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BRITISH MUSEUM (NATURAL HISTORY).

BRITISH ANTARCTIC (“TERRA NOVA”) EXPEDITION, 1910.

NATURAL HISTORY REPORTS.

ZOOLOGY. VOL. I:

VER Piibiveebe: |

ty

fiona) Muse

LONDON:
PRINTED BY ORDER OF THE TRUSTEES OF THE BRITISH MUSEUM.

Sold by B. Quaxrircu, Lrp., 11 Grafton Street, New Bond Street, W. 1;
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AND AT THE
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[All rights reserved.|

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508.44
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Si

PREFACE

Tur ‘Terra Nova” Expedition (1910-13), under the command of the late Captain
R. F. Scott, R.N., 0.V.O., will ever be remembered for the journey to the South
Pole which ended in the death of the leader and his four brave companions. But,
in spite of this disaster, the expedition accomplished all that it set out to do. It
was thoroughly equipped for scientific purposes, in both men and material, and the
programme of scientific work was carried out in full.

The biologist in charge of operations on the ship was Mr. D. G. Lillie, M.A., to
whose skill and energy the large and valuable marine collections are mainly due.
On the outward and homeward voyages from England to New Zealand fine-meshed
tow-nets were put overboard whenever possible, and seventy plankton samples were
obtained ; in addition two hauls were made with the trawl, one near the Falklands,
at a depth of 125 fathoms, and one off Rio de Janeiro, at a depth of 40 fathoms.
The winter cruise (July 10th to October 10th, 1911) round the Three Kings Islands
and to the north of New Zealand produced biological results of great importance ;
eighty plankton samples were obtained, and the seven hauls made with trawl and
dredge at depths of 15 to 300 fathoms revealed a bottom-fauna of extraordinary
variety, including a great number of forms new to science. Between New Zealand
and McMurdo Sound one hundred and thirty-five samples of plankton and fifty of
muds and oozes were obtained ; in the Ross Sea and in McMurdo Sound fifteen rich
hauls with the trawl, at depths of 40 to 300 fathoms, produced a collection which has

added greatly to our knowledge of the Antarctic marine fauna.

iv PREFACE

In this work on the ship several officers helped. Mr. Lillie also wishes to
acknowledge the assistance in trawling and in winding up plankton nets given, often
out of working hours, by Seamen A. S. Bailey, A. Balson, W. L. Heald, J. Leese,
M. McCarthy, T. F. McLeod, and T. S. Williamson. Seaman McLeod also mended
nets; others who should be mentioned are Boatswain A. Cheetham, who helped
with the trawl and supplied tackle, Carpenter F. E. C. Davies, who made frames for
nets, and Chief Engineer W. Williams, who repaired trawl frames and other iron
gear.

In addition to the trawling and tow-netting Mr. Lillie paid special attention to
whales, recording all those seen from the ship. In October 1911 he visited the
Whaling Station at Whangamumu, near the Bay of Islands, and in 1912 he spent
four months (July to October) in the same region on two floating factories belonging
to the New Zealand Whaling Company, He is the author of the Report on the
Cetacea (Zoology, Vol. I, No. 8).

That accomplished naturalist and skilful artist, the late Dr. E. A. Wilson, was
the chief biologist of the shore party; he was mainly responsible for the collection
of Birds, the report on which will include many reproductions of the sketches he
made. Dr. Wilson, with the late Lieutenant H. R. Bowers and Mr. A. Cherry-Garrard,
undertook a most arduous and hazardous winter journey to the Emperor Penguin
Rookery at Cape Crozier, to secure eggs with a series of early embryos, material that
is now in the hands of Professor J. Cossar Ewart, F.R.S. Staff-Surgeon G. Murray-
Levick, R.N., made numerous observations on the habits of the Adélie Penguin, and
has written a report on this subject (Zoology, Vol. I, No. 2) illustrated by photographs
that he took, Surgeon-Captain E. L. Atkinson, R.N., parasitologist to the expedition,
collected and studied numerous parasites, the Antarctic material being taken chiefly
from seals and fishes during the winter of 1911. The results of his work appear in
the Report on Parasitic Worms (Zoology, Vol. II, No. 8) prepared by him in con-
junction with Dr. R. T. Leiper, F.R.S.

PREFACE v

Further details of the scientific work done by the expedition will be found in
“Seott’s Last Voyage” (Smith, Elder & Co., 1913).

When the collections reached England arrangements for working out many of the
groups were made, and it was decided to issue the Zoological Reports from time to
time, as they were ready for publication. The subject-matter was distributed
provisionally among as many volumes as seemed likely to be required, the first to
include the Vertebrates, the second the Molluscs and Worms, and the third the
Arthropods. These three volumes have now been closed, and the outstanding

reports on these groups will appear in other volumes of the series.

C. TATE REGAN,
Keeper of Zoology.

19th January 1924.

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ZOOLOGY, VOL. I.

CONTENTS:

1. Re@an, C. T. Fishes. [Issued June 27, 1914.]
Pp. 1-54, 18 pls.

2. Levicx,G.M. Natural History of the Adélie Penguin. [Jsswed February 27, 1915.]
Pp. 55-84, 21 pls.

3. Linu, D. G. Cetacea. [Issued March 27, 1915.]
Pp. 85-124, 8 pls.

4, Regan, C. T. Larval and Post-larval Fishes. [Issued March 25, 1916.]
Pp. 125-156, 10 pls.

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BRITISH MUSEUM (NATURAL HISTORY).

BRITISH ANTARCTIC (“TERRA NOVA”) EXPEDITION, 1910.

NATURAL HISTORY REPORT.

ZOOLOGY. VOL. I—No. 1, pp. 1-54.

Fishes:

BY

C TATE REGAN, MA.

WITH EIGHT FIGURES IN THE TEXT AND PLATES I-XIQ1,

LONDON :

PRINTED BY ORDER OF THE TRUSTEES OF THE BRITISH MUSEUM,

Sold by Lonemays, Gremy & Co,, 39, Paternoster Row, London, E,C,; B. Quarrroug, 11, Grafton Street, New Bond Street,
London, W.; Dunau & Co,, Lrp., 87 Soho Square, London, W.; and the Mmpnanp Epvucarionan Co.,; Lrp.,
41 and 43, Corporation Street, Birmingham;
OR aT THR
Britis Mouszum (Narurnan History), Cromwell Road, London, 8,W.

1914
[All rights reserved.]

Price Ten Shillings and Sixpence.
Issued 27th June; 1914.J

(British Museum (Caturaf History)

‘Terra Nova” Report.

This zs No.
printed on Special Paper, of the Report on

“ Fishes.”

BRITISH MUSEUM (NATURAL HISTORY).

BRITISH ANTARCTIC (“TERRA NOVA”) EXPEDITION, 1910.

NATURAL HISTORY REPORT.

ZOOLOGY:

Tue Trustees of the British Museum have undertaken the publication of the Natural
History results of the British Antarctic Expedition, 1910, conducted by the ss. “ Terra
Nova,’ under the command of the late Capt. R. F. Scott, R.N., C.V.O. Arrangements

for working out most of the groups collected have already been made.

Mr. C. Tate Regan’s Memoir on Fishes is the part which is ready for publication
first. It may be useful to explain that it is proposed to issue the Zoological Reports,
from time to time, as they are ready for publication. By adopting this method the
delay which results from waiting until a volume is completed may be avoided ; but, on

the other hand, a natural sequence of subjects can hardly be maintained.

The plan that will be adopted will be to distribute the subject-matter provision-
ally among as many volumes as seem likely to be required. No attempt will be made
to complete a volume before commencing the publication of its successor; and two or
more volumes may accordingly be in process of publication concurrently. When a
volume has reached a convenient size it will be completed by the issue of a title-page
and table of contents.

SIDNEY F. HARMER,
Keeper of Zoology.
British Museum (Narurat History),

Lonpon, S.W.
June 27, 1914.

PiIShi ES:

BY C. TATE REGAN, M.A.

I.—Systematic.

1. The Antarctic Fishes ; ; : : d 1
2. Fishes from New Zealand ; ; : : ! : 13
3. Fishes from Brazil. : : . F : F ; 21

11.— General.

1. The Distribution of Antarctic and Subantarctic Fishes . : 24
2. The Antarctic Continent during the Tertiary Period. : 40

I—SYSTEMATIC PART.
1. THE ANTARCTIC FISHES.

THE collection includes examples of twenty-five species, twelve of which have been
described as new to science in a preliminary note (Ann. Mag. Nat. Hist. (8) xq,
1914, pp. 11-14), four of these being new generic types. This large proportion of new
forms is doubtless due to the fact that the bulk of the collection was dredged at depths
varying from about 50 to 250 fathoms. All but three of the species belong to the
group Nototheniiformes, and the additions to our knowledge of the genus 7rematomus,
the Harpagiferinae and the Chaenichthyidae, are of considerable importance. A new
genus of the Bathydraconidae resembles the northern Cottid Icelus in its armature of
spinate bony plates, and the first Antarctic species of Paraliparis is of interest.

MycToPHIDAE.
1. Myctophum antareticum, Giinth. >
55° 6’ S., 120° 3’ W., surface.

MURAENOLEPIDAE.
2. Muraenolepis microps, Lénnb. (PI. 1, fig. 2).
Muraenolepis marmoratus microps, Linnberg, Swedish 8. Polar Exped. Fish., p. 43 (1905).
Depth of body 6 in the length, length of head 43. Length of snout 3, diameter
of eye 5 in length of head, equal to or a little less than interocular width, much greater
B

2 “TERRA NOVA” EXPEDITION.

than interorbital width. Barbel + or § length of head. Maxillary extending to below
anterior part or middle of eye. Dorsal filament (absent in one specimen) longer than
diameter of eye, inserted a little behind base of pectoral. Anal origin only a little in
advance of middle of length of fish. Pectoral $ to 3, pelvics 3 length of head.
Greyish.

Here described from two specimens, 130 and 140 mm. in total length, from off
new land south of the Balleny Islands; depth 200 fathoms.

These seem to belong to the same species as a small fish (55 mm.) from the South
Sandwich Islands, which is doubtless identical with Lénnberg’s JZ mierops from South
Georgia. J marmoratus, Giinth., from Kerguelen, has a somewhat deeper body and
shorter head, larger eye, shorter barbel and dorsal filament. JL. orangiensis, Vaill.,
from Magellan, seems to be more slender and to have a smaller head and longer barbel
than either of the other species, from which it differs also in the much longer tail, more
than 13 as long as the rest of the fish.

NoToTHENIIDAE.
Nototheniinae.
Trematomus, Bouleng.

The known species of this genus number fourteen (or thirteen if 7. dubius be the
young of 7. vicarius), from the coasts of the Antarctic Continent and South Georgia.
In addition to three new species the “Terra Nova” obtained examples of two formerly
placed in Notothenia, enabling me to examine the pectoral arch and to assign them to
their correct position.

Synopsis of the Species.
I. Upper surface of head naked.
A. Cheeks and opercles fully scaled.

Interorbital width 31 to 5 in length of head. D. vi-vint, 32-38. <A. 32-36 . . newnesit.
Interorbital width 8 or 9 in length of head. D. 1v, 37. A. 32-33 : F . nicolai.

B. Cheeks and opercles scaly above, naked below.
D. v—v1, 34-37. A. 31-33. ; : : ‘ ‘ F . borehgrevinkii.
D. 1v—v, 30-33. A. 29-30 ; : : ; A : : j brachysoma.

II. Occiput scaly ; cheeks and opercles fully scaled.
A. Interorbital region naked, or incompletely scaled. D. 1v-v1, 33-38. A. 31-35.
Interorbital region naked, or with a single median series of scales ; diameter of eye 3 (young)
to 41 (adult) in length of head ; 60 to 75 scales in a longitudinal series bernacchit.
Interorbital region with two or three series of scales in the middle ; diameter of eye 43 (adult)
in length of head ; 56 to 59 scales in a longitudinal series . : . vicarius.
Interorbital region naked ; diameter of eye 33 (young) in length of Tend: 55 scales in a
longitudinal series : ‘ : ; 3 : ; : ‘ . dubius.

B. Interorbital region fully scaled ; praeorbital naked.
1. 60 to 75 scales in a longitudinal series, 34 to 46 in upper lateral line.
D. vi-vu, 36-41. A. 33-36 . . ; : : : ; ‘ . hansoni.
D. v-vu, 31-35. A. 31-35 . : : 4 ; ¢ : : loennbergii.

FISHES—REGAN, 3

2. 52 to 56 scales in a longitudinal series, 30 to 36 in upper lateral line.
D. v—vi, 32-35, A. 29-32.
Dorsal spines flexible d : : 2 : : : ; é . pennellii.
Dorsal spines pungent. ; : ; : : ; é : centronotus.
C. Interorbital region fully sealed ; praeorbital scal y:
Snout and lower jaw naked. D. 1vy—vi, 31-34. A. 29-32. 46 to 54 scales in a longitudinal
series ¢ 3 5 ° : : : o b é . 0 ; scotti.
Snout and lower jaw scaly. D. vi-vi, 31-33. A. 54-36. 70 to 80 scales in a longitudinal

series : ; : . : : é : lepidorhinus.
Snout and lower jaw scaly. D. vi, 35-36. A. 33-34. 70 scales in a longitudinal
series 5 5 5 5 : : c 5 5 eulepidotus.

3. Trematomus brachysoma, Pappenh. (PI. 1, fig. 3).
A specimen of 170 mm. in total length, stranded on an ice floe in Ci 2 AS ivi
34’ W. A water-colour sketch made by Mr. Lillie shows the body purplish gray, and

the fins blue, with the dark spots of the same tint as the colour of the body.

4. Trematomus bernacchii, Bouleng. (PI. 1, fig. 1).

Cape Evans and off Cape Adare, 45 to 50 fathoms ; bottom shingle.

5. Trematomus hansoni, Bouleng. (PI. 1, fig. 2).

A large number of examples of this and the preceding species were caught in 1911
at the winter quarters, Cape Evans, by means of a fish trap made of wire netting
stiffened by iron hoops and bars, which was lowered through holes in the ice. It was
noted that when a new hole was tried one or two good catches would result, and then
no more at all, perhaps because the attention of Weddell Seals had been attracted.
The fish were eaten and had a distinctly sweetish taste.

Three water-colour sketches were made by Dr. Wilson to show the natural

coloration ; two of these are reproduced on Plate I.

6. Trematomus loennbergiu, Regan.
Trematomus loennbergii, Regan, Trans. R. Soc. Edinburgh, xu1x, 1913, p. 263, pl. vm, f. 4.

Depth of body 4 to 5 in the length, length of head 3 to 32. Diameter of eye
3 to 34 in leneth of head, interorbital width 6 to 10. Maxillary extending to below
anterior 4 or 4 of eye; upper surface of head to nostrils, cheeks and opercles scaly.
10 to 13 gill-rakers on lower part of anterior arch. Dorsal v—vit, 31-35. Anal 31-35.
Pectoral nearly as long as head ; pelvics reaching anal. Caudal rounded or subtruneate.
Caudal peduncle as long as or longer than deep. 60 to 75 seales in a longitudinal
series, 34 to 46 in upper lateral line; lower lateral line, when developed, sometimes
with as many as 15 tubules. Body with irregular dark cross-bars.

Several examples up to 190 mm. in total length, from off new land south of
the Balleny Islands, at a depth of 200 fathoms, from near Inaccessible Island, 222 to
241 fathoms, and from the entrance to McMurdo Sound, 77° 13’ 8, 164° 18’ W.,
207 fathoms.

4 “TERRA NOVA” EXPEDITION.

7. Trematomus pennellii, Regan (Pl. 11, fig. 2).
Trematomus pennellii, Regan, Ann. Mag. Nat. Hist. (8) xm, 1914, p. 12.

Depth of body 43 to 5 in the length, length of head 33 to 33. Diameter of eye
34 to 33 in length of head, interorbital width 8 to 10. Maxillary extending to below
anterior +.of eye; occiput, interorbital region, cheeks and opercles scaly. 13 to 16
gill-rakers on lower part of anterior arch. Dorsal v—v1, 32-34; spines flexible. Anal
30. Pectoral 3 or + length of head; pelvics extending to origin of anal. Caudal
subtruneate. Caudal peduncle as long as deep. 52 to 56 scales in a longitudinal series
from above base of pectoral fin to caudal, 32 to 36 in upper lateral line; lower lateral
line without tubules. Olivaceous, with two or three series of large dark spots which
may unite to form irregular cross-bars.

Off Cape Adare; 45 to 50 fathoms; bottom shingle. Five specimens, 100 to 140
mm. in total length.

This species is named after Commander H. L. L. Pennell, r..

8. Trematomus centronotus, Regan (PI. mt, fig. 1).
Trematomus centronotus, Regan, Ann, Mag. Nat. Hist. (8), x11, 1914, p. 12.

Depth of body 4 in the length, length of head 34 to 33. Diameter of eye 34 in
the length of head, interorbital width 10. Maxillary extending to below anterior + or
! of eye; occiput, interorbital region, cheeks and opercles scaly. 14 gill-rakers on lower
part of anterior arch. Dorsal v—v1, 32-35; spines stiff, pungent. Anal 29-32.
Pectoral 2 or % length of head; pelvics extending to origin of anal. Caudal peduncle
as long as deep. 52 to 56 scales in a longitudinal series from above base of pectoral
fin to caudal, 30 to 36 in upper lateral line; lower lateral line without tubules. Large
dark spots uniting to form irregular cross-bars.

Two specimens, 175 and 210 mm. in total length, from MeMurdo Sound, 100 to
200 fathoms.

The pungent dorsal spines distinguish this species from all others of the genus, but
it so closely resembles 7. pennellii in other characters, scarcely differing except in the
larger eye, that it is undesirable to place it in another genus.

9. Trematomus seotti, Bouleng. (PI. rv, fig. 2).
Notothenia scotti, Bouleng. Nat. Antarct. Exped. Nat. Hist. 1, Fish, p. 2, pl. 1, f. 1 (1907);
Regan, Trans. R. Soc. Edinb. xtrx, 1913, p. 271.

Depth of body 4 to 55 in the length, length of head 3 to 33. Diameter of eye

2% to 34 in length of head, interorbital width about 12. Maxillary extending to below
anterior 4 of eye; occiput, interorbital region, praeorbitals, cheeks and opercles scaly.
10 to 13 gill-rakers on lower part of anterior arch. Dorsal tv—vi, 31-34. Anal
29-32. Pectoral $ to # length of head; pelvies reaching vent or anal fin. Caudal
rounded. Caudal peduncle about as long as deep. 46 to 54 scales in a longitudinal
series from above base of pectoral fin to caudal, 11 to 23 in upper lateral line; lower

FISHES—REGAN. 5

lateral line without tubules. Body with dark spots or irregular cross-bars; spinous
dorsal blackish ; soft dorsal and anal with a blackish band which is broadest and most
intense posteriorly ; caudal, pectorals and pelvics barred. Vertebrae 15 +32.

This species was known previously only from the type, a specimen of 110 mm.
taken off the Ross Barrier at a depth of 300 fathoms. Numerous examples up to
180 mm. in total length were obtained by the “Terra Nova” in the following
loealities :—

Off new land south of the Balleny Islands, 200 fathoms.

Ross Sea, 74° 25’ S., 179° 3’ E., 158 fathoms.

Entrance to McMurdo Sound, 77° 13’8., 164° 18’ E., 207 fathoms, and 76° 56'S...
164° 12’ E., 160 fathoms.

McMurdo Sound, near Inaccessible Island, 222 to 241 fathoms.

10. Trematomus lepidorhinus, Pappenh. (PI. tv, fig. 1).
Notothenia lepidorhinus, Pappenheim, Deutsche Siidpolar-Exped. x11, Zool. v, p. 169, pl. rx, f. 1,
and pl. x, f. 1 (1912).

Three examples, 135 to 150 mm. in total leneth, from off new land south of the
Balleny Islands, depth 200 fathoms. “These have vi—vir, 31-33 dorsal and 34-35 anal
rays, 75 to 80 scales in a lateral longitudinal series, and 14 to 16 gill-rakers on the
lower part of the anterior arch.

I have examined the pectoral arch on both sides in all three specimens and find
that the foramen is well within the hypereoracoid. I can only conclude that in
exposing the foramen Pappenheim must have enlarged it ventralwards, which may
easily occur unless care be taken.

11. Trematomus eulepidotus, Regan (PI. tv, fig. 3).
Trematomus eulepidotus, Regan, Ann. Mag. Nat. Hist. (8) x, 1914, p. 12.

Depth of body 33 to 43 in the length, length of head 3? to 33. Diameter of eye
34 to 34 in length of head, interorbital width 43 to 5. Maxillary extending to below
anterior } of eye; head covered with scales, only the lips naked. 14 or 15 gill-rakers
on lower part of anterior arch. Dorsal vi, 35-36. Anal 33-34. Pectoral 2 to 4
length of head; pelvics extending to vent or origin of anal. Caudal truncate.
Caudal peduncle as long, or nearly as long as deep. 70 scales in a longitudinal series
from above pectoral fin to caudal, 42 to 46 in upper lateral line, 10 to 15 in lower.
Sides of body with dark spots connected to form a network; dorsal fin with oblique
dark stripes.

Two specimens, 140 and 165 mm. in total length, the smaller taken at the entrance
to McMurdo Sound, 76° 56’ 8., 164° 12’ W., at a depth of 160 fathoms, the larger from
near Inaccessible Island, depth 222 to 241 fathoms. There are two larger examples,
220 and 250 mm., in bad condition, also from McMurdo Sound, taken off Granite
Harbour at a depth of 50 fathoms.

6 “TERRA NOVA” EXPEDITION.

Compared with examples of 7. lepidorhinus of nearly the same size this species
has a shorter snout, more oblique mouth and broader interorbital region. . The tail is
shorter, as the vent is nearly equidistant from tip of snout and end of anal fin, whereas
it is much nearer the snout in 7. lepidorhinus. The much shorter lower lateral line
and the more numerous dorsal rays are further important differences.

12. Pleuragramma antarcticum, Bouleng.

Ross Sea, 158 fathoms. Cape Evans, frozen on an iceberg.

Harpagiferinae.

The subfamily Harpagiferinae includes Nototheniids with the body naked, the
gill-membranes broadly united to the isthmus and not forming a fold across it, the
operculum hooked so that its upper edge is deeply concave, and the upper lateral line
with tubules, the lower reduced to a series of pores.

The genera with a mental barbel were previously represented in the British
Museum only by an example of Dolloidraco longipinnis recently received from Professor
Roule, but the ‘Terra Nova” collection includes a large series of fishes of this group,
representing six species.

Synopsis of the Genera of Harpagiferinae.

I. A mental barbel ; opercles not spinate.
A. Post-temporal not prominent; head not or scarcely broader than deep ; interorbital region

narrow.
Spinous dorsal fin above base of pectoral. : : : ‘ Artedidraco, Loennb.
Spinous dorsal fin above operculum . : 5 ; b : . Dolloidraco, Roule.
B. Upper limb of post-temporal projecting as a prominent curved ridge; spinous dorsal above
operculum.
Head longer than broad, scarcely broader than deep; _ interorbital region
narrow ; : z : : : : 5 : Histiodraco, gen. noy.
Head as long as broad, much broader than deep; interorbital region
wide . é 5 : : : ; : : ‘ Pogonophryne, Regan.
II. No barbel ; operculum and suboperculum each forming a prominent spine . Harpagifer, Richards.

Artedidraco, Linnberg.
Artedidraco, Linnberg, Swedish South Polar Exped. Fish. p. 39 (1905).

Head without ridges or tubercles, covered with loose, smooth skin, longer than
broad, not or scarcely broader than deep; interorbital region narrow. Post-temporal
not prominent. Opercles not spinate. A mental barbel. Body compressed. Spinous
dorsal fin above base of pectoral.

Synopsis of the Species.

I. Barbel club-shaped, with papillose terminal knob.
Depth of body 5 to 54 in the length, length of head 25 to 3. D. mi-tv, 25. A. 17-18. Anterior
rays of soft dorsal longest —. ¢ : 2 3 ; : : : orianae.
Depth of body 4 in the length, length of head 22. D. 1m, 23-24. A. 17. Middle rays of soft dorsal
longest 5 ; ; : c . : ; : 0 F : mirus.

FISHES—REGAN.

~I

II. Barbel without distal expansion, either smooth, finely papillose or slightly fringed.
A. Caudal rounded or subtruncate.

D. 1-tv, 24-26. A. 17-20 . : : : 3 F ; : . skottsbergii.
D. 1v-v, 27-28. A. i9-20 . : : : ‘ ‘ , ; . shackletoni.
B. Caudal slightly emarginate. D. 11-111, 26-28. A. 18-21. 5 ; ‘ . loennbergii.

13. Artedidraco orianae, Regan (PI. v1, fig. 2).
Artedidraco orianae, Regan, Ann. Mag. Nat. Hist. (8) xin, 1914, p. 12.
Depth of body 5 to 53 in the length, length of head 22 to 3. Diameter of eye
+ in the leneth of head, interorbital width 10. Maxillary extending to below anterior

+ of eye. Barbel + length of head, club-shaped, with papillose distal expansion.

Seven short gill-rakers on lower part of anterior arch. Dorsal 111—1v, 25; rays of soft
dorsal decreasing from third or fourth; last adnate to caudal peduncle. Anal 17-18,
separated by an interspace from caudal fin. Pectoral 17-rayed, 7 or ¢ length of head,
extending to third or fourth ray of anal; pelvics as long, extending to vent or origin
of anal. Caudal subtruncate. Five dark bars across back, extending upwards on the
dorsal fins, the first at base of spmous dorsal, the second and fourth stronger than
the rest; a spot on praeorbital, another on cheek; lower part of body irregularly
spotted ; fins with series of spots.

Two specimens, 80 mm. in total length, from off Cape Adare; depth 45 to 50
fathoms ; bottom shingle.

The species is named in honour of Mrs. E. A. Wilson.

A, mirus, from South Georgia, described and figured by Lénnberg (Swedish South
Polar Exped. Fish. p. 40, pl. iv, f 14), from a specimen of 114 mm., appears to differ
from A. orianae in the deeper body (depth 4 in the length), larger head (2% in the
length), longer barbel (4 the length of head), shorter paired fins (pectoral 3, pelvics
3 to 2 length of head), higher soft dorsal with the middle rays longest and the last
joined to the caudal fin, and in the absence of an interspace between anal and caudal.

14. Artedidraco skottsbergii, Lonnberg (Pl. v, fig. 1).
Artedidraco skottsbergii, Linnberg, Swedish South Polar Exped. Fish. p. 48, pl. rv, f. 15
(1905); Vaillant, Expéd. Antarct. Frangaise, Poiss. p. 46 (1906).

Depth of body 43 to 53 in the length, length of head 3 to 3}. Diameter of eye

3 to 32 in the length of head, interorbital width about 14 to 20. Maxillary extending
to below anterior 4 of eye. Barbel simple, smooth or finely papillose, less than $
diameter of eye. Seven gill-rakers on lower part of anterior arch. Dorsal mtv,
24-26. Anal 17-20. Pectoral with 15-17 rays, to # length of head, extending to
vent or origin of anal; pelvies shorter, barely reaching vent in young, not in adult.
Caudal rounded or subtruncate. Body with numerous irregular dark spots; a series
of blotches at base of dorsal fin sometimes continued on sides as bars; vertical fins
with series of spots on the rays, those on posterior part of dorsal and anal and
near upper and lower edges of caudal deep black ; pectorals barred. Vertebrae 15 + 23.

8 “TERRA NOVA” EXPEDITION.

Here described from 18 examples, up to 120 mm. in total length, from the
entrance to McMurdo Sound, 76° 56’ S., 164° 12’ E., 160 fathoms ; 77° 13’ 8., 164° 18’
E., 207 fathoms; and off Granite Harbour, 50 fathoms. The species was known
previously from Graham Land.

15. Artedidraco shackletoni, Waite.
Artedidraco shackletoni, Waite, Brit. Antarctic Exped. Fish. p. 15, pl. m (1911).

Depth of body 4 in the length, length of head 23 to 2%. Diameter of eye
34 to 32 in length of head, interorbital width about 14. Maxillary extending to
below middle of eye. Barbel smooth, tapering, } to 2 (4) leneth of head. Seven
oill-rakers on lower part of anterior arch. Dorsal iv (v) 27-28; middle soft rays
longest. Anal 19 (20). Pectoral with (15) 16-17 rays, + length of head, extending
to vent or origin of anal; pelvics shorter, not reaching vent. Caudal subtruncate.
Uniform or finely speckled ; caudal and pectorals spotted.

Here described from two examples, 132 and 142 mm. in total length, from
the entrance to McMurdo Sound, 77° 13’8., 164° 18’ E., 207 fathoms, and 76° 56’ S.,
164° 12’ E., 160 fathoms. The type, 146 mm. in total length, from off Cape Royds
at a depth of 30 to 80 fathoms, had v, 27 dorsal and 20 anal rays. It had the
barbel a little longer (slightly more than 4 the length of head), the spinous dorsal
Jess, and the soft dorsal somewhat more elevated than in the specimens here
described, but these differences are well within the limit of variation as shown by
other species.

16. Artedidraco loennbergii, Roule (Pl. v, fig. 2).

Artedidraco mirus (part) Lénnberg, Swedish South Polar Exped. Fish. p. 40, pl. 1, f. 4 (1905).
Artedidraco loennbergii, Roule, Deuxieme Expéd. Antarct. Frang. Poiss. p. 13, pl. rv, £. 4
(1913).

Depth of body 5 to 6 in the length, length of head 2% to 34. Diameter of
eye 3 to 3} in length of head; interorbital space very narrow. Maxillary extending
to below anterior + of eye. Barbel simple, or shghtly fringed distally, + to + length
of head. Six or seven gill-rakers on lower part of anterior arch. Dorsal 11-111, 26-28.
Anal 18-21. Soft dorsal and anal usually highest posteriorly. Pectoral with 14 to
16 rays, 3 to 2 length of head, reaching anal in young, but not in adult; pelvies
reaching vent in young, but not in adult. Caudal slightly emarginate. A series of
dark blotches at base of dorsal fin and sometimes at base of anal; a dark lateral band,
made up of irregular spots, from eye to caudal fin; pale bands above and below it
are continued along the upper and lower margins of the caudal fin ; dorsal and middle
of caudal with spots on the rays ; pectorals usually barred.

Previously known from South Georgia and from Graham Land; here described
from twenty-two examples measuring up to 110 mm. in total length from :—Ross
Sea, 74° 25'8.,.179° 3' E., 158 fathoms ; off new land south of the Balleny Islands,

FISHES—REGAN. 9

200 fathoms; McMurdo Sound, entrance, 76° 56’ S., 164° 12’ E., 160 fathoms and
77° 18'8., 164° 18’ E., 207 fathoms, and near Inaccessible Island, 222 to 241 fathoms.
fHistiodraco, gen. nov.

Differs from Dollotdraco in that the upper limb of the post-temporal projects as a
prominent curved ridge, as in Pogonophryne.

17. Histiodraco velifer, Regan (Pl. v, fig. 3).
Dolloidraco velifer, Regan, Ann. Mag. Nat. Hist. (8) x1, 1914, p. 12.

Depth of body 4 in the length, length of head 2?. Diameter of eye 34 in the
length of head, mterorbital width 12. Maxillary extending to below middle of eye, or
beyond. Barbel fringed in its distal half, 2 the length of head. Seven very short gill-
rakers on lower part of anterior arch. Dorsal 111, 26; spinous dorsal short and
high ; soft dorsal elevated anteriorly, the longest rays # to 4% the length of head.
Anal 17. Pectoral 19-rayed, } length of head, extending to origin of anal; pelvics
shorter. Caudal truncate above, rounded below, not more than 2 length of head.
Body marbled or irregularly barred; fins more or less spotted ; caudal crossed by a
dark band.

Two examples, 180 and 190 mm. in total leneth, from the entrance to MeMurdo
Sound, 77° 13’ 8., 164° 18’ E., 207 fathoms.

Pogonophryne, Regan.
Pogonophryne, Regan, Ann. Mag. Nat. Hist. (8) xu, 1914, p. 13.
Differs from /7istiodraco in the wide interorbital region and the strongly depressed
head, as broad as long and much broader than deep.

18. Pogonophryne scotti, Regan (PI. v1, fig. 1).
Pogonophryne scotti, Regan, Ann. Mag. Nat. Hist. (8) x1m, 1914, p. 13.

Depth of body 4 in the length, length of head 25. Diameter of eye 53 in length
of head, interorbital width 43. Maxillary extending to below middle of eye; lower
jaw strongly projecting. Barbel blunt, papillose, shorter than eye. 10 very short
eill-rakers on lower part of anterior arch. Dorsal m1, 25; spines low. Anal 18.
Pectoral 19-rayed, $ length of head; pelvics short, rounded. Caudal rounded. Body
finely spotted and marbled ; fins with series of dark spots on the rays; caudal with a
dark cross-bar.

A single specimen, 290 mm. in total length, from the Ross Sea, 74° 25’ S.,
179° 3' E., 158 fathoms.

A coloured drawing of a fish from the Bransfield Straits reproduced by Loénnberg
(Swedish 8. Polar Exped. Fish. pl. 1, fig. 7) as Artedidraco skottsbergii seems rather
to represent a Pogonophryne, perhaps even the species here described.

This species is named in memory of Captain R. F. Scott, R.N., C.v.o.

10 “TERRA NOVA” EXPEDITION.

BATHYDRACONIDAE.

Gymnodraco, with its depressed naked body, pointed snout, and compressed
uniserial teeth with strong anterior canines, is connected with the genera with the
body subeylindrical and more or less scaly, the snout spatulate, and the teeth
villiform or cardiform, in bands, without canines, by the little known Parachaenichthys.
ixamples of P. georgianus Fisch. recently brought back from South Georgia by Mr. P.
Stammwitz, show that Parachaenichthys has nothing to do with the Chaenichthyidae,
but is a member of the Bathydraconidae, with the depressed form and naked body of
Grymnodraco, but the mouth and teeth of Bathydraco and its allies.

In Parachaenichthys and Gymnodraco the feeble ribs are attached to the long
epipleurals at some distance from the centra,* but in Prionodraco the ribs are stronger,
are inserted directly on the short parapophyses and bear the epipleurals near their
proximal ends.

Prionodraco, Regan.
Prionodraco, Regan, Ann. Mag. Nat. Hist. (8) xi, 1914, p. 13.

Body elongate, compressed, quadrangular, with a series of V-shaped, serrated,
bony plates at each angle; each plate with a backwardly directed spine. Lower series
of plates ending in a group of nearly normal serrated scales behind pectoral fin ;
usually a series of similar scales along middle of side; body otherwise naked. Lateral
line single, incomplete. Snout spatulate; mouth slightly protractile ;. teeth small,
villiform, in bands. Vertebrae 16 + 34.

Prionodraco evansii, Regan (Pl. vu, fig. 1).
Prionodraco evansii, Regan, Ann: Mag. Nat. Hist. (8) xi, 1914, p. 13.

Depth of body 7 to 8 in the length, length of head 3 to 33. Snout as long as or
a little longer than diameter of eye, which is 3 to 34 in length of head ; interorbital
width 15 or more in length of head. Lower jaw a little projecting; maxillary
extending to below anterior margin of eye. Operculum ending in a flat antrorse hook.
He gill-rakers on lower part of anterior arch. Dorsal 34-37. Anal 29-31. Pectoral
; to 4 length of head, extending beyond origin of anal; pelvics not reaching the vent.
Caudal rounded or subtruncate. About 50 plates in upper series; lateral line ending
below anterior part of dorsal fin. Dark spots on body ; usually a well-defined lateral
series of large oblong or squarish spots; a blackish spot on base of anterior part of
dorsal ; dorsal, caudal and pectorals with series of spots on the rays.

al en specimens, measuring up to 132 mm. in total length, from the Ror Sea,
74° 25'S., 179° 3’ E., 158 fathoms; and the entrance to MeMurdo Sound, 76° GIS),
164° ie E., 160 Sia and 77° 13'S., 164° 18’ E., 207 fathoms.

This species is named after Commander E. R. G. R. Evans, R.N., C.B.

* As in the Gobiesocidae, which also have the body depressed.

FISHES—REGAN. 11

CHAENICHTHYIDAE.

The “Terra Nova” collection includes examples of two species of a new genus of
this family, and also new species of Chionodraco and Cryodraco, genera new to the
British Museum collection. The synopsis of the genera given in the “ Scotia” report
may be modified as follows :—

I. Middle rays of pelvic fin longest ; two lateral lines.

A. Lateral line without bony plates.

D. rx-x, 33-40, A. 32-38. no rostral spine ; : ; : Champsocephalus.
D. xm-xv, 28-31. A. 25-27. a rostral spine : : 5 ; 6 Pagetopsis.
B. Lateral line with bony plates ; a rostral spine : : : : 5 . Chaenichthys.

II. Two outer soft rays of pelvic fin longest.
A. Sub- and inter- operculum not spinate ; rostral spine reduced or absent.
Pelvic fins comparatively short, with the rays branched or bifid ; spinous dorsal large, of
seven or eight spines, separated by a short interspace from soft dorsal; two lateral
. Chaenocephalus.

lines 6 2 5 7 : : : :
Pelvic fins long, with the rays simple ; spinous dorsal reduced, of three to five spines,
separated by a long interspace from soft dorsal ; three lateral lines : Cryodraco.

B. Sub- and inter- operculum bearing a pair of spines behind angle of praeoperculum ; rostral spine
well developed ; three lateral lines.

Pelvies 1 5; gill-rakers reduced to a few vestiges near the angles of the arches ( ‘hionodraco.

Pelvics 14; gill-rakers developed as dentigerous knobs or patches . . Chaenodraco.

20. Pagetopsis macropterus, Bouleng.

Three examples from McMurdo Sound, 100 to 200 fathoms.

