops) (Pis. IV. fig. 2, V. fig. l) varies in length
from 40 to 50 feet. Its oil contents may be reckoned from 25 to 35 barrels.

The Cachalot or Sperm whale (Physeter macrocephalus] has a length of from 50 to
60 feet. It has no baleen or finners, but from 58 to 64 teeth in its lower jaw. Its
head constitutes about one-third of the whole animal, and contains about 30 barrels of
spermaceti and sperm oil, the body yielding a similar quantity of oil. This whale is
seldom met with in the waters round the Falkland Islands and dependencies, as its
natural habitat is in warmer zones.

REDUCTION.

With whaling by the method just described, a separate factory for the reduction of
the whale carcase is a necessity. The whaling steamers have no reducing plant on
board, and tow the carcases to the harbour where the company's factory is situated.
These factories are of two kinds — the shore factory and the floating factory.

When the whale is towed to a fully equipped shore factory (P)s. V. fig. 2, VI. figs.
1, 2) it is hauled up a slip on to the Henching platform (PI. III. fig. 1). The baleen or
whale bone is first removed, after which the whale is flenched by cutting off the blubber
in strips, which are passed through a cutting machine and then conveyed by elevator to
large open iron vats, where the oil is extracted by means of steam. The tongue and
kidney fat are dealt with in a similar way, but the flenched carcase has to be chopped
and sawn into suitable pieces before being conveyed and stowed into large boilers, where
the contents are subjected to a steam pressure of 60 Ibs. per square inch for 12 to 15
hours continuously. During this period the oil contained in the bones and flesh is run
off into clearing tanks. The digesters thereafter are emptied of their contents, which
are passed into drying kilns fired by coke, and finally the dried residue is run through

WHALE FISHERIES OF THE FALKLAND ISLANDS AND DEPENDENCIES. 483

a disintegrator, then sifted, and ultimately bagged as a powder or meal and ready
for the market.

A floating whaling factory (Pis. VII. figs. i, 2, VIII. figs. 1, 2) consists of a steamer
or sailing ship up to 7000 tons burden, fitted with all the necessary reducing plant and
with accommodation for storing the oil. These vessels must be moored in a harbour,
as smooth water is required to enable the whales being flenched alongside the vessel.
The blubber and carcase are cut up and hoisted on board, where the treatment is very
similar to that adopted at the shore stations. A few steamers have been fitted with
drying kilns for the manufacture of guano, but so far the results have not been quite
satisfactory, and it is open to doubt whether the expenses of manufacturing and
marketing guano manufactured on board are covered by the prices realised.

PRODUCTS.

Three products may be obtained from the dried meat and bones. Firstly, whale
meat meal, manufactured exclusively from absolutely fresh whale flesh. This is a most
nutritious and wholesome foodstuff, containing 17^ per cent, proteid, and is largely used
for feeding cattle. Secondly, whale guano, made from the remaining flesh and about
one-third of bones. The analysis shows about 8 '50 per cent, ammonia and about
21 percent, tribasic phosphates. Thirdly, bone meal, made exclusively from bones,
and analysing about 4 per cent, ammonia and about 50 per cent, phosphates. The
whole of the dried carcase may also be made into one product, forming a rich guano
and containing from 10 per cent, to 12 per cent, ammonia and 17 per cent, to 24 per
cent, phosphates.

The present value is about 13s. 6d. per unit of ammonia and 6d. per unit of
phosphates per ton delivered at a European port.

The value of whalebone, baleen, or finners, as the whalebone of the Finner
species of the whales is commonly called, is now very small compared to the prices
formerly obtained. The baleen of the southern Right whale is at present only worth
about £750 per ton, and the finners of the Fish whale fetch about £85 per ton, of the
Blue and Finner whales only about £30, and of the Humpback whale about £10 per
ton. These prices are so low that it is questionable whether the expenses of marketing
are not heavier than the value realised. There can be no doubt that it does not pay to
ship Humpback finners at the prices mentioned.

Whale oil is usually graded into four qualities, Nos. I., II., III., and IV., although
some companies add a fifth — No. 0. Xos. 0 and I. are made entirely from blubber,
No. II. from the tongues and kidney fat and from the residue of the blubber boilings,
No. III. from the flesh and bones, and Xo. IV. from refuse.

The present value of Nos. 0 and I. is about £24 per ton net weight, No. II.
£22, No. III. £20, and No. IV. £18 per ton, all delivered U.K. or Continental port,
barrels included, or proportionate reduction if delivered naked.

484 MR THEODORE E. SALVESEN ON THE

WHALING COMPANIES.

Undernoted is a list of the Companies presently operating : —
South Georgia.

1. Compania Argentina de Pesca of Buenos Aires, started in 1904 ; shore station

in King Edward's Cove, Cumberland Bay (PI. VIII. fig. 2); licensed for 4
steam whalers. With this concern is affiliated a floating factory sailing vessel
for the reduction of flenched carcases.

2. Sandefjord Whaling Co. of Sandefjord, Norway, started in 1907 ; shore station

in Stromness Bay ; also floating factory steamer and 3 steam whalers. This
Company was founded in 1906, but the floating factory steamer was totally
wrecked outside of Stromness Bay on an uncharted rock.

3. Toensberg Whaling Co. of Toensberg, Norway, started in 1907 ; shore station in

Husvik Harbour, Stromness Bay. Three steam whalers.

4. Bryde & Dahl's Whaling Co. of Sandefjord, started in 1908 ; floating factory

steamer in St George's Bay. Three steam whalers.

5. The South Georgia Co. Ltd. of Leith, started in 1 909 ; shore station in Leith

Harbour, Stromness Bay (Pis. V. fig. 2, VI. fig. 1, IX. fig. 1). Operates 4
steam whalers.

6. The Ocean Whaling Co. of Larvik, Norway, started in 1909 ; shore station in

New Fortune Bay (PI. IX. fig. 2). Two steam whalers.

7. The Southern Whaling and Sealing Co., Ltd., of North Shields, started in 1911 ;

floating factory steamer in Possession Bay (PI. VIII. fig. l). Two steam whalers.
South Shetlands and Graham Land.

1. Oernen Whaling Co. of Sandefjord, Norway, started January 1 906. Two floating
factory steamers and 6 steam whalers.

3. Sydhavet Whaling Co. of Sandefjord, started 1908. Floating factory steamer

and 3 whalers.

4. Sociedad Ballenera de Magallanes, Punta Arenas, started in 1908. Floating

factory steamer and 3 whalers.

5. Laboremus Whaling Co

„,, ,. All oi ISanderiord, Norway, and started in 1910.

6. Hvalen Whaling Co.

„ , , ,,T1 ,. .bloating factories and 3 whalers each.

7. Odd Whaling Co.

8. Hektor Whaling Co. of Toensberg, Norway, started in 1910. Operates 2 floating

factories, a factory ashore on Deception Island, and 5 steam whalers.