21. Cryodraco atkinsoni, Regan (Pl. v1, fig. 2).

Cryodraco atkinsoni, Regan, Ann. Mag. Nat. Hist. (8), x11, 1914, p. 13.

rc

Depth of body 73 in the length, length of head 33. Diameter of eye 5 in length
of head, interorbital width 42. Snout 1 the length of head. Maxillary nearly reaching
middle of eye. Rostral spine and. symphysial tubercle of mandibles vestigial. Dorsal
1H, 42 ; spinous dorsal above base of pectoral, its spines connected by membrane, the
first and second subequal, 4 the length of head. Anal 46; origin a little in advance of
that of dorsal, nearer to end of snout than to base of caudal. Pectoral nearly 3 length
of head, extending to seventh ray of anal ; pelvies 14 as long as head, extending to
middle of dorsal fin. Caudal slightly emareinate. Dark spots on head and cross-bars
on body ; spinous dorsal blackish ; pelvics dusky.

A single specimen, 292 mm. in total length, from the Ross Sea, 74° 25’ S., 179° 3’ E.,
158 fathoms.

This species is named after Surg. E. L. Atkinson, R.N. ; it differs from C. antareticus,
Dollo, in the smaller eye, shorter pelvic fins, ete. C. pappenheimi, Regan, has the
pelvic fins still shorter, and in its large head and 5-rayed dorsal fin seems to approach

Chaenocephalus.
Cc 2

12 “TERRA NOVA” EXPEDITION.

Chionodraco, Loéunberg.

Body naked, elongate ; three lateral lines without bony plates. Eye somewhat
behind middle of head; supraorbital ridges crenulated ; a rostral spme; teeth bi- or
tri- serial ; gill-rakers absent except for a few vestiges near the angles of the arches ;
sub- and inter- operculum bearing a pair of spines just behind angle of praeoperculum.
Spinous dorsal well-developed, of six or seven spines ; pelvics comparatively short, of a
spine and five branched or bifid rays, the two outer the longest and enveloped in thick
skin. Skeleton essentially similar to that of Champsocephalus. Vertebrae 64.

22. Chionodraco kathleenae, Regan (Pl. vt).
Chionodraco kathleenae, Regan, Ann. Mag. Nat. Hist. (8) x1, 1914, p. 13.

Depth of body about 5 in the length, length of head 2? to 3. Diameter of eye
5 to 6 in length of head, interorbital width 33 to 4. Snout a little less than 3} length
of head. Rostral spine erect or retrorse. Maxillary extending to below anterior part
or middle of eye. Dorsal vi-vi, 38-42, the two fins separated by an interspace.
Anal 34-38. Pectoral } length of head or a little more, extending to above vent or
anterior rays of anal; pelvics 2 to 3 length of head, extending to origin of anal or
beyond. Head and body with dark spots and bars ; spinous dorsal blackish.

Five specimens, 250 to 500 mm. in total length, from the Ross Sea, 74° 25'5.,
179° 3' E., 158 fathoms, and McMurdo Sound, 100-200 fathoms. Also an example in
bad condition from off Granite Harbour, McMurdo Sound, 50 fathoms.

C. hamatus, Lénnberg, known from a specimen of 330 mm., has a larger eye (43 in
length of head, including opercular flap) and shorter pelvic fins, not reaching the vent.

This species is named in honour of Lady Scott.

Chaenodraco, Regan. :
Chaenodraco, Regan, Ann. Mag. Nat. Hist. (8) xm, 1914, p. 13.
This genus differs from Chionodraco in having the supraorbital mdges not
crenulated, the gill-rakers developed as dentigerous prominences, and the pelvic fins
each formed of a spine and only four rays.

23. Chaenodraco wilsoni, Regan (Pl. rx, fig. 1).
Chaenodraco wilsoni, Regan, Ann. Mag. Nat.. Hist. (8) xim, 1914, p. 14.

Depth of body 6 in the length, length of head 34. Snout 22, diameter of eye 4,

in length.of head. Maxillary extending to below anterior 4
of eye. Rostral spine antrorse. Dorsal vit, 39, the two fins continuous at the
base. Anal 33. Pectoral 2 length of head, extending to third or fourth ray of
anal; pelvics 2 leneth of head, extending to seventh ray of anal; rays bifid) A
large dark spot on the spinous dorsal.

A single specimen, 250 mm. in total length, from McMurdo Sound, 100 to 200
fathoms.

This species is named in memory of Dr. E. A. Wilson.

2
3

interorbital width 32

FISHES—REGAN. 13

24. Chaenodraco fasciatus, Regan (PI. 1x, fig. 2).
Chaenodraco fasciatus, Regan, Ann, Mag. Nat. Hist. (8) xi, 1914, p. 14.

Depth of body 7 in the length, length of head 23. Snout 21 in length of
head, diameter of eye 4%, interorbital width 44. Maxillary extending to below
anterior + of eye. Rostral spine antrorse. Dorsal vu, 40, the two fins separate.
Anal 34. Pectoral a little more than # length of head, extending to seventh anal
ray ; pelvics as long as head, reaching fourteenth ray of anal. Body with five
blackish cross-bars.

A single specimen, 92 mm. in total length, from McMurdo Sound, 77° 13/ §
164° 18’ ., 207 fathoms.

The pelvic fin-rays are simple, doubtless a juvenile character. As the snout is
proportionately longer and the eye smaller than in the much larger specimen of C.
wilson, this cannot be the young of that species. The much greater distance
between the last ray of the spinous dorsal and the first ray of the soft fin is also
an important character.

*)

CYCLOPTERIDAE.
25. Paraliparis antarcticus, Regan (PI. 1, fig. 1).

to}

Paraliparis antarcticus, Regan, Ann. Mag. Nat. Hist. (8) xim, 1914, p. 11.

Depth of body equal to or greater than length of head, which is 4 to 43 in
the length of the fish. Snout obtuse, projecting a little beyond mouth, longer than
diameter of eye, which is about 5 in the length of head. Maxillary extending to
below posterior edge of eye; teeth villiform, in bands. Lower end of gill-opening
opposite middle of base of pectoral. Dorsal 60; origin above extremity of operculum.
Anal 55; origin below about ninth ray of dorsal. Pectoral of an upper portion,
19-rayed, 2 to # the length of head, extending a little beyond the origin of anal,
connected by a low membrane including three or four short, spaced rays with a lower
four- or five- rayed portion, as long as the upper and reaching the vent. Skin loose,
transparent.

Three examples, 90 to 140 mm. in total length, from off new land south of the
Balleny Islands, 200 fathoms.

2.—FISHES FROM NEW ZEALAND.

There are examples of thirty-four species, ten of which have been described as new
to science (Ann. Mag. Nat. Hist. (8) xir, 1914, pp. 14-16).

The fishes are from Cape North, unless otherwise stated.

ScyLIORHINIDAE.
1. Scyliorhinus laticeps, Duméril.

14 “TERRA NOVA” EXPEDITION.

CARCHARIIDAE.
2. Galeus australis, Macleay.
SQUALIDAE.
3. Squalus megalops, Waite.
STOMIATIDAE.

4, Idiacanthus niger, Regan (Pl. x, fig. 2).
Idiacanthus niger, Regan, Ann. Mag. Nat. Hist. (8) x1u, 1914, p. 14.

Depth of body 22 in the length, length of head 133. Snout longer than diameter
of eye, which is 8} in length of head ; interorbital width 5. Barbel twice as long
as head. Dorsal 59 ; origin above posterior part of pelvic fins. Anal 38; origin a
little nearer to base of caudal than to that of pelvics, which is equidistant from
head and origin of anal. Photophores in ventral series about 37 from isthmus to
pelvies, 21 from pelvies to anal. Blackish.

A single specimen, 400 mm. in total length, from the stomach of a ‘“ Groper.”

Brauer (Valdivia Tiefsee Fische, p. 60) has given a synopsis of the four species of
this genus hitherto described, in all of which the origin of the dorsal fin is above or in
advance of the base of the pelvies. J. niger is nearest to J. ferov, Giinth., which has
a shorter barbel, the dorsal origin above the base of the pelvics, and a longer anal fin
commencing at a point nearer to the insertion of the pelvics than to the caudal fin.

The following measurements, in millimetres, are taken from the types of J. niger

and J. feroa :—
I. niger. I. ferox.

Lower jaw. : 5 5 : ‘ : ; : : 28 14
Barbel . ‘ : ; ; ; : ; ; : 58 24
Length to base of caudal . F . 5 : : : : 380 192
> » Origin of dorsal : ; : , : ie VN 176 66
3 => Wase Of pelvis - : E ; : : ; : 150 66
,, from base of pelvics to origin of anal : : : ; 120 58
Y ,, origin of anal to base of caudal : J F : 110 68
GONORHYNCHIDAE.

5. Gonorhynchus gonorhynchus, Linn.
A small example from a rock-pool at Whangaroa.

MURAENIDAE.
6. Muraena nubila, Richards

HEMIRHAMPHIDAE.

7. Hemirhamphus intermedius, Cant.
Lyttelton Harbour.

FISHES—REGAN. 15

i)

EXXOCOETIDAR.

8. Hxocoetus spilonotopterus, Bleek.

MaAcRORHAMPHOSIDAR,

Notopogon, Regan.

Notopogon, Regan, Ann: Mag. Nat. Hist. (8) xm, 1914, p. 14.

This genus is distinguished from Macrorhamphosus by the deeper form, the dorsal
fins continuous at the base, the strong second spine followed by five nearly
equidistant and gradually decreasing in length backwards, and by the development of
a patch of. bristles on the nape in the adult fish. From Centriscops it differs in the
last character and in having only three well-developed plates in each dorso-lateral series.

9. Notopogon lilliei, Regan (PI. xu, fig. 4).
Centriscops humerosus (non Richards) McCulloch, “ Endeavour,” Fish. p. 24, fig. 5, and pl. 1x
(1911).
Notopogon lillie’, Regan, Ann. Mag. Nat. Hist. (8) xt, 1914, pp. 14, 20.

Depth of body 22 in length, leneth of head 2. Distance from base of dorsal spine
to vent 12 in that from head to caudal fin. Diameter of eye 33 in length of snout,
a little more than postorbital part of head, twice depth of cheek. Interorbital space
convex, with blunt median ridge ; width ? diameter of eye. Back slightly, belly more
strongly convex. Two series of bony plates on each side of back ; only three large plates
in each series ; ventral plates keeled in front of, spimate behind pelvic fins. Dorsal v1,
14, the two fins continuous ; second spine strong, serrated, inserted above middle of
anal fin, its length a little more than $ distance from opereulum to caudal fin. Anal 19.
Pectoral as long as head without snout. Caudal truncate.

A single specimen, 125 mm. in total length, from New Zealand, caught by
fishermen ; the species is also known from the south coast of Australia ; it is named

after Mr. D. G. Lillie.

10. Notopogon wenosoma, Regan (Pl. x1, fig. 5).
Notopogon xenosoma, Regan, Ann. Mag. Nat. Hist. (8) xm, 1914, pp. 14, 20.

Body elevated, the distance from base of dorsal spine to vent a little greater than
that from head to caudal fin. Head 2} in the length. Diameter of eye a little longer
than postorbital length of head, ? length of snout, twice depth of cheek. Interorbital
space flat, its width 2 diameter of eye. Upper profile with a moderate hump ; belly
convex. Only three large plates in each dorso-lateral series ; last of dorsal series bearing
a spine ; ventral plates spinate ; a pair of spines at posterior end of lower jaw ; a patch of
scales behind dorsal hump modified into short bristles. Dorsal vi, 15, the two fins sub-
continuous ; second spine rather slender, serrated, inserted above caudal peduncle, its
leneth # distance from operculum to caudal. Anal 17. Pectoral as long as head

without snout. Caudal truncate.

16 “TERRA NOVA” EXPEDITION.

A single specimen, 80 mm. in total length, from Cape North; 70 fathoms.

Compared with J. schoteli this species is still more aberrant in form, with the
insertion of the dorsal spine higher and further back, behind instead of in front of the
soft dorsal fin. It agrees more closely with N. fernandezianus, which has heen figured
by Delfin on the cover of his Catalogue of Chilean Fishes. The type of NV. fernandezianus
measures 167 mm., and it may be owing to its larger size that the eye is smaller (53 in
length of head), the snout longer (12 instead of 14 rest of head) than in NV. wenosoma ;
also the greater length of the dorsal bristles is doubtless due to age. The difference in
structure and position of the dorsal spine may be more important, and in all probability
direct comparison would reveal other differences between the two forms.

SYNGNATHIDAE.
L1. Solenognathus spinosissimus, Giinth.
12. Stigmatophora macropterygia, Dumeéril.
D’Urville Island.
TRACHICHTHYIDAE.

13. Paratrachichthys trailli, Hutton.
Elmsley Bay, South Island.

GEMPYLIDAE,
14. Thyrsites atun, Euphyras.
TRICHIURIDAE.
15. Lepidopus caudatus, Kuphras.
SERRANIDAE.

16. Caprodon longimanus, Giinth.

Serranops, Regan.
Serranops, Regan, Ann. Mag. Nat. Hist. (8) xm, 1914, p. 15.

Related to Plectranthias, Bleek, but with the serrations of the lower praeopercular
limb weak, not antrorse, and the scales spinulose. Distinguished externally from
Lepidoperca by the larger mouth, naked maxillary, and almost naked spinous dorsal
fin. Skeleton as in Plectranthias, the frontals smooth and convex behind the orbits
and narrow between them, with the mucous canals in contact and the supraorbital

flanges little developed. Vertebrae 10 + 16.

17. Serranops maculicauda, Regan (PI. x1, fig. 3).
Serranops maculicauda, Regan, Ann. Mag. Nat. Hist. (8) xm, 1914, p. 15.
Depth of body 22 to 3 in the length, length of head 23 to 23. Snout shorter
than diameter of eye, which is 3 in length of head ; interorbital width 6. Upper

*

FISHES—REGAN. 17

surface and sides of head scaly; praeorbital and maxillary naked; lower jaw scaly.
Maxillary extending to below middle or posterior part of eye. 16 gill-rakers on lower
part of anterior arch. 33 or 34 scales in the lateral line. Dorsal x, 15; middle
spines longest, 4 to 3 length of head ; second soft ray more or less produced. Anal
1 7; second spine longest, longer than highest spmes of dorsal fin. Pectoral 3 length
of head. Caudal truncate ; ray at upper angle sometimes produced. A large dark
spot on each side of caudal peduncle, usually another below spinous dorsal.

Eight specimens, 60 to 100 mm. in total length, from seven miles E. of Cape
North ; depth 70 fathoms ; bottom sand.

Lepidoperca, Regan.
Lepidoperca, Regan, Ann. Mag. Nat. Hist. (8) x11, 1914, p. 15.

External characters of Caesioperca, except that the interorbital region is flat
instead of convex, the caudal fin truncate instead of emarginate, the scales larger and
the dorsal and anal rays shghtly less numerous. Skeleton differimg from that of
Caesioperca in the absence of a transverse ridge in front of the occipital crest and in
having the mucous canals of the frontals separated by ‘a narrow groove, whereas in

Caesioperca they border a fossa which broadens out anteriorly. Vertebrae 10 + 16.

18. Lepidoperca inornata, Regan (PI. x1, fig. 4).
Lepidoperca inornata, Regan, Ann. Mag. Nat. Hist. (8) xm, 1914, p. 15.

Depth of body 24 in the length, length of head 23. Diameter of eye 25 in the
length of head, interorbital width 4. Interorbital space flat; praeorbital scaly ;
maxillary not quite reaching vertical from middle of eye; 25 gill-rakers on lower
part of anterior arch. Dorsal x, 16; fourth spine longest, nearly 3 length of head,
13 as long as last spine. Anal 1m 8; second spine a little longer than longest of
dorsal. Pectoral slightly shorter than head; ninth ray from above (eighth from below)
longest. Caudal truncate. 38 scales in a lateral longitudinal series, 41 in the lateral
line. Traces of reddish longitudinal stripes.

A specimen of 135 mm. from near Cape North, 70 fathoms, bottom sand ; also a
second much smaller example, 60 mm. in total length.

This species is closely related to the recently described L. coatsi/, Regan (Trans.
R. Soc. Edinburgh, x1x, 1913, p. 237, pl. vi, f 1), from Gough Island, but is
distinguished by the deeper body, fewer scales, scaly praeorbital, shorter maxillary,
higher last dorsal spine and immaculate dorsal fin.

PLESIOPIDAE.
19. Acanthoclinus littoreus, Forst.

CARANGIDAE.
20. Seriola lalandii, Cuy. and Val.

18 “TERRA NOVA” EXPEDITION.

CEPOLIDAE.
21. Cepola aotea, Waite.
CHILODACTYLIDAE.

22. Chilodactylus macropterus, Forst.

PARAPERCIDIDAE.

23. Parapercis gilliesii, Hutton.
HEMEROCOETIDAE.

24. Ilemerocoetes pauciradiatus, Regan (Pl. xm, fig. 1).
Hemerocoetes pauciradiatus, Regan, Ann. Mag. Nat. Hist. (8) xm, 1914, p. 15.

Depth of body about 8 in the length, length of head 33 to 32, distance from
tip of snout to origin of dorsal fin 34 to 32, to origin of anal about 23. Snout as
long as or shorter than diameter of eye, which is 3} to 32 in the length of head ;
interorbital space narrow; maxillary extending to below middle of eye; 13 very
short gill-rakers on lower part of anterior arch. About 45 scales in a longitudinal
series. Dorsal 36; origin above anterior } of pectoral. Anal 32. Pectoral 2 length
of head ; pelvics reaching vent. Caudal subtruneate. A lateral series of dark spots.

Two specimens, 50 and 62 mm. in total length, from seven miles east of Cape
North ; depth 70 fathoms ; bottom sand.

25. Hemerocoetes macrophthalmus, Regan (PI. xt, fig. 2).
Hemerocoetes macrophthalmus, Regan, Ann. Mag. Nat. Hist. (8) x1, 1914, p. 15.

Depth of body about 10 in the length, length of head 3? to 4, distance from
tip of snout to origin of dorsal fin 34 to 34, to origin of anal 22 to 23. Snout shorter
than diameter of eye, which is 22 to 3 in length of head ; interorbital space very
narrow ; maxillary extending to below middle of eye; 15 very short gill-rakers on
lower part of anterior arch. About 47 scales in a longitudinal series. Dorsal 39 ;
origin above anterior 4 of pectoral. Anal 36. Pectoral 3 length of head; pelvies
reaching vent. Caudal truncate; upper rays sometimes produced. A series of dark
spots along middle of side, another series on back.

Two specimens, 90 and 120 mm. in total length, from seven miles east of Cape
North ; depth 70 fathoms ; bottom sand.

I propose the new name J//emerocoetes waitei for the species described and figured
by Waite (Rec. Canterbury Mus. 1, 1911, p. 245, pl. trv, f. 1) as Hemerocoetes
acanthorhynchus. The dorsal has 42 rays and the anal 40, and these fins originate
somewhat further forward than in H. macrophthalmus, which it closely resembles in
other characters.

The true J. acanthorhynchus, Forster, is the species described and figured by
Waite (t.c., p. 247, pl. xrv, f. 2) as ZZ. microps. The original description, that the

FISHES—REGAN. 19

eye is less than 4 of the length of the head in a specimen of 200 mm., leaves no
doubt on this point. In examples of this species I count 40 to 42 dorsal and 38
to 40 anal rays.

STROMATEIDAR.
26. Centrolophus maoricus, Ogilb.

Depth of body 34 in the length ; length of head 34 to 34. Snout from a little
shorter to a little longer than diameter of eye, which is 34 to 41 in the leneth of head
and less than interorbital width. Maxillary extending to below anterior } af 1 of eye.
14 oill-rakers on lower part of anterior arch. Dorsal 43 ; origin above base of pectoral.
Anal 27. Pectoral 3 length of head; pelvies a little shorter. Caudal emarginate.
Caudal peduncle nearly twice as long as deep. 180 scales in a longitudinal series ;
lateral lime curved anteriorly, becoming straight above origin of anal. Purplish ;
longitudinal series of oblong pale spots more or less distinct on sides of body ; two
broad dusky bands across the body, one in front of and one above the anal fin.

Three Kings Islands, north of New Zealand ; surface.

Here described from two specimens, 150 and 280 mm. in length, which have the
body a little deeper and the head a httle larger than in young examples of the Atlantic
C. niger, Gmel. ; also the fin-rays are somewhat more numerous and the origin of the
dorsal fin is further forward ; it is probable that these specimens pertain to CL maoricus,
Ogilby (Rec. Austral. Mus. 11, 1893, p- 64), described from a larger fish.

Since my revision of the genus Centrolophus (Ann. Mag. Nat. Hist. (7) x, 1902,
p- 194) Waite* has described a new species, (. huttoni, from New Zealand, well
distinguished by the large number of fin-rays. C. britannicus has been rediscovered,f
and I have examined the type of the Californian [cichthys lockingtoni, Jord. and Gilb.,
in the Smithsonian Institution ; this is a Centrolophus, very similar to CL niger.

27. Cubiceps caeruleus, Regan.

Cubiceps caeruleus, Regan, Ann. Mag. Nat. Hist. (8) x11, 1914, p. 15.

Depth of body nearly equal to length of head, 3? to 3% im length of fish. Snout
as long as or a little shorter than diameter of eye, which is 35 to 33 in length of head
and a little less than interorbital width. Maxillary not quite reaching vertical from
anterior margin of eye; praeorbital narrowed posteriorly, not completely concealing
maxillary. 14 or 15 gill-rakers on lower part of anterior arch. Dorsal x1, 123. Anal
mi 21. Pectoral as long as head, extending to origin of anal. Caudal forked.
Probably not more than 50 scales in a longitudinal series. Bluish.

Three Kings Islands.

Two specimens, 100 and 110 mm. in length, from the stomach of a Seriolella,

* Trans. N. Z. Inst. xm, 1910, p. 388.
+ Pellegrin, Bull. soc. zool., xxxvu, 1912, p. 20,

D 2

20 “TERRA NOVA” EXPEDITION.

In the Atlantic C. gracilis, Lowe, the body is less deep, the snout shorter, and the
eye larger; also the maxillary is completely hidden, and the pectoral fin extends
beyond the origin of the anal. Owing to the imperfect condition of the types of
C. caeruleus the number of scales cannot be certainly stated, but they seem to be
larger than in C. gracilis, which has 58 to 66 in a longitudinal series.

Since my revision of the genus Cubiceps (Ann. Mag. Nat. Hist. (7) x, 1902,
p- 122), when four species were recognised, Jordan and Snyder have described a
species from Honolulu under the name Ariomma lurida. This is figured by Jordan
and Evermann (Bull. U. 8S. Fish. Comm. xxim, 1905 p. 217, pl. xxxvm), and is
evidently closely related to Cubiceps pauciradiatus, Giinth. and C. brevimanus, Klunz.

BROTULIDAE.
Pyramodon, Radcliffe.
Pyramodon, Radclifie, Proc. U.S. Nat. Mus. xniv, 1913, p. 175.
Cynophidium, Regan, Ann. Mag. Nat. Hist. (8) x1, 1914, p. 16.

Head and body naked, compressed ; tail tapering ; no lateral lime. Head unarmed,
without large muciferous channels, smooth and convex above ; eyes well-developed ;
no barbels. Mouth large, protractile, terminal, with the lower jaw included ; teeth
subconical, in a narrow band in praemaxillaries, uniserial and unequal in lower jaw and
on palatines; upper jaw with a rather wide toothless interspace between a pair of
canines, which are outside the mouth when it is closed ; lower jaw with a pair of
strong recurved anterior canines ; vomer with a very strong curved canine followed by
a few smaller teeth. Gill-membranes separate, free; 7 branchiostegals; 4 gills;
pseudobranchiae very small. Vent and origins of dorsal and anal fins a short distance
behind head ; vertical fins confluent at end of tail; no distinct caudal fin ; pectorals
well-developed ; pelvics jugular, a pair of simple filaments.

This genus differs from Snyderidia, Gilbert, 1905 (Bull. U.S. Fish. Comm. 1903,
p- 654), im the presence of pelvic fins. In many respects these two genera seem to
connect the Brotulidae with the Fierasferidae, but I have ascertained that Pyramodon
agrees with the Brotulidae in the structure of the upper surface of the skull, the
supraoccipital separating the rather small parietals.

28. Pyramodon punctatus, Regan (PI. xu, fig. 3).
Cynophidium punctatum, Regan, Ann. Mag. Nat. Hist. (8) x1, 1914, p. 16.

Depth of body nearly equal to length of head, which is 6 in the length of the
fish ; distance from end of snout to origin of dorsal fin 47. Snout, diameter of eye
and interorbital width subequal, about } the length of head. Maxillary extending
beyond eye. Three gill-rakers and several rudiments on lower part of anterior arch.
Origin of dorsal fin shghtly in advance of vent ; pectoral * length of head ; pelvics
1 length of head, or $ distance from their base to origin of anal. Olivaceous, powdered
with little dark spots.

FISHES—REGAN. 21

A single specimen, 185 mm. in total length, from seven miles east of Cape North ;
depth 70 fathoms; bottom sand.

The other known species, ?. ventralis, Radcliffe, was described from a specimen
taken near Doworra Island, Dutch E. Indies, at a depth of 205 fathoms.

SCORPAENIDAE.

29. Sebastes percoides, Richards.
30. Scorpaena cardinalis, Richards.

TRIGLIDAE.

31. Trigla kumu, Less. and Garn.
BorTHIDAE.

32. Pseudorhombus boops, Hect.

33. Arnoglossus mongonuiensis, Regan (PI. x1, fig. 2).

Arnoglossus mongonuiensis, Regan, Ann. Mag. Nat. Hist. (8) x1, 1914, p. 16.

“4

eye 3 to 34 in length of head; lower eye a little in advance of upper ; interorbital
space narrow. Maxillary extending to below anterior edge of eye ; males with a blunt
spine on snout and a prominent knob below symphysis of lower jaw. Six or seven
gill-rakers on lower part of anterior arch. About 70 scales in a longitudinal series.
Dorsal 86-90; second to fifth rays prolonged in males. Anal 72-76. Pectoral of
eyed side 2 to 2 (f) or 2 (%) of the length of head, of blind side about 3 (2%) or
less (2). Caudal rounded or obtusely pointed. Olivaceous, with darker spots.

Four examples, 75 to 85 mm. in total length, from off Cape North; depth

14 to 30 fathoms: bottom sandy.

Depth of body 23 to 2% in the length, length of head 4 to 43. Diameter of

GZOBIESOCIDAE.

34. Crepidogaster simus, Hutton.

3.—FISHES FROM BRAZIL.
These were taken off Cape Frio, in 22° 56’ 8., 41° 34’ W., at a depth of
40 fathoms.
RAIIDAE.
1. Psammobatis cirrifer, Regan (Pl. xt).
Malacorhina cirrifer, Regan, Ann. Mag. Nat. Hist. (8) xm, 1914, p. 16.
Dise about as broad as long (to posterior end of pelvic fins), terminating in a
barbel borne by a small triangular prominence ; anterior margins of disc shghtly
e us .
undulated, outer angles rounded. Vent nearer to tip of snout than to end of tail.

bo
bo

“TERRA NOVA” EXPEDITION.

Snout twice as long as longitudinal diameter of eye with spiracle ; interorbital width
equal to transverse diameter of eye. Internasal width less than } praeoral length of
snout, less than distance of either nostral from edge of disc. Teeth obtuse, close-set ;
about 40 series in upper jaw, which has a median emargination. Disc smooth below,
above with scattered spines with radiating bases, set more closely near the anterior
and posterior margins ; a series of spines at inner margin of each orbit; a triangular
patch on scapulary region ; tail with three series posteriorly, five anteriorly, continued
forward on the disc as two only ; two spmes between the well-separated dorsal fins.
Brownish, with scattered darker and paler spots.

A single specimen, a young female, 220 mm. in total leneth. In the spimation
and in the separation of the dorsal fins this species is nearer to the Chilean one
deseribed by Garman from an adult male as Malacorhina mirus than to Psammobatis
rudis, Giinth. Malacorhina seems to be a synonym of Psammobatis, and the adult male
of Ps. cirrifer may have the anterior margins of the dise notched as in Ps. mirus ;
similar differences in form due to sex and age are known in feaia fyllae, Liitken.

MURAENIDAE.
2. Muraena ocellata, Agass.
CONGRIDAE.

3. Congromuraena balearica, Delaroche.

SYNODONTIDAE.

4. Trachinocephalus myops, Forst.

SYNGNATHIDAE. =

5. Hippocampus punctulatus, Guichen.

SERRANIDAE.
6. Serranus auriga, Cuv. and Val.
TRIGLIDAE,

7. Prionotus brachychir, Regan (Pl. xt, fig. 1).
Prionotus brachychir, Regan, Ann, Mag. Nat. Hist. (8) xu, 1914, p, 16.

Depth of body about 335 in the length, length of head (without opercular spine)
23 to 3. Diameter of eye nearly equal to length of snout or postorbital part of head ;
interobital space a little concave, its width ? diameter of eye. Maxillary extending to
below anterior } of eye. Opercular and praeopercular spines strong ; no other spines
on head; a short cleithral spme; 10 gill-rakers on lower part of anterior arch. 50 to
60 scales in a lateral series, 45 to 50 in lateral line; chest naked. Dorsal vui—xt,

FISHES—REGAN. 2é

10-12 ; second or third spine longest, nearly 4 length of head. Anal 10-12.
Pectoral shorter than head, extending to origin of anal or slightly beyond. Caudal
emarginate. A blackish spot near edge of spinous dorsal, between fourth and sixth
spines ; soft dorsal with series of small dark spots; caudal with three dark cross-bars ;
pectoral blackish.

Several specimens, 70 to 80 mm. in total length.

8. Prionotus tribulus, Cuv.

OPHIDIIDAR.

9, Ophidium brevibarbe, Cuy.

30THIDAR.

10. Etropus microstomus, Gall.
11. Paralichthys oblonqus, Mitch.

12. Nystreurys brasiliensis, Regan (Pl. x, fig. 1).
Xystreurys brasiliensis, Regan, Ann. Mag. Nat. Hist. (8) x1, 1914, p. 17.

Depth of body 24 in the length, length of head 4. Snout 2% diameter of eye,
which is 3 in length of head. Eyes separated by a narrow ridge. Maxillary
extending a little beyond anterior + of eye; lower jaw $ length of head. Gill-rakers
moderately elongate, 10 on lower part of anterior arch. 85 scales in a longitudinal
series. Dorsal 83. Anal 66. Left pectoral nearly as long as head, right scarcely more
than $ length of head. Caudal pointed. A large double ocellus at end of arch of
lateral line ; a small ocellus between it and dorsal fin ; a large dark spot posteriorly on
lateral line.

A single specimen, 170 mm. in total length.

The only other known species of this genus is YX. //olepis, Jord. and Gilb., from the
coast of southern California.

CYNOGLOSSIDAE.

13. Symphurus plagiusa, Linn.
BATRACHOIDIDAE.
14. Porichthys porosissimus, Cuv. and Val.

LOPHIIDAE.

15. Lophius piscatorius, Linn.

“TERRA NOVA” EXPEDITION.

Il—GENERAL PART.

THE DISTRIBUTION OF ANTARCTIC AND
SUBANTARCTIC FISHES.

A. COAST FISHES.

Ir is convenient to distinguish between coast fishes and oceanic fishes, Including im the
former not only the littoral forms but also fishes that may occur at no great distance

oe

Tristan da Cunhe
-Gough A

AFRICA

|

| Bouvet / «Mana Is

~ Crozet Is

SCeeega 5 Sandwich bs

~ S.Orkney Is
(O" pakland ES "y

Sere HSS

VAerguelen |
2 St. Rul

Anenoete |
‘ SEA >

be ,
Heara i.

90° w

VICTORIA

ROSS SEA ALAND
p fe

x Macguane &.

Gaptel f

s. zi ,
2 Auckland |.
Antinodesk:

2 Stewart /
}
Cheathan? ‘As. dhe
ee ZEALAND

TASMANIA

Oi

Fic. 1—Map showing the principal localities mentioned in the text.

from the coasts in water down to two or three hundred fathoms deep, and are not

pelagic or bathypelagie.

FISHES—REGAN. 25

For the first time our knowledge of the fishes of the coasts of the Antarctic

continent is sufficiently advanced to make it worth while to attempt to delimit an
antarctic zone, and to divide it into districts.
For the purposes of reference it may be stated at once that the conclusions I have

>

& RQ ane

~*~,
~

Rog
ryt Ta

“Ne

Fic. 2.—Map showing the mean annual surface isotherms of 6° C, — ——— — and 12° C.
_—-— -—.- , as calculated by Dr. Schott, respectively approximating to the northern boundaries

» of the Antarctic and Subantarctic Zones; the extreme limit of pack-ice ~ ~ ~ bounding the
Glacial District (G); and the Kerguelen (K), Magellan (M) and Antipodes (A) Districts.

reached are that south of the tropical zone the distribution of coast fishes is best
illustrated by the following classification :—
1. South Temperate Zone, with seven districts: Chile, Argentina, Tristan

da Cunha, Cape, St. Paul, Australia, and New Zealand.
E

26 “TERRA NOVA” EXPEDITION.

2. Subantarctic Zone, with two districts: Magellan and Antipodes.

3. Antarctic Zone, with two districts: Glacial and Kerguelen.

About 90 per cent. of the species of fishes known from the coasts of the Antarctic
continent belong to the division Nototheniiformes; therefore it is evident that the
delimitation of an Antarctic region or zone, so far as the fishes are concerned, must be
based on the distribution of this group. Recognising this when working out the
Antarctic fishes of the “Scotia” expedition about two years ago, I took the opportunity
of monographing the Nototheniiformes. Since then Professor Roule’s report on the
fishes of the second Charcot Expedition has appeared, adding to our knowledge of the
species and their distribution. Of the 25 Antarctic species of the “Terra Nova”
collection 22 are Nototheniiformes, half of these bemg new species, and several others
new records for Victoria Land. This being so, the list of the species and their
distribution given below, whilst agreeing in the main with my monograph, includes
several additions and alterations. The South Temperate localities are given first, the
subantaretic next, and the strictly Antarctic last. ‘ Magellan’
northwards to Chiloe and Cape Blanco, and “ Antipodes” includes the extreme south

?

includes the coast

of New Zealand and the neighbouring subantarctic islands.

DISTRIBUTION OF THE NOTOTHENIIFORM FISHES.

S. Teuperare |SUBAST
Wows ARCTIC ANTARCTIC ZONE.
ag ZONE.
| 4 = | ale -| a] .| |
| 5 Sig a) S| al si sole
S| |slz a) |b 2) 8] 8] 8).
saa gga elale ac eMaele
af 2) g/221$|2a)e1\elalglaleals
‘a| |-8| 3/5i| 3/3 S13] S| S| el el elelel sl
S| 4| aa) 0/44) S| Rl a|s| a) alo EIE|S|
BovicHTHYIDAE.
Pseudaphritis, Casteln. |
uryillii, Cuv. & Val. we |ocdiec| se] cae} eal eal saieaies|es| sles! | oo) ekunesh=walberiiabe
Cottoperca, Steind.
gobio, Giinth. as et (ead (Sed fess os lois, Ps fos eto Sel ie! es Peel fos
macrophthalma, Regan |..||..|<-)s-)--| oe) .- (sel leet osleciecles| eat esis}
macrocephala, Roule . & 21 yal sol oa) 0a Sap eh eats olla] ee acaba
Bovichthys, Cuv. & Val. |
variepatus, Richards. . |. .))..|<<|0<)4]-e| Ae] eal -~| elias wale] oes] oo fos
angustifrons, Regan Saleeltta eens
diacanthus, Carmich. .|..|..|-#)..|--t.|.. Bey (eae, aed be Rel fos Weal fae
chilensis, Regan . . |+|..|-.|..|--|..]..]..]..|..|..|--]--|--].-]--|-.] B. patagontcus may be the same as
patagonicus;-Berg = = |i. |edacl cate ac eae alae ealea sel Satee eokee B. chilensis, which may range
veneris, Sauvage . .|..|..|.-|#/--|..]..|..]..|.-|..| ]--[..]. |--|..| ~Southward to Magellan.
decipiens, Regan Fey FS | Ses AS On We tel Io fed Fo feel eal fat fae) fe
psychrolutes, Gunth. —..|...|-<|<>|--}s<}. [oe] < <pecl2<|--] <<]

roseopictus, Hutton

FISHES—REGAN. 27

S. TEMPERATE

ARCTIC AN! NE
Wires) | TARCTIC ZONE.

Marion Island.
Wilhelm Land.

| Coats Land.

Argentina.
Tristan da Cunha.
St. Paul.
S.E. Australia.
New Zealand S

; Antipodes.
South Orkneys.
Graham Land.
Victoria Land.

Chile.

| Falklands.
| Kerguelen.
| South Georgia.

| Magel

|
7
|
|

NovroTHENIDAE. | |

Trematomus, Bouleng.
MewMesiy Boulenge aileel-e|--|-«|\2-] 6. leo |2 |e. |eol orl. of. «| ele]
nicolai, Bouleng. > loa}[de{lool lool lor larl leon el lex tol eel (el eal aur
borchgrevinkii, Bouleng.|..|..|.-|-.|--|..|.. ata [Jel |+
brachysoma, Pappenh. |..|..|..|..|.-|--|--].-|..|--|--|.-|--]--|#
bernacchimeBoulenga selves ince |e > =e le -|locll- |e. [ele oe
womans, batts » 0 ||ee\/oolleallealaelleelleallee|| eee Sine
Ginlonins, BAe, o & ||palloollaaledeealledll lee heelly Wiotelbea evaliorst lee
HABBGUINE BONING Mma vale |e. |-|--\.-|2-|..ha.|-ofo.|F]..(F
loenmberci piegameme cn |eelietilia |erelecls.le%|eetea/=-)/.-| lle <|at
joemacilhins Tag 5 a |loallau|jool\6ol/o5| ea a6] eel (el oelee| lool aollee
Gentronotus, Regan .|..|..|--|--|--|--|..|..|..|--|--|-
scotti, Bouleng. ae | tesa
lepidorhinus, Pappenh. |..
eulepidotus, Regan. |:

Pleuragramma, Bouleng

antarcticum, Bouleng. leh salls0llo>\lan|(oallae)ladlipelleciloniieo| aur

ee wats
+ ++4++4+4+4:

Notothenia, Richards.
trigramma, Regan .
canina, Smitt leollog! eelhe Alle
ramsayi, Regan . .|..|..|--|--|--| |..
tessellata, Richards.
wiltoni, Regan |
brevicauda, Lénnb.
longipes, Steind.
sima, Richards.
squamifrons, Giinth.
larseni, Lénnb.
gibberifrons, Lénnb.
acuta, Giinth.
vaillanti, Regan
mizops, Giinth.
nudifrons, Lénnb. .
marionensis, Giinth.
angustifrons, Fisch. a|laalloe well eealteee
elegansy Guintlicn she) ele alte \oc|\-rills(| =.
cornucola, Richards. seal sen\ ara ee
cyaneobrancha, Richards|..|..|.....|..|..
coriiceps, Richards.
rossi, Richards. ocd Nate ol elioel lee
macrocephala, Giinth. . |+|..|..|..|..|+
microlepidota, Hutton .|..)..|..)..)-
colbecki, Bouléng.. .|..|..|..|..|.. ieee
filholiySaueage, = |. -|..)--|--|-«|- - ailgallaciee alle

Dissostichus, Smitt | | | Ie
eleginoides, Smiti . .|..|..|..|.-|--].-|--|#]..]--|--]-- Aral velo

2 Gi | |

See Cun, & Val. [+ |+).-|.-[.-[o-lec[-tl#|.-feofce{--[o-{--|-c|--| Ranges to northern Chile andthe

Artedidraco, Lénni. | ete ee Ve alae || La Plata.

orianae, Regan. 5 | oollee| | alloel/ Sele al el fess heels] [ae } teen fake
minus, onnv) 2 = © |... celloalteo|aelleallocligslies|(ao|(ar|es|taclloc healer

+t +++ ++:
+. t+ 44+: 4+

Northward to Milford Sound
and Chatham Island.