9. Norge Whaling Co. of Larvik, Norway, started in 1910. Floating factory

steamer and 3 steam whalers.
10. Chr. Salvesen & Co. of Leith, started in 1911. Two floating factory steamers

and 3 whalers.
Falkland Islands.

1. New Whaling Co. of Leith, started in 1908 ; shore station on New Island
(PI. VI. fig. 2). Operates 3 whalers.

WHALE FISHERIES OF THE FALKLAND ISLANDS AND DEPENDENCIES. 485

South Orkneys.

1. Whaling Companies, Rethval & Thule of Christiania, Norway, started in January

1912. Operate 2 floating factory steamers and 4 whalers.

2. Normanna Whaling Co. of Sandefjord, Norway, started in 1912. One floating

factory and 2 whalers.

3. Corral Whaling Co. of Bergen, Norway, started in 1912, with 1 floating factory

steamer and 2 steam whalers. The floating factory steamer was lost in the ice
in the spring of 1913.

u

7O

John Xathitton, del.

The Falkland Islands and Dependencies.

PRODUCTION.

During last season, from say 1st November 1912 till the end of April 1913, the
catch of the 21 steam whalers operating at South Georgia was about 5000 whales, of
which about 52 per cent, were Humpbacks, about 42 per cent. Finuers, and about 6 per
cent. Blue whales. This catch yielded about 200,000 barrels of whale oil, and about
8000 tons of whale guano.

The season at the South Shetlands and Graham Land is much shorter, and it is

486 WHALE FISHERIES OF THE FALKLAND ISLANDS AND DEPENDENCIES.

seldom that there is open water for more than four and a half months. The 32 whalers
working in these waters killed also about 5000 whales, yielding about 200,000 barrels
of oil, and about 100 tons of guano at the only shore station there, on Deception Island.
The percentage of the whales caught was as follows : about 43 per cent. Finners, about
35 per cent. Blue, and only about 22 per cent. Humpback whales.

At the South Orkneys the season is still shorter, open water being seldom met with
before the end of December, and ice setting in again about the middle or end of March.
The 6 steam whalers at work from these Islands caught about 800 whales, yielding
about 27,000 barrels of oil. The proportion of the various species is much the same as
for the South Shetlands and Graham Land.

At the Falkland Islands the 3 whalers operating brought in only 87 whales, com-
posed of 43 Fish whales, 36 Finners, and 8 Humpbacks, yielding 2128 barrels of oil and
275 tons guano.

The total production in the Falkland Islands and dependencies is thus about
430,000 barrels of oil — which is more than half of the world's output during last
season — and 8375 tons of guano, the gross value of which is about £1,350,000 sterling.
The industry gives employment to about 3500 men composing the crews of the whaling
and transport vessels and working at the factories ashore and afloat.

EXPLANATION OF PLATES.

PLATE I.
Steam whaler Sedna of Leith.

PLATE II.

Fig. 1. Steam whaler Rainna of Leith.

Fig. 2. Southern Right whale (Bcdaena australis).

PLATE III.

Fig. 1. Finner whale (Balxnoptera musculus).
Fig. 2. Finner whale (Balxnoptera musculus).

PLATE IV.

Fig. 1. Finner whale (Balsenoptera musculus).
Fig. 2. Humpback whale (Megaptera loops).

PLATE V.

Fig. 1. Humpback whale (Megaptera boops).

Fig. 2. Whale factory, Leith Harbour, South Georgia.

PLATE VI.

Fig. 1. Whale factory, Leith Harbour, South Georgia.
Fig. 2. Whale factory, New Island, Falkland Islands.

PLATE VII.

Fig. 1. Whale factory, New Fortune Bay, South

Georgia.
Fig. 2. Floating whale factory, s.s. Neko of Leith.

PLATE VIII.

Fig. 1. Floating whale factory, s.s. Horatio of Leith.
Fig. 2. Floating factory, s.s. Restitution of North
Shields, Possession Bay, South Georgia.

PLATE IX.

Fig. 1. Floating factory, ship Nor of Sandefjord,
King Edward's Cove, South Georgia.

Fig. 2. S.S. Coronda of Leith, discharging stores at
Leith Harbour, South Georgia.

PLATE X.

Female Cachalot or Sperm whale (Physeter mac,ro-
cephalus), captured 1913, on flenching
platform, Leith Harbour, South Georgia.

Scot. A'at. Ant. Exp.

SALVESEX : Whale Fisheries. — PLATE III.

VOL. IV.

Fir.. 1. — Firmer Whale (Baiienoptera musculus).

FIG. 2.— Finner Whale (Balienopttra mitsculws).

Scot. Nat. Ant. Exp.

SALVESE.N : Whale Fisheries. — PIRATE IV.

VOL. IV.

FIG. 1. — Finner Whale (Balsenoptera mtisculus).

FIG. 2. — Humpback Whale (Megaptera boops).

Scot. Nat. Ant. Ejcp.

VOL. IV.

SALVESEX : Whale Fisheries. — PLATE V.

FIG. 1.— Humpback Whale (Megaptera boops).

FIG. 2. — Whale Factory, Leith Harbour, South Georgia.

Scot. Nat. Ant. Exp.

SALVESEX : Whale Fisheries. — PLATE VI.

VOL. IV.

FIG. 1.— Whale Factory, Leith Harbour, South Georgia.

V

FIG. 2.— Whale Factory, Xew Island, Falkland Islands.

Scot. Nat. Ant. Exp.

SALVKSEX : Whale Fisheries. — PLATE VII.

VOL. IV.

FIG. 1. —Whale Factory, New Fortune Bay, South Georgia.

FIG. 2.— Floating Whale Factory, s.s. Neko of Leith.

Scot. A~at. Ant. Exp.

SALVESEX : Whale Fisheries. — PLATE VIII.

VOL. IV.

FIG. 1.— Floating Whale Factory, s.s. Horatio of Leith.

FIG. 2. — Floating Factory, s.s. RvtUutivn of North Shields, Possession Bay, South Georgia.

Scot. Nat. Ant. Exp.

SALVESEX : Whale Fisheries. — PLATE IX.

VOL. IV.

Flo. 1 — Floating Factory, ship Nor of Sandefjord, King Edward's Cove, South Georgia.

Fin. 2.— S.S. Coranda of Leith, discharging stores at Leith Harbour, South Georgia.

PART XX.
CETACEA.

SCOTTISH NATIONAL ANTARCTIC EXPEDITION.
XX.-SOME OBSERVATIONS ON ANTARCTIC CETACEA.

BY WM. S. BRUCE, LL.D., F.R.S.E.,

Director of the Scottish Oceanographical Laboratory.
(WITH TWO PLATES AND ONE TEXT FIGURE.)

( 491 )

Some Observations on Antarctic Cetacea : Scottish National Antarctic
Expedition. By William S. Bruce, LL.D., F.R.S.E. (With Two Plates and
One Text-figure.)

(Handed in October 21, 1914. Issued separately March 31, 1915. )

The study of the Cetacea forms one of the most interesting and most difficult
problems for zoologists, and at the same time one of the most important from the
industrial standpoint.

The main difficulty encountered is the costliness of the investigation, both on
account of the size of the animals, the difficulty of following out their migrations,
and the great commercial value of the products of the carcase : the financial
resources of zoologists have so far proved entirely inadequate to meet the amount
of money required to make whalers devote sufficient time towards assisting them
in these important researches. Because, while attending to the requirements
of zoologists in handling the animals for anatomical investigation, they would be
losing the chance of catching many other whales each of great commercial value.

Even an institution like the Natural History Museum (British Museum), though
receiving a large annual Government grant, has never been able to offer sufficient money
to Dundee whalers for the skeleton of a Greenland whale, and is in consequence without
an example of it, in spite of the enormous number that have been killed, ever since
the foundation of the Museum, by Scottish and other whalers. The offer of £100 for
the skeleton of Balsena mysticetus, when the whalebone in its mouth alone was
worth £2700, only produced a smile on the faces of those hardy Arctic skippers and
owners. Even now, with much reduced prices, which have for the time killed the
Dundee whaling trade on account of severe competition by Norwegian methods,* a
sum many times greater than that would not induce any owner, skipper, or crew to
consider the proposal.

The new Norwegian methods, t however, of fishing whales other than the Green-
land or Bowhead whale in almost all seas outside the limits of, but close up to, the
compact pack ice of Arctic and Antarctic seas, which entails hauling the carcases of
the whales up on landing slips ashore or alongside large tank ships anchored in a
sheltered harbour, have enabled closer investigation to be made during recent
years by trained naturalists ; and in view of the fact that detailed investigations
of these southern whales has thus been more possible during the last few years,
and that a special effort has recently been made in this direction by the despatch to

* " The Whale Fisheries of the Falkland Islands and Dependencies," by THEODORE K SALVESEN, F.R.S.E. (with
ten plates), ridt Part XIX. pp. 475-486.

t On the return of Mr BCRS MURDOCH and myself from the Antarctic voyage of 1892-93, in conjunction with
Captain LARSEX and Mr H. J. BULL, we endeavoured to raise interest in Britain to develop whale fisheries in
connection with the Falkland Islands and Dependencies. Our project, however, was not supported by business men,
or by the Colonial Office. Ten years later Captain LARSEX induced Argentine capitalists to form the " Conipania
Argentina de Peaca " of Buenos Aires, and erected a whaling station at South Georgia, with four whalers, which lias
yielded, and is still yielding, magnificent returns.

492 DR W. S. BRUCE

South Georgia of Major BARRETT HAMILTON, whose untimely death zoologists so deeply
deplore, I have thought it well to publish for what they are worth, without alteration
by an outsider and with my own interpretation, the brief notes I made over twenty
years ago, answering criticism only when it is misleading and not merely derisive.

In his " Cetaces " in the Resultats du voyage du S. Y. " Belgica," published in 1903,
Mr E. G. RACOVITZA passes criticism on certain notes made by myself and others on
board the Dundee whalers in 1892-93. The criticism is so wild as to be worthless,
and for that reason has been ignored.

I recorded Cetacea south of latitude 50° S. on board the Balsena* on 29 t days
out of 85 days, viz. from llth December 1892 to 23rd March 1893; and the
naturalists of the Scotia recorded Cetacea south of latitude 50° S. on 38 days out
of 82 days, viz. from 7th January 1903 to 23rd March 1903, 27th November 1903
to 3rd December 1903, as well as on 31 days out of 56 days, viz. from llth February
1904 to 7th April 1904.

On some days when Cetacea might have been in the vicinity of the Balsena,
I was unable to record their presence on account of heavy ship's duties which were
not those of a naturalist,} but which by contract I had agreed to undertake and
accordingly fulfilled.

The species of whales,§ as far as they are known, were :—

1. BAL^NA ATJSTRALIS :

Balsena australis, Desmoulius.
(The Southern Right Whale.)

2. BAL^NOPTERA MUSCULUS :

Balsena musculus, Linn.

Balsena tripinnis, R. Sibbald.

Balsenoptera musculus, Van Beneden and Gervais.

Balaenoptera physalus, True.

(The Finner, the Common Rorqual, Fin Whale, or Razor Back.)

3. BAL.ENOPTERA SIBBALDI :

Balaenoptera sibbaldii, Flower.

Balsena tripinnis, R. Sibbald.

Balsena maximus borealis, Knox.

Physalus sibbaldii, Gray.

Balsenoptera musculus (Sulphur Bottom), True.

(The Blue Whale, Sibbald's Rorqual, or the Sulphur Bottom.)

* From Edinburgh to the Antarctic,})? W. Q. BURN MURDOCH, with chapter by W. S. BRDCK. Longmans, Green
& Co., 1894. t Not 7 days, as recorded by Mr RACOVITZA.

I Mr RACOVITZA sailed as official naturalist on board a scientific ship. I signed articles on board a whaler.
Those early observations, however, taken under highly disadvantageous circumstances by my scientific colleagues
and myself, stimulated the despatch of the Belgica, on which Mr RACOVITZA sailed, and in turn all the many scientific
expeditions sent out from Europe during the past twenty years.

§ The classification is according to that of Sir WILLIAM TURNER in his Catalogue entitled The Marine Mammals
in the Anatomical Museum of the University of Edinburgh, 1912.

SOME OBSERVATIONS ON ANTARCTIC CETACEA.

493

4. BALSENOPTERA BOREALIS :

Balsenoptera borealis, Lesson.
Balsenoptera laticeps, Gray.

(The Seihval, or Rudolphi's Rorqual.)

5. MEG AFTER A BOOPS :

Balsena boops, Linn.

Balsena longimana, Rudolphi.

Megaptera boops, Van Beneden and Gervais.

(The Humpback, the Hunchback, or Long-armed Whale.)

6. HYPEROODON ROSTRATUS :

Balsena rostrata, 0. H. Miiller.
Hyperoodon B'utskopf, Lacepede.
Hyperoodon bidens, Fleming.
(The Bottlenose Whale.)

7. ORCA GLADIATOR :

Delphinus orca, Linn.
Orca gladiator, Gray.

(The Grampus, Killer Whale, or Sword Fish.*)

The Sperm Whale or Cachalot (Physeter macrocephahis), though captured more
recently at South Georgia, was not seen either by the Balaena or the Scotia south
of the fiftieth parallel of latitude. It is interesting to note that all the sperm whales
captured at South Georgia have been males except one (vide SALVESEN, "The
Whale Fisheries of the Falkland Islands and Dependencies," PI. X., Vol. IV., Part
XIX.)- Besides whales, black fish and porpoises of doubtful species were recorded,
and one especially, seen frequently in the vicinity of the Falkland Islands and
in Stanley Harbour, appears to be of considerable interest.