4 + bh . eile a i libelee Northward to Concepcion and
aN Ales eailee Lyttelton.

tt+++i 5:

1D}

28 “TERRA NOVA” EXPEDITION.

S. TEMPERATE
ZONE.

Tristan da Cunha.
St. Paul. ———
S.E. Australia.
New Zealand.

Chile.
Argentina.

ANTARCTIC ZONE.

Marion Island.

Antipodes.

| Falklands.

| Magellan.

Graham Land,

| Victoria Land.
Wilhelm Land.
Coats Land.

South Orkneys.

South Georgia.

Kerguelen. |

Artedidraco—cont.
skottsbergii, Lénnb.
shackletoni, Waite .
loennbergii, Roule .

Dolloidraco, Roule
longidorsalis, Roule

Histiodraco, Regan
velifer, Regan .

Pogonophryne, Fe, gan
scotti, Regan ae Ue

Harpagifer, Richards.
bispinis, Forst.

BATHYDRACONIDAB,
Bathydraco, Giinth.

antarcticus, Giinth.
macrolepis, Bouleng.
scotiae, Dollo .
Gerlachea, Dollo
australis, Dollo
Racovitzaia, Dollo
glacialis, Dollo
Prionodraco, Regan
evansii, Regan .
Parachaenichthys,
Bouleng.
georgianus, Fisch. .
Gymnodraco, Bouleng.
acuticeps, Bouleng.

CHAENICHTHYIDAE.

Champ socephalus;@i! |
esox, Giinth. mre eee
gunnari, Lénnb.

Pagetopsis, Regan
macropterus, Bouleng. .

Chaenichthys, Richards.
rhinoceratus, Richards. a
rugosus, Regan

Chaenocephalus, Regan| Be Sole eral fea eel lel lb hae Ie

aceratus, Lénnb.
Cryodraco, Dollo
antarcticus, Dollo .
atkinsoni, Regan
pappenheimi, Regan
Chionodraco, Lénnb.
hamatus, Loénnb.
kathleenae, Regan
Chaenodraco, Regan
wilsoni, Regan .
fasciatus, Regan

Jl t)4)4[4)4)4)..)..)..

..|..|+|..| S.E.of Heard Isd. 1,260 fathoms.
.|../+)..|..| S.W. of Balleny Is. 252 fathoms.
..).... +) Off Coats Land, 1,410 fathoms.

FISHES—REGAN.

The supposed relationships of the Nototheniiform families and of the cenera of

Ss

the Bovichthyidae are shown by the following diagram :—

Chaenichthyidae Bathydraconidae
Nototheniidae
| Bovichthys
Cottoperc:
Pseudaphritis
FA

Bovichthyidae.

In summarising the distribution of the Nototheniiformes the Bovichthyidae, the
most generalised family, may be considered first. Psewdaphritis most nearly represents
the prototype of the whole group and is the only genus peculiar to the South
Temperate Zone ; but the single species also differs from other fishes of the family in
that it is chiefly, or perhaps entirely, fluviatile. It may perhaps be regarded as a
relict form, analogous to the Galaxias of New Caledonia, dating from a time when the
seas of Australia were colder. The related Cottoperca is marine and is confined to the
Magellan District, whilst Bovichthys, which seems to be Cottoperca specialised, is
principally South Temperate, but is Subantarctic also in the Antipodes District.

The remaining families include 22 genera; one (/leginops) is monotypic and
occurs in the Magellan, Chilean and Argentine Districts; the rest are Antarctic,
16 peculiar to the Glacial District, one peculiar to the Kerguelen District, and four
found also in the Subantarctic Zone. Of these last, two (Dissostichus, Harpagifer)
are monotypic, the third (Champsocephalus) has only two species, and the fourth is the
large genus Notothenia, well represented in both Antarctic and Subantarctic Zones.

The species of Notothenia form five natural groups :—

(1) the tessellata group, comprising the first eight species in the list and confined
to the Magellan District ; these have the interorbital region rather broad, the
upper surface of the head except the snout, and its sides except the praeorbital
fully scaled.

(2) the sguamifrons group, including only NV. squamifrons from Kerguelen and
N. larseni from Graham Land and South Georgia; distinguished from
the preceding by the narrow interorbital region and the scaly snout and
praeorbital.

(3) the acuta group, including the next five species, all belonging to the Antarctic

“TERRA NOVA” EXPEDITION.

These have the interorbital region narrow, but resemble the fessellata

Zone.
group in the scaling of the head.
(4) the marionensis group, differing from the last in that the cheeks and opercles

are scaly above, naked below. This includes only N. marionensis, from
Marion Island, and N. angustifrons, from South Georgia.

Fic. 3.—Distribution of Bovichthyidae : P. Pseudaphritis urvillii ; C1, Cottoperca gobio ; C 2, C. macroph-
thalma ; © 3, C. macrocephala ; B1, Bovichthys variegatus ; B 2, B. angustifrons ; B3, B. diacanthus ;
B4, B. chilensis; B5, B. patagonicus; B6, B. veneris ; B7, B. decipiens; B8, B. psychrolutes ;

B9, B. roseopictus.
(5) the coriiceps group, with the upper surface of the head naked and the opercles
scaled only on the upper part of the operculum. This includes the last eight

species and is the only group found in both zones.

FISHES—REGAN. 31

Next to the Notothentiformes the Zoarcidae are of importance. This is principally
a northern family and includes both oceanic and coast fishes, many of the latter
frequenting rather deep water. South of the tropics the Zoarcidae are represented by

fROGELY
b7e. 7%
acdef kouwv
AS§ 27.3.5,

Fic. 4.—Distribution of Trematomus and Notothenia: 1, T. newnesii; 2, T. nicolai; 3, T. borchgrevinkii ;
4, T. brachysoma ; 5, T. bernacchii ; 6, T. vicarius; 7, T. dubius; 8, T. hansoni; 9, T. loennbergii ;
10, T. pennellii; 11, T. centronotus; 12, T. scotti; 13, T. lepidorhinus; 14, T. eulepidotus.
a, N. trigramma; b, N. canina; c, N. ramsayi; d, N. tessellata; e, N. wiltoni; £, N. brevicauda ;
g, WN. longipes; h, N. sima; i, N. squamifrons; j, N. larseni; k, N. gibberifrons; 1, N. acuta ;
m, N. vaillanti ; n, N. mizops; 0, N. nudifrons ; p, N. marionensis ; q, N.-angustifrons ; x, N. elegans ;
s, N. cornucola; +, N. cyaneobrancha; u, N. coriiceps; v, N. rossii; w, N. macrocephata ;
x, N. microlepidota ; y, N. colbecki ; 2, N. filholi.

a number of genera from the coasts of America and Antarctica. The following lst 1s

based on my revision in the “Scotia” report, modified by the study of a paper by

32 “TERRA NOVA” EXPEDITION.

Prof. Lahille* that I had overlooked, as it had been omitted from the Zoological
Record.

DISTRIBUTION OF THE SOUTHERN ZOARCIDAE.

SUBANT-
8. Re tage wae Antarctic ZONE.
: ONE.
E epee
IPPECPER EE EEERAE
al 2] 8)2|4)S| Sia) 4) 2] g/ele|a/e\3|3
Al Be) 3/2) o| a) 8) 3/8] 5] 8] 3) B)8)e| 8
B\<|S)ala/a)a/S\ Sl) a|s/alalolelelsS
| | | | |
|
Lycenchelys, Gill | |
antarcticus, Regan .|..| |..|..|.-]--|-- waltore lta allcte |ieaifece ior |O2ay LOIS: rls Oem Wiel varie)
Ophthalmolycus, Regan | | fathoms; other species from
NOR Nh ES G6 \eolee|lenl) olloo|/ecl/es)|ar\oe|loulfoc aol alters deep water N. of the equator.
(omen ie Tete gs Ilxe|lGallanitoclfeaiioe|oallalaolleollosilec eo lP lo olleallax
Caneolepis, Lahille = | ie | | |
acropterus, Lahille . Bd el [eee lee el ee Lee foes [ete alles neal SAN rel
Iluocoetes, Jenyns | he aie |
fimbriatus, Jenyns . | le larllsellaol lsc vefeefesfes
elongatus, Smitt é Alles alle |telleal sella sl trsllo seiltsrell bare ti
Lycodichthys, Pappenh. ) | | | -
antarcticus, Pappenh. |.....|..|.. 1 Neclfeelaol|aeljoc|ico|fec|foulles|{co/or|
Austrolycichthys, Regan | Ia | | I
brachycephalus, Papph.|..|..|..|..)..|.. Me wefeefeades ..|-.]+]--
bothriocephalus, Papph.|..|..|..|..|..|..]..| Jeofoefeefeafeefee]eo[ lee
Austrolycus, Regan | | bee |
moreno, Hanille ys 2 )\\ oats) -|/e7-1) 12) (-10)| ell ene 155] [eolleallao||colloollaslloe
depressiceps, Regan -.|..|..|..|..|..|.-|.-|4/#]..].-|..[--[--|-- bal
ibis G 6 6 Ilbailgolloniisal[oalnallen||Wlolloc|co||sal|s4|loollasiooj|oc
Phucocoetes, Jenyns | |
lntitans, Jenyns/ =. |. |/2-|--|--|--|--|--|--|atleofeeforles|=atastecter
Crossostomus, Lahille | | |
fimbriatus, Lahille ae |eeallceal ioral lave poe be
fasciatus, Lénnb. ed tecol [eel eesllanes Lavelle sallsrilc | Ie leealers
Platea, Steind. | al
insignis, Steind. SA ate es| lee Isalleel hoe
Maynea, Cunningh. | Eile | |
patagonica,, Cunningh.. |..|..|..|..|.-)--|--||#E]--|--)--}--le-|--]=<]--
Ehama nen Js & | a lleolladiloallaalwelloalaotloolloa|(aallac| (Ue llool|ecllouljeoljoa
Melanostigma, Ginth. |
gelatinosum, Giinth. .|..|. |..)..|. |.-].- +..|..]..]..]..|..].-)..).-| Magellan Straits, 24 fathoms ;

other species from deep water
of N. Atlantic and Pacific.

Two species are the southern representatives of the northern deep-water genera
Lycenchelys and Melanostigma. The rest are all generically distinct from the northern

* Lahille, ‘‘ Nota sobre los Zoarcidos Argentinos,” An. Mus. Nac. Buenos Aires xvi, 1908, pp. 403-441.
The net result of this paper is to add Austrolycus morenoi and Caneolepis acropterus and to replace
Crossolycus by Crossostomus, unless the latter is regarded as preoccupied by Crossostoma. Lycodalepis
laticinctus is probably a synonym of Austrolycus plater.

FISHES—REGAN. 33

members of the family ; two genera are peculiar to the Antarctic, two common to the
Antarctic and Magellan, four peculiar to the Magellan District, one common to
Magellan and Argentina, and one peculiar to the last-named.

The Magellan District is the headquarters of the southern Zoareidae, which may
have reached it originally from the north along the American coast, perhaps migrating
in rather deep and cold water. It is of interest to note that no Zoarcidae are known
from the South Temperate and Subantarctic Zones outside the Argentine and Magellan
Districts. Evidently the southern coast fishes of this family have been able to extend
their range only along a nearly continuous shore line ; this is in harmony with what
is known as to their breeding and development.

THE ANTARCTIC ZONE.

This includes the coasts of the Antarctic continent and the islands that lie to the
south of the isotherm of 6° C., with the probable exception of Macquarie Island.

It is characterised by the complete absence of South Temperate types, by the
absence of Bovichthyidae and the great development of the other Nototheniiformes.
It is more sharply marked off than any other zone, the percentage of peculiar genera
being extremely high and that of species that range beyond its limits very low.

GLACIAL Disrricr.

This ineludes the coasts of the Antarctic continent and neighbouring islands,
together with South Georgia, the South Sandwich Islands, and probably Bouvet Island,
all lying within the extreme limit of pack-ice. About 90 per cent. of the coast fishes
of this district are Nototheniformes and most of the rest are Zoarcidae.

NororHentipagz. The Nototheniinae are represented by Trematomus (14 spp.),
Pleuragramma (1 sp.), Notothenia (6 spp.), and Dissostichus (1 sp.). Eleginops is
absent. The two first-named genera are confined to this area; five of the six species
of Notothenia are peculiar, the exception being .V. coriiceps, which ranges to Kerguelen.
Dissostichus eleginoides is common to Graham Land and Magellan.

The Harpagiferinae (Artedidraco, Dolloidraco, Histiodraco, Pogonophryne, Harpa-
yifer) ave confined to this district, with the exception of Marpayifer bispinis, which
extends also to the Kerguelen and Magellan Districts.

BATHYDRACONIDAE are confined to the district. Lathydraco must be regarded
as oceanic, and probably the related genera Gerlachea, Racovitzaia and Prionodraco
should go with it ; but none of these has been found far from the coast of Antarctica,
the most northerly being B. antarcticus, taken midway between Wilhelm Land and
Heard Island. Parachaenichthys georgianus from South Georgia and Gymnodraco
acuticeps from the coasts of the Antarctic continent may be reckoned as coast fishes.

CHAENICHTHYIDAE are characteristic, the only extralimital forms being the two
species of Chaenichthys (Kerguelen) and Champsocephalus esox (Magellan).

34 “TERRA NOVA” EXPEDITION.

; ‘ ee
ZOARCIDAE are represented by a species of Maynea from South Georgia, the
only other species of the genus inhabiting the Magellan District, and by three species
from the coasts of Antarctica, belonging to two endemic genera, Lycodichthys and
Austrolycichthys, velated to the Magellan J/uocoetes and Austrolycus respectively.

Fic. 5.—Distribution of Harpagiferinae: A 1, Artedidraco orianae ; A 2, A. mirus; A3, A. skottsbergii ;
Ad, A. shackletoni; A5, A. loennbergii; D, Dolloidraco longidorsalis; V, Histiodraco velifer ;
P, Pogonophryne scotti; H, Harpagifer bispinis.

MtrAENOLEPIDAE. This family comprises but a single genus with one species
from Magellan, another from Kerguelen, and a third, Muraenolepis microps, from the
district now under consideration.

Corrmpakr, Sclerocottus schraderi, Fisch., from South Georgia parallels the

FISHES—REGAN,

35
Antarctic Zoarcidae as the representative of a northern family generically distinct
from the northern members of the group.

In the present state of our knowledge subdivision of the Glacial District would

be premature: many of the species are known to have a circumpolar distribution,

\
Ca

Fig. 6.—Distribution of Chaenichthyidae: Cs, Champsocephalus: 1, esow; 2, gunnari; P, Pagetopsis

rt)
macropterus; C, Chaenichthys: 1, rhinoceratus; 2, rugosus; Cn, Chaenocephalus aceratus ;
Cr, Cryodraco: 1, antarcticus; 2, atkinsoni; 3, pappenheimi ; Ci, Chionodraco: 1, hamatus ;
2, kathleenae ; Ch, Chaenodraco: 1, wilsoni ; 2, fasciatus.

and further researches will doubtless increase their number. It may be noted that
from Victoria Land 12 species of Trematomus and only one of Notothenia have been
described, from Graham Land six of 7rematomus and five of Notothenia, and from
South Georgia three of Trematomus and six of Notothenia. That South Georgia is

1a We

36 “«“TERRA NOVA” EXPEDITION.

rightly included can hardly be questioned; of its 15 species of Nototheniiform
fishes nine are known also from Graham Land, and the remaining six include
species of Trematomus and Artedidraco, whilst only one (Champsocephalus gqunnar’)
has Magellan affinities.

KerGueLen District.

This includes the island of Kerguelen and also Heard Island, the Crozet Islands,
and Marion and Prince Edward Islands.

The fish-fauna of Kerguelen is known from the collections made there by the
“Erebus” and “Terror” and the “Challenger”; recently the “Gauss” collected
there, but obtained no additional species.

There are two species of the cosmopolitan genus Rava ; Muraenolepis marmoratus
is related to M. microps and M. orangiensis of the Glacial District and of Magellan
respectively, and Zanclorhynchus spinifer is a genus and species peculiar to the island,
its nearest relative being the south temperate Congiopus (Agriopus).

The rest are Nototheniiformes, viz., five species of Notothenia, one of Harpagifer
and two of Chaenichthys. Of these Notothenia corticeps and Harpagifer bispinis are
found on the coast of the Antarctic continent ; the rest are peculiar, but show relation-
ship to species known from South Georgia and Graham Land, for \. sqguamifrons is
closely related to V. larseni, N. acuta to N. gibberifrons, N. mizops to N. nudifrons,
and N. cyaneobrancha to N. coriiceps, whilst Chaenichthys, a genus peculiar to the
island, belongs to the Antarctic family Chaenichthyidae. Little is known of the other
islands that may pertain to this area, but it may be noted that at Marion Island
Notothenia marionensis is the representative of the South Georgian .V. angustifrons.

At the first glance it may seem that as so many characteristic Antarctic genera
appear to be absent and most of the Nototheniidae belong to Notothenia, which is well
represented in the Subantarctic Zone, the Kerguelen District might be included in the
latter. But a more critical examination shows that the tessel/ata group, characteristic
of Magellan, is absent, that the squamifrons, acuta and marionensis groups are present
and are found elsewhere only in the Glacial District, and that the coriiceps group is
represented by .V. coriiceps, an Antarctic species, and by the related NV. cyaneobrancha.

The only way to mark the dissimilarity of the fish-fauna of Kerguelen from that
of Magellan or of the subantaretic islands of New Zealand, and to express its affinity to
that of Antarctica, is to include it in the Antaretic Zone as a separate district, small
and impoverished, but with well-marked characters.

THE SUBANTARCTIC ZONE.

This includes the Falklands and the southern extremity of South America,
northwards about to Chiloe and Cape Blanco, and the extreme south of New Zealand,
with the adjacent islands, Stewart, Auckland, Campbell, Antipodes, and perhaps
Macquarie. Its limits correspond approximately to the isotherms of 12° C: and 6° C.

FISHES—REGAN. 37

A number of South Temperate types extend into this zone, Galaxiidae are
characteristic, the Nototheniiformes are represented chiefly by Notothenia, and there are
several peculiar Zoarcidae.

MaGeELLAN Disrricr.

This includes Tierra del Fuego, the coasts of Patagonia northward to Chiloe on
the west and Cape Blanco on the east, and the Falkland Islands.

BovicHTHYIDAE are represented by the genus Coftoperca (3 spp.), which is
confined to this district.

NoroTHENIDAE: Zrematomus, Pleuragramma and the Harpagiferinae except
Harpayifer bispiis are absent. Eleginops mactovinus ranges northward to Valparaiso
and Buenos Aires, and Dissostichus eleginoides southwards to Graham Land. Peeuliar
to this area is a natural group of eight species of Notothenia (tessellata group)
comprising all the species with the upper surface and sides of the head, except the
snout and praeorbital, scaly, and with the interorbital region rather broad. Two other
species (LV. cornucola and N. macrocephala) of quite another type occur also in the
Antipodes District, whilst a third (.V. e/egans), not found elsewhere, may be related
to NV. cornucola.

CHAENICHTHYIDAE are represented by a species of Champsocephalus, a genus that
includes one other species from South Georgia.

ZOARCIDAE: Llwocoetes (2 spp-)s Phucocoetes (1 sp.), Crossostomus (2 spp.); and
Platea (1 sp.) are peeuliar ; Ophthalmolycus and Maynea each has a species here and
another in the Antarctic, whilst Austrolycus comprises two species from this district
and one from Buenos Aires.

MuRAENOLEPIDAE. Muraenolepis orangiensis is confined to this district, and
represents therein the two Antarctic species of the genus.

GALAxuDAE. Of six species of Galavias in this district three are known to be
marine; G. maculatus and G. gracillimus have been found in the sea off the Falklands,
and the former at Puerto Montt also. The range of the species in the sea is not well
ascertained ; in fresh water they extend at least to Arauco and to the lakes at the head
of the Rio Negro.

The majority of the fishes mentioned above are not known to range outside the
district as delimited ; the remainder of the fishes that oceur within its boundaries are
Chilean or Argentine forms.

AntipoprEs Disrricr.*

This comprises the southern extremity of New Zealand and the islands southward
to Campbell Island ; whether Macquarie Island should be cluded is unknown. Most
of the few fishes known are New Zealand species ; Zoarcidae are absent, and the only

* Subantarctie Islands of New Zealand, Wellington, 1909. Art. xxv. pp. 585-598, Pisces by

E. R. Waite.

3

io)

“TERRA NOVA” EXPEDITION.

distinctive feature is the presence of five species of Notothenia of the coriiceps group,
three of them peculiar, and the other two found also in the Magellan District and
extending in New Zealand somewhat to the north of the boundaries assigned above.
Galazias attenuatus is marine south to Campbell Island and at least as far north as
Cook’s Straits and Tasmania ; in fresh water the genus extends to New Caledonia.*

B. OCEANIC FISHES.

It has been mentioned above that the Glacial District, for coast fishes, comprises
the islands that he within the extreme limit of pack-ice. It may be of some interest to
consider the known distribution of the oceanic fishes, whether pelagic, bathypelagic or
abyssal, that have been recorded from within this limit.

ARGENTINIDAE.

The widely distributed genus athylagus is represented by three species—B.
antarcticus, Giinth., B. gracilis, Loennb., and B. glacialis, Regan. Of these the first
ranges north to 37° 8., the second to 49° 8., and the third is known only from
specimens taken off Coats Land.

(GONOSTOMATIDAE.

Cyclothone microdon, Giinth., is cosmopolitan.

STOMIATIDAE.

Stylophthalmus paradoxvus, Brauer, is a larval form recorded from near Bouvet
Island, and also from the South Atlantic and Indian Oceans. -

SUDIDAE.

Notolepis coatsii, Dollo, is a genus and species that may be peculiar to the area
within the extreme limit of pack-ice. It is cireumpolar (cf. Regan, Trans. R. Soe.
Edinb. xi1x, 1913, p. 233); a larva from Graham Land has recently been figured by
Prof. Roule (Deuxitme Expéd. Antarct. Frang. Poiss. pl. m1).

MycrTopHIDAE.

The cosmopolitan genera Myctophum and Lampanyctus are each represented by a
single species. MZ. antarcticum, Giinth., is crcumpolar, and ranges north to 28°S. L.
braueri, Loennb., is known from a specimen from off Coats Land, and another from
49° 56’ S., 49° 56’ W.

* Weber and Beaufort, Zool. Anz. xt, 1913, p. 172.

FISHES—REGAN. 39

SCOPELARCHIDAE.

Dissoma anale, Brauer, is known from near Bouvet Island, and from the South
Atlantic and Indian Oceans.

ALEPIDOSAURIDAE.

Eugnathosaurus vorar, Regan, is a genus and species known only from off
Coats Land.

GADIDAE.

Melanonus gracilis, Giinth., originally described from the Antarctic, is now known
to range in the Atlantic to 36° N.

MACRURIDAE.

Cynomacrurus piriei, Dollo, is a genus and species known only from the type

taken off Coats Land. The other species belong to widely distributed genera :—

o
Chalinura ferrieri, Regan, and C. whitsoni, Regan, are from Coats Land, and the
latter from 48° S., 10° W. also. Nematonurus lecointe’, Dollo, has been taken off
Graham Land, near the South Orkneys, and in 48°8., 10° W. NV. armatus, Hect., is
known from near the Crozet Islands and to the north of Wilkes Land, also from the
Pacific as far north as 87° N. Lionwrus jfilicauda, Giinth., ranges north to the latitude
of Valparaiso and the La Plata, and has been taken in the Antarctic Ocean to the
north of Wilkes Land.

BaATHYDRACONIDAE.

As mentioned above, bathydraco certainly and three related genera probably
should be classed as oceanic; they are not known to occur beyond the limits of
the Glacial District.

ZOARCIDAE.

Lyecenchelys antarcticus, Regan, from near the South Orkneys, is the first southern
species of Lycenchelys.

BROTULIDAE.

Holcomycteronus brucei, Dollo, is from near the South Orkneys; a second species
has been described by Garman from 2°-26° N., 82°-110° W.

CYCLOPTERIDAE.

Liparis steineni, Fisch., and Careproctus georgianus, Loennb., from South

‘Georgia, and Paraliparis antarcticus, Regan, from Victoria Land, are the Antarctic

representatives of genera that seem to be cosmopolitan, but may be best represented
in northern seas and include many deep water species.

40 “TERRA NOVA” EXPEDITION.

Our knowledge of the distribution of the pelagic and deep sea fishes of the
Antarctic Ocean is obviously not suttciently complete to permit of an attempt to
co-ordinate the results so far obtained.

2. THE ANTARCTIC CONTINENT DURING THE
TERTIARY PERIOD.

Many authorities believe that in the early Tertiary the Antarctic continent
was connected with Australia and with South America; some think with Africa
also. The distribution of the Fishes, and of other groups of animals, has been
considered to support this hypothesis.

FISHES.

The distinctive features of the fish-fauna of the coasts of Antarctica are that
nearly all the genera and species are peculiar and that they nearly all belong to a
single group, the Nototheniiformes, which is characteristic of and almost restricted
to the Antarctic and Subantarctic Zones. In the Antarctic Zone this group has
developed into a large number of types that differ greatly in structure, appearance
and habits. These facts seem to point to the conclusions that Antarctica may have
long been isolated and that its coasts may have been washed by a cold sea for a
long time, probably throughout the Tertiary Period.

It has been suggested that identical or related species of different parts of the
South Temperate Zone are part of the fauna of the coasts of an Antarctic continent
that formerly connected America, Africa and Australia. It is very difficult to make
this view harmonise with the facts. Tristan da Cunha and St. Paul, 4,500 miles
apart, have at least two species of shore fishes (Labrichthys ornatus and Chilodactylus
monodactylus) im common. Have these persisted unchanged whilst the coasts of
Antarctica have receded to their present position, and whilst the fauna of Kerguelen
has become differentiated? It seems far more likely that the distribution of these
species is due to present conditions; Tristan da Cunha and St. Paul are nearly in
the same latitude and on the same isotherm, and the Antarctic Drift runs directly
from one to the other. In all probability it will be found that the species common
to both islands have floating eggs and larvae that swim at the surface; it is even
possible that the adult fish may occasionally migrate from one island to the other.
The case of related species, as for example Seriolella velaini from St. Paul, and
S. antarctica from Tristan da Cunha, may be explained on similar lines; it is
evident that the distance between the islands is too great to be traversed by these
species, but under somewhat different conditions the parent form may have lived
at both and either at the Cape or at Marion and the Crozets, or at other islands
that may have existed and served as stepping stones, but have now disappeared.
Whatever may be the true explanation, it is certain that the construction of

FISHES—REGAN. 41

Tertiary land-bridges to account for all cases of this sort would reduce the oceans to
a few puddles.

Much has been made of the distribution of the Galaxiidae and Haplochitonidae, for
some time regarded as fresh-water fishes found in Southern Australia and Tasmania,
New Zealand, and the southern part of America. It is now known that Galaaias
attenuatus, the only species common to all these regions, breeds in the sea. In the
“ Scotia” report | have shown that these two families are Salmonoids related to the
Osmeridae, and their marine origin may be regarded as certain. Like the northern
Salmonoids they are establishing themselves in fresh water, and it is interesting to note
that Galaxias occurs at the Cape and even in New Caledonia, where, like the Trout of
Algeria, it remains as the witness of a glacial epoch.

None of the families of true fresh-water fishes of either South America or Africa
occurs in Australia, except the Osteoglossidae, a generalised and ancient type. Even in
this case the relationship is not with America or Africa, but with Asia, Scleropages
comprising one species from Queensland and New Guinea, and one from Borneo and
Sumatra.

Thus neither marine nor fresh-water fishes support the theory that the Antarctic
continent connected America with Australia during the Tertiary Period. This being
the case, I have been led to examine somewhat critically the other zoogeographical
evidence in support of this theory. Some of this is derived from the similarity of
marine faunas, or from the distribution of fresh-water organisms that may have had a
marine origin. Of more importance are the land animals, and Dollo, in his monumental
report on the “ Belgica” fishes, comes to the conclusion, “ C’est (Antarctide Tertiaire
de M. Osborn—ou une Antarctide analogue, indispensable pour les Marsupiaua et
Miolania—qui explique le mieux la Biogéographie des Poissons Antaretiques et Sub-
antarctiques.” Unable to accept this for the Fishes, I have looked into the question of
the Marsupials and JMiolania.

MARSUPIALS.

It has been suggested that Caenolestes and the extinct Patagonian members of the
Epanorthidae may be related to the Australian group Diprotodontia. This is by no
means generally accepted, and several authorities believe that the Epanorthidae may
have been derived from a primitive Didelphoid type which has evolved a diprotodont
dentition independently. This view is supported by the fact that the Epanorthidae are
eleutherodactyle, whereas the Australian Diprotodonts are syndactyle.*

It has recently been shown by Gidley f that the Multituberculates are “ Dipro-
todonts,” so that Marsupials of this type date back to the Triassic, and even if they be
monophyletic, their occurrence in Australia and South America loses its significance.

* Thomas, Proc. Zool. Soc. 1895, p. 870 ; Gregory, Bull. Amer. Mus. xxvut, 1910, p. 211.
+ Proc. U. 8. Nat. Mus. xxxvi, 1909, p. 611.
G

“TERRA NOVA” EXPEDITION.

42
Recently some American writers * claim to have proved the relationship of the

Sparassodonts of Ameghino, Borhyaena, Prothylacinus, ete., of the Miocene of
I to} ? > e b >
Patagonia to the Tasmanian 7’hylacinus, and even unite them in a family Thylacinidae,

distinct from the Dasyuridae ; for this there seems to be but little justification.

Fie. 7.—Skulls of A, Borhyaena tuberata (after Sinclair) ; B, Phascologale dorsalis (after Thomas), and C,
Thylacinus cynocephalus ; seen from above. Phascologale has been selected for comparison as a small
insectivorous type, differing from the large carnivorous Thylacinus in the form of the cranium and

zygomatic arch, but resembling it in many important structural characters.

Tomes} has shown that in Borhyaena the enamel of the teeth resembles in
structure that of Creodonts and Carnivores, lacking tubules, and with the groups of

* Sinclair, Rep. Princeton Exped. Patagonia, 1v, 1901, p. 333; Gregory, Bull. Amer. Mus. xxvm,

1910, p. 207.
tT Proc. Zool. Soc. 1906, 1, p. 45.

FISHES—REGAN, 43

prisms interlacing to produce a characteristic pattern in cross-section. In Thylacinus
and Dasyurus, on the contrary, the enamel is typically Marsupial in that it is
penetrated by tubules continuous with those of the dentine and has the prisms straight
and parallel. The two genera mentioned show a special agreement in that the tubules
end at some distance from the surface. Ameghino states that in the Sparassodonts not
only the third praemolar, but the canine and sometimes the second praemolar, have
deciduous predecessors. In Thylacinus, as in the other Dasyuridae, only the third
praemolar is precéded by a milk tooth. I have examined the skulls of Thylacinus and
several other Dasyuridae (Fig. 7, B,C) and find that they agree in havine the orbits
well backward, the nasal processes of the praemaxillaries long, the nasals but moderately
expanded posteriorly, the maxillary and frontal meeting in a suture, the jugal emitting
a postorbital process just before its junction with the squamosal, the occipital region
triangular in outline, the basisphenoid foramina paired and palatal vacuities present
(said to be absent in some species of Phascologale),

The Sparassodonts, as described and figured in Sinclair's admirable monograph,
differ in the more anterior position of the orbits, shorter nasal processes of the prae-
maxillaries, nasals strongly expanded posteriorly, meeting the lachrymals and separating
the maxillaries from the frontals, the absence of a distinct postorbital process, the
semicircular occipital outline, the unpaired basisphenoid foramen, and the absence of
palatal vacuities (Fie. 7, A).

The dentition of Thylacinus is readily derivable from the primitive Dasyurid type
(Phascologale) ; the teeth of the Sparassodonts correspond closely to those of Thylacinus
in form and number, except that the metacone of the fourth upper molar is vestigial or
absent, whereas in Thylacinus it is well-developed.

There appears to be no escape from the conclusion that Thylacinus is a true but
aberrant member of the Dasyuridae, and that it has nothing to do with the
Borhyaenidae, a family well characterised by peculiarities in the skull, and in the
structure and perhaps in the succession of the teeth. The specialised carnivorous
dentition, superficially similar to that of Zhylacinus, has been independently evolved.

MToLaANtIA.

The family Miolaniidae includes some large extinct Pleurodiran Tortoises that are
remarkable for the development of a caudal sheath of bony rings and the presence
of dermal bony bosses on the head.

There are two genera, Miolania, Owen, and Niolamia, Ameghino. The former
includes two species, J. platyceps, Owen,* from the Pleistocene of Lord Howe Island,
and M. oweni, A. S. Woodward, from the Pleistocene of Queensland. Niolaimia

* Owen, Phil. Trans., 1886, pp. 471-480, pls. xxix, xxx ; and 1888, pp. 181-191, pls. xxxI—xxxvit.
ft Woodward, Ann. Mag. Nat. Hist. (6) 1, 1888, p. 89.
G 2

44 “TERRA NOVA” EXPEDITION.

comprises a single species, VV. argentina, Ameghino,* from Patagonian deposits that
were at first stated to be Cretaceous, but may prove to be Miocene.

In Miolania (Fig. 8, B) the skull, seen from above, is somewhat oblong in form,
with the snout broadly rounded. The dermal bosses all have the appearance of separate
elements. On the upper surface may be recognised a large but low parietal pair ;
behind them is an occipital pair that project backwards, and in front of them a
smaller frontal pair mesially, and ‘a postfrontal pair laterally. A pair of subconical
bosses, rounded or ovate in transverse section, on each side of the parietal pair, project
as lateral “horns”; a much smaller pair are placed directly in front of them. Other
features of the genus that may be mentioned are that the praemaxillaries have a
median pit for the reception of the symphysial beak of the mandible, that the
palatal extensions of the praemaxillaries and maxillaries bear two sharp ridges within

Fie, 8.—Skulls of A, Niolamia argentina, and B, Miolania oweni, seen from above. In B the position of
the anterior margin of the praemaxillaries is indicated by a dotted line.

and parallel to the margin of the upper jaw, that the nasals project beyond the
praemaxillaries, and that there is a bony internasal septum.

In Miolamia (Fig. 8, A) the skull is nearly triangular in outline, with the snout
more acute than in Miolania. The bosses differ considerably from those of Miolania,

expansions, the lateral “horns” are broad and flat, triangtlar im section, and have no
smaller pair in front of them. Further differences from J/iolania are that the prae-
maxillaries are not pitted, the mandible is not beaked, the upper jaw has a single
blunt intramarginal ridge, the nasals do not project beyond the praemaxillaries, and
there is no internasal septum.

These differences have already been pointed out by Dr. Smith Woodward (Proc.
Zool. Soc. 1901, 1, pp. 174-176), but he has not insisted on them so much as on

* Woodward, Proc. Zool. Soc., 1901, 1, p. 170, pls. xv—xvuil.

FISHES—REGAN. 45

the essential similarity of the American and Australian fossils, which he included in
one genus, Miolania. However, admitting their relationship and their distinctness
from all other Chelonians, it seems to me that these two well-marked types may he
regarded as different genera. If it be considered that several recent genera of the
Chelonia are represented in the Eocene by species quite like the modern ones, the
occurrence in Australia and South America of types so dissimilar—one of which
dates back at least to the time when Lord Howe Island was part of the continent
of Australia, and the other is known from deposits that may be Miocene or perhaps

earlier—can be explained without inventing a Tertiary land-bridge.

The conclusion is that neither the fresh-water fishes, nor the marine fishes, whether
Antarctic or South Temperate, support the theory that Antarctica has connected
Australia with South America in Tertiary times, nor, so far as I can read the evidence,

does the distribution of other groups of animals confirm this view.

46 “TERRA NOVA” EXPEDITION.

BIBLIOGRAPHY.
PrincrpaAL MEMOIRS DEALING WITH ANTARCTIC AND SUBANTARCTIC FISHES.