Cetacea observed South of 50° Latitude on board the " Balsena."

Date. Locality.

Temp.

Colour.

1892.

Dec. 11 51' 49' S., 57° 35' W.,

40-7' F.

...

Piebald porpoises

Lagenorhynchus cruciger?

3 miles off Cape

Pembroke.

12 53° 38' S., 55° 17' W.

40-6° F.

Two large finners

Balsenoptera musculus.

16 59° 24' S., 51° 01' W.

34-8" F.

...

Great numbers of

» >>

finners

17 61° 06' S., 50° 03' W.

31-8* F.

...

Finners; but

i> ii

none resem-

bling the Bow-

head whale.

First ice.

* The term " sword fish," though confusing, is the usual name given to the northern grampus hy British and,
I believe, American whalers.

494

DR W. S. BRUCE

Cetacea observed South of 50° Latitude on board the " Baliena " —continued.

Date.

Locality.

Temp.

Colour.

1892.

Dec. 21

62° 21'S., 54° 03' W.

31-5° F.

Dirty

Many finners •

Balaenoptera musculus.

green

„ 22

63° 02' S., 54° 31' W.

32-0° F.

Many whales

sp. ?

,, 24

64° 10' S., 55° 40' W.

32.2° F.

Many whales and

,, ,, and Orca

grampuses

gladiator.

„ 30

In the vicinity of

. ..

Dirty

Finners

Biiliunoptera musculus.

Erebus and Terror

dark green

Gulf, Graham Land

„ 31

V >i

...

Blue whales

,, sibbaldi.

1893.

Jan. 4

)> '>

...

...

Finners

,, musculus.

6

» ii

Many finners

» ii

7

>» »

. . .

Dark blue

Finners

» »

8

)) i»

. . .

Green

Whales

sp.?

12

64° 40' S., 56° 40' W.

. . .

Olive green

Finners

,, musculus.

„ 13

In the vicinity of

Green and

A humpback

Megaptera boops.

Erebus and Terror

dark blue

Gulf, Graham Land

14

i »

Whales

Balsenoptera sp. ?

15

> »

32-6° F.

Blue

Finners

,, musculus.

20

J M

31-5° F.

...

Finners

jj »

25

I >»

...

Finners

u »>

26

» ) >

Finners, includ-

»> "

ing a white

finner

H 27

>J »

Finners

» jj

28

)> >i

...

Finners

I* »>

Feb. 2

» 11

30-8° F.

Finuers

IT >J

6

i» i>

Blue and

Finners

)5 ))

clear

7

ii ii

...

>i

Finners

)» »>

„ 17

62° 33' S., 54° 35' W.,

n

Finners

J» )1

Joinville Land,

S. x W.|W.

Mar. 6

50° 35' S., 53° 53' W.

...

i>

Finners and

„ „ and

Porpoises (pie-

(Lagenorhynclius cru-

bald?)

ciger 1).

Observations on board the " Scotia," First Voyage.

Date.

Locality.

Temp.

Colour.

1903.

Jan. 7-25

Port Stanley, 51° 41'
S., 57° 51' W.

53-1° F.

...

Piebald porpoises
seen several

Lagenorhynclius cruciger?

times

„ 27

52° 55' S., 55° 00' W.

44-6° F.

Light
greenish
blue

Several piebald
porpoises seen

n >i

„ 31

58° 14' S., 45° 15' W.

35-4° F.

Blue

Finners and
bot 1 1 en oses
plentif u 1

Balaenoptera musculus and
Hyperoodon restrains.

SOME OBSERVATIONS ON ANTARCTIC CETACEA.

495

Observations on board the " Scotia" First Voyage — continued.

Date.

Locality.

Temp.

Colour.

1903.

Feb. 1

59° 32' S., 43° 10' W.

34-0° F.

>i

Finners were seen

Balaenoptera musculus.

in great num-

bers

*

60° 47 .S., 44° 00' W.

29-9° K.

H

Finners

ii i>

3

61'06'S., 43° 40' \V.

29-8° F.

11

Finner whales

j» ii

very conspicu-

ous

6

60° 10' S., 42° 35' W.

33-r.° F.

)?

Several fanners

ii »i

8

59° 44' S., 36° 40' W.

31-0° F.

»

Finners conspicu-

»i »

ous

9

59° 42'S.,34* 13' W.

30-0° F.

II

Finners with

ii ii

calves

„ 10

60° 05' S., 32* 10' \V.

30-2° F.

II

Two bottlenoses

Hyperoodon rostratus.

„ H

60° 03' S., 3-J° 31' \V.

30-0 F

n

Two small whales

Balsenoptera (borealis?).

about 20 feet

long, head and

back resemb-

ling finner,

probably the

fish whale.

„ 12

59° 49' S., 31° 32' W.

29-9° F.

ii

A large finner

,, musculus.

„ 13

59° 43' S., 30° 44' W.

30 0° F.

n

Two finners

ii >i

,, H

59° 33' S., 27° 37' W.

30-8° F.

19

Several finners

» ii

„ 15

61° 37' S., 26° 10' W.

31-0*F.

n

Several finners

» »

„ 16

62° 52' S., 25° 00' W.

30-9° F.

j»

Plenty of finners

ii ii

all day, especi-

ally in the

morning

„ 17

64° 18' S., 23° 09' W.

31-7° F.

...

Finners seen in

ji >i

afternoon

„ 20

69° 39' S., 22' 58' W.

29-0° F.

Dark

Finners

ii n

slatey blue

„ 21

69° 46 S., 19° 10' W.

29-8° F.

it

Finiiers and

„ ., and

bottle noses

Hyperoodon boops.

have been ob-

served, the

former scarce

„ 22

70° 21' S., 17° 00' W.

29-0° F.

Blue

Whale heard

Ba'aenoptera sp. t

blowing about

6 p.m.

„ 25

69° 44' S., 18° 02' W.

29-2° F.

Very dull

Two grampuses

Orca gladiator.

blue

„ 26

69° 36' S., 20° 20' W.

29-1° F.

Light dull

A "spout" of a

Balsenoptera sp. ?

blue

whale seen

» -<

69° 32' S., 24° 00' W.

29-2° F.

Dull blue

A grampus

Orca gladiator.

„ 28

(39° 22' S., 26° 36' W.

29-4° F.

Light blue

A few grampuses

» >i

Mar. 1

69° 03' S., 28° 02' W.

29"2° F.

Dull blue

Grampuses num-

» i>

erous

3

: 68° 35' S., 31° 56' W.

29-4° F.

. -.

Three grampuses

>j »»

5

68° 11' S., 34° 17' \V.

29-0° F.