1842.—Junyns, L. The Zoology of the voyage of H.M.S. Beagle. Fishes. 172 pp. 29 pls.
1844—-8.—Ricuarpson, J. The Zoology of the voyage of H.M.S. Erebus and Terror. 1. Ichthyology.
139 pp. 60 pls.
1880.—GiinrHer, A. Report on the Shore Fishes procured during the voyage of H.M.S. Challenger.
82 pp. 32 pls.
1881.—Giinrner, A. “ Account of the Zoological Collections made during the survey of H.M.S. Alert in’
the Straits of Magellan and on the coast of Patagonia. Fishes.” Proc. Zool. Soc, 1881.
pp- 19-22. Pls. I, U1.
1885.—Fiscner, J.G. “Uber Fische von Siid-Georgien.” Jahrb. Hamburgisch. Wissensch. Anst. IT.
pp. 49-62. Pls. I, IT. fig. 9.
1887.—Ginruer, A. Report on the Deep-Sea Fishes collected by H.M.S. Challenger. 335 pp. 66 pls.
1889.—Ginruer, A. Report on the Pelagic Fishes collected by H.M.S. Challenger. 47 pp. 6 pls.
1891.—Vaintant, L. Mission scientifique du Cap Horn. VI. Zoologie. Poissons. 35 pp. 4 pls.
1897-8.—Smirt, F. A. “ Poissons de l’expédition scientifique & la Terre de Feu sous la direction du Dr.
O. Nordenskjéld.” Bihang. Svensk. Vet.-Akad. Hand]. XXIII, IV, No. 3. 37 pp. 3 pls.
and XXTV,IV, No.5. 80pp. 6 pls.
98.—SreInDACHNER, F.—‘ Die Fische der Sammlung Plate.” Zool. Jahrb. Suppl. IV. pp. 281-334.
Pls. XV-XXTI.
1901.—Detrin, F. T. Catalogo de los Peces de Chile. Valparaiso. 133 pp.
1902.—Boutencer, G. A. “ Pisces” in The Natural History Collections of the Southern Cross. pp.
174-189. Pls. XI-X VIII.
1904.—Dotto, L. Résultats du voyage du Belgica. Poissons. 240 pp. 12 pls.
1905.—Linnperc, E. The Fishes of the Swedish South Polar Expedition. Wissensch. Ergebn.
Schwedisch, Siidpolar-Exped. V,6. 69 pp. 5 pls.
1906.—Lonneera, E. “ Contributions to the Fauna of South Georgia. 1. Taxonomic and biological notes
on Vertebrates.” K. Svensk. Vet.-Akad. Handl. XL. No.5. 104 pp. 12 pls.
1906.—Braver, A. Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition auf dem Dampfer
Valdivia. Die Tiefsee-Fische. I. Systematischer Teil. 420 pp. 16 pls.
1906.—Vaitiant, L. Expédition antarctique frangaise. Poissons. 51 pp.
1907.—Boutencer, G. A. National Antarctic Expedition. Natural History. II. Fishes. 5pp. 2 pls.
1909.—Warrs, E. R. “Pisces” in Subantarctic Islands of New Zealand. pp. 585-598.
1911.—Waire, E. R. “Antarctic Fishes” in British Antarctic Expedition, 1907-9. Biology. pp. 11-
is dee TNE
1912.—PapprenHeiM, P. ‘‘ Die Fische der Antarktis und Subantarktis” in Deutsche Siidpolar-Expedition,
1901-1903. XIII. Zool. V. pp. 163-182. Pls. [X—X.
1913.—Reaan, C.T. “The Antarctic Fishes of the Scottish National Antarctic Expedition.” Trans,
R. Soc. Edinburgh, XLIX. pp, 229-292. Pls. I-XI.
1913.—Routr, L. Deuxiéme Expédition antarctique frangaise. Poissons. 24 pp. 4 pls.

1

io)

FISHES—REGAN.

EXPLANATION OF THE PLATES.

PLATE I.
Fig. 1.—Trematomus bernacchii.
» 2— as hansoni.
PLATE II.

Fic. 1.—Paraliparis antarcticus.
», 2.—Muraenolepis microps.
» 93—Trematomus brachysoma.,

PLATE III.
Fic, 1.—Trematomus centronotus.
9 - pennellii.
PLATE IV.
Fic. 1.—Trematomus lepidorhinus.
» 2— 5 scotti.
eo a eulepidotus.
PLATE V.
Fie. 1.—Artedidraco skottsbergii.
» 2— 5 loennbergii.

» 93.—Histiodraco velifer.

PLATE VI.
- Fie. 1.—Pogonophryne scotti (reduced to 3).
» 2.—Artedidraco orianae. :
PLATE VII.

Fig. 1.—Prionodraco evansii.
» 2.—Cryodraco atkinsoni (reduced to @).

PLATE VIII.
Chionodraco kathleenae (reduced to 3)

PLATE IX.
~ Fic. 1.—Chaenodraco wilsoni.
a i fasciatus.
PLATE X.
Fic. 1.—Xystreurys brasiliensis.

,, 2.—Idiacanthus niger.

AT

“TERRA NOVA” EXPEDITION.
PLATE XI.
Fic. 1—Prionotus brachychir.

+ 2.—<Arnoglossus mongonuiensis.
.» 3—Serranops maculicauda. — 3

> 4.—Lepidoperca inornata. a
PLATE XII. os
Fic. 1—Hemerocoetes pauciradiatus. . “

” 2.— » macrophthalmus.

Acanthoclinus littoreus, 17.
acanthorhynchus, Hemerocoetes, 18.
aceratus, Chaenocephalus, 28.
acropterus, Caneolepis, 32.
acuta, Notothenia, 27, 29, 36.
acuticeps, Gymnodraco, 28, 33.
Agriopus, 36.
Alepidosauridae, 39.
anale, Dissoma, 39.
angustifrons, Bovichthys, 26.

3 Notothenia, 27, 30, 36.
antarctica, Maynea, 32.

3 Seriolella, 40.
antarcticum, Myctophum, 1, 38.

5 Pleuragramma, 6, 27.
antarcticus, Bathydraco, 28, 33.

3 Bathylagus, 38.

_ Cryodraco, 11, 28.

= Lycenchelys, 32, 39.

“A Lycodichthys, 32.

rr Paraliparis, 13, 39.
aotea, Cepola, 18.
argentina, Niolamia, 44.
Argentinidae, 38.
Ariomma lurida, 20.
armatus, Nematonurus, 39.
Arnoglossus mongonuiensis, 21.
Artedidraco, 6, 27, 33, 36.

% loennbergii, 7, 8, 28.
[ mirus, 6, 7, 8, 27.
= orianae, 6, 7, 27.
3 shackletoni, 7, 8, 28.

9 skottsbergii, 7, 9, 28.
atkinsoni, Cryodraco, 11, 28.
attenuatus, Galaxias, 38, 41.
atun, Thyrsites, 16.
auriga, Serranus, 22.
australis, Galeus, 14.

a Gerlachea, 28.
Austrolycichthys, 32, 34.

Fr bothriocephalus, 32.
Py brachycephalus, 32.
Austrolycus, 32, 34, 37.
~ depressiceps, 32.
= morenoi, 32.
$5 platei, 32.

FISHES—REGAN. 49

INDEX

balearica, Congromuraena, 22.
Bathydraco, 10, 28, 33, 39.

3 antarcticus, 28, 33.
- macrolepis, 28.
” scotiae, 28.

Bathydraconidae, 10, 28, 29, 33, 39.
Bathylagus, 38.

” antarcticus, 38.
» glacialis, 38.
” gracilis, 38.

Batrachoididae, 23.
bernacchii, Trematomus, 2, 3, 27.
35. 36.

boops, Pseudorhombus, 21.
borchgrevinkii, Trematomus, 2, 27.
Borhyaena, 42.
Bothidae, 21, 23.
| bothriocephalus, Austrolycichthys, 32.
| Boyichthyidae, 26, 29. /

Bovichthys, 26, 29.

5 angustifrons, 26.
FA chilensis, 26.

hi decipiens, 26.

B diacanthus, 26.
Pa patagonicus, 26.
Pr psychrolutes, 26.
i roseopictus, 26.
PA variegatus, 26.
Pr veneris, 26.

brachycephalus,, Austrolycichthys, 32.
brachychir, Prionotus, 22.
brachysoma, Trematomus, 2, 3, 27.
brasiliensis, Xystreurys, 23.
braueri, Lampanyctus, 38.
brevibarbe, Ophidium, 23.
brevicauda, Notothenia, 27.
brevimanus, Cubiceps, 20.
britannicus, Centrolophus, 19.
Brotulidae, 20, 39.
brucei, Holcomycteronus, 39.
Caenolestes, 41.
caeruleus, Cubiceps, 19.
Caesioperca, 17.
Caneolepis, 52.

5 acropterus, 32,
canina, Notothenia, 27.

50 “TERRA NOVA”

Caprodon longimanus, 16.
Carangidae, 17.
Carchariidae, 14.
cardinalis, Scorpaena, 21.
Careproctus georgianus, 39.
caudatus, Lepidopus, 16.
Centriscops, 15.

5 humerosus, 15
Centrolophus, 19.
A britannicus, 19.
3 huttoni, 19.
rn lockingtoni, 19.
rr maoricus, 19.
5 niger, 19.

O7

centronotus, Trematomus, 3, 4, 27.
Cepola, 18.
5 aotea, 18.

Cepolidae, 18.
Chaenichthyidae, 11, 28, 29, 33, 36, 37.
Chaenichthys, 11, 28, 33, 36.

4 ARerocorntus 28.

0 rugosus, 28.
Chaenocephalus, 11, 28.

A aceratus, 28.
Chaenodraco, 11, 12, 28.
3 fasciatus, 13, 28.
“ny wilsoni, 12, 13, 28.

Chalinura ferrieri, 39.

55 whitsoni, 39.
Champsocephalus, 11, 12, 28, 29, 37.
esox, 28, 33, 36.
gunnari, 28.

”
»
chilensis, Boyvichthys, 26.
Chilodactylidae, 18.
Chilodactylus macropterus, 18
, ati 40.
Chionodraco, 11, 12, 28
hamatus, 12, 28.
_ kathleenae, 12, 28.
cirrifer, Malacorhina, 21.
8 Psammobatis, 21.
coatsii, Lepidoperca, 17.
» Notolepis, 38
colbecki, Notothenia, 27.
concolor, Ophthalmolycus, 32
Congiopus, 36.
Congridae, 22.
Congromuraena balearica, 22.

”

coriiceps, Notothenia, 27, 30, 33, 36, 38.

cornucola, Notothenia, 27, 37.
Cottidae, 34.
Cottoperca, 26, 29.
3 gobio 26.
PA macrocephala, 26.
3 macrophthalma, 26.

EXPEDITION.

Crepidogaster simus, 21.
Crossolycus, 32.
Crossostoma, 32.
Crossostomus, 32, 37.
5 fasciatus, 32.
ma fimbriatus, 32.
Cryodraco, 11, 28.
os antarcticus, 11,
rs atkinsoni, 11, 28.
© pappenheimi, 11, 28
Cubiceps, 20.

98

ss brevimanus, 20.

3 caeruleus, 19.

> gracilis, 20.

A luridus, 20.

n pauciradiatus, 20.

cyaneobrancha, Notothenia, 27, 36.
Cyclopteridae, 13, 39.
Cyclothone microdon, 38.
Cynoglossidae, 23.
Cynomacrurus piriei, 39.
Cynophidium, 2

* punctatum, 20.
Dasyuridae, 42.
Dasyurus, 43.
decipiens, Bovichthys, 2
depressiceps, Austrolycus, 32.
diacanthus, Bovichthys, 26.
Diprotodontia, 41.
Dissoma anale, 39.
Dissostichus, 27, 29, 33.

- eleginoides, 27, 33, 37.
Dolloidraco, 6, 28, 33.

“ longidorsalis, 28.

op velifer, 9.
dubius, Trematomus, 2, 27.
elegans, Notothenia, 27, 37.
eleginoides, Dissostichus, 27, 33, 37.
Eleginops, 27, 29, 33.

. maclovinus, 27, 37.
elongatus, Iluocoetes, 32.
Epanorthidae, 41.
esox, Champsocephalus, 28, 33, 36.
Etropus microstomus, 23.
Eugnathosaurus vorax, 39.
eulepidotus, Trematomus, 3, 5
evansii, Prionodraco, 10, 28
Exocoetidae, 15.

Exocoetus spilonotopterus, 15.
fasciatus, Chaenodraco, 13, 28.
- Crossostomus, 32.
fernandezianus, Notopogon, 16.
ferox, Idiacanthus, 14.
ferrieri, Chalinura, 39.
filholi, Notothenia, 27.

-

“A

filicauda, Lionurus, 39.
fimbriatus, Crossostomus, 352.
5 Tluocoetes, 32.
fyllae, Raia, 22.
Gadidae, 39.
Galaxias, 29, 37, 41.
on attenuatus, 38, 41.

5 gracillimus, 37.
5 maculatus, 37.

Galaxiidae, 37, 41.

Galeus australis, 14.
gelatinosum, Melanostigma, 32.
Gempylidae, 16.

OF] ay “C < 2¢
georgianus, Careproctus, 59.

oH Parachaenichthys, 10, 28, 33.
Gerlachea, 28, 33.
5p australis, 28.

gibberifrons, Notothenia, 27, 56.
glacialis, Bathylagus, 38.

35 Racovitzaia, 28.
Gobiesocidae, 21.
gobio, Cottoperca, 26.
Gonorhynchidae, 14.
Gonorhynchus gonorhynchus, 14.
gonorhynchus, Gonorhynchus, 14.
Gonostomatidae, 38.
gracilis, Bathylagus, 38.

5 Cubiceps, 20.

FA Melanonus, 59.
gracillimus, Galaxias, 37.
gunnari, Champsocephalus, 28.
Gymnodraco, 10, 28.

5 acuticeps,'28, 33.
hamatus, Chionodraco, 12, 28.
hansoni, Trematomus, 2, 3, 27.
Haplochitonidae, 41.

Harpagifer, 6, 28, 29, 33, 36, 37.

a bispinis, 28, 33, 36, 37.
Harpagiferinae, 6, 33, 37.
Hemerocoetes, 18.

op acanthorhynchus, 18.
5 macrophthalmus, 18.
Ee microps, 18.

3 pauciradiatus, 18.

- waitei, 18.

Hemerocoetidae, 18.
Hemirhamphidae, 14.
Hemirhamphus intermedius, 14.
Hippocampus punctulatus, 22.
Histiodraco, 6, 9, 28, 33.

FA velifer, 9, 28.
Holcomycteronus brucei, 39.
humerosus, Centriscops, 15.
huttoni, Centrolophus, 19.
Tcichthys lockingtoni, 19,

FISHES—REGAN.

Idiacanthus ferox, 14.

ro niger, 14.
inornata, Lepidoperca, 17.
insignis, Platea, 32.
intermedius, Hemirhamphus, 14.

or

Tluocoetes, 32, 34, 37.

“ elongatus, 32.

7 fimbriatus, 32.
kathleenae, Chionodraco, 12, 28.
Labrichthys ornatus, 40.
lalandii, Seriola, 17.

Lampanyctus, 38.

3 braueri, 38.
larseni, Notothenia, 27, 29, 36.
laticeps, Seyliorhinus, 13.
laticinctus, Lycodalepis, 52.
latitans, Phucocoetes, 32.
lecointei, Nematonurus, 39.
Lepidoperca, 16, 17.

Pr coatsii, 17.

7 inornata, 17.
Lepidopus caudatus, 16.
lepidorhinus, Notothenia, 5.

5 Trematomus, 3, 5, 6, 2
lilliei, Notopogon, 15.
liolepis, Xystreurys, 23.

Lionurus filicauda, 39.
Liparis steineni, 39.

littoreus, Acanthoclinus, 17.
lockingtoni, Centrolophus, 19.

5A Icichthys, 19.
loennbergii, Artedidraco, 7, 8, 28.

5 Trematomus, 2, 3, 27.
longidorsalis, Dolloidraco, 28.
longimanus, Caprodon, 16.
longipes, Notothenia, 27.
Lophiidae, 23.

Lophius piscatorius, 23.
lurida, Ariomma, 20.
Lycenchelys, 32.

" antarcticus, 32, 59.
Lycodalepis laticinctus, 32.
Lycodichthys, 32, 34.

Dy antarcticus, 32.
maclovinus, Eleginops, 27, 37.
macrocephala, Cottoperca, 26.

es Notothenia, 27, 57.
macrolepis, Bathydraco, 28.
macrophthalma, Cottoperca, 26.
macrophthalmus, Hemerocoetes, 18.
macrops, Ophthalmolycus, ows
macropterus, Chilodactylus, 18.

3 Pagetopsis, 11, 28.
macropterygia, Stigmatophora, 16.
Macrorhamphosidae, 15.

lie

4

52

Macruridae, 39.

maculatus, Galaxias, 37.
maculicauda, Serranops, 16.
Malacorhina cirrifer, 21.

3 mirus, 22.
maoricus, Centrolophus, 19.
marionensis, Notothenia, 27, 30,
marmoratus, Muraenolepis, 36.
Maynea, 32, 34, 37.

af antarctica, 32.
* patagonica, 32.
megalops, Squalus, 14.
Melanonus gracilis, 59.
Melanostigma, 32.

.S gelatinosum, 32.
microdon, Cyclothone, 38.
microlepidota, Notothenia, 27.
microps, Hemerocoetes, 18.

PA Muraenolepis, 1, 34, 36.
microstomus, Etropus, 23.
Miolania 43, 44.
owenl, 45.

rs platyceps, 43.
mirus, Artedidraco, 6, 7, 8,
Malacorhina, 22.

», Psammobatis, 22.
mizops, Notothenia, 2

”?

97

Al.

”

7, 36.
mongonuiensis, Arnoglossus, 21.
monodactylus, Chilodactylus, 40.
morenoi, Austrolycus, 32.
Muraena nubila, 14.

5 ocellata, 22.
Muraenidae, 14, 22.
Muraenolepidae, 1, 34, 36, 37.
Muraenolepis marmoratus, 36.
microps, 1, 34, 36.

5 orangiensis, 36, 37.
Myctophidae, 1, 38.
Myctophum, 38.

a antarcticum, 1, 58.
myops, Trachinocephalus, 22.
Nematonurus armatus, 39.

5 lecointei, 39.
newnesil, Trematomus, 2, 27.

”

nicolai, Trematomus, 2, 27.
niger, Centrolophus, 19.

» Idiacanthus, 14.
Niolamia, 43, 44.

5 argentina, 44.
Notolepis coatsii, 38.
Notopogon, 15.
fernandezianus, 16.
liliei, 15.
schoteli, 16.
xenosoma, 15, 16,

”

”

“TERRA NOVA”

36.

EXPEDITION.

Notothenia, 27, 29, 35, 36, 38.
9 acuta, 27, 29, 34.
5

7
a

brevicauda,
canina, 27.
colbecki, 27.

O7

cornucola, 27, 37.

cyaneobrancha, 27, 36.
elegans, 27, 37.
filholi, 27.
gibberifrons, 27, 36.
larseni, 27, 29, 36.
lepidorhinus, 5.
longipes, 27.
macrocephala, 27, 37.

microlepidota, 27.
mizops, 27, 36.
nudifrons, 27, 36.
ramsayl, 27.
rossi, 27.
scotti, 4.
sima, 27.
tessellata, 27
trigramma, 27.
vaillanti, 27.

a wiltoni, 27.
Nototheniidae, 2, 27, 29, 33.
Nototheniinae, 2, 33.
nubila, Muraena, 14.
oblongus, Paralichthys, 23.
ocellata, Muraena, 22.
Ophidiidae, 23.

Ophidium brevibarbe, 23.
Ophthalmolycus, 32, 37.
concolor, 32.
es macrops, 32.
orangiensis, Muraenolepis, 36, 37.
orianae, Artedidraco, 6, 7, 27.
ornatus, Labrichthys, 40.
Osteoglossidae, 41.

oweni, Miolania, 43.

Pagetopsis, 11, 28.

macropterus, 11, 28.

”

”

”
pappenheimi, Cryodraco, 11, 28.
Parachaenichthys, 10, 28.

”
paradoxus, Stylophthalmus, 38.
Paralichthys oblongus, 23.
Paraliparis antarcticus, 13, 59.
Paratrachichthys trailli, 16.
patagonica, Maynea, 32.
patagonicus, Bovichthys, 26.

angustifrons, 27, 30, 36.

coriiceps, 27, 30, 33, 36, 38.

marionensis, 27, 30, 36.

squamifrons, 27, 29, 36.
, 29, 36, 37.

: ; AR OR
georgianus, 10, 28, 33.

pauciradiatus, Cubiceps, 20.

a Hemerocoetes, 18.
pennellii, Trematomus, 3, 4, 27.
percoides, Sebastes, 21.
Phascologale, 42.
Phucocoetes, 32, 37.

. latitans, 32.
piriei, Cynomacrurus, 39.
piscatorius, Lophius, 23.
plagiusa, Symphurus, 23.
Platea, 32, 37.

» insignis, 32.

platei, Austrolycus, 32.
platyceps, Miolania, 43.
Plectranthias, 16.
Plesiopidae, 17.
Pleuragramma, 27, 33, 37.

3 antarcticum, 6, 27.
Pogonophryne, 6, 9, 28, 33.

Ay scotti, 9, 28.
Porichthys porosissimus, 23.
porosissimus, Porichthys, 23.
Prionodraco, 10, 28, 33.

as evans, 10, 28.
Prionotus brachychir, 22.
- tribulus, 23.
Prothylacinus, 42.
Psammobatis cirrifer, 21.

; 29
5 mirus, 22.
5 rudis, 22.

Pseudaphritis, 26, 29.
5 urvillii, 26.
Pseudorhombus boops, 21.
psychrolutes, Bovichthys, 26.
punctatum, Cynophidium, 20.
punctatus, Pyramodon, 20.
punctulatus, Hippocampus, 22.
Pyramodon, 20.
punctatus, 20.
ventralis, 21.

bP)

”
Raia, 36.

» tyllae, 22.
Raiidae, 21.
Racovitzaia, 28, 35.

7 glacialis, 28.
ramsayi, Notothenia, 27.
rhinoceratus, Chaenichthys, 28.
roseopictus, Bovichthys, 26.
rossii, Notothenia, 27.
rudis, Psammobatis, 22.
rugosus, Chaenichthys, 28.
schoteli, Notopogon, 16.
schraderi, Sclerocottus, 34.
Sclerocottus, 34.

oh schraderi, 34.

FISHES—REGAN.

Scleropages, 41.
Scopelarchidae, 39.
Scorpaena, 2

3 cardinalis, 2

Scorpaenidae, 2

scotiae, Bathydraco, 28.
scotti, Notothenia, 4, 27.

5, Pogonophryne, 9, 28.

» Lrematomus, 3, 4, 27.
Seyliorhinidae, 13.
Scyliorhinus laticeps, 13.
Sebastes percoides, 2
Seriola lalandii, 17.
Seriolella antarctica, 40.

3; velaini, 40.
Serranidae, 16, 22.
Serranops, 16.
. maculicauda, 16.

Serranus auriga, 22.
shackletoni, Artedidraco, 7, 8, 28.
sima, Notothenia, 27.
simus, Crepidogaster, 21.
skottsbergii, Artedidraco, 7, 9, 28.
Snyderidia, 20.
Solenognathus spinosissimus, 16.
Sparassodontia, 42.
spilonotopterus, Exocoetus, 15.
spinifer, Zanclorhynchus, 36.
spinosissimus, Solenognathus, 16.
Squalidae, 14.
Squalus megalops, 14.
squamifrons, Notothenia, 27, 29, 36.
steineni, Liparis, 39.
Stigmatophora macropterygia, 16.
Stomiatidae, 14, 38.
Stromateidae, 19.
Stylophthalmus paradoxus, 38.
Sudidae, 38.
Symphurus plagiusa, 23.
Syngnathidae, 16, 22.

Synodontidae, 22.
2

tessellata, Notothenia, 27, 29, 36, 37.

Thylacinidae, 42.
Thylacinus, 42.

Thyrsites atun, 16.
Trachichthyidae, 16.
Trachinocephalus myops, 22.
trailli, Paratrachichthys, 16.
Trematomus, 2, 27, 33, 35, 36 37.
bernacchii, 2, 5, 27.
borchgrevinku, 2,
brachysoma, 2, 3, 2
centronotus, 3, 4, 2
dubius, 2, 27.
eulepidotus, 3, 5,

x

uv

54 «TERRA

2

Trematomus hansoni, 2, 3, 27.
3

ap lepidorhinus, 3, 5, 6, 27.
# loennbergii, 2, 3, 27.
<5 newnesii, 2, 27.
5 nicolai, 2, 27.
3 pennellii, 3, 4, 27.
: ai
5 scotti, 3, 4, 27.
ey vicarius, 2, 27.

tribulus, Prionotus, 23.
Trichiuridae, 16.
Trigla, 21.

ekcumus ile
Triglidae, 21, 22.
trigramma, Notothenia, 27.
urvillii, Pseudaphritis, 26.
vaillanti, Notothenia, 27.
variegatus, Bovichthys, 26.

INOWEAS2

EXPEDITION.

velaini, Seriolella, 40.
velifer, Dolloidraco, 9.

», Histiodraco, 9, 28.
veneris, Bovichthys, 26.
ventralis, Pyramodon, 21.
vicarius, Trematomus, 2, 27.
vorax, EHugnathosaurus, 39.
waitei, Hemerocoetes, 18.
whitsoni, Chalinura, 39.
wilsoni, Chaenodraco, 12, 13, 28.
wiltoni, Notothenia, 27.
xenosoma, Notopogon, 15.
Xystreurys brasiliensis, 23.

és liolepis, 23.
Zanclorhynchus spinifer, 36.

Zoarcidae, 31, 34, 37, 39.

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BRITISH MUSEUM (NATURAL HISTORY).

B BRINISH ANTARCTIC (“TERRA NOVA") BXPRDITION, 1910

3 NATURAL HISTORY REPORT.

e ZOOLOGY. VOL. 1, No. 2. Pp. 55-84.
ey HORLINES wilh

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BY
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Che Trustees

oF

THE BRITISH MUSEUM.

British Museum (Natura? History).

Tica Nig of 25 copies of

“Terra Nova” Natural History of the

Adélie Penguin, printed on Special paper.

or
or

NATURAL HISTORY OF THE
ADELIE PENGUIN.

By Staff-Surgeon G. MURRAY LEVICK, RN.
(Member of the Expedition).

WITH PLATES I-XXI, FROM PHOTOGRAPHS TAKEN BY THE AUTHOR.

CONTENTS.
PAGE
Be iogeephy : ; , ; ‘ 55
- Natural History of the Adélie Penguin eve se ane nae)
1. Arrival at the rookery . : : : : : : ; : 57
2. Mating . ; : ; : 59
3. Nesting, eges, eabation, aetine and aahiine : : 61
4. The games of Adélie Penguins —. : : ; : 70
5. Enemies of the Adélie Penguin. : f : : - : 74
6. Nesting on cliffs : : : : : ; ; ; 76
7. Mortality . : : : : : ; 78
8. Rate of growth and care oy youne . : ; ; Be
9. Departure from the rookery . : ; j ; 80
10. The “ ecstatic” attitude : ’ ‘ : : ; 82

BIBLIOGRAPHY.*

HE habits of Adélie Penguins have been dealt with from time to time by different
iters, but the information to be had from these 1s fragmentary and often misleading.
fact, the only extensive work on this subject that may be accepted as entirely
rrect is that of Dr. Edward A. Wilson, published in Vol. II of the British Museum
sport on the Natural History of the National Antarctic (“ Discovery”) Expedition,
1901-4 (Aves, pp. 36-58).

. a) As the present work deals simply with the habits of Pygoscelis adeliw, and is
veo for those who are already acquainted with the subject, the reader is advised

t in type, under the title “ Antarctic Penguins: a Study of their Social Habits.” (London: William
nemann, 1914).—S. F. H.
I

. Kennan Inetitus7
fx
oA
<
fo Ty 1915

Nee ; y&
< Mational Muse

Pace Dr: Levick has published an account of his observations in popular forms, since these pages were’

56 “TERRA NOVA” EXPEDITION.

to study Dr. Wilson’s more thorough and more scientific pages. Reference may also
be made to :—

‘“ Report on the Collections of Natural History made in the Antarctic Regions
during the Voyage of the ‘Southern Cross.” (British Museum, 1902.)
III. Extracts from the private Diary of the late Nicolai Hanson.
IV. Aves, by R. Bowdler Sharpe.

Information is here meagre, owing to the sad death of the Expedition’s zoologist,
N. Hanson, at an early date. His work was carried on by Mr. H. Evans, another
member of the Expedition, but both Hanson’s and Evans’ notes were handed over to
the commander, and have not been published. Mr. Bernacchi also contributes some
notes in this volume.

“To the South Polar Regions.” (Hurst & Blackett, 1901.)

Beginning on page 190, Mr. Bernacchi tells us a good deal about Adélie
Penguins, some of his observations differmg somewhat from those of the present work.
The photographs are very interesting.

“ Report on the Scientific Results of the Voyage of S. Y. ‘Scotia, 1902-1904,” Part I,
“ Zoological Log.” Scottish Nat. Antarctic Exp., Vol. IV, Zool. (1908).

These notes are brief and incomplete.

“ The Penguins of the Antarctic Regions.” Smithson. Rep. for 1912, pp. 475-482,
with plates 1-9 (1913).

Here Dr. Louis Gain, the zoologist of Dr. Charcot’s Expedition (1908-1910), gives
a brief sketch of the Adélie Penguin’s habits. He proves that some at any rate,
and probably all, return to the same rookeries year after year to breed. This is of
great importance, as it definitely settles the question of the penguins’ “sense of
direction,” which alone guides them over their hundreds of miles of migration. It
is surprising that he names Lobodon carcinophagus and Leptonychotes weddelli as
formidable enemies of the Adélie Penguin. His photographs are particularly good.
The subject is also treated by the same author under the title :—

“ La Vie et les Meurs du Pingouin Adélie.”
1913, pp. 501-521, figs. 1-13 (1914).

With the exception of Dr. E. A. Wilson’s account, referred to above, perusal of
these publications, as well as the casual references to Pygoscelis adeliw to be found
among the other works dealing with antarctic exploration, conveys the impression that
the writers have either lacked opportunity, or have not done full justice to the habits
of these wonderful birds.

IX* Congres Internat. Zool., Monaco,

ADELIE PENGUIN—LEVICK. 5

bt |

THE NATURAL HISTORY OF THE ADELIE PENGUIN
(Pygoscelis adeliae).

1—ARRIVAL AT THE ROOKERY.

On October 13th, 1911, one Adélie Penguin was seen on the rookery at Cape Adare.
As a blizzard came on then, with thick drift, nothing more could be seen until the next
day, when no birds were visible.

On October 15th two were walking about the beach: they were separate in the
forenoon, but later they kept company, and hung about the south-east corner of the
rookery, under the cliff, where they were sheltered from the cold wind that was blowing.

On October 16th, at 11 am., about twenty penguins arrived, all remaining very
inactive, and quietly sitting or walking about. By 4 p.m. their numbers had increased
to about a hundred, and for the most part they still squatted listlessly about the
rookery. They were well scattered, some being solitary, and others in small groups,
but all keeping to the hollows and making no attempt to start work on the nesting sites
up on the knolls. There was no sign of any pairing taking place. All the birds were
in fine plumage and condition.

During the night of October 16th the number of arrivals increased greatly, so that
on the morning of the 17th there was a thin sprinkling scattered over the rookery, a
few in pairs or threes, but most in groups of a dozen or more, and all were very
phlegmatic,.lying on their breasts with outstretched beaks, apparently asleep, and still
away from the nesting sites.

During October 17th the arrivals became very much more frequent, and soon, as
we looked across the sea-ice to the northwards, we could see a long line of Adélies
approaching, tailing out in snake-like fashion, as far as the horizon (Pl. 1). On this
date some few birds took possession of old nests on the higher grounds. They merely
squatted in these, making no attempt to repair them in any way, and afterwards I|
found that they were unmated hens, waiting for mates to come to them, and that this
was the usual custom among them.

About 9 p.m. a light snowstorm came on, and those few who had taken possession
of nests left them to join the other birds in the hollows. One group, which had
arrived at the ice-foot in the morning, halted on the sea-ice, without ascending the little
slope leading to the rookery, and stayed there all day.

From their general appearance and behaviour it was evident that some of the
penguins were fatigued after their long journey, and although when the weather cleared
on October 18th a fair number paired and started to build nests, the great majority still
sat quietly about.

The first birds to build all took their stones from old nests, as so many lay
unoccupied at this time.

58 “TERRA NOVA” EXPEDITION.

In every case where I was able to distinguish between the cock and the hen,
the latter remained on or by the nest, and built it with the stones which the cock
brought her, and I think the work is always divided between them in this way.

After watching very carefully for some days, I came to the conclusion that
pairing never takes place before the birds arrive at the rookery. All those in the
long line that approached the breeding ground seemed quite independent of one
another.

During the march across the sea-ice on their way to the rookery, they use
both the methods by which a penguin travels over solid ground; viz. : by walking
and tobogganing (PI. II). It was most interesting to walk some way out along the
approaching line, and then to stand and watch it stream past.

First would come a string of birds toddling along, their little legs enabling
them to advance only about six inches at each step; but, goimg at the rate of about
130 steps per minute, they covered some two-thirds of a statute mile per hour. In
the still aw their little wheezy respiration could be heard distinctly, and they
seemed to be somewhat out of breath.

Close on their heels would come another string, tobogganing on their breasts,
and using their legs as propellers, the rate of progression being exactly the same as
when they walked, so that the procession kept its formation as it passed over the
ice. Every now and then those that walked would flop forward on to their breasts
and toboggan, and those that tobogganed would as suddenly pop up on to their legs
and walk.

On October 19th nest-building was in full swing, with all the squabbling and
thieving that has so often been described by different writers.

Some of the birds seemed inveterate thieves, and collected most, if not all, of
their stones in this way. Depredators, when caught, were driven off with great fury
by those whom they attempted to rob, and were sometimes chased for some distance.
It was curious to see the difference in the appearance of the fleeing thief and his
pursuer. As the former raced away among the nests, doubling on his tracks, and
attempting by every artifice to get lost in the crowd, and so rid himself of his pursuer,
his feathers lay close back on his skin, giving him a sleek aspect which made him
appear half the size of the irate bird who sought to catch him, with feathers ruffled in
indignation.

To the human eye there is not the slightest difference in the outward appearance
of the cock and the hen. Even when a particular pair have been watched for many
days, so that they are quite familiar to the observer, not the least distinction can
be seen, though after some experience differences in their behaviour may be noted
at once, whilst the muddy marks of the cock’s feet on the hen’s back, when these
are present, serve to distinguish one from the other at a glance.

Without cessation the stream of arrivals continued until October 30th, when it
became intermittent, and two days later had stopped altogether. By this time the

ADELIE PENGUIN—LEVICK. 59

rookery was crammed with nesting birds to the number of some three-quarters of
a million, the occupants in many places spreading right up the cliff of Cape Adare,
a large colony having established itself at the very summit, at a height of some
1,000 feet above the sea.

2.—MATING.

As I have already said, mating takes place after the birds arrive at the
rookery, and the following customs are prevalent.

Usually the hen establishes herself on an old nest, or, very often, scoops out
a hollow in the ground, and sits in this, waiting for a cock to propose himself (PI. IT).

Whilst so waiting, she makes no attempt to build. At the beginning of the
season, when the great stream of arrivals continually pours into the rookery, she
does not have long to wait. Later, however, as the rookery gets filled up, and
comparatively few birds in that vast crowd remain unmated, her chances are naturally
not so good.

For example, on November 16th there was a hen so established in a little
scoop she had made in the ground. Looking thinner and more forlorn as day after
day passed by, she made no attempt to build a nest, until November 27th, when a
cock was found to have joined her, and the little scoop she had made more than ten
days before was lined with stones which she was placing in position, as her mate, steadily
at work, brought them to her one by one. Her forlorn appearance of the past ten days
had entirely given place to an air of occupation and happiness.

Wishing to learn more of the penguin’s mating habits, on October 23rd I went to
the place where the stream of arrivals entered the rookery, and singling out one
particular bird, which afterwards proved to be a cock, followed him to see what he
would do.

Threading his way through nearly the whole length of the rookery, he avoided the
tenanted knolls where the nests were, though occasionally he seemed to glance up at
them in passing.

Every hundred yards or so he stopped, ruffled his feathers, closed his eyes and
remained so for about half a minute, when he would rouse himself and continue on his
way. Evidently he was tired after his long journey and struggling against a desire to
sleep.

Arrived at length at the southern end of the rookery, he seemed to make up his
mind, and boldly ascending a knoll which was well tenanted and covered with nests,
walked straight up to one of these on which a hen sat.

A cock stood by her side, but the newcomer either did not see him, or refused to
notice him. Sticking his beak into the ground in front of the nest, then lifting his
head, he made as if to place an imaginary stone in front of the hen: a most obvious
piece of dumb show.

She took not the slightest notice of him, neither did her mate, to my surprise.

60 “TERRA NOVA” EXPEDITION.

Abandoning this quest, he went straight up to another couple who were established
near by, but seeing him coming, and evidently knowing what he was after, the cock
immediately flew at him, and after a sharp fight im which each used his flippers
energetically, the interloper was driven down the side of the knoll and away from the
nests, when the victor returned to his hen. With the persistence of his kind, the
newcomer came straight back to the nest, but weariness seemed to overcome him, for
he settled himself for a doze, in which he continued until I was too cold to await
further developments.

Little knots of cocks were to be seen about the rookery thirsting for battle, and
keeping a jealous guard on each other’s movements.

Seeing that this was gomg on all around, one day I took out my camera and
selected a typical case for illustration.

A group of three cocks were engaged in bitter rivalry round a hen who was
cowering on her scoop, in which she had been waiting. She appeared to be bewildered
and agitated by the desperate behaviour of the cocks. In a further development of the
scene, two of the cocks are shown squaring up for battle (Pl. IV). The combatants,
hard at it, used their weight as they leant their breasts against one another, whilst
they rained in the blows with their powerful flippers. At the end of the fight the
victor rushed the vanquished cock before him, out of the crowd on to a patch of snow,
on which he held him down and gave him a terrible hammering. When his conqueror
at length left him he lay for some two minutes or so on the ground, his heaving
breast alone showing that he was alive, so completely exhausted was he ; but recovering
himself, he at length arose and crawled away, a damaged flipper hanging limply by
his side, and he took no further part in the proceedings.

It is only occasionally that one sees a cock so thoroughly beaten as this. As a
rule when one of the combatants is too exhausted to go on, or has been knocked down,
the other is content, and runs back to the hen or to the other cocks who may be
gathered near her. When he has vanquished these, or got rid of them in some way,
he generally goes up to the hen, who then feels or feigns some reluctance to take him,
and I have seen a victorious cock run up to the hen’s scoop and squat in it whilst she
pecked him cruelly, he merely hunching himself up with closed eyes, until she desisted,
after which, with pretty overtures and soothing sounds, he pacified her, and soon they
appeared to be mated permanently.

As time went on, and the proportion of unmated to mated birds became smaller
and smaller, the cocks watched each other more jealously, and began to go about in
little batches in consequence, squabbling and fighting continually, and hindering one
another in the quest for mates.

Desperate as their encounters are, | think that one penguin never kills another.
In many cases blood is drawn. I saw one with its eye put out, and that side of its
beak (the right side) covered with blood, whilst the crimson mark of a blood-stained
flipper across a white breast was no uncommon sight.

ADELIE PENGUIN—LEVICK. 61

It is evident that during the first days of his wedded life a cock only keeps his
mate by dint of constant vigilance and some further battles; I often saw errant cocks
making overtures to mated hens. I do not think, however, that this ever takes place
after the eggs are laid and regular family life has begun.

With red paint I marked the breasts of a good many couples, renewing the paint
as it faded, through the whole breeding season, and in every case the couples remained
perfectly faithful to one another.

3.—NESTING, EGGS, INCUBATION, FEEDING AND FIGHTING.

Just outside our hut door our geologist left some chips of white quartz lying on
the ground. Shortly afterwards, I found two of these chips in a nest about thirty yards
away, and they showed up brightly and distinctly against the black basalt of which all
the pebbles on the rookery are composed. As a rule the penguins are careful to select
rounded stones for their nests (Pl. V), and as these fragments of quartz were jagged and
uncomfortable, they were most unsuitable for nest building, and it was evidently the
brightness of the stones which attracted them. As I looked on, the owners of the
pieces of quartz were wrangling with some neighbours, and a penguin in the nest
behind them shot out its beak and stole one of the pieces, which it placed in its own
nest. Later, both pieces were stolen from nest to nest, till I lost them.

This incident suggested an experiment which I tried immediately. I painted
some pebbles a bright vermilion, and had others covered with a bright green material,
as I had no other coloured paint. Mixing a handful of these coloured stones together, I
placed them in a little heap near a nest-covered knoll. Some hours later, | returned to
find nearly all the red stones and one or two of the green ones gone, and afterwards
found them in nests. Later still, the rest of the red stones vanished, and, last of all,
the green ones.

All these coloured stones were taken to nests, and some time later, like the pieces
of quartz, they were stolen from nest to nest, and soon were distributed widely in all
directions.

On other occasions I saw pieces of tin, pieces of glass, half a stick of chocolate, and
the head of a bright metal teaspoon in various nests near our hut, the articles evidently
having been taken from our scrap heap. It then became evident that penguins like
bright colours, and they seem to prefer red to green, as instanced by the selection of
the coloured pebbles.

Though most nests are built of a mixture of pebbles varying in size from very
small to as big as the birds can carry, some individual couples make theirs entirely with
very big stones, and some entirely with very small ones (PI. VI); and a large stoned
nest and a small stoned nest may be seen side by side, in places where pebbles of all
sizes may be collected. Some of the penguins choose stones so large that it is a matter

62 “TERRA NOVA” EXPEDITION.

for wonder how they can lift and carry them. As a rule, fairly large stones are chosen,
their comparative sizes being well shown in some of the photographs.

In selecting sites for their nests, the penguins at Cape Adare have shown a very
remarkable instinct.

The beach on which is the rookery is raised into a series of undulations and knolls,
and some of the lower lying ground is covered by little lakes of thaw water, rendered
slimy and muddy by the guano that is blown into it by the frequent gales.

Rising from one of these lakes is a big knoll that appears to be in every way
suitable for nesting. When the birds arrive at the rookery, and for the first half of the
breeding season, the water of the lakes is frozen hard, and like most of the solid ground
is covered by a thin layer of wind-blown snow, so that the surface of the ice is scarcely
to be distinguished from the land, and the island knoll in question is perfectly accessible.
Yet not a nest is to be seen on it, nor any sign of an attempt having been made in
past years to build one.

It is evident from this that the penguins realise that in some six weeks’ time they
will only be able to reach the spot by wading through the muddy water. They not only
realise that there will be water where the solid ice now is, but that it will be slimy, and
make them in a mess, and accordingly the island is taboo. It is not that they object
to fresh water, as in other places they are frequently to be seen wading through clean
fresh-water pools.

Not far from the above-mentioned islands there is another mound rising from the
lake, but connected with the mainland by a narrow pathway of stones rising Just above
the water. This mound is covered with nests. While the lake is frozen they approach
it across the ice ; later, when this thaws, they have the narrow path by which to reach
their nests without covering themselves with mud.

The barren island was a very distinet feature of the rookery when the available
ground was inconveniently crowded with nests.

Where selection has brought about such remarkable uniformity as is to be seen in
the whole species, individual traits of character are hardly to be expected in any great
degree, but some differences exist nevertheless.

The sizes of the stones selected for nests by different birds have already been
commented on. Also, some of them prefer to build their nests at a great height,
whilst others prefer the lower ground. This is evident, as both high and low sites are
chosen when there is still ample room on the flat beach as well as up the cliff of Cape
Adare, to a height of 1,000 feet ; yet in order to get at many of the nests up the cliff,
their occupants have to make frequent journeys involving a difficult and arduous climb
to the top of the cliff, and then a walk across a precipitous snow slope hanging over the
brink of a clear drop of many hundreds of feet on to the ice below.

Again. some have less alertness and general force of character than others, a
difference evidenced in a striking manner in some of the little colonies.

Nests are seen which are incessantly losing stones from their walls owing to the

ADELIE PENGUIN—LEVICK. 63

pilfering of neighbours, any lack of vigilance on the part of their occupants being at
once noted and taken full advantage of, and sometimes, though rarely, weak characters
allow their more aggressive fellows to huff them of their stones before their very eyes,
without offering any resistance. As fast as they acquire stones and place them in
position, these are taken from them, the actual floor on which the hen sits being the
only portion of their home which remains permanent, whilst a more aggressive
neighbour sits upon a noble pile, not a pebble of which is ever stolen.

If watch is kept on a colony busily engaged in building their nests, some of the
cocks are seen to make fairly long journeys for stones, working steadily forth to and
back from some patch of loose ground where building material is to be had.
Occasionally, however, a cock may be seen to steal a// his stones from neighbouring
nests, craftily making his way among these until he sees his chance, when he quickly
grabs a pebble, and triumphantly returns to his mate with the spoil in his beak. Again
and again he will return to the same nest, of which most probably the cock is away on
a similar quest, while the hen has all her attention taken up in pecking at the
head of the lady sitting nearest to her.

Here, then, are seen, side by side, a cock who prefers the quieter but more
laborious method of gathering his own material, and another who chooses the more
exciting but easier method of stealing it. The latter, of course, is not always

successful, being occasionally caught in the act, when he may he seen tearing off

with an irate bird in pursuit, often tripping up over the stones and other inequalities
of the ground, and getting a good hammering when caught. In a small colony
which I had under special observation there was a particular cock who was an
inveterate thief. I think that every stone on his nest had been stolen from
neighbours. As he slunk about the colony, his guilty conscience made him smooth
his feathers close against his skin, and this made him look much smaller than the
other birds, who seemed to know him, as most of them aimed a peck at him as he
passed their nests.

General aggressiveness is a quality found in different degrees in different
individuals.

Close to the path from our hut to the meteorological screen a good many
couples had made their nests, and consequently our meteorologist passed them many
times a day on his way to take his observations. There was one particular cock
that invariably flew at him, whenever he happened to be at the nest, though the
others took little or no notice when they were passed.

When making observations of the incubative period of the eggs, and other
matters, there were certain nests I visited twice in each twenty-four hours, always
gently lifting the hen for a moment to examine the nest beneath her.

Some of the hens pecked and swore at me savagely, seeming never to get used
to the operation, while others minded scarcely at all.

One hen whose chick I removed at intervals in order to weigh it, instead of

K

64 «TERRA NOVA” EXPEDITION.

getting more used to the process as time went on, became more exasperated with
me each time I went to her nest, though I was very gentle indeed, both with her
and the chick, and as she saw me approach, when I was still some way oft, would
rise from the nest and ruffle her feathers, trembling with indignation, and making
a great noise. I always gave her a fur mit to peck at while I temporarily borrowed
her chick.

The instinct which causes a bird to procure food and bring it to the nest for
its offspring is, as we know, common to all species, this duty being equally shared
by both cock and hen, im the case of the Adélie Penguin, from the time when the
chicks first appear.

On one occasion, however, I noticed an unusual trait of character on the part
of a cock. As the season advanced and the sun’s altitude increased, almost all the
snow which had covered the rookery during the early spring disappeared owing to
thawing and ablation, leaving the ground bare and dry.

Whilst the snow remained the birds, as they sat on their eggs, used to quench
their thirst by gobbling that which lay within reach. Afterwards, however, they
seemed to suffer greatly for want of water, and were to be seen panting with their
beaks open and tongues exposed. Those who were not engaged in incubating used
to journey frequently to the various drifts that still remaimed, gobbling great
quantities of snow ; one drift in particular, which had formed in the lee of our hut,
being visited by crowds who came incessantly to quench their thirst.

One day a cock was seen to pick up a lump of snow in his beak and carry it
a considerable distance to his mate as she sat on the nest. He deposited it on the
ground in front of her, and she ate it at once. When I mentioned this to
Mr. Priestley, he told me that when he was at Cape Royds with Sir Ernest
Shackleton’s Expedition, he had seen the same thing occur. As this was seen only
once at Cape Adare, it is evidently a very rare occurrence, and I mention it here as
a characteristic development possessed by only a few individuals. The cock, when
away from his mate, evidently had in his mind the fact of his hen being thirsty
and unable to get snow as he could.

Owing to our having had several snowfalls without wind, and to the action of
the sun’s rays falling through that clear atmosphere on to the black rock, there were
in some places masses of slush and then actual floods as the thaw water trickled
down into the hollows. Some of the penguins having made their nests in these low-
lying positions, these were threatened with destruction by the floods (Pl. VII). Here
the occupants were to be seen doing everything they could to avert this calamity,
and from each nest the cock worked busily, making journey after journey in quest
of stones, with which the hen built the little castle higher and higher, and so kept
the eggs above water, so that some of the pools were dotted with little islands on
which the hens sat.

[ noticed one nest in particular, by the side of a pool, which still remained a

ADELIE PENGUIN—LEVICK. 35

6d

foot or so clear of the water and on dry ground, but nevertheless its inmates quite
realised the emergency that threatened them, and that it must be provided against,
and for hours the cock passed to and fro, wading across the little lake to the far
side, from which he got his stones.

This scene is shown in Pl. VII. In the right hand corner of the picture the
cock is seen in the act of delivermg another stone to the hen, who is waiting to
receive it, whilst some of the nests are seen actually surrounded by water.

Here I quote from my notes :—

“Nov. 10th. This evening I saw a hen penguin trying to sit on a nest with
two eggs. The nest had no stones, as they had all been stolen by neighbours, and
as it was scooped deeply in the ground, and in a slush of melting snow, the eggs
were nearly covered with water. The poor hen stood in this, and kept trying to sit
on the eggs, but each time she did so, sat in the water and had to get up again.
She was shivering with cold and all bedraggled. I took the two eggs out of the
nest, and Browning and I collected a heap of stones (partly from her richer neighbours !)
and built the nest well up above the water. Then I replaced the eggs, and the hen at
once gladly sat on them, put them in position, and was busily engaged in arranging
the new stones around her when we left.”

When one egg has been laid the hen still sits on the nest, as the egg would
be frozen if it were left uncovered, besides which skua gulls are always ready to
pounce on any that lie exposed, and it is not until the second egg is laid that she
goes to feed, and the cock takes his turn on the nest for the first time.

As nearly as I could ascertain, pairing began on October 16th or 17th, and we
found the first egg on November 3rd.

In order to determine the period between the laying of the two eggs, | numbered
seven nests with wooden pegs, writing on the pegs the date on which each egg was
laid, and the following was the result obtained :—

Date of appearance Date of appearance
of first egg. of second egg. Interval.

No. 1 nest. . . November 14th = Only 1 laid.
No. 2 nest. . . November 13th November 16th 3 days.

No. 3 nest. . . November 14th November 17th 3 days.
INoy4enéshs = =

No. 5 nest. . . November 12th November 16th 4 days.

No. 6 nest. . . November 8th November 12th 4 days.

No. 7 nest. . . November 24th — Only 1 laid.

the average interval in the four cases where two eggs were laid being 3°5 days.
The only notes I have on the lapse of time between the laying of the egg and the
hatching of the chick are that the first chick appeared in No. 5 nest on December 19th
K 2

66 “TERRA NOVA” EXPEDITION.

(37 days), and in No. 7 nest on December 27th (33 days), whilst in another case,
in which observations were most carefully made, the period was only 31 days. This
seems to show that the incubation of the embryo does not invariably begin as soon
as the egg is sat upon, as the observations were most carefully made and recorded,
each nest, as I have said, being marked and visited daily.

The following table enables a comparison to be made between my own results and
those of Wilson and Bernacchi :—

Wilson. Bernacchi. Levick.
First egg laid : . 5 : Middle of November Nov. 2nd Noy. 3rd
Chick hatched. é : : Middle of December Dec. 9th Dec. 4th
Nestling’s down moulted : : Jan. 9th to 16th.

Not until the eggs have been laid does either of the birds go to feed. Then one
of the pair goes off to the water for this purpose, and stays away in many cases
for some days (about 7 to 10), after which it returns to relieve the other, who goes
off for about the same period. Then, when the chicks are hatched, they relieve one
another at more frequent intervals, as seen by the time-table given on pp. 67, 68.

A most astonishing fact is the long fast which the birds undergo between
their arrival at the rookery and the hatching of the chicks, the shortest period of
this total abstinence being about 18 days; but as the first of the pair to go off
to feed remains away in many cases for ten days, the other must fast for about
28 days.

This fact, occurring as it does during the most arduous period of the penguins’
year, furnishes a most surprising proof of the wonderful endurance possessed by these
birds. They arrive tired after their journey to the rookery. In the case of the
cocks, they go through a long period of repeated battle and continual anxiety. They
propagate their species, and work to gather stones for their nests, yet for 28 days
they eat nothing at all, and at the end of that time, though dirty and bhedraggled,
they seem little the worse for it.

The reason for this fasting is, perhaps, partly that they dare not leave their nests
unprotected during the early part of the season, when building is in progress all
around, and stones are in great request ; and also because, as I have already remarked,
until the eggs have come, strange cocks frequently make overtures to hens who are
already mated, and thus the cock can only ensure the safety of his home by his
constant protection.

Consequent on the penguins’ abstinence from food, no guano is deposited about
the rookery until the eggs have been laid, and so the brick-red colouring of the rookery
described by previous authors is not seen during the early part of the season.

Instead of this, bright green watery excreta are dropped, consisting of bile, water,
salts, and epithelial cells (the bile of penguins is bright green). The bird as it sits on
its nest never fouls this, but squirts its excreta well clear of its walls. This taking

ADELIE PENGUIN—LEVICK. 67

place in different directions as the bird shifts its position, the nest assumes the
picturesque appearance of a bright green flower, as the penguin adds its decorative
offerings petal by petal. °

Even after the chicks have come, the nest is kept clean, as they lie with their
heads towards the centre and their little sterns protruding.

As soon as feeding began again, the whole appearance of the rookery changed, and
from green, its colour was changed to brick-red, owing to the colouring-matter derived
from the HKuphausia on which the Adeélies entirely subsisted.

The manner in which the incubating and nursing duties are shared, in order to give
each of the birds a periodical chance to go and feed, has been well described by former
writers. I kept a watch on a good many couples, and their arrangements seemed to

73

tally very closely with the following “ watch bill” kindly kept for me by Mr. Priestley.
As the nest in question was close to the meteorological screen, Mr. Priestley
passed it several times during the day as well as in the night, on his way to take his
observations ; and from his notes, so obtained, I made out the following time-table.
Only one egg was seen.
Nov. 14th—Ege laid. Hen sitting.

, 27th—A cock seen to join the hen for the first time since the 14th. He
took her place on the nest. This was the first day on which any
red guano was seen about the nest.

Dec. 10th.—The hen returned between 8 p.m. and 10 p.m., having been absent
since Nov. 27th. Fresh red guano: the first for many days.
14th.—The cock relieved the hen between 8 a.m. and 10 am. Between
6 p.m. and 8 p.m. the chick was hatched, the hen remaining on
the nest.
» 17th—At 8 am. the cock was found to have relieved the hen.
, 18th.—Hen mounted guard between 6 p.m. and 8 p.m.
, 20th—Cock relieved guard about 8 am. At 8 p.m. both cock and hen
were at the nest, the hen standing by it, the cock on it.
,, 21st.—The hen relieved guard at 8 p.m.
,, 28rd.—Cock came back at noon and relieved guard.
,, 24th.—The cock remained on guard all day. The hen was gone from 1 till
6 p.m., when she returned and relieved guard.
. 25th—8 a.m. Both at nest, hen still on. 10am., changed guard. Hen
gone.
., 26th.—Hen on nest. Cock standing near.
» 27th—8 a.m. Cock on nest.
, 28th—8 am. Hen on nest.
», 29th.—Cock relieved guard.
,, 30th.—Hen arrived at 3 p.m., and relieved guard.
, 3lst.—10 p.m. to midnight, changed: cock on. Both there at 10 p.m.

68 “TERRA NOVA” EXPEDITION.

Jan. Ist.—10a.m., both at nest.

12 noon, both at nest: the youngster complicating matters by
running away every time he was passed by the observer, thus
getting himself and his parents embroiled with their neighbours.

2 p.m. Hen on nest: cock gone.

= 2nd.—10a.m. Hen on nest.

12 noon. Chick disappeared.

2p.m. Nest deserted.

4p.m. Cock on nest: no chick.

xs 3rd.—Cock on nest, with chick.

From the above table it will be seen that the hen was not relieved by the cock
until a fortnight after she had laid her egg, so that she must have been without food
fora month. Then she left, and only returned to relieve the cock after the lapse of
another fortnight, it being worth remembering that each was absent for the same
length of time.

On the appearance of the chick, a different régime began, the chick having to be
fed, and journeys made at regular intervals for the purpose of getting food. All over
the rookery, as the young birds appeared, there was a marked change in the appearance
of the parents as they came up from the water.

Hitherto they had been merely remarkable for their spotless plumage, in contrast
to their former dirty state, but now their shape too was greatly altered, for their meals,
in place of merely satisfying their own individual wants, had now to provide for the
offspring as well, and they were in consequence so distended with their heavy load of
Euphausia that they were obliged to lean back to counterbalance the weight of their
bellies that bulged before them as they walked. Frequently they would find to their
cost that they had attempted too much, and overcome by the labour of their journey
over the rough ground, they would be sick, depositing the whole load on the ground, and
having perforce to return to the sea for more. Little red heaps of mashed and partly
digested Huphausia were consequently to be seen about the rookery. Once I saw a
diligent parent, who, having actually arrived at the nest, could not contain himself
long enough for the chicks to help themselves in the manner common to them—of
thrusting their heads down his throat—and vomited the meal on to the ground. Seeing
what was coming, I had my camera ready, and one of my photographs, not here
reproduced, represents him in the act. The dismay shown by the hen, when she
saw what was taking place, is indicated plainly by her expression.

Neither chicks nor adults ever attempt to eat food of any sort from the ground,
the chicks always feeding directly from the throat of the parent (Pl. VIII), and
the adults always from the water, and hence these little heaps of vomited food are
invariably wasted.

Near our hut was a scrap heap, on which were thrown remains of all kinds of
food, and the penguins’ hatred of the skuas (Pl. IX) was well shown here.

ADELIE PENGUIN—LEVICK. 69

None of the food was of the least use to the penguins, but we noticed after
a time that one or two penguins were almost always there, guarding the heap against
the skuas. In fact, a constant feature of this heap was the sentry penguin, making
little runs hither and thither at the skuas, who would then simply rise a yard or
two into the air out of reach, the penguin squalling in its anger at being unable to
follow its enemy. At this time, the penguin would imitate the flying motion with
its flippers, seeming instinctively to attempt to mount into the air as its remote
ancestors did before their wings had been adapted solely to swimming.

Close to the scrap heap there was a large knoll crowded with nests, and it
was this colony which supplied the sentries. Very rarely did one of these leave the
heap until another came to relieve it, so long as there were skuas about, but when
the skuas went so did the penguins. The instant the skuas returned, a penguin
would be seen to run from the knoll to the heap. It seemed that there was some
primitive understanding about the matter amongst the penguins, as there was never
a crowd of them on the heap, the rest appearing satisfied as long as one of
their number remained on euard.

As custodian of the nest, there is no doubt that the hen is very much more
efficient and reliable than the cock. When the former is doing duty on the eggs,
no ordinary circumstance induces her to leave them for a moment. She wrangles
very frequently with her immediate neighbours, and she and they spend hours on
end in pecking at each others’ heads, but this only happens between those who can
reach each other without leaving the nest. The cocks, on the other hand, behave very
differently. Starting to squabble, they wax hotter and hotter, and frequently end by
leaving their nests and going for one another in a proper battle with their flippers,
fighting backwards and forwards over their nests and often scattering the eggs, large
numbers of which are lost annually in this way.

On occasions we saved eggs by replacing them in the nests and stopping the
fight, when the combatants would quickly forget the quarrel, and again settle down
to their duties.

Two cocks, fighting like this in the midst of a crowded colony, were a danger
to their neighbours, as they not only incommoded them by bumping into them and
fallme over them, but were apt to cause misunderstandings that ended in further
dissension. Perhaps the other birds realised this, as they would evince every sign
of anger when two of their number started a fight.

On this subject I find a note in my diary for November 24th, 1911, which
seems important, and I give it word for word.

“This afternoon I saw two cocks (probably) engaged in a very fierce fight
which lasted a good three minutes. They were fighting with flippers and bills, one
of them being particularly clever with the latter, frequently seizing and holding his
opponent just behind the right eye, whilst he battered him with his flippers. After
a couple of minutes, during which each had the other down on the ground several

70 ; “TERRA NOVA” EXPEDITION.

times, three or four other penguins ran up, and apparently tried to stop the fight.
This is the only construction I can put on their behaviour, as time after time they
kept running in when the two combatants clinched, pushing their breasts in between
them, but making no attempt to fight themselves, whilst their more collected
appearance and smooth feathers were in marked contrast to the angry attitudes of
the combatants.

“The fight which had started on the outskirts of a knoll crowded with nests,
soon edged away to the space outside, and it was here that I (and Campbell, who
was with me) saw the other penguins try to stop it.

“The last minute was a very fierce and vindictive ‘mill, both fighting with
all their might, and ended in one of them trying to toboggan away from his opponent,
but he was too exhausted to get any pace on, so that just as he got into the
crowd again he was caught, and both fought for a few seconds more, when the
apparent victor suddenly stopped and ran away.

“The other picked himself up and made his way rapidly among the nests,
evidently searching for one in particular. Following him, I saw him run up to a nest
near the place where the fight had begun. There was a solitary penguin waiting
by this nest, which was evidently new, and not yet completed, being without eggs.

“The cock I had followed, ruffed and battered with battle, ran up to the
waiting bird, and the usual side-to-side chatter in the ecstatic attitude began,
and continued for half a minute, after which each became calmer, and I left them
apparently reconciled, and arranging stones in the nest. This incident was after the
usual nature of a dispute between two mates for a hen, but the pacific interference
of the other birds was quite new to my experience. That it was pacific I am quite
convinced, and Campbell agreed with me that there was little doubt of it. All the
nests about had two eges under incubation, and the pair in question must have
been newcomers.”

The above note, | am afraid, gives a rather meagre impression of an astonishing
scene, of which I have a very distinct recollection.

As the two birds fought, several couples stood around them, who from time to
time turned to one another, making sounds and gestures as if they were arguing some
point, when one of them would turn and run in between the opponents, literally
pushing itself in between them, and though striking no blow itself, doing everything
possible to hinder them. As is often the case during a fight, many others looked on
with apparent interest in the proceedings, pausing in their own affairs to do so.

4—_THE GAMES OF ADELIE PENGUINS.

That Adéhes have developed a taste for playing certain primitive games seems
perfectly evident. They never play on the ground of the rookery itself, but on the
sea-ice, on their way to and from their bathe.

,ADELIE PENGUIN—LEVICK. ral

For a great part of the season there was some half mile or so of this sea-ice
intervening between the rookery and the open water, and this stretch presented a lively
scene as the thousands of birds, in parties, passed to and fro over it, the outward bound

parties of dirty birds from the rookery passing the spruce bathers, homeward bound
after their meal and frolic in the sea.

To watch these parties was interesting and instructive. On leaving their nests
the birds made their way down to the ice-foot (Pls. X, XVI, XVII) on ie themueacice:
Here generally they would wait about and join up with others until enough had
gathered together to make a little party, which would then gaily set off for fin: water.
They were now in the greatest possible spirits, chattering loudly and frolicking with
one another from time to time, playfully chasing each other about, or indulging in a
little friendly sparring with their flippers. Arrived at length at the water's edge, the
same procedure was almost always gone through.

The object of every bird in the party seemed to be to get one of the others to
enter the water first, and they would crowd up to the very edge of the ice, trying by
various tricks to push each other in. As a closely packed bunch of birds stood on
the brink, those behind would try to rush the front rank over, who then, barely
recovering themselves, would rush round to the rear and endeavour to turn the tables
on the others. Occasionally one would actually get pushed in, only, by a lightning
turn under water, to bound out again on to the ice, with a movement so quick as to
resemble an indiarubber ball hitting the water and bouncing out again.

Then for some time they would chase one another about, seemingly bent on
having a game, but each bird intent on avoiding being the first in. Sometimes this
would last a few minutes only, but often for an hour or so, until suddenly one of
the number started to run at full tilt along the edge of the ice, the rest following
closely on his heels, until at last he took a clean header into the water. One after
another the rest of the party followed him, each one taking off from the spot where he
had entered, and following one another so quickly as to have the appearance of a lot
of shot poured out of a bottle into the water. The accompanying photographs
(Pls. XI, XII) show this latter scene being enacted. Then for a few seconds not a
ripple would be made, and a dead silence ensue, till they would all appear some twenty
or thirty yards out and start rolling about and splashing in the water, cleaning them-
selves and making a noise that resembled a lot of boys calling out and chatting one
another. Seemingly reluctant as they had been to enter the water, when once there
they evinced every sign of enjoyment, and would stay in for hours at a time.

From what I have said it will be seen that the penguins spent a great deal of time
on their way to and from the water, especially during the earlier period before the
sea-ice had broken away from the ice-foot, as they had so far to walk before arriving
at the open leads.

As a band of spotless bathers, returning to the rookery, their white breasts and
black backs glistening in the sunlight, met a dirty and bedraggled party on its way

L

*

~1
bo

“TERRA NOVA” EXPEDITION,

from the rookery, frequently both would stop, and the clean and the dirty mingle
together and chatter with one another for some minutes. If they were not speaking
words in some language of their own their whole appearance belied them, and as they
stood, some in pairs, some in groups of three or more, chatting amiably together, it
became evident that they were sociable animals, glad to meet one another, and like
many men, pleased with the excuse to forget for a while their duties at home, where
their mates were waiting to be relieved for their own spell off the nests. After a
variable period of this intercourse, the two parties would separate and continue on their
respective ways, a clean stream issuing from the crowd in the direction of the rookery,
a dirty one heading off towards the open water, but here it was seen that a few of
those who had bathed and fed, and were already perhaps half-way home, had been
persuaded to turn and accompany the others, and so back they would all go, over the
way they had come, to spend a few more hours in skylarking and splashing about in
the sea.

On this strip of sea-ice the penguins would spend hours, and, gathered in small
parties, play a sort of ‘touch last.” Games of this sort are often seen to be played by
the young of different species of mammalia, but I believe that among birds they are, at
any rate, uncommon.

Another very favourite game of theirs is worth recording.

The tide in the vicinity of the rookery flowed at a considerable rate (some six
knots at times), and on it there drifted a succession of ice-floes of different sizes. As
one of these floes arrived at the top end of the rookery it would be boarded by a party
of Adélies, who sometimes crowded on to it until it would hold no more. The
“excursion boat,” as we got into the habit of calling it, borne by the stream, would
then drift the whole length of the rookery, its occupants showing every sign of
enjoyment in the ride. Sometimes they stood silently contemplating the scenes on the
ice-foot as they were borne past them; at others, especially when passing close to the
shore, they would shout remarks to the other penguins that stood on the ice-edge, who
would in turn shout back at them, so that a running fire of chaff seemed to pass
between those on the bank and those on the floe as the crowded “ excursion steamer ”
passed on its way.

Occasionally a knot of penguins who stood hesitating on the shore, on being
shouted at by those on the floe, would make up their minds suddenly, and all plunge
into the sea and swim off to the floe to board it; and if, as often happened, it were
already crowded, many of those on the floe would be pushed off one side, as the fresh
party boarded it on the other.

Arrived at the lower end of the rookery, every bird would suddenly plunge into
the tide and swim all the way back against the stream, only to board a fresh floe for
another ride down. Some of them must have spent many hours of the day in this
mannet.

During the nesting-season, at any rate, the clubbing of the penguins into parties

ADELIE PENGUIN—LEVICK. 73

takes place only after they have left the nesting-ground. Parties are never seen
making their way among the nests, and when, after bathing, they reach the rookery,
they invariably break up and go their several ways. Again, when on their way out
from the rookery, groups are formed on the ice-foot, they consist of strangers from
different parts of the rookery, not of neighbours from any particular spot. }

The manner in which penguins swim in the water has already been described by
many writers, and it is enough merely to allude in these pages to their two methods Of
progression, which are by swimming on the surface as a duck does, and secondly by
“porpoising” (Pl. XIII)—a method distinctive of their order. When swimming on
the surface they sit low in the water, the upper part of the back being submerged,
so that the neck sticks up out of the water. In this position they make ‘fair progress,
attainmg a speed of some six knots. This, however, is their slowest method of
progression in the water, and when they travel quickly they always use the
“ pnorpoising ” method.

It is beneath the surface that they are most agile. Here they use their powerful
flippers for propulsion, the action of these appearing to be exactly that of a bird’s
wings in flying. Their speed and agility under water may be compared to those of
fishes, and they can turn to either side, or completely double in their tracks in the flash
of a moment. Their power of leaping from the water merits a special description, and
IT am able to show photographs taken whilst they are doing this.

After the sea-ice had broken away from the ice-foot on the shores of the
rookery, a ledge of ice varying from three to twelve feet high rose precipitously
from the water in many places, and here their leaping powers were to be seen at
their best. The highest leap I saw was exactly five feet from the surface of the
water.

When about to land on a high ledge a party of Adéhes swim to within twenty or
thirty yards of it, when all may be seen to stretch up their necks and survey their
landing-place. Then in a moment all disappear beneath the surface, not a ripple
showing the direction they are taking, till suddenly they all shoot clean out of the
water, either together in a cloud, or in a stream, one after the other, and land with the
greatest precision on to the top of the ice (Pl. XIV). The fact that this is very
frequently undercut by the waves, and projecting some feet towards them, makes their
accurate judgment all the more remarkable, as from the moment they disappear beneath
the surface after their preliminary survey of their landing-place, twenty or thirty yards
off, or sometimes more, to the moment they leap from the water, they must carry in
their minds the exact distance from the spot where they are to rise at the ice, yet I
never saw one of them misjudge the distance so far as to rise under the overhanging
ledge, or jump short of the landing-place. My photographs show them leaving the
water, in the air, and landing on the ice. When they land on a slippery surface
(Pl. XV) they generally fall forward in the tobogganing position and slide forward a
short distance before rising to their feet; but when they land on snow they throw

L 2

74 “TERRA NOVA” EXPEDITION.

their feet and heads well forward, finishing up in a “ hollow back” position as they
come to a standstill.

All their feeding is done beneath the surface. At Cape Adare the tall ice-foot,
projecting sheer from the water, afforded good opportunities for observing them at
this occupation, and in the clear water beneath they could be seen flashing hither
and thither in quest of Luphausia. Every now and then a milky cloud is seen to issue
from the mouth of one of them, and to float away down the tide, and this is made
by their vomiting the whole meal into the water as they swim, so that they may
have the pleasure of catching another one. After vomiting they never pause in their
career for an instant, going straight on with the hunt as if nothing had happened.

Once as we were watching some Adeélies at play in a narrow “lead” im the sea-ice,
one of them suddenly leapt from the water and landed on the ice, holding in its
beak a large pebble which it dropped on the snow, diving back into the water
immediately.

The depth of the sea here was ten fathoms, the lead was a quarter of a mile
from the shore; but it is difficult to believe that a bird could dive to such a depth!
There were, however, no pebbles to be seen in the sea-ice here, which had been
formed in situ.

In the art of diving the Adélies are very perfect. Some of the crags along the
ice-foot rose to a height of some fifteen to twenty feet above the water (Pl. XVI).
They did not commonly dive from such a height as this, and though very often they
hesitated on the brink, generally descended to some lower part before entering the
water. Twelve feet, however, was no uncommon height for them. In some places
the water shoaled up to the edge, and they were to be seen diving (Pl. XII) from a
height of six feet into a foot or so of water, in which case they generally fell fairly flat.

When diving into deeper water their positions were often very graceful and
perfect.

5.—ENEMIES OF THE ADELIE PENGUIN.

Evidence goes to show that in the water the Adélie has only one enemy, the
Sea-leopard (Hydrurga leptonyx), and that out of the water the adult has absolutely
no deadly foe, except when man and his dogs are about.

It has been suggested that the Killer-whale (Orcinus orca) preys upon them,
but this has been a doubtful point, and the following incident distinctly points the
other way. One day as I watched a large number of Adeélies at play in a wide open
water-lead some half mile from the shore, a large Killer appeared suddenly from
beneath the ice on one side of the lead, and coming up to breathe brought the upper
half of his head and body above the surface as he crossed the open space, finally
disappearing beneath the ice on the farther side. In spite of the swirl and commotion
that he made, and of the fact that the water around him was crowded with penguins,

ADELIE PENGUIN—LEVICK. 75

none of these appeared to take the least notice of him, and went on with their play
as if nothing had happened.

In distinct contrast to this is their behaviour in the presence of the Sea-leopard.
Should one of these appear amongst them, as we often saw, they sped off in the
utmost terror, never pausing till they had put some hundreds of yards between
them and their arch-enemy. The Sea-leopards congregate in the sea in the
neighbourhood of the rookeries during the breeding-season, and the number of
Adélies they kill and eat is almost incredible.

On cutting open one of these seals which I had shot, I found its stomach
distended by the carcases of no fewer than eighteen penguins in different stages of
digestion, whilst its intestines were stufted with the feathers of many more.

As one became familiar with the fish-like activity of the Adélies in the water,
and compared this with the clumsier movements of the huge Killers, it became
evident that the birds had a great advantage over the whales, and could avoid
these as easily as a terrier can avoid a cow.

The Sea-leopards, on the other. hand, proved actually to be faster than the
penguins. Their favourite practice was to lie motionless beneath the overhanging
ledges of the ice-foot, their heads only above water, not a ripple betraying their
presence to the penguins gathered on the ledge above them. When these dived
into the water they came almost on the top of their enemy, which with a rush
and a snap of its huge jaws would seize one of them in a moment. I was very
fortunate in securing the photograph reproduced in Pl. XVIII, which depicts this scene
very finely. The penguins are hesitating on the ledge above the Sea-leopard.

They cannot see him, but very well know of the danger which so often meets
them when they enter the water, and the tricks they all play on one another in
the attempt of each to get one of his neighbours to be the first to go into the
water are fully explained. The only possible way in which they can make their
enemy betray himself is to push one of their number in, and in this endeavour they
employ every artifice.

When a party, actually in the water, is attacked by a Sea-leopard, they
“porpoise” off in a mad panic, not in a clump, but spreading themselves out in a
fan-shaped formation as they go, and continue their flight until, as I have said, a
safe distance is interposed between them and their pursuer. Frequently, however,
one is overtaken by the seal, showing that the latter swims the faster, When
overtaken the bird abandons the attempt to get away, and adopts different tactics,
invariably swimming round and round in a circle of some dozen yards diameter,
and whilst he can keep this up, his ability to turn quicker than his enemy saves
him, but in the end he becomes exhausted, his efforts feebler and feebler, until the
great jaws of the Sea-leopard appear above the water, and the penguin is no more.

When they are hungry, the Sea-leopards swallow the penguins whole, feathers and
all, but when they are well fed they skin them first. This they do by seizing the

76 “TERRA NOVA” EXPEDITION.

bird by the feathers and shaking it from side to side till a large portion of the skin
comes away, when they drop this, take a fresh hold, and tear another piece off, and
so on till, at any rate, the greater part of the skin and feathers is removed from
the body.

It is evident that sometimes a penguin escapes, as occasionally we saw them
making their way along the ice-foot, terribly injured, and these generally had the
skin of the whole of their breasts peeled away and hanging from them like an apron,
and their breast-muscles were bared and bleeding.

Greatly as they are terrified of the Sea-leopard in the water, they probably pay
little attention to him when he is on the ice, as it is very common to see them
walking about on a floe in close proximity to Weddell Seals, and it is hardly to be
expected that they would distinguish the two species without close mspection. On
one occasion, too, we shot two Sea-leopards on the same floe, and whilst I was skinning
one, and the dead body of the other lay in a life-like position near by, a large crowd
of penguins landed on the floe, and with their usual curiosity came right up to see
what was going on, showing not the least fear of the enormous carcases of their
enemies, one of which measured eleven feet in length.

6.—NESTING ON CLIFFS.

Hitherto I have made only passing remarks on the fact that the penguins built
their nests far up the precipitous side of Cape Adare, but now I am going into this
matter at some length, as so much has been said on the subject by other authors.

The cliff up which they build rises almost perpendicularly along the eastern side
of the rookery, but is a good deal broken in places, affording foothold to the birds
who have climbed to all its accessible parts, making their nests on ledges and in
niches of the rock, whilst several colonies of nests have been made on the flat ground
at the top.