Turquoise

A few grampuses,

>! 1)

one with its

dorsal fin

broken

6

67* 39' S., 36° 10' W.

3] -4° F.

Blue

A school of gram-

>» ?i

puses

496

DR W. S. BRUCE

Observations on board the " Scotia," First Voyage — continued.

Date.

Locality.

Temp.

Colour.

1903.

Mar. 12

65" 29' S., 44° 06' W.

29-0° F.

Light blue

Bottlenoses seen

Hyperoodon rostratus.

twice

,, 13

64° 48' S., 44° 26' W.

29-4° F.

Blue | Grampuses

Orca gladiator.

14

64° 30' S., 43° 45' W.

29-0° F.

»

A finner's blast

Balsenoptera musculus.

seen

„ 16

63° 51' S., 40° 50' W.

29-9° F.

» )

Grampuses seen

Orca gladiator.

» 21

Leathwaite Strait, S.O.

• > •

Several finners

liixlaenoptera musculus.

„ 22

Between Saddle Island

30-0° F.

Blue

Finners

jj »i

and Cape Bennett

» 23

Leathwaite Strait, S.O.

29-8* F.

»

Grampuses and

Orca gladiator and Balsen-

finners

optera musculus.

Nov. 27

Off north coast of

29-0° F.

A few grampuses

Orca gladiator.

Coronation Island

were sighted

„ 28

59° 43' S., 48° 10' W.

32-7° F.

Blue

Bottlenoses and

Balsenoptera musculus and

finners were

Hyperoodon boops.

conspicuous

during the day

„ 29

58° 28' S., 51* 56' W.

33-3° F.

Pale blue

Many finners were

Baltenoptera musculus.

seen early this

morning

Second Voyage of the " Scotia."

Date.

Locality.

Temp.

Colour.

1904.

Feb. 11

55° 47' S., 54° 19' W.

39-8" F.

Blue

A school of por-

...

„ 13

59° 56' S., 49° 30' W.

36-0° F.

»

poises seen in
the evening
A large whale

Balsenoptera (sibbaldi 1).

seen

„ 23

,, 24
„ 25
„ 26
„ 29

61° 28' S., 41° 55' W.
62° 49' S., 38° 12' W.
64° 29' S., 35° 29' W.
65° 59' S., 33° 06' W.
68° 08' S., 27° 10' W.

33-3° F.
31-6° F.
31-0° F.
30-2° F.
29-2° F.

i

»
j
i

>

vSeveral finners
Several tinners
Several finners
Some finners
Many grampuses
and the blast
of a whale

,, musculus.

»5 M
)» J)
>' »»

Orca gladiator and Balsen-
optera sp. ?

Mar. 1
2

68° 43' S., 24° 15' W.
71'04'S., 23° 10' W,
off Coats Land

31-0° F.
30-8° F.

I*
»

Some grampuses
Grampuses

Orca gladiator.
»i i)

3
4

72° 18' S., 17° 59' W.,
off Coats Land
72° 22' S., 18° 13' W.,
off Coats Land

30-9° F.
30-9" F.

n

j»

Many grampuses
and whales
Some whales were
seen in the

,, ., and Balaen-
optera sp. ?
Balsena (australis t).

evening ; Cap-
tain Robertson

considered their

blast resembled

thatof the Bow-

head. A few

grampuses were
also sighted

SOME OBSERVATIONS ON ANTARCTIC CETACEA.
Second Voyage of the " Scotia " —continued.

497

Date.

Locality.

Temp.

Colour.

1904.

Mar. 5

72° 31' S., 19° 00' W.,

29-2° F.

Indigo

Grampuses and

Orca gladiator and I'alaen-

off Goats Land

blue

whales

optera sp. ?

„ 12

74* 01' S., 22° 00' W.,

A grampus

Orca gladiator.

off Coats Land

13

74' 01 'S., 22° 00' W.,

Several bottle-

Hyperoodon rostratus and

off Coats Land

noses and two

Orca gladiator.

grampuses

1*

73° ll'S., 23° 53 \V.

28-9° F.

Indigo

Two grampuses

Orca gladiator and Balaen-

seen and finner

optera musculus.

heard blowing

„ 15

71°50' S., 23° 30' W.

29-1° F.

Bright

A finner

Balaenoptera musculus.

indigo

.

16

71° 28' S., 22° 32 W.

29-0° F.

Indigo

Many finners and

Balaenoptera musculus and

grampuses

Orca gladiator.

„ 17

71° 22' S., 18* 15' \V.

29-6° F.

Blue

Many grampuses

» » »

and finners

„ 18

71° 22' S., 16° 34' \V.

29 9° F.

M

Grampuses

Orca gladiator.

19

71' 32' 8., 17° 15' \V.

29-8* F.

>i

Grampuses and

Balaenoptera musculus and

finners

Orca gladiator.

„ 20

71° 17'S., 18*50' VV.

30-4° F.

Antwerp

Numerous gram-

» j» »

blue

puses and a

few finners

, 21

69C 33' S., 15° 19' W.

31 -0' F.

19

Some finners

Balaenoptera musculus.

22

68° 32' S., 10° 52' W.

31-8° F.

Cobalt

One or two finners

» »

, 23

68° 32' S., 12* 49' W.

31-1° F.

Indigo

Many grampuses

„ „ and

and finners

Orca gladiator.

, 24

68° 41' S., 12° 36' W.

...

• • •

Finners

Balaenoptera musculus.

, 25

68' 26' S., 11° 11' W.

31-3° F.

Antwerp

A few finners

i» D

blue

„ 27

66* 57' S., 11° 13' W.

30-5° F.

Blue

Some grampuses

Balaenoptera musculus and

and finners

Orca gladiator.

„ 28

65° 58' S., 11° 24' W.

31-2° F.

H

Some grampuses

i> »> »

and finners

Apr. 3

56° 55' S., 10° 00' W.

32-9° F.

J)

Many whales all

Megaptera boops and

day, mostly

Balaenoptera sp. ?

humpbacks

6

54°33'S., 11° 47' \V.

...

»

Whales were seen

Balaenoptera sp. ?

7

53° 58' S., 10° 10' W.

...

Dull

A whaleand three

Balaenoptera sp. and Orca

Antwerp

or four gram-

gladiator ?

blue

puses

No whales seen

between this

position and

Gough Island,

but porpoises

seen in 46° 31'

S., 10° 10' W.

and 43° 21' S.,

8° 30' W.