There is one colony at the very summit, whose inhabitants can only reach it
by a long and trying climb to the top, and then a walk of several hundred yards
across a steep snow-slope hanging over the very brink of a sheer drop of 700 feet
into the sea.

During the whole of the time when they are rearmg their young, these moun-
taineers must make several journeys during each twenty-four hours, to carry their
enormous bellyfuls of Huphausia all the way from the sea to their young on the
nests—a weary climb for their little legs and bulky bodies, each upward journey taking
them some two hours of strenuous climbing. The greater number who had undertaken
this did’ so at a time when there were ample spaces unoccupied in the most eligible
parts of the rookery.

There is evidence to show that Adéhe Penguins have in them a strong inclination
to climb heights. Already I have mentioned that large masses of ice have heen

ADELIE PENGUIN—LEVICK.

17
stranded by the sea along the shores of the rookery. These fragments of bergs, some
of them fifteen feet or so in height, have formed a miniature mountain-ranee along the
shore. All day, and every day, parties of penguins are to be seen assiduously climbing
the steep sides of this little range. Time after time they get half way, and have
to descend to try a different route. Frequently, having with much pains scaled a
slippery incline, one is seen to miss his footing, and come shding down to the bottom
again, only to pick himself up and have another try.

This climbing was generally undertaken by small parties who had clubbed together,
as they generally do, from social inclination. Sometimes a little band of climbers
would take an hour or more in finding a way to the summit, if they had chosen a
difficult place. Arrived at the top, they would spend a variable period there, some-
times descending at once, sometimes spending a considerable time there, gazing
contentedly about them, or peering over the edge to chatter at other parties below.

Again, about half a mile from the beach, a large berg, some 100 feet in height,
was grounded in fairly deep water; it was accessible at first over the sea-ice, but later,
when this had gone, was surrounded by open water. Its sides were sheer except on one
side which sloped steeply from the water's edge to the top. From the time when they
first went to the sea to feed, until the end of the season, there was a continual stream
of penguins ascending and descending this berg.. As I watched them through glasses,
I saw that they had worn deep paths in the snow from base to summit. They had
absolutely nothing to gain by going to all this trouble but the pleasure they seemed to
derive from the climb, and when at the top merely had a good look round and came
down again.

When the stream of penguins was pouring into the rookery at the beginning of
the season, I kept a look-out for those who were to nest up the cliff, and several times
saw birds, on arriving at the rookery, make for the heights without any hesitation,
threading their way almost in a straight line through the nests to the screes at the
bottom of the cliff, and up there to one or other of the paths leading up its side.

Probably these birds had been hatched there, or had nested there before, and were
making for their old haunts, and I suppose they must have found mates when they
got there.

But this lack of hesitation in making for a certain spot gives rise to some
interesting speculation, because if one sex made for a particular nesting-site, why not
the other, in which case cock and hen who had mated the previous year would meet
again, and so possibly in many cases the same pairs may mate year after year, should
both survive.

Battles took place up these heights, as they did below, and many times we saw
one of the combatants, knocked off his feet, roll bumping down the slope from ledge
to ledge until he recovered his footing, only to climb straight to the spot he had left.
and give fresh battle to his opponent.

78 “TERRA NOVA” EXPEDITION,

7.—MORTALITY.

The mortality among Adeélies is very high during the breeding season, both among
the young and the old, whilst an enormous number of eggs are wasted.
Waste of life is due to the following causes :—

The eggys.
Skuas.
Cocks fighting among the nests.
Flood from thaw-water.
Death of parents.
Snow-drifts.
Land-slides.
The young chicks.
Skuas.
Land-slides.
Interference with by cocks.
Getting lost.
Death of parents.
The adults.
Sea-leopards.
Land-slides.
Snow-drifts.

In the above lists I have left out the wanton depredations committed by some of
the expeditions which visit the Antarctic, owing chiefly to the hcence given to ignorant
seamen, but such visits, being made at rare intervals, can have little effect on the
population.

Two of the causes in my list which are not dealt with elsewhere in my text are
land-slides and snow-drifts.

At Cape Adare, the screes at the foot of the cliff are perhaps the most thickly
populated part of the rookery. As the thaw proceeds, boulders of different sizes are
continually rolling down the cliff, some of them falling many hundreds of feet before
they plunge in among the nests on the screes, doing terrible damage and often rolling
some distance out into the rookery.

At other times, owing to the bursting out of thaw-water which has been dammed
up at the top of the cliff, large land-slides are caused, which bury many hundreds of
nests beneath them. Indeed, these great screes on which the nests are built have been
formed by these land-slides taking place from year to year, and no doubt form the
graves of thousands upon thousands of former generations.

One such slide took place whilst we were at the rookery, doing terrible damage,

4

ADELIE PENGUIN—LEVICK. 79

and we worked for many hours rescuing the unhappy birds that had been buried alive.
The mass of rocks and rubble that fell was followed by an avalanche of snow, and as
we worked in this, we saw here a flipper, and there a leg, just projecting from under
the snow, and waving disconsolately from time to time as if appealing for assistance.

We dug out some birds who had been buried for hours, yet were still alive and
uninjured ; but those which had been struck by rocks were worse off, and hundreds
crawled about the rookery for days after in a terribly injured condition, some of them
literally having been flayed alive.

During a snowstorm the birds sit tight on their nests, and in some places are
completely covered up by a drift. As a rule, the bird on the nest kept a space open
above it by holding its head well up, but sometimes it was quite buried. As air diffuses
well through snow, death never takes place by suffocation, and the bird can live for
weeks beneath a drift, sitting on its nest in a little chamber thawed by its own warmth
in the snow. As a rule, a few hours only elapse before the snow settles down and
abates sufficiently to expose the nest once more, but sometimes a breeze springs up
which is not strong enough to blow the snow away, but simply hardens the surface of
the drift into a crust, which lasts many weeks, and the bird is imprisoned, and after
this little black dots are seen about the ground, which are the heads of penguins
thrust through little breathing-holes that they have kept open. One of my photographs
showed an Adélie engaged in an altercation with its mate, who was very angry, and
aimed savage pecks at the head of the imprisoned one each time it was protruded.

8.—RATE OF GROWTH AND CARE OF YOUNG.
(Pls. XIX-XXI.)

All that I have said hitherto, in describing the way in which the parents foster
and feed their young, applies only to the earlier part of the season, when the offspring
are weak and comparatively helpless. The chicks, however, grow at a great rate, eating
with avidity all the food that is brought them. On weighing a chick at intervals I
obtained this astonishing result :—

Ounces.
ihe egg. ; : : ‘ . : : . 4°56
The chick when hatched 3°00
Five days old . 13°00
Six a Las
Eight _,, BAT 5
Nine ,, 28°50
Eleven ., BY OS
Twelve ., Biante

When the chicks are about a fortnight old an entire change takes place in the
social arrangements of the colony, and in place of the individual care bestowed by
M

80 “TERRA NOVA” EXPEDITION.

each couple upon their clutch, the chicks are gradually massed into little groups,
or eréches, and each créche is guarded by a few birds who keep a good look-out on
the chicks to prevent them from straying and so hecoming an easy prey to the
skuas, while the rest of the parents make journeys to the water for food.

The reason why the parents pool their offspring in this way is evidently because
the voracious appetites of the chicks, as they get bigger, impose too great a demand
on the industry of the parents, who like to spend a certain part of the day in
playing games and enjoying themselves in the water. Hence, instead of one or
other of each couple remaining constantly in charge, three or four remain to guard
groups of chicks varying in number from twelve or so to large masses of many
dozens, who are kept in a close clump by those who remain for the purpose, the
rest of the parents being free to leave them and share the duty of providing food
for the whole.

Before these créches are formed, | think no parent feeds any chicks but its
own, though it may pass the whole length of the rookery on the way from the
water to its nest; and later, the old birds appear to remain faithful to their own
créches, because although they are seen to be pestered by wandering and starving
chicks, piping shrilly and plaintively for food, it is rare to see them yield their
loads of food to the poor little beggars, many of whom, growing weaker, daily fall
a prey to the ever-watchful skuas. In fact, if each colony of parents did not work
solely for its own créche, it is obvious that the chicks up the heights and at the back
of the rookery would come off second best, and many of them starve, whereas these are
just as well nourished as those nearer the water.

It is worth remarking here that until the créches are formed, parents, as a rule,
rigidly exclude the offspring of other birds from their nests, and I have seen little
chicks who had lost themselves, mortally pecked by strange mothers whose protection
they sought.

9.—DEPARTURE FROM THE ROOKERY.

One morning Priestley came into the hut and told me that the penguins were
“ drilling on the sea-ice,” and that I had better come out and look at them. I went
with him to the ice-foot, and this is what we saw.

Many thousands of Adélies were on the sea-ice between the ice-foot and the open-
water leads, then some quarter of a mile distant. Near the ice-foot they were
congregating into little bodies of a few dozen, whilst farther out near the water, massed
bands, some thousands strong, stood silent and motionless. Both the small and the
large bands kept an almost rectangular formation, whilst in each band all the birds
faced the same way, though different bands faced in different directions.

As we watched, it became evident that something very unusual was going on.
First, from one of the small bands a single bird suddenly appeared, ran a few yards in

{

aes

ADELIE PENGUIN—LEVICK. 81

the direction of another small band, and then stopped. In the flash of a moment the
entire band from which he came exec vuted the movement “left turn,” this bringing
them all into a position facing him. So well ordered was the movement that we could
scarcely believe our eyes. Then from the small band our single bird had approached,
another single bird ran out, upon which his own party did exac tly as the first had done,
so that the two stood facing one another, some fifteen yards apart. Then n spontaneously
the two bands marched straight towards one another, and joined to form one body.

After this, we saw the same procedure being enacted in many other places, the
penguins coming down from the rookery and forming small bands which massed
together. Then the augmented body would join another augmented body, forming
a still larger one, which then joined another, and so on, until a great mass of
birds stood together in rows, all facing in one direction like a reciment of soldiers.
One of these masses stood not far from us, a compact rectangular mass.

They stood thus for many minutes, quite motionless and silent, when suddenly, as
before, a single bird darted out from among the crowd and ran a few yards toward the
open water, when, as if it had received a word of command, every bird faced left.
Then the whole regiment marched for the water, keeping its formation almost
unchanged, till it arrived at the edge of the ice, where it halted, and subsequently
entered the water in batches.

This procedure continued for many hours, the penguins that day observing this
extraordinary behaviour, the most astonishing part of which lay in the accuracy of
their drill-like movements, so that we might have been watching a lot of soldiers
executing movements on a field day. Probably the sudden motions of these bodies of
birds was brought about by a sound uttered by the single bird which acted as leader,
though we did not hear this.

The actual reason for their departure from their usual customs is beyond my
knowledge. There was nothing to be seen to account for it, but the penguins evidently
obeyed some instinct which affected them all on this and two subsequent occasions
when the same thing took place. The only suggestion I have to offer is that at some
remote period the Adélies used to mass together in great numbers as the time for their

- migration approached, and that this phenomenon was some sort of reversion to bygone

habits; but against this theory is the fact that the scenes were enacted long before any
thought of leaving the rookery could have possessed them.

Having now given an account of the habits of Adélie Penguins during their
breeding season, when they are found at their rookeries, there remains the question of
their habitat during the winter; when they leave the Antarctic shores for the Pack.

Unfortunately, only a rough idea can be formed on this subject, as so few data are
to hand, and the movements of the pack ice itself are so little known. For the
following information concerning the limits of this mass on and about the meridian of
Cape Adare (170° 10’ E.), Iam indebted to Commander Harry L. L. Pennell, R.N.,
who commanded the “ Terra Nova” from 1910 to 1913, and probably that part of the

82 “TERRA NOVA” EXPEDITION.

subject which more nearly concerns the penguins of Cape Adare rookery will be found
in Table A, whilst for the benefit of zoologists of future expeditions, who may be
carrying these studies further, I append Table B.

The birds from Cape Crozier Rookery (77° 30! §., 169° 30’ E.) must have some
400 miles further to travel when they go north in the autumn than those at
Cape Adare.

TABLE A.
| Northern Limit Miles from Southern Limit | Miles of Pack :

Mean Date. | of Pack. C. Adare. of Pack. N. and §. Remarks,
Reb: 3, 1839) e 68° S. 190 2 q | Balleny
Jans Gla Sale Wor 66° 30! | 280 69° 150 | Ross
Reb: 1 1895)- = 66° 15’ 300 69° 45! 210 _ Kristensen
Heba O89 ee. 66° 0! | 315 Gs Wy Fa Borchgrevink
Feb. 27,1904 . . q | ay 70° 30! ? Scott
Titelo), ts, WOO 5 nil i nil a “Terra Nova”
Marslos O25 2) 2 nil a3 nil ae “Terra Nova”
Jane oO LOS een ae nil a nil oo “Terra Nova”

Norr.—Ross, Kristensen, Scott, Shackleton and Pennell, all, however, found Pack late in the

season while trying to work west along the coast when only some 45 to 75 miles north of Cape Adare, and
all were turned by this Pack.

According to Commander Pennell it appears probable that there is a great hang of Pack in the sea
west of Cape Adare and south of the Balleny Islands, and most probably it is here that the Adélies repair
when they leave the Cape Adare rookery in the autumn. I think, however, it is safe to conclude that
they seek the northernmost limits of the Pack during the winter, as these would offer the most favourable
conditions.

TABLE B.
| ; re Extends Minutes of Latitude
Date. Longitude. Northern Limit. Ne anal S Northern Limit is N. of
| : : Cape Adare.
| Miles
Vem, 3 EO go | 178° E. 67° 8. 140 250 (‘ Discovery ”)
IDE, GHIA IEPA GS 180° 66° 30' 60 280 (‘* Morning”)
Second belt 69° 30 130 (“Morning ”)
Wee, 20, 103 2 . ee WAY 66° 30! 60 270 (‘ Nimrod”)
IDS WAAC 5 ome SS We 64° 45' 300 390 (“ Terra Nova”)
IDEs Pi, WOM 5 5 ge We 65° 20! 160 360 (“Terra Nova”)
Jany 2840 en 166° E. 64° 30! i 400 (Wilkes)
Wim, Gh I 6 5g | 162° E. 64° 30! 270 400 (“ Terra Nova”)

THE “ECSTATIC? APEITUDE:

Both cocks and hens at times are seen to take up a very curious position, holding
their bodies absolutely erect, their necks being stretched perpendicularly to their
utmost length. On assuming this attitude, they raise their flippers horizontally, and
with eyes half closed, make a guttural, croaking sound for a few seconds, whilst a

ADELIE PENGUIN—LEVICK. 83

spasm seems to possess them. This procedure appears to be induced under various
circumstances by a feeling of satisfaction. It is seen most commonly at the rookery
during the breeding season, but also at other times and places, and may be compared
to the crowing of a cock, except that it is exhibited by both sexes, and provokes no
reply from the other birds. It also suggests the braying of asses. Many times when
one of the two mates has started this procedure, I have seen the other at once walk
up to it and emit a low, soothine sound, which at once has caused it to relax its effort
‘and return to the normal state. Called by Dr. Wilson the “ ecstatic” attitude, I see
also that it has been named by the members of the French Antarctic expedition,
“ Brayant”” and ‘“ Chant de satisfaction.”

At the Zoological Society’s Gardens, in Regent's Park, was a male King Penguin
which performed the same sort of antic when its keeper stroked its neck. This bird
accompanied the exhibition with a sort of singing note, far more musical than that of
the Adéle. The Black-footed. Penguins, however, never do it, even when they are
breeding.

Out on the sea-ice, far from their rookeries, | have seen Adélies seized by this
ecstasy, as well as in the neighbourhood of their nests, while both cocks and hens not
uncommonly rise from their recumbent position on the eggs, during the incubating
period, to assume the attitude, impelled by some impulse for which it is difficult to
find a meaning.

LIST OF THE PLATES.

PLATE 1.—A long line of Adélie Penguins approaching their breeding-ground.
TI.— Walking and “ tobogganing” over sea-ice to the Rookery.

I1T.—The proposal: the female on the right in her scoop.

TV.—Two cocks squaring up for battle.

5 V.—Nests made of stones.

VI.—Nests of stones: some large and some small.

» WII.—Floods.

, VIIT.—Method of feeding the young.

TX.—McCormick’s Skuas fighting over Seal-blubber.

* X.—-Adélie Penguins on the ice-foot. .

A XI.— Diving flat into shallow water.

> <i 5; i * eS

» XNIIT.Adélie Penguins “ porpoising.”

XIV.—-Leaping from the water: one bird jumped 4 feet high and 10 feet long.
_ XV.—J umping from the water on to slippery ice.

» 'VI.—Adélie Penguins on the ice-foot.

., XVII. —A knot of Adélie Penguins on the ice-foot.

,, XVIII.—Sea-leopard lurking beneath the overhanging ledges of the ice-foot.

» Y'IX.—A couple with their chicks.

» X%X.—Adélie Penguin with chick 12 days old.

» X'XI.—Adélie Penguins and chick.

LONDON: PRINTED BY WILLIAM CLOWES AND SONS, LIMITED, DUKE STREET, STAMFORD STREET, S.E., AND GREAT WINDMILL STREET, W.

“Le “d

‘puno18-Surpaoigq srayy Suryovoidde sumnsudg ol[ppy jo oul] suo, Vv

Adélie Penguin, Pl. I.

‘ASO[00Z (I8IH ‘YeN) ‘snIA *:

a ‘Id ‘um3ueg o19PV T TOA -
- Taste | ata cae ards

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_ tale, Gath, Piha em -

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Adélie Penguin, Pl.

‘A1aYOOY IY} 0} VdI-"as 19AO

Suruvssoqoz,, pue Suryle MA

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‘Il ‘Id ‘umsueg 219py

Siding It

‘T JOA ‘Aso[00Z
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--

(StH ‘J8N) ‘SDI JE

‘6g “a ‘dooos 19y Ul 3YSI1 94} uO a[eWay ayy : [esodoid oy]

Adélie Penguin, Pl. III.

‘III ‘Id ‘uinsueg a12pyv ‘Il ‘1OA ‘AS0]007 (ISIE, JEN) ‘Sn ‘IW
‘ ‘O16| ‘podxq (AON essa) 91}01eJUY “Ig

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Adélie Penguin, Pl. IV.

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‘uINSUsd 119PV

‘ajqzeq i103 dn Sutrenbs sy909 om]

‘T ‘JOA ‘ASOjOOZ
016] ‘podxy (BAON eiso,) oTjoreJUY “Wg

(ISTH “JBN) ‘Sn “Ja

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Adélie Penguin, Pl, V.

‘A ‘Id ‘umsusq 9119PV

‘soguojs fo gpelu SISION

‘TL JOA ‘AsolOoZ
‘O16| ‘pedxq (BAON eso) o1FOIeJUY ‘Ig

(4SIH ‘J8N) ‘sn

Wea

‘Ig ‘d ‘[]eus 9Wos pur asiIv| 9UIOS +SIUOJS JfO SISIN

Adélie Penguin, Pl. VI.

‘TA ‘Id ‘uinsusd 91[9PpV ‘T ‘1OA ‘AS0]00Z (ISIH “eN) ‘sn ‘wg
‘O16! ‘pedxq (eAON eso) o1oIeJUY “IWI_

| aS as V .

*,
(9
>.

i 3) ‘a

Adélie Penguin, Pl. VII.

TA ‘Id ‘uinsueg 9119PV

‘Spool

‘T JOA ‘ASO[OOZ
‘0161 ‘pedxq (eAON eo L) oNoreyUy ‘We

(ISIH ‘J8N) ‘SMW ‘Ja

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: ; :
7 ; .
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f a

:
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‘99 ‘d

Adélie Penguin, Pl. VIII.

TIA ‘Id ‘umnsusg sappy

‘Sunod oy} Surpoaj jo poyryy

‘TT ‘JOA ‘ASO]OOZ7
‘0161 ‘pedxgq (eAON eiJo]) o1]01eJUYy “JIG

(StH ‘3BN) ‘sn

Bact

=) Q ae = 7
‘sg ‘d ‘Iaqqnyiq-[vVaS JIAO SulzysSly senyS s,YOUWIODVIW

Adelie Penguin, Pl. IX.

5 : ‘sn ‘ys
‘XI ‘Id ‘umnsueg s19PpVv ‘I ‘OA ‘AS0]007 (ISTH JBN) ‘sn ‘Tg
- 016| ‘podxy (BAON eito) o1jO1ejUy 11g

"thd

X.

Adélie Penguin, Pl.

‘X ‘Id ‘umBuaq e119PV

‘JOOJ-991 JY} UO SUINSudg I]PPV

‘T ‘JOA ‘As8o0]00Z
‘0161 ‘podxy (BAON edo L) o01RJUY “HE

(STH ‘YEN) ‘S0IN

ata

Adélie Penguin, Pl. XI.

“19JVM MOT[LYS OJUL BIJ SULATG

‘Id ‘uNsusq 9119PV

‘I TOA ‘Asoj00Z
‘0161 ‘pedxy (eAON essay) oMorejUYy ‘WIG

(CISHH “JN) ‘sn, Fe

Adélie Penguin, Pl. XII.

IX Td

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STTPPV

‘TT JOA ‘ASoO[007
‘0161 ‘podxq (eAON ea) o1jOIeJUYy ‘JIG

(ISTH ‘JeN) ‘sn

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‘ond

Adélie Penguin, Pl. XIII.

TIX ‘Id ‘Uinsusg s1]9pVv

_ sutstodiod,, suinsudg ollppy

‘l TOA ‘ASo[007
‘0161 ‘podxgq (eAON ea) oNoIelUYy ‘Ig

(SIH ‘JEN) ‘SNA ‘I

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Adélie Penguin, Pl, XIV.

‘I ‘1OA ‘AS80007 (ISIH “FEN) ‘SN “Fd

: ‘Id ‘uinsuag 9I[9p
eee ee ‘O16| ‘podxq (eaon Bila) o1o1eyuy “jg

‘Ld ‘gar Ataddijs 0} uo sajem ayy wosy Surdun {

Adélie Penguin, Pl. XV.

‘AX ‘Id ‘uUINSuUsqg 9I19PV ‘T ‘OA ‘ASo0[007Z (SIH ‘VEN) ‘sn ‘Wg
016] ‘podxq (BAON eiioL) ororejuy “lg

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Adelie Penguin, Pl. XVII.

TIAX ‘Id ‘uInsued s1]9pyv

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‘I ‘IOA ‘ASo][00Z
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‘oud "JOOJ-99I

Adélie Penguin, Pl. XVIII.

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jo saspoa, Sursuvysaao ayy yyeousq Ssurysny piedoa-vas

‘I [OA ‘ASOTOOZ
‘0161 ‘podxq (eAON esa) o1jorejUy ‘Wg

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We

"6h ‘99 ‘did ‘'Syoryo Itayy yy ajdnoo y

Adélie Penguin, Pl. XIX.

“XIX ‘Id ‘UInsueg sIT9py ‘T ‘JOA ‘A80]007 (SIH ‘TeN) ‘Sn ‘Wd
‘O16| ‘pedxq (eAON eo) O1OIeJUY “Ig

6h ‘pjo sAep ZI Yoyo YIM uInsuag al[ppy

KX.

Adélie Penguin, Pl.

‘XX ‘Id ‘umMsueqg 9119PV ‘| ‘JOA ‘AB0T00Z (ast ‘TeN) ‘su ‘WBE
‘0161 ‘podxy (eAON eiIo L) o1OrelUy ‘WIG

Adélie Penguin, Pl. XXI.

TXX ‘Id ‘UINsuIg 21T9py

‘yoryo puv sutnsusg dII[9pVv

‘I ‘JOA ‘A8o0[007
‘O16| ‘podxy (eAON eilo) oForejuy ‘jM_

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SH ANTARCTIC ( “TERRA NOVA’) EXPEDITION, 1910,
i > NATURAL HISTORY REPORT.

ZOOLOGY. VOL. |, No. 3. Pp. 85-124,

CETACEA.

D. G,, LILLIE, M.A.
~- (St, John’s College, Cambeiddes Biologist to the Expedition).

ITH FOURTEEN FIGURES IN THE TEXT AND PLATES I-VILL.

os LONDON:
‘ED BY ORDER OF THE TRUSTEES OF THE BRITISH MUSEUM.

eee ie pear anno earnest Sirect,
Soho Londo: 5 thi ‘ON.
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nF Rane ‘Sane Sl
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VOL, I.

Hi 1915

85
*
CETACEA.
Bre De Ge EME TE, “Mla.
(Biologist to the Expedition).
CONTENTS.
PAGE PAGE
T.—INtRopuction . 85 Mystacoceti (continued).
II.—Descriptions OF SPECIES . 86 BALAENOPTERIDAR (continued).
Mystacoceti — Megaptera nodosa, Bonnaterre
(continued. )

BaLAENIDAE Ph Distribution and Migration in
Balaena glacialis, OTe 86 the Southern Hemisphere 110
Neobalaena marginata, Gray . Si Balaenoptera acutorostrata, Lacépéde 113

BALAENOPTERIDAE— 45 musculus, Linnaeus . 115
Megaptera nodosa, Bonnaterre 87 3 physalus, Linnaeus. 116

Introduction 87 " borealis, Lesson 3 lites
Size 88
Body Form 89 Odontoceti—

Egiour d 20 PHYSETERIDAE

3. o C 3 Va

ool prea Physeter catodon, Linnaeus . euis
Baleen 95 ZIPHIIDAE—

Hae 95 Hyperoodon rostratus, Miiller 6 lis)
Jacobson’s Organ i 97 Mesoplodon australis, Flower 5 tals
Throat-grooves 5 : 97 DELPHINIDAE—

Movement of the Lower Jaw . 99 Oreinus orca, Linnaeus . 5 tS)
Mammary Glands. ; ~ LOL The “ High-finned” Whale . . 120
Auditory Organ ? : . 103 Globicephala melaena, Traill . 5 PAL
Scapula and Vertebrae — . . 108 Delphinus delphis, Linnaeus . . 121
Foetuses : : ; ~ Log Tursio peronii, Lacépéde j . 121
Food . : : : . 109 Lagenorhynchus obscurus, Gray . 122
Habits , : : . 109 3 wilsoni, sp.nov. . 123

I—INTRODUCTION.

A CAREFUL watch was kept for Cetaceans on board the “Terra Nova” during all her
voyages ; and although we did not succeed in capturing any specimens of this group,
accurate records and notes were taken of nearly every whale and dolphin which we
saw. Great care was exercised to determine the species, as far as this was possible from
an examination of the animals as they swam in the sea.

In writing this Report I have had access to the notes made by the late Dr. E. A.

O

“86 “TERRA NOVA” EXPEDITION.

Wilson and by Commander H. L. L. Pennell, R.N., and other officers of the “Terra
Nova.” The zeal shown by the Officers of the Watch and the whole ship's company
to notify the biologist whenever a Cetacean was sighted, deserves to be highly com-
mended ; indeed, it must be confessed that the cries of “‘ Whale” were often more
numerous than the responses on the part of the biologist.

During the winters of 1911 and 1912 I spent several months at whaling stations
near the Bay of Islands, New Zealand ; and was thus able to examine three species of
large whales in detail. Opportunities were taken of talking to the whalers; and some
information was thereby obtained, concerning the distribution and migration of whales
in the Southern seas. This memoir contains a record of the Cetacea seen by us on
board the “Terra Nova”; and also some anatomical and other notes on the species
examined in New Zealand, together with any new information which I have been able
to collect with regard to the Cetacea of the Southern Hemisphere.

I have to thank Messrs. Spurling & Son, of Tasmania, for their permission to
republish the photograph shown in Plate IV., fig. 1. Professor W. B. Benham, F-.R.S.,
of Dunedin, N.Z., has kindly allowed me to make use of four previously unpublished
photographs in Plates IV. and V. I am indebted to Dr. W. G. Ridewood for the
photograph of the embryo Humpback Whale in Plate IV., fig. 4. The photographs in
Plate VI. were taken by Paymaster F. R. H. Drake, R.N., a member of the Expedition,
and to him I tender my best thanks. My sincere thanks are due to the Editor, Dr. 8.
F. Harmer, for helping me in many ways during the preparation of this Report.

Il DESCRIPTIONS OF =SEEGIES:

MYSTACOCETI.
BALAENIDAE.
1. Balaena glacialis,* Bonnaterre.

No individuals of this species were observed by the “Terra Nova.” It would
seem to be doubtful if this whale ever penetrates into the ice-covered seas of the
Antarctic, although it still appears to be fairly plentiful in sub-antarctic regions.

This species is caught by the whalers at the South Shetlands, usually towards
the end of the season, about February and March. At South Georgia it is taken
all through the season, although March was said to be the best month. It is found
off Campbell Island, to the south of New Zealand, throughout the year. Thirteen
Southern Right whales were captured at the last-mentioned locality by Mr. Cook

* Or Balaena australis, Desmoulins. Collett, R., ‘“‘ Norges Pattedyr,” Kristiamia, 1912, p. 545.
Bull, H. J., “The Cruise of the ‘ Antarctic,” 1896, p. 214. Racovitza, E., “ Résultats du voyage
du 8.Y. ‘ Belgica,’ 1897-99, Zoologie, Cétacés,” 1903, p. 19. Wilson, E. A., National Antarctic Exped.,
1901-4, Nat. Hist. Reports, Vol. II., 1907, Mammalia, p. 1. Liouville, J., Documents Scientifiques,
Deuxiéme Expéd. Charcot, 1908-1910, Cétacés, 1913, pp. 5-27. Turner, Sir W., Proc. Roy. Soc. Edinb.,
XXXV., 1914, Pt. I. (No. 2, p. 18).

CETACEA—LILLIE. 87

between March and August, 1911; and seventeen were taken during the same
season in 1912.
2. Neobalaena marginata, Gray.

We did not see a single whale which could be certainly determined as belonging
to this species. Before he left the “Terra Nova,” in January, 1911, the late
Dr. Wilson came to the conclusion that the whales which he had identified as
Neobalaena marginata,* when on board the ‘ Discovery ” in 1901-4, were in reality
a species of Balaenoptera, which I was afterwards able to identify as being
Balaenoptera acutorostrata, Lacépede. (See page 113.)

BALAENOPTERIDAE.
3. Megaptera nodosa, Bonnaterre. Plates I., II?, III., 1V., fig. 4.

INTRODUCTION.

During the month of October, 1911, I visited Messrs. Jagger and Cook’s
Whaling Station at Whangamumu, near the Bay of Islands, New Zealand, and
through the kindness of the manager, Mr. H. F. Cook, I was able to examine seven
individuals of the New Zealand Humpback Whale. In the winter of 1912, four
months (July to October) were spent on board the two Norwegian floating
factories belonging to the New Zealand Whaling Company, who were also exploiting
the waters in the neighbourhood of the Bay of Islands. And here again, thanks
to the extreme courtesy and generous hospitality shown to me by the manager,
Mr. L. 8. Hasle, and all on board the 8.8. “Rakiura” and 8.V. “ Prince George,”
I was enabled to investigate some thirty more specimens.

It will be seen in the account which follows that, after a fairly close study of
thirty-seven Humpback Whales caught in New Zealand waters, I came to the
conclusion that they were identical with JJegaptera nodosa, Bonnaterre,f of the
Northern Hemisphere, and that no evidence was obtained for the existence of the
two species, Megaptera lalandii, Fischer, § and M. novae zealandiae, Gray. ||

It has been shown by True §J that the Humpbacks of the Eastern and Western

* Wilson, E. A., op. cit., p. 4.

+ This name is employed in accordance with True’s paper “ On the Nomenclature of the Whalebone
Whales of the tenth edition of Linnaeus’s ‘Systema Naturae.’” Proc, U.S. Nat. Mus., Vol. XXTI., 1899,
pp. 617-635.

t Or Megaptera longimana, Rudolphi.

§ MM. Van Beneden et Gervais, “Ostéographie des Cétacés,” 1880 (1868-1879), p. 130. Gray,
J. E., 1866, Brit. Mus. Cat. Seals and Whales, pp. 126-128. Giglioli, E. H., 1874, Napoli, “ Cetacei
osservati duranti il viaggio della ‘Magenta,’ ” pp. 30-44.

|| Gray, J. E., Proc, Zool. Soc., 1864, p. 208. Hector, J., Proc. New Zealand Inst., 1872, Vol. V.,
p- 156; 1877, Vol. X., p. 335.

q True, F. W., “ Whalebone Whales of the Western North Atlantic,” 1904, Smithsonian Contribu-
tions to Knowledge, Vol. XXXIII., pp. 211 and 271, Pls. 29-41. See also Collett, R., “ Norges
Pattedyr,” Kristiania, 1912, p. 606.

0 2

88 “TERRA NOVA” EXPEDITION.

shores of the North Atlantic, and of the North-Eastern Pacific, are probably all
members of the one species, M. nodosa, Bonn. As far as our present knowledge
goes, it would appear that this widely distributed species is the sole representative
of the genus Megaptera. It is of interest to note that the same world-wide identity
seems to occur also in the case of the four generally admitted species of the genus
Balaenoptera, although a fifth species of this genus, B. brydei, recently described,
has at present only been recorded from the seas off the coast of South Africa.*

SIZE.

Tt was very dithcult to obtain exact measurements of whales at the floating
factories, as the carcases were flensed and cut up in the water and taken on board
the ship in pieces. When the whales were lying in the sea, previous to the
dissecting operations, they were generally partially submerged, which prevented
accurate measurement of their length from being made. Careful measurements
were collected, however, in the case of the seven specimens seen at the shore station

Taste I.—Megaptera nodosa, Bonnaterre.

Length of Males. | Length of Females.
|

Feet.
35

Average length of Males— | Average length of Females—

35 feet 40 feet

at Whangamumu during October, 1911. This compensated to some extent for my
inability to obtain more than two length-measurements from the many individuals
which I examined at the floating factories.

The average length of these nine whales was 38 feet, which is rather too short
to be typical of all the Humpbacks captured off the Bay of Islands in the season
of 1912. The largest individual of this species seen during my stay was about
51 feet long; and I feel vertain that, if it had been possible to measure the
lengths of a larger number, the average would not have been far short of 40 feet.

The length-measurements obtained are given in Table 1, and it will be noticed

* Olsen, Orjan, Proc. Zool. Soc., 1913, pp. 1073-1090, Pls. CIX.-CXITI.
} Giving suck to calf when shot.

CETACEA—LILLIE. 89

that the average length of the females is greater than that of the males. The
measurements were made in a straight line from the tip of the snout to the notch
between the tail-flukes.

Mr. A. H. Cocks’ estimate of the average lengths of 94 Humpbacks, taken off
the Northern Coasts of Norway and Russia in the years 1885 and 1886, is 351 feet
in the case of the males, and 401 feet for the females.* From this it will be seen
that the Southern members of the species are almost identical in size with their
confreres in the Northern Hemisphere.

Bopy Form.

In general appearance, the New Zealand Humpback (Plates [. and II.) agreed
exactly with the photographs and descriptions of Northern types.+

Proportional measurements, taken in the case of four adults and a large foetus,
are shown in Table II.

Taste II.—Megaptera nodosa, Bonnaterre.

Measurement. Male. Male. | Male, Female, | Female
Foetus.
| ft. ims. | ft. ins. | ft. ins. | ff. ins. | ff. ins.
Total length . ‘ . ; . ; : -|30 8)31 —|34 -|42 -|]13 6
Tip of snout to eye. ; >: | : 2 10
Tip of snout to angle of mouth 8 8
Eye to ear : i 1
Nostrils in front of eyes . ; ; peal ce een Ol eee. ew || ea me
Tip of snout to anterior end of mmbilicus F F elon von) shi 08 9G) 2 5h ae fw
Length of umbilicus : = - 8 1 1 1 on ee
Posterior end of umbilicus to cata ior end of the urino- |
genital groove é 2 2 2 2 ne 2B} - tii
Length of urinogenital groove . : eee a i ee TH
Posterior end of urinogenital groove to anterior end | |
Of anus; ‘ : F 3 ; : Auli) Pied teh yee 2 1 i) 1h =| oat
Length of anus er | ee ees rae Gy ee
Posterior end of anus to Hise edge of tail: fines ca ere AST Gili Dad R= 1 6
Greatest breadth of tail-flukes . 3) ALN og tt Zion || ome i 2!
Length of one tail-fluke . ; ; ; F | 53 S: BD 1. 129
Length of pectoral fin (axil to tip) 3 ; : o |) UO = 10° 9 4. -
Greatest width of pectoral fin . ; ; ‘ PQ Bs! |
Eye to anterior edge of pectoral fin ; ill | Gl ua
Posterior end of dorsal fin to notch of flukes ; | a to) a) a st A
Height of dorsal fin ; ; , ae “ts is x - 4
Width of body between axils of pectoral fins 3 4

In Table HI. some of these measurements have been reduced to percentages of
the total length, for comparison with similar measurements, in European and American
specimens, given by True.f

* Cocks, A. H., The Zoologist, 1887, Vol. XI., p. 213.

+ Struthers, J., Journal of baniiers and Physiology, 1887, pp. 109-125. True, F. W., ‘“‘ Whalebone
Whales of the Western North Atlantic,” Smithsonian Contributions to Knowledge, 1904, Vol. XX XTII.,
pp- 211-231, Pls. 37-41. Haldane, R. C., Annals of Scottish Nat. Hist., April 1905, p. 66, Pl. IIT.

{ True, F. W.., loc. cit., p. 223.

90 “TERRA NOVA” EXPEDITION.
Taste LIT.—Megaptera nodosa, Bonnaterre. European, American, and New Zealand.
|
E B e e B 3 : ; —
a A= = oI = a |H$s8|s8h/ga8 | Se |s &
a OD DOD DO DB o> 2 o> 4D HS | 2S | oa ano § o
SH SH SH oH SH a @ A538 ideeladge: no a4 2
A cites ane cee =| = ao | “3a Ba |t5a tS | BH
Sg | So Ciks) Sr Sd ts[e=! aS HSE eis Sa |) ea) TS Ie8 Toe | Pao
= ol —] = = a DQ © wy
ad aa | ad | qo so" |4onq mg BS ae ae Ie,
Veale eral t Melee acest eta et EM I Syne isies || lee Te ||| Sie) as
Sg | 68 | ce | Sg | os | Me aes a oo Re eee
=N mN mN mN mN ZS ee SAS | ea
Q ~Q ica) isa AQ
Sex | |S |S Pe Neeteaa |) Oy (eee) Nh eae eran ean
: e ; Foetus
, ” , ” , ” , ”" , ” , ” , ey NP we, ” , ” , " , ”
Total length . . | 30 8 | 31 0} 34 0} 42 0] 13 6 | 46 71] 46 6 | 45 5 | 42 2 | 51 63) 44 OF
Notch of flukes to || sé 4 y % %, Vs VE Naa %, vs
anus ; . | 25°0 | 25:2 | 23-7 | 20°0 | 20-9 | 25.82) 24:5 | 24-0) 22-9) — —
Notch of flukes to |
navel 3 . | 45:1 | 43°5 | 39-4 | 37°8 | 41°3 | 44°42) 43-3 | 41°8 | 42.4) — _
|
Length of pectoral fins |
(axil to tip) - | d2°6 | —* | 31-6 | — | 29-6} 31-0 | — | 2821) 28:9 |) 29a else
1 Straight, from lower jaw. ? From figure. 5 Danish measure. ‘ Approximate.

The cut-water, mentioned by Struthers as occurring beneath the chin in
the Tay whale, was a very constant feature in the New Zealand specimens. The
number of nodes or bosses on the anterior margins of the pectoral fins were

counted in several individuals, and each fin was found to

oe have two larger and seven lesser nodes, as in Northern

Megaptera.

is The shape of the dorsal fin varied a good deal,* as is

shown in text-fig. 1; but in each case a basal portion could

be distinguished which was surmounted by the fin proper.

gia, The variations in the outline of the dorsal fin were quite
independent of the variations im the amount of pigmentation.

Se a end The same type of fin was found in individuals which varied

Fig. 1.—Outline diagram of greatly in colour.

15% variations in the shape Behind the dorsal fin, in most of the specimens, the
ees ane ddoraal ridge of the tail was crenulated, as described by

Zealand Humpback whale
(Megaptera nodosa, Bonn.). Andrews in Lehachianectes glaucus.f

CoLour.

In the case of thirty individuals the colour was carefully noted, and it was found
that every gradation occurred in the amount of black and white, as shown in text-
fig. 2, Nos. 1-4.

* Cabrera, A., “Fauna Ibérica. Mamiferos,” Madrid, 1914, Cetacea, p. 398.
} Andrews, R. C., “Monographs of the Pacific Cetacea,’ Mem. American Mus. Nat. Hist., N.S.
Vol. I., Pt. V., 1914, p. 259.

CETACEA—LILLIE. 91

It was more or less possible to group the colours of the thirty specimens into
seven divisions, as indicated in the first column of Table IV.

Taste IV.—Megaptera nodosa, Bonnaterre.

Colour.
No. 1
Between No. 1 and No. 2
No. 2 ;
Between No. 2 and No, 5
No. 3°!
Between No. 3 and No. 4
No. 4

|
Number of Number of |

Males. Females. Total.
: = l
| el 7
3 1 | 4
3 il 4
2 4 6
2 1 6
9 K

The numbers in this table refer to
the four types shown in text-fig. 2,
Nos. 1—4, and three intermediate types
have been added.

It will be noticed that the two
extreme types were the rarest, and that
the females tended to be darker than
the males.

The variable coloration of Hump-
back whales has been known for some
time*, and a few observers have tried
to attach thereto a specific significance.
This. however, appeared to be quite
unwarrantable in the case of the New
Zealand specimens, since the differences
in colour were not accompanied by any
other distinguishable characters.

It has been suggested by Rawitz

* Racovitza, E., 1903, ‘Résultats du
voyage du S.Y. ‘ Belgica,’ Cétacés,” p: 20.
Rawitz, B., Sitzb. Ges. Nat. Fr. Berlin, 1897,
pp. 146-150. True, F. W., ‘“ Whalebone
Whales of Western North Atlantic,” Smith-
sonian Contributions to Knowledge, 1904, Vol.
XXXIIL., p. 216, Pls. 37-41. Haldane, R.C.,
Annals of Scottish Nat. Hist., April 1905, p.
a, TAG MB

um

— inant

AEN, RETR ie
ER. |
Sk

4

4

Fia. 2.—Diagram to show the variation in the amount
of pigmentation in the New Zealand Humpback
whale: 1—4, colour varieties (see text).

92 “TERRA NOVA” EXPEDITION.

that these whales may possibly get whiter with age; but this idea was refuted by
the fact that some of the whitest whales, seen at the Bay of Islands, were very
small and showed signs of immaturity. Moreover, an unborn foetus (133 feet in
length) was found to possess exactly the same coloration as the mother. They were
both coloured as in text-fig. 2, No. 2.

All the specimens examined were black on the dorsal surface, and most of them
had some white on the ventral surface. Two were almost entirely black all over.
The regular way in which the dorsal pigmented area encroached upon the white
ventral region was very marked in the various individuals: the three points of black
in the posterior half of the animal, which were barely indicated im No. 1 type
(text-fig. 2) became more pronounced in No, 2, and in No. 3 formed bands round
the abdomen ; in No. 4 they coalesced and spread out over the entire ventral surface,
with the exception of two spots below the dorsal fin and a small area below the
mandible.

Fic. 3.—Ventral surface of a New Zealand Humpback whose pigmentation is between
Nos. 3 and 4 in text-fig. 2.

The mode of spreading of the pigment on the ventral surface is shown in
text-fig. 3, which represents the type between Nos. 3 and 4 (text-fig. 2).
Over the lighter areas the pigment occurred in oblique lines of flecking, which
converged on either side towards the mid-ventral line. All the individuals,
including the foetus 135 feet in length, were dappled like grey horses. The dapple-
markings, which were particularly noticeable on the shoulders, pectoral fins, flanks
and tail, consisted of white spots, rmgs and streaks on the black pigmented areas,
and the same marks in black on the white areas. These markings are shown in
text-fig. 4, A; they were quite distinct from the scars made by rocks and barnacles.

In the whiter individuals there were often patches of grey on the white ventral
surface, and especially at the junction of black and white areas in the neighbourhood
of the dapple markings. The grey colour was due to a diminution in the amount
of black pigment in the Malpighian layer of the epidermis.

The interior of the mouth had a grey appearance, with the exception of the
narrow strip of pink palate which made a keel-lke partmg between the greyish-
white hairs of the ranges of baleen-plates.

The tongue was of a darker grey colour than the hairs of the baleen-plates.

CETACEA—LILLIE.

-
oo

EXTERNAL PARASITES.

Barnacles.—All the specimens examined had barnacles growing upon them, and
the mode of distribution of these parasites over the body of each whale was found
to be remarkably constant. The ventral surface appeared to be the favourite place
for barnacles, although a few were occasionally found on the dorsal surface of the
head. The back was generally quite free from parasites. A thick patch of
barnacles always occurred on the anterior, ventral surface of the throat, in the
middle line, just behind the chin. A smaller patch was generally to be found in
the region of the genitalia. Besides these areas, barnacles seemed to show a
preference for projecting knobs; the bosses on the anterior margins of the pectoral
fins were nearly always crowned with barnacles, also the extreme tips of the
pectoral fins and tail-flukes; occasionally a barnacle was found surmounting one
of the hair-tubercles on the snout. It was noticed that barnacles nearly always
occupied pigmented areas of the whales’ skin; very few were found on a white
surface.

The extremely characteristic patch of barnacles behind the chin was always
situated on an equally characteristic patch of grey or black skin, and the number
of individuals constituting the patch varied with the size of this pigmented area.*
(See Pls. I. and IL., figs. 2 and 7.) Only two species of barnacles were found on the
Humpbacks: Coronula diadema and Conchoderma auritum. The former was always
fastened directly to the skin of the whale, while Conchoderma only grew upon the
shells of Coronula. As many as fourteen individuals of Conchoderina were found
on one Coronula. Barnacle-scars were often to be seen on the skin of these whales.
Each scar marked the spot from which a Coronula had been removed, either by
the host rubbing itself against a rock, or by some predatory fish. These marks,
formed by the base of the Coronula shell, were very easily distinguished from other
scars, as they consisted of a sunken dome-shaped area, surrounded by a circular
depression. The surface of the dome was impressed by some 18 small furrows,
which radiated from a ring near its apex. (Text-fig. 4, C and D, b.)

Whale-lice—-Every specimen was infested with Paracyamus boopis. These
“lice” were found all over the body of each whale, sticking into the epidermis by
their sickle-shaped claws. They were most plentiful among the barnacles and in
the throat-grooves on the ventral surface, in which places the young “lice”
appeared to seek shelter.

Dr. Calman tells me that the Cyamids have no free-swimming stage, and there-
fore they probably pass from whale to whale by actual contact, during copulation and
lactation. When the young and adult “lice” were removed from a whale and put
into a glass vessel containing sea-water, they sank to the bottom and caught hold of
one another with their claws in their endeavours to get a foot-hold in a soft surface.

* Shown in one of True’s plates, loc. cit., Pl. XXX VIII, Fig. 1.

VOL. I:

94 “TERRA NOVA” EXPEDITION.

They were quite unable to swim. Whale-lice should be of value in helpimg to
determine the specific identity of their hosts. It was of interest to find that the New

Zealand Humpbacks were infested with a Northern form of Paracyamus.

SCARS.

Wounds, in all stages of healing, occurred irregularly over the surface of the
skin, though they were perhaps more often found at the sides of the animal and on
the head. The fresh scars were like little pits, usually about three inches long, two
inches broad, and 13 inches deep. They presented the appearance of having been
cut out with a sharp instrument, and the piece of skin and blubber entirely removed.

D
Fig. 4.

A—Dapple-markings on skin of New Zealand Humpback whale.

B— Marks on skin due to wounds: a, open scar ; b, scar partially healed; c, scar
completely healed.

C—Barnacle-scar.

D—Section of blubber to show: a, open wound caused by rock ; b, barnacle-sear.

The cut ends of the fibres and blood-vessels, which penetrated the dermis or blubber,
could be seen on the walls of the cavity.

The open sores were frequently infested with Paracyamus boopis. The healed
sears had a characteristic oblong shape (text-fig. 4, B, c). When such scars occurred
on pigmented areas of the skin they were usually lighter in colour than the surrounding
epidermis. They could not, however, be mistaken for the white dapple-markings
referred to above, since they had a very characteristic rayed appearance. These
wounds were, almost certainly, made by the whale coming in contact with sharp
ledges of rock while swimming near the coast. Off the coast of New Zealand on
several occasions | have seen these shore-loving whales swim through narrow channels
between isolated rocks, and play in their vicinity. It would be very difticult for the

CETACEA—LILLIE. 95

animals to avoid touching the rocks in such places. Whalers maintain that Hump-
backs consciously rub themselves against rocks in order to remove the barnacles which
invariably adhere to this species. There may be some truth in this belief, but I found
open wounds on the body of the specimen of Balaenoptera borealis Lesson, which was
caught off the Bay of Islands; and the scars on this whale were identical in appearance
with those on the Humpbacks. Healed scars, such as are represented in text-fig. 4,
B, c, have been seen in other species of Balaenoptera.* The members of this genus
are known to be free from barnacles, yet, if we may judge by the scars, they would
appear to be in the habit of occasionally brushing up against rocks. It is possible that
some of the scar-markings may be due to parasites beneath the epidermis. Mr. Cook,
who has watched the Humpbacks off the north of New Zealand for twenty years, told
me that he noticed more scars on the animals when they were on their southward
migration. (Sve p. 110.)
BALEEN.

The whalebone-plates were uniformly black in colour, with the exception of a few
plates on either side of the anterior end of the snout (PI. I). About three or four of
these plates on each side were entirely white ; the others were black on the outer edges,
with bands of white on their inner sides.

The hairy inner edges of all the plates were greyish-white in colour. The
individual bristles were coarse, and when seen en masse they had a slight appearance
of waviness. The largest baleen-plates seldom exceeded 20 inches in length. The
whalebone agreed in every character with that of the Northern members of the species.t

In an embryo, 133 feet in length, the baleen-plates were relatively narrow and
only covered the outer edges of the palate. It would appear that after birth the
baleen-plates increase in width until they cover the whole surface of the palate, with
the exception of the median crest or keel.

Hatrs.

Short hairs or vibrissae were found to occur on the facial region in Jlegaptera
nodosa, very much as they do in Balaenoptera,t with the exception that many of them
were situated upon dome-shaped humps or tubercles. On the dorsal surface of the
rostrum or beak there were generally three hair-tubercles on each side of the blowholes.
About five formed a median line down the snout, from the nostrils to its tip. (See
text-fig. 5, A., and PI. II, figs. 4 and 6.) Some eight or more occurred on either side of

* Collett, R., Proc. Zool. Soc., 1886, p. 265. Lillie, D. G., Proc. Zool. Soc., 1910, p. 783.
+ Struthers, J., Journal of Anatomy and Physiology, 1888, Vol. XXII., p. 121. True, F. W.,“° Whale-
bone Whales of the Western North Atlantic,” Smithson. Contr. to Knowledge, 1904, Vol. XX XTITI., p. 230.
t Lillie, D. G., Proc. Zool. Soc., 1910, p. 773. [Nors.—This paper appears in the “ Proceedings”
with text-fig. 70, p. 775, turned upside down. The mistake was entirely the fault of the printers, who
took the figure out to repack, after the proofs had been passed for press, and put it back reversed. |
Burfield, 8. T., Report on Belmullet Station, British Ass. Rep., Dundee, 1912, p. 177.
Pp 2)

=

96 “TERRA NOVA” EXPEDITION.

the beak, arranged approximately in two alternate series of four: an outer row on the
extreme edge of the beak, and an inner row situated a short distance away from the
edge; making in all about twenty-seven tubercles on the upper jaw. On either side
of the lower jaw there were about eight hair-tubercles, also arranged in two more or
less alternating rows of four. On each side of the blunt extremity of the mandible,
pointing directly forwards, there was a cluster of two or three larger tubercles and a
few smaller ones. These clusters contained the largest of all the hair-tubercles, and
constituted the foremost part of the animal.

A— Anterior view of the mouth of a Humpback, to show the movement of the rami of
the lower jaw (semi-diagrammatic) : a, free ends of the baleen-plates ; b, tongue
seen through the space between the lips; c, rami of mandible in the position
which they occupy when the mouth is open ; d, upper jaw ; e, hair-tubercle.

B—Transverse section of the symphysis between the rami of the lower jaw: a, white
fibrous tissue ; b, central pulpy substance.

There were, altogether, about twenty-eight tubercles on the lower jaw. The
number and character of the tubercles corresponded very closely with those described
as occurring on the Northern MJegaptera.*

The lengths of the hairs when withdrawn from their follicles were, on an
average, about 32 mm. The longest observed was 45 mm. The free portion of

True, F. W., “ Whalebone Whales of the Western North Atlantic,” Smithsonian Contributions
to Knowledge, 1904, Vol. XX XIII, p. 225,

“hh:

CETACEA—LILLIE. 97

each hair which projected beyond the surface of the tubercle was about 10 mm.
Some of the hairs situated at the tip of the snout, and which were without
tubercles, only had 5 mm. of their length projecting from the surface of the skin.
The height of the tubercles varied from half an inch to two inches.

These hairs have been shown to possess a sensitive function,* and in all
probability they serve to indicate to Whalebone Whales the presence of the small
plankton animals upon which these whales feed. The sensitive feeler, or barbell,
which occurs beneath the chin of certain fishes, which feed upon plankton, has
probably a similar function to that of the vibrissae in the Mystacoceti.

JACOBSON’S ORGAN.

The remains of the ventral ends of Stenson’s duct could be seen in several
specimens. They consisted of two shallow depressions on the ventral surface of the
tip of the snout, very similar to those of Balaenoptera musculus, Linn.t

THROAT-GROOVES.

About twenty-four furrows were counted between the pectoral fins in several
individuals, and they were found to agree in number and character with those of
Northern members of this species.{

The pink coloration which was noticed by the present writer,§ a few years
ago, in the throat-grooves of two species of Lalaenoptera was also observed in some
of the New Zealand Humpbacks. Careful investigation showed that it was caused
by an effusion of blood, which evidently takes place after the death of the animal,
as pointed out by Mr. Burfield.j| There is, therefore, no truth in the surmise that
the grooves may serve to aerate the blood.

Their function is to give elasticity to the floor of the mouth, for the purpose
of increasing the capacity of the mouth-cavity. Since the grooves extend backwards
to the navel, it would seem possible that they also serve to increase the power of
expansion of the lungs. On account of the reduced condition of the sternum and
the mobility of the ribs, the dilatation of the thorax takes place laterally, and
would be greatly facilitated by the extra elasticity imparted to its ventral wall by
the furrows. It was of interest to note that the large superficial muscles, which

* Japha, A., 1904, Zool. Jahrbiicher Abt. f. Anat. und Ont., Bd. XXIV., pp. 1-40. Rawitz, B.,
Tnternat. Monatschrift f. Anat. und Physiol., 1906, Bd. XXIII., Hefte 1-3.

{ Kiukenthal, W., “ Vergl-anatom. u. Entwick. Unt. an Walthiere,” in Denkschr. med. naturw.
Gesellsch. Jena, Bd. III, Theil 2, 1893, pp. 321, 349. Lillie, D. G., Proc. Zool. Soc., 1910, 2, p. 774.
Burfield, 8. T., Report on Belmullet Station, British Ass. Rep., Dundee, 1912, p. 177.

{ True, F. W., ‘‘ Whalebone Whales of the Western North Atlantic,’ Smithsonian Contributions to
Knowledge, 1904, Vol. XXXIII., p. 224. Struthers, J., Journ. Anat. and Physiol., 1888, Vol. XXII,
p- 118.

§ Lillie, D. G., Proc. Zool. Soc., 1910, 2, p. 784.

|| Burfield, 8. T., Report on Belmullet Station, British Ass. Rep., Dundee, 1912, p. 176.

98 “TERRA NOVA” EXPEDITION.

occur beneath the blubber of the throat-groove region in Balaenoptera, were also
present in Megaptera.

When the coat of blubber was removed from the carease of a whale, the
more superficial muscles had a tendency to come away with the blubber. These
muscles were confined to the ventral anterior three-quarters of the animal, in the
regions occupied by the throat-grooves and the mammary glands. (See text-fig. 7.)

There appeared to be two pairs of superficial muscles in the throat-groove
region; the anterior pair may possibly be the mylohyoid, and the posterior pair
may represent the pectoralis major. Delage,* who described these muscles in
Balaenoptera physalus Linn., regarded the anterior pair as modified superficial
muscles of the neck. He mentioned that the fibres of the muscles were arranged
more or less longitudinally in B. physalus, but in Megaptera they were oblique.

The probable function of the superficial muscle in the mandibular region is to
diminish the mouth-cavity, after each mouthful, by the contraction of its floor.

Delage suggested that the function of the posterior superficial muscles might
be to compress the air in the lungs, after each act of inspiration, in order to render
the animal heavy enough to sink without effort. The Right Whales and the Sperm
Whale float on the surface of the sea after they are shot, whereas the Balaenopteridae
tend to sink. The latter family have ventral grooves and an associated system of
muscles, which are absent in the former types. It is possible that the contraction
of the ventral superficial muscles, after death, presses the air out of the lungs, and
so causes the Balaenopteridae to sink. The elasticity of the ventral body-wall in
these whales would allow of a great expansion of the lungs after inspiration,
which would make the animal so light that “ sounding ” would be difficult, unless
it could alter its density at will by means of the muscles of the ventral grooves.
The superior mechanism for increased lung-capacity and the regulation of density
which seems to be possessed by the Balaenopteridae, lend some support to the
widespread belief that they are capable of remaining under water for a longer
period than other whales.

I made further inquiries of whalers as to the length of time it was customary
for the Balaenopteridae to remain below water, and was given to understand that
the Humpback rarely remained under water for more than a quarter of an hour
between each two acts of respiration. Seven minutes was considered to be the
normal interval between each two breaths in this species.

The belief, which is held by some whalers,t that the larger Balaenoptera can
remain below the surface for twelve hours, was discredited by those whom I met in
New Zealand. The latter were of opinion that the Balaenoptera were never in the

* Delage, Y., Arch. Zool. Exp., 2 Sér., Tome IIT. 3 bis, 1885, pp. 29-38.

+ Andrews, R. C., ‘“‘Monographs of the Pacifie Cetacea,” Mem. Amer. Mus. Nat. Hist., New
Series, Vol. 1, Pt. V., 1914, p. 256.

t Lillie, D. G., Proc. Zool. Soc., 1910, p. 790.

CETACEA—LILLIE. 99

habit of staying below for more than one or two hours at most. This estimate would
tend to coincide with the period arrived at by Kiikenthal * on the following ingenious
line of argument. The normal interval between each two respiratory acts in man is
three and one-third seconds ; in the pearl-diver this interval can be extended to one
and a half minutes ; the normal interval in whales is seven minutes; therefore, by rule
of three, the interval in Balaenoptera can be extended to three hours. Whales which
can swim under water for two or three hours would easily evade the most vigilant
whalers, and lead the latter to suppose that their quarry had gone below for a
whole day.
MovEMENT OF THE Lower Jaw.

Some years ago Mr. Lydekker noticed in a skull of Balaenoptera borealis, Lesson,
that when the mouth was closed the rami of the lower jaw appeared to be turned
upwards and inwards, whereas when the mouth was open they were turned down, so
that the convexity became outwards instead of upwards.

In order to test this observation in a fresh specimen, and to ascertain if it
occurred in the case of the Humpback, I persuaded the whalers at Whangamumu to
open the mouth of a freshly killed whale, which measured forty feet in leneth, by
fastening a noose round the snout and raising the upper jaw by means of the winch.

When the jaws were closed the rami of the lower jaw fitted closely against the
edges of the upper jaw throughout its entire margin, so that the blade of a knife could
not be inserted between the lips at any point.

As the mouth was opened, I distinctly saw the convexity of each ramus turn
slightly downwards and outwards, causing the width of the space, enclosed by the two
rami, to increase. This action of the rami of the lower jaw, in the process of opening
and closing the mouth, is well illustrated by the rim round the mouth of an ordinary
hand-bag. When the bag is fully open, it corresponds to the lower jaw of a Whalebone
Whale whose mouth is wide open. Close the bag rather less than a quarter, by raising
each side some fifteen degrees from the horizontal, and it gives an idea of the position
of the rami when the whale’s mouth is completely closed.

Before permittmg the mouth of the whale upon which I was experimenting to
close, the masseter and temporal muscles were severed in order to keep the lower jaw
in the position which it had taken up when the mouth was open. The upper jaw was
then lowered upon the mandible, and a space of one and a quarter feet occurred on
each side between the inner edges of the mandible and the outer rim of the beak.
(See text-fig. 5, A).

Thus, when the mouth was open, the width of the space between the rami was
shown to be increased by two and a half feet.

* Kiikenthal, W., Denkschr. med. naturw. Gesellsch. Jena, Bd. TIT., Theil 2, 1893, pp. 312-317.
t True, F. W., ‘“‘ Whalebone Whales of the Western North Atlantic,’ Smithsonian Contributions to
Knowledge, 1904, Vol. XXXIII., Pl. 41, figs. 1, 2 and 4. Liouville, J., op. cit., Pl. IV., fig. 2.

100 “TERRA NOVA” EXPEDITION.

This alteration was brought about by the raising and lowering of the convexity of
each ramus, simultaneously, and respectively with the shutting and opening of the
mouth. The movement of the rami necessarily involves the elasticity of the
symphysis, which joins the two rami together at their anterior extremities. The
structure of the symphysis, upon examination, was found to be almost identical with
that of the intervertebral cartilages. '

It has been suggested that a whale swims by means of movements of the tail-
flukes, somewhat after the manner of a steamer which is forced through the water by
its propeller. If this were true it would imply that the flukes of the tail had
independent muscular movements of their own. Now this is surely impossible, since
the flukes are entirely composed of white fibrous tissue and blubber, and contain no
trace of muscular tissue or tendons.

Moreover, I have often watched the tail-flukes of the Piked Whale and of various
dolphins in the act of swimming, and have seen no other movement in the flukes than
that which takes place in the caudal fin of fishes. In the Cetacea it would seem that
the tail as a whole moves up and down when the animal is progressing, and the motion
appears to be very similar to that which takes place in the tail of a fish, except that
in the latter the movement is from side to side.

For this reason, the backbone of a whale has to be especially flexible at the
posterior end, in order to allow the powerful tail, which is the principal organ of
propulsion, to have free play, Consequently the vertebrae have imitated, as it were,
those of fishes, in substituting large intervertebral discs for the imterlocking, bony
processes of ordinary land vertebrates. These discs consist of a broad ring of white
fibrous tissue surrounding a central core of jelly-like pulpy substance, the nucleus
pulposus.

Precisely the same structure occurs in the symphysis of the lower jaw (text-
fig. 5, B). The thick ring of tough, elastic tissue surrounding a ball of jelly forms an
admirable hinge, and permits the necessary movement of the rami when the whale
opens and shuts its mouth. The amount of hinge-movement required in this case is
about equal to that of the intervertebral discs of the lumbar region.

A similar symphysis occurs in the mandible of Balaenoptera; and the above-
mentioned movement of the rami is probably characteristic of Whalebone Whales in
general.

The reason for this movement of the lower jaw is undoubtedly to allow of an
increase in the straining surface of the baleen. As shown in text-fig. 5, A, a, the
plates of baleen curve outward at their distal ends, and project beyond the margin of
the upper jaw to the extent of about a foot. By this means some eight square
feet are added to the straining area in a full-sized Humpback.

The outward movement of the convexity of the rami enables the mandible to
clear the projecting baleen-plates, when the mouth is opened and closed.

CETACEA—LILLIE. 101

Mammary GLanps.

The mammae of Megaptera were found to agree very closely with the descriptions
of these organs in other members of the Balaenopteridae given by previous observers ;*
so that it will only be necessary here to mention a few points which do not appear to
have been hitherto recorded. There were two furrows, about 13 feet long and 3 inches
deep, situated one on either side of the urinogenital opening, both of which contained
a nipple.

Near each nipple-groove, in the female, there was usually a smaller furrow without
a nipple. This accessory groove either occurred on the outer side of each nipple- .
groove or between the latter and the vulva. (See text-fig. 6, A.)

In several dead females the nipples were protruded beyond the general level of

js Saas os Ske

Fic. 6.—A—Diagram to show the position of the nipple-grooves in the Humpback :
a, anus ; b, vulva; ¢, nipple-groove ; d, secondary furrow ; ¢, alternative position
of latter. B—Lateral view of walls of nipple-groove everted : a, nipple.

the ventral surface of the whale, by the eversion of the walls of the nipple-grooves.
(See text-fig. 6, B.)

When everted, the walls of each furrow formed a dome-shaped swelling, with the
nipple on its posterior side. The latter was thus raised about 3 inches above the
general level of the surface of the body. When unprotruded it was nearly 3 inches
beneath the surface.

Iam unable to say whether this protruded condition of the teats was a natural
occurrence, or was produced by abnormal causes after death. It was of interest, however,
to note that it could occur, and could undoubtedly facilitate the process of lactation.
That it was a natural condition of the teats is suggested by the latter function, and by
the presence of erectile tissue around the nipple-region. Moreover, the looseness of the
skin in this locality, together with the secondary furrows, would allow of the necessary
expansion of the nipple-grooves for the protrusion of their walls.

An elongated compressor muscle covered the posterior two-thirds of each
mammary gland.

The anterior bundles of this muscle arose in the superficial fascia between the two
breasts, and passed across each gland in an oblique direction to their insertions in the

* Hunter, J., Phil. Trans. Roy. Soc., 1787, p. 75, Pl. xxi. Turner, W., Trans. Roy. Soc. Edinb.,
Vol. XXVL., 1870, pp. 197-251. True, F. W., ‘‘ Whalebone Whales of the Western North Atlantic,”
Smithsonian Contributions to Knowledge, 1904, Vol. XX XTIT., Pl. 20.

VOL. I. Q

102 “TERRA NOVA” EXPEDITION.

superficial fascia on the outer side of each breast. The posterior bundles arose at a
point immediately below and in front of the nipple, and were inserted laterally, as in
the case of the anterior bundles (text-figs. 7, e, 8, A). Each mammary gland consisted
of a flattened oblong body, which passed from the nipple in an anterior and slightly
dorsal direction. (See text-fig. 8, A).

In a female 40 feet in length, which was suckling a calf when killed, the gland
was 53 feet long, by 14 feet broad at its widest part, just im front of the nipple. The

Fic. 7.—Ventral view of a Humpback, with the coat of blubber removed, to show
superficial muscles (diagrammatic) : a, mylohyoid muscle ; b, pectoralis major ;
c, umbilicus ; d, vulva ; e, compressor muscle of left breast.

ren IN ARN \ << <<

Fie. 8..-A—Ventral view of left mammary gland of a Humpback: the thick line
marks outline of gland ; compressor muscle indicated with fine lines: a, nipple.
B—Longitudinal section through gland: a, nipple with narrow duct; b, wide
portion of central canal.

nipple contained the narrow mouth of a relatively wide canal, which traversed the
centre of the gland throughout its whole length. In the broad portion of the breast
above-mentioned, the canal was dilated into a spacious reservoir. A large number of
ducts united with the central canal and ramified in the substance of the gland. (See
text-figs. 8, B, and 9.) It would appear that the central canal is an elongated
false teat.

The milk collects im the reservoir, and by the contraction of the compressor
muscle a stream of the fluid is forced through the nipple into the mouth of the calf.
The latter is quite unable to draw off the milk from the breast by sucking, after the

CETACEA—LILLIE. 103

manner of an ordinary mammal, because there is a complete absence of air in its
mouth at any time.

When the mother feels the lips of her calf at her teat she pumps a supply of milk
into its mouth. It is said that the calf holds the teat in the angle of the mouth near
the eye.

A whaling captain told me that he once shot a female Balaenoptera physalus
Linn. when im the act of giving suck to her calf. After death the ventral surface
turned upwards, as it always does, and the nipple-grooves were seen to be protruded.
On inflating the whale with air, in order to render it more buoyant for towing home,
two jets of milk from either breast rose into the air, like small fountains ; some of the
milk was collected in pans. The amount of milk which issued from this animal was
estimated at 18 or 20 gallons. The discharge of milk, in this case, was evidently due to
the pressure of air within the body-cavity causing the tissues of the abdominal wall
to tighten, and so squeezing out the contents of the mammary glands.

TITIAN

{HL

Fic. 9.—Sagittal section through mammary gland and adjacent structures of a
Humpback: A, nipple; B, blubber; C, compressor muscle of gland ;
D, substance of mammary gland; E, its central canal; F, muscles of body-
wall; G, body-cavity.

The milk was like thin cream in appearance, and has been used for cooking
purposes by whalers. When eaten raw it has a slightly sweet flavour, and leaves a
faint taste of oil in the mouth after it is swallowed.

In the males, the mammae were similar to the description given by Struthers, in
his account of a male Humpback caught off the coast of Scotland.* The compressor

muscles were less developed in the males than in the females.

Aupirory ORGAN.

The auditory apparatus was examined in several adult Humpbacks, and also in
the foetus 13$ feet in leneth. In every case I was surprised to find the external
auditory meatus completely closed up for a few inches of its course, on the inner side
of the blubber,

* Struthers, J., Journ. Anat. and Physiol., 1888, Vol. XXII, p. 117.
Q 2

104 «TERRA NOVA” EXPEDITION.

The position of the small, slit-like earhole on the surface of the head of this whale
has been noted by Struthers.*

From this orifice a narrow tube, about 5'5 inch in diameter, traversed the blubber,
which was about 34 inches thick in this region of the head. The tube was continued
through the underlying tissue for about 2 inches, and gradually decreased in diameter
until it ended blindly.

The meatus was here closed up for 3 inches of its course. (See text-fig. 10, N.)
It then widened out again, somewhat abruptly, to a diameter of rather more than

Fic. 10.—Diagram of the outer and middle ear of a Humpback (Meyaptera).
A, passage of nares leading to blowhole ; B, opening of pterygoid fossa tube
into nasopharynx ; C, opening of Eustachian tube into nasopharynx ; D, velum
palati; E, epiglottis tube; F, pharynx leading to mouth; G, opening of air-
passage into pterygoid fossa; H, opening of Eustachian tube into tympanic
cavity ; 1, tympanic bulla ; J, coat of yellow elastic tissue and fat ; K, spong
layer containing air; L, tympanic membrane; M, wide inner portion of
external auditory meatus containing plug of ear-wax; N, portion of meatus
closed up ; O, narrow portion of meatus opening on surface of head.

an inch, and maintained a fairly uniform size for the remainder of its passage to
the tympanic bulla.

The total length of the canal was about 1 foot 9 inches in a Humpback whale
40 feet in length.

The walls of the wide innermost portion of the meatus (text-fig. 10, M) were
invariably pressed together. The sole contents of this tube consisted of the finger-like

* Struthers, J., Journ. Anat. and Physiol., 1888, Vol. XXIL., p. 122.

at

CETACEA—LILLIE. 105

external surface of the tympanic membrane and the plug of ear-wax, exactly as they
oceur in Balaenoptera.

Mr. Burfield and Mr. Erik Hamilton noticed that in several of the Balaenoptera
examined by them off the Irish coast, the meatus was closed up for a part of its course,
as described here in the case of the Humpback.

I overlooked this closed area when working at the ear in Balaenoptera musculus
Linn. and B. physalus Linn. a few years ago.* The presence of water in the imner
portion of the meatus of one or two of these specimens led me to think that the
meatus was open to the exterior throughout its whole length, and was normally full of
sea-water. Iam now inclined to believe, however, that the occurrence of the water in
the meatus was accidental, and due to the whalers playing the hose over my dissections
when washing the “ flensing slip” during my absence. Sea-water hoses were kept
constantly running when whales were being cut up on the “slip.”

Carte and Macalister reported the meatus as open in Balaenoptera acutorostrata,
Lacépéde, and it may be so in that species; but unless the canal is very carefully
sectioned throughout its entire length it is easy to overlook the few inches which are
closed up.

In Balaena mysticetus the meatus has been figured by Gray,f and was recorded
by him as open, though of a very small diameter in its outer part. The investigation,
however, does not appear to have been sufficiently close to preclude the possibility of
his having overlooked the closed area.

The plug of ear-wax which occurs in the meatus of the Humpback has been
recently figured and described by Sir William Turner.§ — It is therefore only necessary
to say that plugs similar to the one described by him were found in all the adult
specimens; but the plug had not begun to form in the embryo of 134 feet in length.

With reference to Turner’s suggestion that the length of the plug may bear a
relationship to the thickness of the external coating of blubber over the head, I would
here point out that the plug only occurred in the wide portion of the meatus, close to
the tympanic membrane, and was therefore more than a foot away from the coat of
blubber, in a Humpback of average size. Moreover, the diameter of that portion of
the meatus which traverses the blubber is only 7/5 inch, whereas the width of the plug
is seldom less than $ inch in its narrowest part.

The tympanic membrane had the same shape as in Balaenoptera. In an adult
whale, the total length of the sac and ligament was 34 inches, and the greatest
diameter was 1 inch.

The walls of the meatus were compressed and closely surrounded the plug of wax,
imprinting upon it the impressions of the ridges and furrows of their inner surfaces.

* Lillie, D. G., Proc. Zool. Soc., 1910, p. 775.

} Carte and Macalister, Phil. Trans., 1867, p. 252.

t Gray, R., Journ. Anat. and. Physiol., Vol. XXIIT., 1889, p. 300.

§ Turner, Sir W., Proc. Roy. Soc. Edinb., 1913, Vol. XXXIV., Pt. 1 (No. 2), p. 11.

106 “TERRA NOVA” EXPEDITION.

With the exception of the above structures, the inner portion of the canal was empty.
The lack of pressure in the meatus enabled the sac-lke tympanic membrane to project
for 3 inches into its lumen. If the canal had been open to the sea-water, this extreme
outward convexity would be very difficult to understand.

The tympanum in Monodon and Phocaena is concave externally, and the meatus
is open. In the following whales the membrane is known to be convex on its outer
surface, and it is extremely probable that the meatus is closed in all these cases,
as it undoubtedly is in the Humpback; Balaena mysticetus and Bb. glacialis ;
Balaenoptera musculus, B. physalus, B. borealis, and B. acutorostrata; Megaptera
nodosa and Hyperoodon. It is possible that this condition of the auditory canal occurs
throughout all the larger Cetacea.

The tympanic bulla was examined in several specimens and was found to be identical
in shape with that of the Northern Meyaptera. The species Megaptera novae zealandiae
was founded by Gray on the ear-bone alone, which was found by him to be shorter
and more swollen than in JL nodosa.

After a careful comparison of the identical bulla
from M. novae zealandiae which Gray * described,
with those of J/. nodosa Boun., | have been unable
to find any point of difference between the specimens,
which could not be amply accounted for by differ-
ences of age and individual variation. For instance,

the bullae of the 135 feet foetus were very much
more rounded and swollen than those of the adults.
(Plate III, figs. 1-3.)

A specimen f of the left tympanic bulla of J/

H

lalandii, Fischer was examined and found to be
Fic. 11.— Humpback :

diagrammatic

1876.

section through the layers of tissue
which surround the tympanic bulla
and separate the tympanic cavity
the pharynx: A, tympanic
cavity; B, bony wall of bulla ;
C, fatty tissue; D, yellow elastic
tissue; E, spongy tissue containing
air; F, muscular layer of pharyngeal
wall; G, mucous membrane; H,
cavity of pharynx.

from

exactly like the bulla of Northern specimens of J/.
nodosa, and those from New Zealand which were
obtained by me.

The bulla was surrounded ventrally by two
distinct coats of tissue, each of which preserved,
more or less, the shape of the tympanic bone.

The inner layer (see text-fig. 11, C and D) was
about 3 inches thick and lay close against the bulla.

* Gray, J. E., Catalogue Seals and Whales, Brit. Mus., 1866, p. 128, and Proc. Zool. Soc., 1864,

p: 207.

The right tympanic and periotic bones described in these papers were presented to the British
Museum by Mr. Stuart in 1864 (No. 1468, now labelled M. lalandii).

Turner, Sir W., ‘“ Marine

Mammals in University Mus. Edinburgh,” 1912, p. 65.
{+ Specimen in British Museum labelled, “ Left Tympanic of Megaptera lalandii, New Zealand,

Presented by the Wellington Museum.”