If reference be made to the Zoological Log of the Scotia (Vol. IV. Part I.), it will
be found that the plentiful appearance of whales is almost always associated with the
presence of large numbers of birds of different species and with phosphorescence of

498 DR W. S. BRUCK

the sea. The presence of Eupliausia, which are themselves phosphorescent, is also a
sign that general conditions are favourable to the presence of whales ; in short, any
sign, direct or indirect, of a plentiful food supply both for whales and birds is signifi-
cant. Mr RACOVITZA throws doubt on my statement that myriads of Cape pigeons
and thousands of finners were seen on December 16, 1892, in latitude 59° 24' S., longi-
tude 50° Ol' W. Twenty years later this statement is more than vindicated by Mr
THEODORE E. SALVESEN * reporting that nearly eleven thousand whales were killed
and captured in that same region from "1st November 1912 till the end of April
1913," even after several years' hunting. The previous season seven thousand whales
were taken at South Georgia alone. The sight of these whales and birds in December
1892 will for ever remain one of the most vivid of my Antarctic recollections.
Whales' backs and blasts were seen at close intervals c[uite near to the ship, and
from horizon to horizon, while Cape pigeons were tumbling over each other after
small pieces of fat thrown over the ship's side, just as do fulmar petrels after scraps of
whale fat in the northern hemisphere. These Cape pigeons were captured with an
ordinary hand landing-net over the side of the ship in such numbers that our
crew of forty-seven hands were furnished with a very full supply of " scouce." t
The sea was swarming with Euphausia. This host of animal life in Antarctic seas
is surely no more impossible than a swarm of locusts, an army of lemmings, or a
flock of owls.

It will be noted that on January 26, 1893, I recorded seeing a white finner
(Balaenoptera musculus}. Doubt has been thrown on this observation of mine, but
I have recently seen a photograph of a piebald humpback whale that was landed in
South Georgia, and am told that this is not a solitary example, and that white hump-
backs have also been taken there. I now have the additional satisfaction of present-
ing herewith an excellent photograph (Plate II.) of a piebald Nordkapper (Balsena
biscayensis) which was taken, and lent to me for reproduction, by the well-known
author Mr J. J. BELL. Mr BELL tells me that out of a dozen whales he saw captured
in the Atlantic, near St Kilda, about seven years ago, " several were piebald."

Globiocephalus species ?

On the 22nd November 1892, on board the Bal&na in 40° Ol' S., 48° 55' W., I
record : " Very green sea. Whales and seaweed. Globiocephalus." These are also
recorded by BURN MURDOCH, \ with the accompanying black-and-white sketch. On
November 24 he describes them thus : " We saw many hundreds of small whales
or porpoises the night before this last gale. They came up from the N.W., and

* Vide Vol. IV., Part XIX. Special reference should also be made, not only to Mr THEODORE E. SALVESEN'S
account of "The Whale Fisheries of the Falkland Islands and their Dependencies" in this volume, but also to his
article entitled " The Whaling Industry of To-day," in The Scottish Hunkers' Magazine, vol. iv., No. 14, July 1912,
pp. 109-119.

t A whaler's term for a thick stew resembling jugged hare, made of guillemots, penguins, etc.

J From Edinburgh to the Antarctic, by W. G. BURN MURDOCH, pp. 141-142.

SOME OBSKRVATIONS ON ANTARCTIC CETACEA.

499

passed us swimming S.E., travelling in companies of seven or eight, plunging half
out of the seas, and tossing up spirts of white water. They were about seven feet
in length, with black round heads and a white patch over the eye. Some had
patches of grey-white on their backs. They resembled the American drawings
(True) of the pigmy sperm, but had a large dorsal fin."

BURN MURDOCH'S sketch and my remembrance of these animals incline me to
the belief that they were Globiocephalus, and that they were Globiocephalus melas
(Traill). BURN MURDOCH'S estimate of their length, the estimate of an extremely
accurate field naturalist, is, however, small, being only a little more than half the
length given by TRUE ; moreover, the white patch over the eye and some grey -white

Globiocephalus. By W. G. Burn Murdoch.

on their backs do not tally with the white markings recorded by TRUE, but these
may be variable. The dorsal fin, however, appears to agree with that of Globio-
cephalus. MURIE'S figure of the head closely resembles that of BURN MURDOCH,
except in regard to the upper lip, the shape of which in the case of a swiftly
moving animal may easily have been missed. The shape and position of the
dorsal fin in both are practically identical, as well as the general contour of the
body.* It may be useful to give TRUE'S description t of the genus and species
Globiocephalus melas : —

" Globiocephalus, Gray.

" Rostrum short and very broad. Rostral portion of intermaxillae flat and very
broad (sometimes covering the entire anterior half of the rostrum). Symphysis of
mandible short. Pterygoid bones large and in contact. Teeth few and large, 7 to 11,
confined to the anterior half of the rostrum. Vertebrae, 57 to 60.

" Head globular, with a rounded protuberance on the lip ; beak wanting ; mouth
oblique. Dorsal fin very long, low, and thick. Pectoral fins narrow and very long.
Colour black.

* MURIB, Trans, of the Zool. Soc. of London, viii., 1873, pi. xxx. fig. 1.

t "Contributions to the Natural History of the Cetaceans : a Review of the Family Delphinidse," by FREDERICK
W. TRUE, Bulletin of the United States National Museum, No. 36, pp. 183, 184, pi. xl. fig. 1.

500 DR W. S. BRUCE

" Globiocephalus melas, Traill. (PI. xl. figs. 1 and 2.)

" Size large ; form stout. Head globose ; forehead protuberant, overhanging the
lip in adult individuals ; body especially deep opposite the dorsal fin. Pectoral fins
very long, slender, and pointed ; length about one-fifth the total length of the body.
Dorsal fin on a long base and strongly recumbent, situated anterior to the middle of
the length of the body. Caudal ridges prominent, extending respectively to the
dorsal fin and to the vent. Flukes large and broad. Mouth oblique.

" General colour uniform black ; a large hastate white area on the breast, extending
from the line of the corners of the mouth to the base of the pectoral fins; from
behind this area a white band, which is much the broadest in the posterior half,
extends backwards along the median line to the vent ; the whole white area has the
general form of an arrow with its head, shaft, and feathers.

"Teeth-. Vertebra: C. 7 ; D. 11; L. 13(-14) ; Ca. 27(-29) = 58-60. Skull

large and massive. Rostrum longer than broad at the base ; its breadth at the base
slightly less than three-fourths its length. Intermaxillae large and flat ; not greatly
broader anteriorly than posteriorly ; rugose in front ; their outer margins not
coinciding with the margins of the maxillae, except at the extremity of the rostrum ;
separated in the median line throughout. Superior nares broad transversely, and
bordered by narrow plates of the intermaxillse. Vomer extending nearly to the
extremity of the rostrum. Pterygoid bones large, obtusely keeled, and closely
approximated in the median line. Temporal fossae moderate, oval.

" Measurements of the Exterior (from Bell). — Total length, 182 inches (462'3 cm.) ;
extremity of snout to corner of mouth, 14'5 inches (36'8 cm.); to dorsal fin, 55 inches
(1397 cm.) ; length of pectoral fin along anterior edge, 50 inches (127 cm.) ; greatest
breadth of pectoral fin, 11 inches (27'9 cm.).

"Measurements of the Skull (U.S. National Museum, No. 12,100, Cape Cod,
Massachusetts). — Total length, 655 cm.; length of rostrum, 33'4 cm.; breadth of
rostrum at base, 24'1 cm. ; at its middle, 19'4 cm. ; breadth of intermaxillse at same
point, 15 '4 cm. ; breadth between orbits, 42 '3 cm. ; length of temporal fossa, 17 '3 cm.

" Habitat. — Atlantic coast of North America to New Jersey ; coast of Europe ;
Cape of Good Hope ; New Zealand."

THE PIEBALD PORPOISE (Lagenorhynchus cruciger'l}.

Considerable interest attaches to the porpoise * seen several times by myself on
board the Balsena in 1892 and 1893 in the vicinity of the Falkland Islands, and
seen again in 1903 and 1904 by the other naturalists of the Scotia and myself in
this locality, the same probably as a school of porpoises that the Scotia fell in with

* This porpoise was called a " ground porpoise " by the crew of the Balsena.

SOME OBSERVATIONS ON ANTARCTIC CETACKA.

501

to the south of Gough Island. For convenience, I have called this animal the
" Piebald Porpoise," on account of its special markings, which are here specially
considered.

I here tabulate the results of those observations in the same way as I

have the larger Cetacea.

The Piebald Porpoise (Lagenorhynchus crucigerf).

Date.

Locality.

Temp.

Colour.

1892.

Dec. 4

46° 41' S., 51° 40' W.

55-3° F.

...

Porpoises.

,, 11

Port William, Falkland

40-7° F.

Piebald porpoises.

Islands, 51° 40' S.

1893.

Mar. 3

Port Stanley, 51° 42' S.,

. . .

Piebald porpoise captured in fisher-

57° 51' AV.

man's net. Skeleton taken and

afterwards presented to University

College, Dundee.

6

50° 35' S., 53° 53' W.

Porpoises (piebald ?).

9

47° 45' S., 47° 52' W.

56-7° F.

...

Porpoises.

10

46° 31' S., 45° 30' \V.

56-8' F.

Porpoises.

12

43° 51' S., 40* 47' W.

57-3° F.

Porpoises.

16

39° 28' S., 36° 48' W.

65-7° F.

Porpoises (piebald 1).

1903.

• Ian. 4

47° 37' S., 57° 25' W.

49-2° F.

Very dark

The following note is from the Zoo-

green

logical Log of the Scotia, Vol. IV.

Part I. p. 6 : "A large school of

porpoises, black on the back and

white on the belly, probably the

same species as those seen by Mr

Bruce in 1892 in about the same

latitude and at Port Stanley, were

seen playing under the bows of

the ship at 8.45 p.m."

„ <--'•"'

Port Stanley, 51° 42' S.,

53-1° F.

Turbid

Piebald porpnises were seen several

57° 51' \Y.

times.

27

52° 55' S., 55° 00' W.

44-6° F.

Light greenish

Several piebald porpoises seen.

blue

1904.

Feb. 1 1

55° 47' S., 54° 19' \V.

39-8c F.

Blue

A school of porpoises (probably

piebald) seen in the evening.

Apr. 14

46° 35' S., 10° 10' \V.

42-9* F.

Lipht cobalt

Porpoises (probably piebald).

„ 18

43° 21' S., 8° 30' W.

52-0' F.

Cobalt

White, piebald, and black porpoises.

On 3rd March 1893, at Port Stanley, Falkland Islands, I purchased one of these
porpoises from a fisherman. It had been drowned by being entangled in one of his
nets in the harbour. This fresh specimen measured 4 feet 4 inches in length, and
before dissecting it for skeletal purposes it was fortunate that the well-known artist,
Mr "VN . G. BURX MURDOCH, who was my companion on that voyage, was able to make
a series of faithful studies of the animal as it lay on the deck of the Bal&na. These
studies are now reproduced fac-simile (vide Plate). After discussion with Mr BURN

502 DR W. S. BRUCE

MURDOCH we have both come to the conclusion that it was better to reproduce
faithfully the drawings of twenty-two years ago rather than to court even slight
errors by redrawing them in finished diagrammatic form. The drawings are a very
faithful representation of the freshly killed specimen.

On my return to Dundee I entrusted the skeleton to Professor D'AECY W,
THOMPSON, C.B., who accepted it for his excellent Museum at University College,
Dundee. Professor D'AnCY THOMPSON has recently looked through the Museum,
but has failed to find the specimen. He suggests that " during a long period some
years ago when the Museum was not well served, possibly this small skeleton,
which had got somewhat injured by rats, was either thrown out or was made
use of for class purposes, in the belief that it was only a common porpoise."
The loss of this skeleton removes the only concrete facts beyond Mr BURN
MURDOCH'S drawings and my measurement, and with Professor D'ARCY THOMPSON
I can only regret its loss.

Consulting with Dr S. F. HARMER, inquiries have been made at Cambridge, but
no trace of the animal is at this time to be found. Neither does any other specimen
appear to exist in British or foreign museums. In his report on the seals of the
Discovery the late Dr WILSON * describes a porpoise which he frequently saw in
subantarctic seas ; but its length, to my mind, precludes the probability of it being
the same species. Dr JACQUES LIOUVILLE f describes a school of eight brown-and-
white " Delphinides," the colour and position of the markings of which appeared to
him to be characteristic.

Dr LIOUVILLE gives the length of those animals seen by him as I'lO metres, i.e.
about 44 inches, but presumably this was estimated and not actually measured. It
may be noticed that this compares somewhat with the size of my Falkland Islands
specimen, which had a measured length of 52 inches. Dr WILSON'S specimens were
estimated at 8 to 10 feet long, i.e. 96 to 120 inches, or fully double the length of my
specimen. Dr LIOUVILLE is of opinion that the animals seen by himself and Dr
WILSON are of the same species but different ages. But in shape as well as in size
my specimen resembles Dr LIOUVILLE'S rather than Dr WILSON'S. The snout is more
truncated than beaked, though it is somewhat depressed above, as can be seen in
Mr BURN MURDOCH'S outline sketch, as well as by the shading in his other drawings.
The pectoral and dorsal fins are more like those in Dr LIOUVILLE'S sketch, and indeed,
considering that the latter was drawn from a distant and active specimen, they may
well prove to be in reality identical. The main difference is in the dorsal fin, which
Dr LIOUVILLE has shown to have a sharp point, whereas Mr BURN MURDOCH indicates
a blunt termination. The characters of the tail in the two drawings are almost
identical. In size and shape, therefore, making allowance for drawings made from

* (British) National Antarctic Expedition, ii. 1907.

t "Sur le polymorphisme d'un Delphinide' des mers australes : Delphinus cruciger, Quoy et Gaymard" : Note de
M. JACQUES LIOUVILLE, Comptes rendus, t. 156, p. 90, stance du Janvier 1913 de 1'Acad. des Sciences.

SOME OBSERVATIONS ON ANTARCTIC CETACEA. 503

a quickly moving animal in the case of Dr LIOUVILLE, the two animals closely
resemble one another in form if they are not actually identical.*

The markings, however, of Dr LIOUVILLE'S animal and mine are decidedly
different. The Balxna specimen is mainly white, and its markings are black
rather than brown. The whole head, dorsally as far as the line of the base of
the pectoral fins, laterally less far back, thence just past the axilla, and extending
back in a central ventral line as far back as the extremity of the pectoral fin,
is black, with the exception of the throat. The snout is black, not white, as in
Dr LIOUVILLE'S drawing, and so are both upper and lower lips. The tail is entirely
black : ventrally this black extends forward to half way between the root of the
tail and the vent ; dorsally forward of the dorsal fin, which is itself black, to the
middle of the back. All the rest of the body is white. There is no lateral stripe
as in the case of Dr LIOUVILLE'S drawing. I am inclined, however, to think that
this is a polymorphic form of the same animal, especially as His Excellency W. L.
ALLARDYCE, formerly Governor of the Falkland Islands, tells me that his impression
of the Falkland Islands porpoise is that it is somewhat striped. Mr LYDEKKER is
of similar opinion to myself that the Balaena specimen is like Dr LIOUVILLE'S
Lac/enorJtyitchus cmciger, Q. and G. ; but in the absence of the skeleton this
determination cannot be absolutely relied on.

The following is TRUE'S description t of the genus Layenorlynchus and of
Layenorlnjnchus wuciger : —

"7. Lagenorhynchus, Gray.

" Rostrum large and broad. Rostral portion of intermaxillse flat (somewhat
convex in obliquidens and electra). Pterygoid bones in contact or separate.
Symphysis of mandible short. Teeth variable in size, 22 to 45. Vertebrae,
73 to 92.

" Head with a short, ploughshare-like beak (not distinctly marked off from the
forehead in obscurus). Dorsal and pectoral fins moderate, falcate. Caudal ridges
very prominent. Sides with two areas of light colour separated by irregular,
oblique dark bands.

" Layenorltyitchiis cruciger (D'Orbigny and Gervais). (PI. xxv. figs. 1 and 2.)

'; Beak short, only slightly marked off from the convexity of the forehead. Muzzle
to the corner of the mouth, forehead, back, dorsal fin, tail, and pectoral fins black.
On the sides, from the eye and base of the pectoral fins to the tail, a broad black
band. This band is broadest above the base of the pectoral fin, and decreases in

* "Cetaces de 1'Antaretique," par le Dr J. LIOUVILLE (Deuxieme Expedition Anhirctique Franfaise, Paris, 1913),
pp. 165-179.

+ " Contributions to the Natural History of the Cetacams : a Review of the Family Delpuinidae," by FREDERICK
\V. TRUE, Bulletin of the United States National Museum, No. 36, pp. 168, 170, 171, Washington, 1889.

504 DE W. S. BRUCE

width posteriorly until a point about opposite the anus is reached, after which it
again increases in width and joins the black of the tail. Between the median and
lateral black bands and on the belly the colour is white, more or less pure.
(D'Orbigny and Gervais.)

28
' Teeth - ; skull smaller than that of L. acutus, but similar in proportions.

28

Rostral portion of premaxillaa flat, not twisted into a vertical position at the distal
extremity ; their outer margins straight ; triangular area high, flat, and smooth ;
temporal fossae moderate, oval directed backwards ; pterygoid bones in contact in
the median line, moderate, very short antero-posteriorly, not keeled laterally ;
the conjoint postero-internal free margin transverse, as in L. acutus ; vomer
extending along three-fourths of the rostrum, appearing on the palate as a
narrow ridge.

"Measurements of the Skull (Mus. d'Hist. Nat., Paris, No. a 3045, type). — Total
length, 34'9 cm. ; length of rostrum, 17'8 cm. ; breadth of rostrum at the base,
10'7 cm. ; at its middle, 6'6 cm. ; breadth of intermaxillae at same point, 4'1 cm. ;
breadth between orbits, 17'1 cm. ; length of temporal fossa, 6'9 cm. ; depth of
temporal fossa, 4'1 cm.

" Habitat. — Seas south of Cape Horn ; Pacific Ocean."

In the museums of Britain and of the Continent there is great lack of material
for the comparative study of the larger forms of Cetaceans.

Through the general interest of some owners, however, front fins of many of
the species and other parts have recently enriched collections, and have enabled
the comparative anatomist to arrange phalanges, carpals, and other bones in
their skeletons and make models in a manner truer to their actual disposition in
the living animal.

What is really required are several scientific Cetacean expeditions with different
classes of ships : (l) one a well-protected ice ship, such as the Scotia, adequately fitted
with all the usual equipment for the fishing of Balsena mysticetus within the limits
of the polar pack ; (2) another a thoroughly modern whaler of the Norwegian type,
for securing different species of whales outside the ice limit, and the free use of a
whaling-station ship for a definite time ; (3) a third ship especially fitted out for the
hunting of smaller Cetacea ; (4) and, fourth, a vessel that will devote itself to following
the tracks of whales, to give us definite information regarding migration and
breeding habits. There should be no question of making these ships pay by hunting
for other than scientific purposes, although other scientific observations and collec-
tions should be made, such as temperature, salinity, and plankton as far as they
are definitely associated with the food supply and life of the whale.

The question of migration is of very special interest. Unlike almost all other
marine animals, there are indications that whales, if not the smaller Cetacea as well,
are not so dependent on the temperature of the water remaining moderately constant,

SOME OBSERVATIONS ON ANTARCTIC CETACEA. 505

but are capable of withstanding very great variations in temperature. There are
indications, for instance, that whales migrate from polar to tropical seas, and that in
consequence northern and southern species are probably in many cases identical, as
has been urged by Sir WILLIAM TURNER and others.*

* " The Right \\Tiale of the North Atlantic, Balxna bucayensis, etc.," by Principal Sir WILLIAM TURNER, K.C.B.,
F.R.S., D.C.L., Trans. Hoy. Soc. Edinburgh, vol. xlviii. pt. iv., 1913.

"The Baleen Whales of the South Atlantic," by Principal Sir WILLIAM TURNER, K.C.B., F.R.S., D.C.L.,
Proc. Roy. Soc. Edinburgh, vol. xxxv. part i. No. 2, 1914

EXPLANATION OF PLATES.

Plate I. Facsimile drawings by Mr W. G. BURN MURDOCH of a piebald porpoise (Lagenorhynchus

l) taken at the Falkland Islands.
Plate II. Photograph of Nordkapper (Balsena biscat/insi*), by Mr J. J. BELL.

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