Turner, W., “ ‘ Challenger’ Report, Cetacea,” pp. 30-31,

Hector, Sir J., Trans. New Zealand Inst., 1878, Vol. X., page 335,

CETACEA—LILLIE. 107

It consisted of a firm case of yellow elastic tissue (D), which enclosed a loose
fibrous layer, heavily charged with fat (C). The outer coat (EK) was formed of
spongy tissue containing air-spaces, and attained a thickness of some 4 inches
or more. The above-mentioned caps of tissue were underlaid ventrally by the
muscular wall of the pharynx (F). The tympanic cavity was thus separated from the
cavity of the pharynx by very nearly a foot of tissue. The tympanic cavity and air-
spaces, such as the pterygoid fossa, were the same as in Balaenoptera, The air-passage
(see text-fig. 10, B-G) which opened on the dorsal wall of the naso-pharynx, close to
the inner extremity of the nasal septum, and communicated with the pterygoid fossa,
was termed by me the Eustachian tube, in the description of the auditory apparatus of
Balaenoptera.* This tube should, however, be called the pterygoid fossa tube, as the
opening of the Eustachian tube proper was situated in a shghtly more posterior
position, on the ventral wall of the naso-pharynx, near the free edge of the velum palati.

The Eustachian tube, which was very much wider than the air-passage supplying
the pterygoid fossa, led directly to the tympanic cavity, and also supplied air to the
coat of spongy tissue surrounding the bulla.

The question naturally arises: how do sound-waves reach the labyrinth in
whales? In the smaller Cetacea, such as Phoeaena and Monodon, the external meatus
is open, and the tympanum is concave externally, as it is in other mammals. It would,
therefore, seem possible that in these whales the sound-waves are transmitted to the
middle-ear through the water in the meatus.f

But in the larger forms, such as JJegaptera, the meatus is closed up, and there is
no way for sound-waves to reach the ear through the mouth.

It is possible, however, that these whales hear through their nasal passages.
Sound-waves could be conducted through the water to the alar fibro-cartilages, which
form the elastic lids of the external nares. Within the blowholes there is a continuous
column of air, leading into the tympanic cavity through the Eustachian tube. Sound-
vibrations could be transmitted through this air to the walls of the dense, cowrie-shaped
bulla, which is attached to the periotic by two slender pedicles, and is surrounded by
air-containing tissue. The bulla should, therefore, act as a sounding-box, and it is
connected to the fenestra ovalis by the chain of ossicles.

It is also possible that sound-waves are conducted from the alae of the external
nares to the bulla, by means of the nasal septum. In this case the mechanism of
audition in the larger Cetacea would resemble, to some extent, that of the diving
Mosasaurians, such as Plioplatecarpus,f although the parts involved are formed out of
very different structures morphologically. The fibro-cartilaginous covers to the nostrils,
together with the attached cartilage of the nasal septum, which occur in whales, would

* Lillie, D.G., Proc. Zool. Soc., 1910, p. 779.
+ Denker, A., Anat. Hefte, Wiesbaden, 1902, Abt. 1, Bd. XIX., Heft. LXII., pp. 423-447.
t Dollo, L., Bull. Soc. Belge Géol., Bruxelles, Tome XTX., 1905, p. 125, pl. IIT.

108 “TERRA NOVA” EXPEDITION.

correspond in function to the ossified tympanic membrane and the extra-columella
expansion in Plioplatecarpus. The bony septum formed by the vomer, together with
the pterygoid and periotic bones, the bulla and ossicles, would perform the function of

the columella in the whale-like reptile.
If neither one nor the other of the above modes of sound-conduction be correct, it

is difficult to understand how these whales hear. Yet it is firmly believed by whalers
that they are sensible of the report of a gun and similar sounds.

SCAPULA AND VERTEBRAE.

The shape of the scapula was identical with that of the Northern JJegaptera.*
There was no sign of an acromion process on the blade-bone of any of the adults, or on
that of the 134 ft. foetus. (Text-fig. 12.) This point was carefully examined, as the
occurrence of an acromion process on the scapula was said to be one of the specific

Fig. 12.—Megaptera nodosa, Bonn. Scapula of 134 feet foetus. Fic. 13.—Left scapula of New Zealand
Bay of Islands, N.Z. Humpback whale (adult): C.P.,
coracoid process.

characters of JZ. lalandi Fischer.t There was a slight trace of the coracoid process.
(See text-fig. 13.) In other osteological characters the New Zealand Humpback
was in agreement with Northern types. The vertebral formula was ascertained in one
case and found to be C. 7, D. 14, L. 10, C. about 20. The cervical vertebrae were
not united in the young specimen. It is extremely probable that the anterior cervicals
in this species become fused in old age.

* Cabrera, A., ‘ Fauna Ibérica. Mamiferos,” Madrid, 1914, Cetacea, p. 398.
{+ Van Beneden et Gervais, “ Ostéographie des Cétacés,’ p. 133. Turner, W., ‘“ Challenger”
Reports, Vol. I., p. 30.

CETACEA—LILLIE. 109
ForrusEs.

Two foetuses only were obtained. One of these measured 134 feet in leneth and
yas taken from a mother of about 44 feet in length, caught on July 24th, 1912. The
measurements of this foetus have been given in Table II. on page 89. It had very
nearly completed its uterine existence. The other embryo was taken from a mother of
about 48 feet in length and some 60 tons in weight, which was killed off the Bay of
Islands, New Zealand, on October 10th, 1912. This specimen (Plate IV., fig. 4)
is 61 mm. in total length, and must be one of the smallest embryos known from any

Whalebone whale. The general measurements are as follows :

Total length to root of tail , : : 5 : : . 43 mm.
Head-length . 5 : ° : : : ; ; . 25 mm.
Taillength . : : : : : : ; . 18mm.
Tip of snout to external nares . : : . F . 10mm.
Length of fore limb (posterior edge, axil to ip) ; ‘ : . 12mm.
iene of longest digit . : : : 2) emm:
nani of eye (between inner angles a ey eli) : : : . 1:5 mm.

Hair-tubercles can be seen on the snout, arranged like those in the adult. There
are four along the median line, between the nostrils and the tip of the snout; and a
row of nine on each side of the beak, towards the outer edge.

The manus has four digits. The third and fourth digits are much longer than the
second and fifth.

The tail-flukes are only just perceptible, when viewed from the dorsal or ventral
aspect. It is hoped that a full account of this embryo will be published later.

Foop.

A careful examination of the stomach was made in many cases, but very few of
the whales had any trace of food in their alimentary canals. This was not altogether
a surprise, when one considered the relatively small amount of plankton obtained by
the “Terra Nova” in her daily hauls off the north of New Zealand, during the months
of July, August, and September, 1911. The only food organisms found in the stomachs
were unidentifiable remains of Schizopoda.

Hasits.

The Southern Humpbacks, like those of the Northern Hemisphere, are somewhat
slower in their movements than species of Balaenoptera. They seldom stay under
water for more than seven minutes, between each two acts of respiration. They show
a distinct partiality for coastal waters; and it was quite a common sight, on the
coast of New Zealand, to see a school of three or more of these whales pass through
the narrow channel between an isolated rock and the mainland (p. 94). This habit
was formerly made use of, during the whaling season, by Mr. Cook of Whangamumu.
He placed a wide-mesh net, made of wire rope, across the channel, some 50 yards
broad, between a rock and the shore. A look-out was kept from the cliffs ; and when

VOL, I, R

110 “TERRA NOVA” EXPEDITION.

a whale became entangled in the net, a boat was immediately despatched to shoot the
captive by means of a Hotchkiss gun.

An average of about eight Humpbacks each season was obtained by these means,
over a period of nearly twenty years. No other species were taken in the net.

DISTRIBUTION AND MIGRATION IN THE SOUTHERN HEMISPHERE.

July 29,1910 . 22 RMS 528 selene : . School of two.

August 4, 1910 5 a0? 2S Tee a} WY : . School of five or six travelling S.W.
*October 26, 1910 5 AUB ISE, TO? 24 1 : . School of five or six travelling south.
*August 11, 1911 . Off the Three Kings Islands, N.Z. » School of three travelling north.
*March 15, 1912 + 68° BUS 69e 45iek: ;
*January 30, 1913 . 67° 'S., 1672 25! EE:

The above list gives the occasions on which Humpback whales were seen by the
“Terra Nova.” On the dates marked with an asterisk the whales showed themselves
very clearly, so that identification was quite certain. In spite of careful observations,
we did not see any of these whales in the Ross Sea; which tends to support the
opinion arrived at by M. Racovitza,* that the Humpback does not penetrate into ice-
covered seas, although it is very common in Antarctic waters wheresoever they are
fairly free from ice. It used to be the whale chiefly taken by the whalers who go
every summer to South Georgia, the South Shetlands, and Graham’s Land, but it has
become less common at some of these localities during the last two or three years. The
Humpback is also captured off the east and west coasts of South America, South Africa,
New Zealand, Tasmania, Norfolk Island, and Australia.

Excellent opportunities were afforded to me for studying the seasonal migrations
of Humpback whales, off the north of New Zealand, during the winters of 1911 and
1912. There seemed to be very good evidence that these whales spend the summer
months in the Antarctic Ocean, down to the northern limit of the pack-ice; and
that at the beginning of winter they migrate northward into the warm seas, in the
neighbourhood of New Zealand and Norfolk Island.

The first whales of the season began to pass the Bay of Islands, on their way
northward, about the middle of April. They continued to go towards the north
until the end of August. The greatest number passed northward of this locality in
May and the early part of June. After the middle of September, at the Bay of
Islands, the first members of the long procession were to be seen going southward, on
their way back to the Antarctic Ocean. The majority passed south of the Bay during
October, and by the middle of December they were all to the south of this place.

The whaling season at the Bay of Islands lasts from May to November. There
is a slack time in the middle of the season, from July to September, when all the
whales are to the northward. These three months are the busy time for the whalers
at Norfolk Island.

* Racovitza, E., “ Résultats du voyage du 8.Y. ‘ Belgica,’ Zoologie, Cétacés,” 1903, pp. 32 and 46.

ay

CETACEA—LILLIE. 111

These regular migrations, which are described here probably for the first time,
have been known to New Zealand whalers for many years, and they adjust their
headquarters in accordance therewith. In the winter, whaling operations are carried
on from the Bay of Islands; and in the summer from Campbell Island, in the
Sub-Antarctic. The plankton samples taken on board the ‘Terra Nova” during
her three summer voyages from New Zealand to the Ross Sea, and by Mr. Nelson
at the winter quarters in McMurdo Sound, tend to show that there is considerably
more food for Whalebone whales in the cold Antarctic waters than in the warm
seas to the north of New Zealand. Diatoms were so abundant in parts of the Ross
Sea, that a large plankton net (18 meshes to an inch) became choked in a few
minutes with them and other members of the Phytoplankton. It is extremely
probable that m such localities whales feed upon the plants as well as the animals
of the plankton. Moreover, as whalers are only too well aware, the blubber is
appreciably thicker when the animals are leaving the cold waters than it is when
they are on their way South after their sojourn in the sub-tropics.

It would therefore seem unlikely that the migration of Humpbacks from South
to North in the winter can be caused by the search for food. .It is far more
probable that this journey is undertaken for the purpose of parturition, in order
that the calves may be born in congenial climes. The latter motive received
support from the fact that mothers, with newly born calves, were constantly to be
seen off the Bay of Islands after the beginning of July. On July 24, 1912, a female
was shot, from which I obtained a foetus 133 feet in length. The length of the
calf at birth is about 15 feet in this species.

On October 10, 1912, I found a foetus 24 inches long, and possibly three months
old, nm a female which was killed while on her way southward.

Foetuses of intermediate length have never been seen by Mr. Cook, during the
twenty years he has been at the Bay of Islands.

The above data suggest that parturition, and possibly copulation, take place in the
warm seas during the Southern winter ; and that the mother carries the calf throughout
her summer visit to the Antarctic Ocean. The period of gestation in this species
is either about twelve or twenty-four months, according to whether the mother
carries the calf for one, or two seasons. Several young, unpregnant females were
seen at the Bay of Islands, which showed that the northward migration was not
confined to females in the last stages of pregnancy; and the apparent absence of
foetuses of intermediate size, at the New Zealand Whaling Station, tends to support
the shorter estimate for the period of gestation.

From information given to me by whaling captains, who have worked at their
trade in various parts of the Southern Hemisphere, and whose veracity of statement
I have tested and found to be reliable, it appears that similar migrations of
individuals of this species are known to take place off the shores of South America,
South Africa, Tasmania, and Australia. This evidence tends to show that in the

in 2

112 : “TERRA NOVA” EXPEDITION.

Southern summer the Humpback whales concentrate in the Antarctic Ocean, where
they are found in great numbers off the South Shetlands, South Georgia, and
similar localities, between the months of November and March. They turn North
in the autumn and radiate outwards, passing up the coasts of the three great
continental areas, towards the equator. But as it is scarcely possible to observe
the Cetacea in the Antarctic Ocean during the winter, we do not know how many
remain there throughout those dark months.

It therefore appears to be possible, as far as the Mystacoceti are concerned, to
divide the Southern whaling grounds into’ three classes. The first class, or best,
‘fisheries,’ are in the cold Antarctic seas, where there is plenty of food for
Whalebone whales. These grounds can only be worked durmg the summer months,
on account of their high latitude.

The second class areas are off the shores of the great continents, between
Lat. 50° S. and the equator. These waters probably owe their favourable character
to cold southern currents, such as those which run up the western shores of South
America and South Africa; and also to the rivers, which carry ammonia into the
sea from the land, and thereby minister to the needs of the Nanno-plankton, upon
which the plankton animals feed, which are in their turn the food of the Whalebone
whales.

The best season for these grounds appears to be during the winter.*

The third class, or poorest, fields are in the warm, open oceans, where the
plankton-supply falls to a minimum. At the present time it does not pay whalers
to exploit the open seas for the Mystacoceti.

The foregoing general scheme of what we may perhaps call the quantitative
distribution of the Whalebone whales, although it would seem to apply to all
members of the group, is perhaps best demonstrated by the case of the Humpback.
The coast-loving habits of this whale cause it to be more readily observed than the
species of Balaenoptera, which tend to keep further out at sea.

As an illustration of the so-called ‘bad luck” which accrues to whalers who
fail to realise the quantitative distribution of whales, | may mention the case of
the Company with whom I spent four months. They exploited the warm seas to
the north of New Zealand, and employed there a “plant” which was on a scale
suitable for a first-class whaling ground, such as South Georgia. The result was they

*The cold currents on the western sides of the continents are due to an up-welling of cold water from
the bottom of the oceans. In the tropics, the deeper layers replace the water carried westward by the
trade winds as the Equatorial Current. In temperate regions the greatest vertical circulation takes place
in the winter, when the surface-temperature is at a minimum, and the denser upper layers sink down and
are replaced by the warmer water from below.

The bottom-water is said to be comparatively rich in ammonia from the decomposition of organic
remains on the sea floor; and consequently when this water comes to the surface, during the winter

months, the plankton reaches a maximum, See “The Depths of the Ocean,” by Sir John Murray and
Dr. J. Hjort, London, 1912, pp. 371 and 378.

CETACEA—LILLIE. 113

could not get whales enough to make the enterprise pay, and had to leave. New
Zealand waters are poor whaling fields, and should only be exploited by small concerns.
These remarks on distribution and migration are put forward tentatively, and
can lay no claim whatever to finality. They are based upon such facts as we
possess ; but a great deal more work will have to be done before we can hope for any
certainty as to the movements of all the different whales over the oceans of the world.

4. Balaenoptera acutorostrata, Lacépede. (PI. IV., figs. 2,3; Pl. V., figs. 1, 2;
PPV ties. 1—65-P), VII, fe. 1.)

December 10, 1910 : ; : : 66° 38’ S., 179° 04’ W.

ij 1S. : ; , : 67° 24 S., kre 34’ W.
- PB. op : : : : 68° 25' S., 179° 11’ W.
sat SLO is : ; . 70°S., 180° W. '
January 2, 1911 é ‘ : ; (sy OBY Shy line}? 24 1d,
5 Shi ey i ; : : Off Cape Crozier, Ross Island.
A Les F : ; : In McMurdo Sound.
3 265° 5 Off Glacier Tongue, McMurdo Sound.
ta Sees 78. 30) S:, 140? 35! W:
February 2, ,, Off King Edward VII. Land.
” 3, ” a a
3 a ss Bay of Whales, Great Ice Barrier.
a i, 55 (AS UOMS§ Mef AAO! VAY
i Gh 55 Ce Oe Shy LUPE Nive
is ie 77° 33' S., 176° 51’ E.
oi oss. (2a OP Seliger bbe
; DI 69° 10' S., 164° 30’ E.
March Lae 67° 25’ S., 160° 40’ KE.
~ By, ig 67° 22'8., 160° 31’ E.
an Use Ss 65a. 02 SS leh 22/5R:
January 2, 1912 WOn O20 Sa litiie ole Bre
m BS 5. U2 WTS, 2 (ayy Ide
A (ema (Asan Ole Semel yileenl Sib;
- irae ee (ie Way tsi, UES Bhi) 1B).
7 We os (OS Misys, a OY 10%
” 15, ” ar a
March IMSy cp 64° 03’ S., 160° 12’ K.
= OTe os Bp MG Si isi Gilly 13.
December 29, _ ,, GORE Se V66r lie We
ls Me 23S. 166273. We

>

January 1, 1913 ; ; ; : Tilero Dass LOOm Ole Wie

* By 5p 5 3 : : 71° 37’ 8,, 166° 55’ W.
” 4; ” c G 2 . fara <a

ines , , 71° 41'S., 166° 47’ W.
e a ee 71° 44’ S., 167° 57! W.
a ae 76° 32’ S., 163° 50’ E.

When sailing in Antarctic waters to the south of Lat. 64° 8., scarcely a day passed
without our getting a sight of one of these whales. The above list gives the occasions

on which members of this species were clearly seen.

114 “TERRA NOVA” EXPEDITION.

This whale was mistaken for Neobalaena marginata by Dr. Wilson when on board
the “Discovery” in 1901-4. On the “Terra Nova” Expedition, although we knew
that Balaenoptera acutorostrata had been identified in New Zealand and Antarctic
waters,* and we were quite prepared to find it in the Ross Sea, yet there was only one
occasion on which these whales showed themselves sufficiently well to put their
determination beyond dispute. This was on March 3, 1911, in Lat. 67° 22) Ss lone:
160° 31’ E., when the ship was ploughing her way through thick pack-ice, in which the
water was freezing between the floes, so that the only open spaces for miles around
were those made by the slow movement of the ship. We saw several of these whales
during the day, making use of the holes in the ice near the ship for the purpose of
“blowing.” There was scarcely room between the floes for the whales to come up to
“low” in their usual manner, which consists in rising almost horizontally, and breaking
the surface of the water with their backs. On this oceasion they pushed their snouts
obliquely out of the water, nearly as far as the eye, and after “ blowing,” withdrew

“several times one

them below the water again. Commander Pennell noted that
rested its head on a floe not 20 feet from the ship, with its nostrils just on the water-
line; raising itself a few inches, it would blow and then subside again for a few
minutes to its original position with its snout resting on the floe. They took no notice
of pieces of coal which were thrown at them by the men on board the ship.” (See
Pl. VL, figs. 1-6.)

They swam close to the surface, in the pool of open water under the stern of the’
ship, and frequently rolled over in play, so that their external characters could be
easily determined.

They were between 20 feet and 30 feet in length. The dorsal fin was shaped as in
Pl. V., fig. 2. The throat-grooves numbered about sixty, as far as we could
ascertain.

In colour they were greyish black above, and most of them were white
throughout on the underside. A few were grey ventrally, with white patches in the
middle of the ventral surface. There was considerable variation in the amount of
pigment on the lower surface of the body. A characteristic triangular patch of
lighter colour oceurred on either side of the back, in the region of the dorsal fin.
(ScecP ie Vii coe)

In the majority of cases, the pectoral “ flippers” had a broad white band on their
dorsal surfaces, but in some, the “ flippers” were almost entirely white above and
below. The posterior dorsal margins of the tail-flukes were white im some cases.

* Hutton, F. W., and Gray, J. E., Ann. Mag. Nat. Hist., 1874 (4), Vol. XIII. J. von Haast,
Trans. New Zealand Inst., 1880, Vol. XIJI., p. 169. Bull, H. J., “The Cruise of the ‘ Antarctic,’”
1896, p. 141. Benham, W. B., Trans. New Zealand Inst., 1901, Vol. XXXIV., p. 151. Racovitza, E.,
« Résultats du voyage du 8.Y. ‘ Belgica,’ 1897-99, Zoologie, Cétacés,” 1903, p. 56. Liouville, J., op. cit.,
p- lll. Stead, D. G., “A Brief Review of the Fisheries of New South Wales,” Sydney, 1910. Trivett,
J. B., Official Year Book of New South Wales Fisheries, 1912. Published 1914.

CETACEA—LILLIE. 115

On January 2, 1911, Dr. Wilson saw one of these whales fairly close to the ship,
and recorded his impressions as follows: ‘This whale was about 30 feet long. Its
nose was sharp and rorqual shaped, and its dorsal fin like BL. musculus [ie B.
physalus)|. The pectorals were rather square and quite white. There was a patch of
white, or yellowish white, on each side running diagonally upwards, abaft the dorsal
fin.” On January 4, 1911, he says: ‘‘ same species seen to ‘ breech’ and shoot its head
and shoulders out of the water upside down.”

The only character in which these whales appeared to differ somewhat from the
descriptions of B. acutorostrata* was in the slightly greater breadth of the snout.
Whether these little Rorquals were identical, or not, with the northern Piked Whale, I
feel convinced that they belonged to the same species as the young Rorqual from New
Zealand which was described as Balaenoptera rostrata by Dr. Benham,¢ who has kindly
allowed me to make use of some unpublished photographs of this specimen. (Pls. LV.,
figs. 2 and 3, V.)

The Norwegian whalers whom I met in New Zealand, and found to be fairly
shrewd judges of the different species of whales, told me that the “‘ Minkehval,” which
is their name for &. acutorostrata, is shot off the South Shetlands, during the whaling
season, for human food. Sir William Turner has recently described a foetus and
tympanic bone belonging to this species, which were sent to him from the South
Atlantic.{ Mr. D> G. Stead includes this species in his list of the whales hunted in

Australian waters.§

Linnaeus. (PI. VIL, fig. 1.)

5. Balaenoptera musculus,|

December 9,1910. . 65° 8'S., 177° 40' W. February 22,1911. . 69° 10'S., 164° 30’ E.
x NO ee On OL Olio. 28) Wie || 0 285037 oe LAS G0 88a b-

84, | 69° 1S, 178°28 W. | March 2, ,, . 67° 25’ S., 160° 40’ E.
ae a a ’ pane 66° 37’ S., 161° 42! E.

a QO en = — — December 27, ,, 64° 56’ S., 175° 30' W.
January, 29,1911. © 78° 12S:, 174° 44° B. January 5, 1912 fr VOSS i205) By:
3 30 e ioe 2405 176- Do! Er. December 22, ,, 55° 34’ S., 174° 35’ W.
Sem OMS LTO od" We January 14,1913. (oe akey Sh, Ile 0! Mie
Hebruanye ii, 2 . (8. 38's. 166° 17’ W. Fr og WAS EX SES ie at oe
be 6 Mim OQeSs line 2a NV. February 2; 5, 90. Ole 18085 lov” 53).
TON =. 9.0729 008s, 171° 56! B. Apal 18, pe ee eet2° 0978, “55e 1ouiwe

”

* Carte and Macalister, Phil. Trans., 1868, pp. 201-261, Pls. 4-7. Turner, W., Proc. R. Soc. Edinb.,
1891-92, pp. 36-75. ‘True, F. W., “ Whalebone Whales of the Western North Atlantic,’ Smith-
gonian Contributions to Knowledge, 1904, Vol. XX XIIT., p. 192, Pls. 22-28 and 49.

+ Benham, W. B., Trans. New Zealand Inst., 1901, Vol. XX XIV., p. 151.

t Turner, Sir W., Proc. Roy. Soc. Edinb., 1914, Vol. XXXV., Pt. 1 (No. 2, p. 12).

§ Stead, D. G., “A Brief Review of the Fisheries of New South Wales,” Sydney, 1910. Trivett,
J. B., Official Year Book of New South Wales Fisheries, 1912. Published 1914.

|| Or Balaenoptera sibbaldii, Gray. Turner, Sir W., Trans. Roy. Soc. Edinb., Vol. XX VI., 1870,

pp. 197-251, pls. 5-8.

116 “TERRA NOVA” EXPEDITION.

The Blue Whale was nearly as common in the Ross Sea as the little Piked Whale.
These two whales were often seen together, and they both appeared to be very fond of
frequenting the pack-ice.

Some of the Blue Whales which we saw were of very large size, and must have
been fully 90 feet in length, or even more.” There was no mistaking these whales, with
their characteristic dorsal fin situated well aft. (See Pl. VIL, fig. 1.)

A young immature male of this species was taken off the Bay of Islands, New
Zealand, during my visit to the Norwegian whaling station. This specimen was
65 feet long. In general appearance it exactly resembled the individuals of this
species which I had examined in the Northern Hemisphere. It was of a bluish grey
colour with lighter flecks all over the body, which has been likened to the appearance
of galvanised iron. The only white on the animal was on the underside of the
flippers. The baleen-plates were entirely black, with coarse, black, curly bristles on
their inner edges. The tongue was dark grey, and the palate was black.

This whale is obtained off the South Shetlands, South Georgia, South America,
South Africa, New Zealand, and Australia.*

6. Balaenoptera physalus,~ Linnaeus.

July igs, USO ge EPO PIN, BIS BO MW | March 10, 1912. 70° 28'S., 174° 39! BH.

October 6, ,, . « 41° 46)S:, 121° 39' H: | December!25, 4, 5 =. 62e 20ES Salo ome

December 5, ,, . . 56° 41’ S., 176° 23’ E.? | -March 24, 1913. 55° 55' S., 152° 39’ W.

March 9 191 . . 6725'S. 160240) Ee |) Atpril Bh gp 52° 03’ S., 63° 46’ W.

January 5, 1912 . . 72° 19'S., 172° 05' B ? | 33 We} 55 ADS 09ESe Dom Dae
acne. 75° 23'S., 164° 39’ E.?

The Common Rorqual, or “ Finner,” appeared to be very scarce in the Ross Sea.
We only saw this whale eleven times during the whole progress of the Expedition, and
on some of these occasions the determination was not certain. The doubtful occurrences
have been marked in the list with a note of interrogation,

I saw a stranded member of this species at Stewart Island, in March, 1911, and
although it was very much decomposed there was no doubt about the determination.
One of these whales was caught off the Kermadec Islands in October, 1912; I did not
see it, but was assured by the whalers that it was indistinguishable from the
“ Finners ” which they are accustomed to take im the Northern Hemisphere.

This whale used to be very common off the Falkland Islands, but it is said to
have become scarce there of late years. It is taken off the coasts of South America,
South Africa, South Georgia, and the South Shetlands. But it is evidently not fond of
ice-covered seas, as has been remarked by M. Racovitza.§

* Racovitza, E., op. cit., p. 54. See also Liouville, J., op. cit., p. 73.

+ Or Balaenoptera musculus, auctt.

{ Parker, T. J., Trans. New Zealand Inst., 1884, Vol. XVII., pp. 3-13.
§ Racovitza, E., op. cit., p. 55. See also Liouville, J., op. cit., p. 86.

CETACEA—LILLIE. N74

7. Balaenoptera borealis, Lesson. (EL avails fio. 22)

June LOS AUSTOT tom OL ONE OCA RO eee . One, travelling eastward.
August AP ie es moO: 49°85 172 13! We ; . One.
» fx ms 5 ob 2S, to DANG . One, small size.
September 3, ,, . oom Dia Sel S24 Ol Hae . Travelling eastward.
Seem = 382 46'S, Oa me a
ie Ae 2) 395 21S. (84° 580 he. wet
October oh ase ae 5 SP UE Shy WHS SP ae . Two adults, one calf.
of ie ae OCS 18201 ike . ? One.
5A Glas ; A Gf See Ollie Ao! sh ee . % Three.
i De ss 43° 41'S., 156° 56’ BE. . . ? Several.
22, 44° 95'S,,160° 3'E. . . One,

We did not see any of these whales in the Antarctic. It is possible, however, that
some of the larger specimens of the whales which we identified as Balaenoptera
acutorostrata may have belonged to this species. We did not see any Antarctic
whales which appeared to be between 37 and 55 feet in length. The specimens of
B. acutorostrata usually varied between 20 and 32 feet in length.

A few whales of about 37 feet were occasionally met with, and we regarded
them as being large individuals of 4. acutorostrata. :

On October 3, 1910, we saw two adults and a calf; their identity was
beyond any doubt. The two adults were estimated at 50 feet long and the calf was
20 feet long.

Of the specimen seen on October 22, 1910, Dr. Wilson wrote: “I have no doubt
in saying it was precisely similar in every respect to the North Sea LB. borealis, so far
as appearance in the water goes.” An individual of this species was taken off the Bay
of Islands, New Zealand, during my visit to the whaling station. This whale was a
young female, 40 feet in length. (See Pl. VIL, fig. 2.)

It will be noticed that this whale is depicted in the plate as bemg of a lighter
colour than those which are usually described as blue-black. The colour was taken
from a fresh specimen. The species of Balaenoptera, when freshly killed, are highter in
colour than is generally supposed. The epidermis only becomes dark when the carcase
is out of the water and exposed to the sun. The whalebone plates were black, with fine
greyish white bristles on their inner edges. The bristles on the posterior third of the
ranges were darker grey than in the anterior two-thirds, where some of the bristles
were quite white. The delicate, silky nature of the bristles of the baleen-plates is
characteristic of this species. The tongue was grey and pink. The palate was pink.
This specimen agreed exactly with the descriptions of Northern members of this
species.* Balaenoptera borealis is taken at the Falkland Islands, South Georgia, South
America, South Africa, and New Zealand. It is said to be scarce at the South
Shetlands, but it was seen on several occasions by the “ Belgica” near Graham

* Collett, R., Proc. Zool. Soc., 1886, p. 243-265, Pls. 25-26.

VOL. I.

118 “TERRA NOVA” EXPEDITION.

Land.* Liouville seems to have referred all the small Rorquals seen by him in the
Antarctic to this species.f

To sum up with regard to the Balaenopteridae of the Southern Hemisphere.
It would appear that all the members of the family which are known in Northern
latitudes occur all over the oceans of the Southern Hemisphere. Lalaenoptera musculus
and B. acutorostrata penetrate in large numbers to the farthest shores of the ice-covered
seas. B. borealis does not appear to be quite so common in the ice, while B. physalus
and Megaptera nodosa do not seem to go further south than the outskirts of the pack.

A new Rorqual has recently been described by Olsen} and called by him
Balaenoptera brydei. Up to the present this whale has only been taken off the
coast of South Africa. We of the ‘ Terra Nova” did not come across it at all.

ODONTOCETI.
PHYSETERIDAE.

8. Physeter catodon, Linnaeus.

Physeter catodon and P. macrocephalus, Linn.

The only occasion on which we saw this whale was on March 31, 1912, when
we were off the south of New Zealand in Lat. 44° 56’S., Long. 172° 53’ E.

A school of twelve were seen very clearly. Towards the end of February, 1911,
thirty-six bulls and one female were stranded on the beach at Perkins Island.
Tasmania. The bulls appear to have been swimming after the cow, who took them
into shallow water when the tide was receding, with disastrous results. A photograph
of this stranded school was published in the Otago Witness for March 15, 1911, and
is reproduced here. (PI. IV., fig. 1.)

ZIPHITDAE.
9. Hyperoodon rostratus, Miiller.
March 10, 1911 ‘ : : 5 BP OSE G2 Osi 13,
December 29, 1912 : 2 69RD Se IG Gs aiiaWe

We saw two specimens of this whale on March 10, 1911, swimming southward
together. They were evidently old males, about 30 feet in length, and of a hght
brown colour with white spots. Although we only saw this whale twice in the
Antarctic, there seems to be no doubt that it occurs in these regions.¢ The whalers
at the South Shetlands regard the ‘“ Bottlenose,” which they obtain in the far South,
as identical with the species which they hunt off the coast of Norway.

* Racovitza, E., op. cit., p. 38.

+ Liouville, J., op. cit., p. 110.

{ Olsen, Mrjan, Proc. Zool. Soc., 1913, pp. 1073-1090, Pls. CIX.-CXIIT.

§ Racovitza, E., op. cit., p. 58. Wilson, E. A., op. cit., p. 5, Fig. 3. Liouville, J., op. cit.,
pp. 140-151, Pl. VI.

CETACHA—LILLIE. 119

10. Mesoplodon australis, Flower.

A skeleton of this whale was found on the beach at Great Exhibition Bay.
near North Cape, New Zealand, in August, 1911. The skull and scapula were
brought home.

This species was founded by Sir William Flower* upon a skeleton from Lyall
Bay, New Zealand. The chief characters of the skull in the type specimen are as
follows :-—The rostrum is 18 inches long and narrow at its base ; basirostral groove
present ; foramina for the infraorbital branch of the second division of the fifth
nerve one behind the other; the palatines lie outside and free from the pterygoids ;
the teeth are missing in the specimen, but the sockets are near the hinder edge of
the symphysis. The species Jf haasti, Flower was founded upon a rostrum and
mandible only, which were found on the east coast of the North Island, New
Zealand.t The only points in which this species differs from the above are as follows :
The rostrum is 225 inches long and more compressed ; the basirostral groove and pit are
deeper. The height of the tooth in this species is 3°3 inches; its ereatest antero-
posterior breadth is 3°9 inches; and its greatest lateral width 3 inch.

The specimen from Great Exhibition Bay is almost identical with the type of
M. australis, except that the rostrum is 20 inches long. The mandible and teeth are of
the same shape as in J/. haasti, but the tooth is somewhat smaller. The height

5

of the tooth is 3 inches ; its greatest antero-posterior breadth is 2} inches; and its
greatest lateral width is $ inch.

It has been pointed out by Dr. H. O. Forbest that Sir W. Flower’s two
provisional species, J/. australis and JM. haasti, are possibly only variations of JL. grayi,
Haast, which is a well-founded species.

Although I am inclined to agree with this author, that the differences between
the three species are hardly specific, I have retained the species I. australis for the
“Terra Nova” specimen until more material is available, without which it is not
possible to arrive at a satisfactory conclusion with regard to the number of species
which should comprise the rare Southern forms of this genus.

DELPHINIDAE.
11. Orcinus orca, Linnaeus.
Orca gladiator, Bonnaterre.
January 3, 1911 6 : - ‘ Off Cape Crozier, Ross Island.
” 4, 5 ; : : Off Cape Evans, MeMurdo Sound.
. 290 5 é 5 : ; 78° 12'S., 174° 44’ BE.
»» 31,. 5, > : : : 78° 30'S., 170° 35’ W.

* Flower, Sir W. H., Trans. Zool. Soc., Vol. X., Pt. IX., 1878, p. 415, Pl. LXXI., Fig. 1,
Pl. LXXILI., Fig. 1.

{, Dbid:, Pll DXOXT., Hig. 3; Pl. UXXIT., Fig. 3.

t Forbes, H. O., Proc. Zool. Soc., 1893, p. 216.

wr

120 “TERRA NOVA” EXPEDITION.

February 1, 1911 3 5 F j TASS Bie) Shy UGS? WAH? VY
es Le aise : ‘ : : Off Coulman Island.

September 6, ,, 3 : 3 : 34° 25'S., 172° 10’ EB.

January 6, 1912 ‘ ; : : UES OS, WG Wey 13h

P 1 OSes : ; 3 : 78> (BY tSi, 16s)” Bays" 1D),

i 1, 55 j , ; : 76° 54’ S., 166° 39! BE.

i We 5 : , ; O° Way Sh, UG Oe

» MNS) 5 ‘ : ; ; —

a 22) Ss : ‘ ‘ : T2608, 1652 Wii Be
March 1, Gp ; ‘ : F 64° 03' S., 160° 12’ E.
January 20, 1913 : 3 ; : 77° 46'S., 166° 8 EB.

Z oie es jo) ae ae oe ue

e Die ayy : ; ‘ : (eh oll So Wek? wy" TB

This was undoubtedly the commonest Cetacean in the Ross Sea. ‘‘ Killer” whales
could be seen near the ship almost every day, so we did not attempt to keep a
record of every occurrence. The above are only a few of the dates on which this
whale was sighted. I have not much to add to the account of this whale given
by*Wilson,* except that, on this expedition, we obtained evidence that the “ Killers”
not only hunt along the edge of the fast ice in search of seals, but occasionally
break up the ice with their backs, when it is not too thick, in order to dislodge
seals which they see lying on the icefield.t

“ Killers” were more common off the coast of the Antarctic lands than in the
pack-ice, out at sea.

THE “ HiGH-FINNED” WHALE.
December 19, 1910 . : ‘ : 5 67° 54’ S., 178° 28’ W.
February 9, 1911. : : : é Off Glacier Tongue, McMurdo Sound.

This whale was first seen by Wilson, when on the “ Discovery” Expedition. We
saw a school of some twelve, or more, very clearly on February 9, 1911. They were
about 30 feet in length, and the majority of them had long, pointed dorsal fins, as
figured by Wilson.f{ In one case the concavity of the fin was pointing forwards. A few
members of the school had shorter dorsal fins, exactly like those of Orcinus orca.

The only points in which these whales appeared to differ from the ‘“ Killer” were
in the uniform black colour of the back, and in the height of the dorsal fin. The
variability in the size and shape of the dorsal fin in Orcinus orca is well known.

On January 22, 1912, we saw a school of what were undoubtedly “ Killer”
whales swimming near the edge of an ice-flow, on which some Adelie penguins were

* Wilson, E. A., op. cit., pp. 6-8, Fig. 7.

+ “Scott’s Last Expedition,” Vol. I., pp. 94-96.

{ Wilson, E. A., op. cit., p. 4, Pl. I.

§ Id., Fig. 7. Flower, Sir W. H., “ Recent Memoirs on the Cetacea,” published by the Ray Society,
1866, pp. 153-188. Collett, R., “ Norges Pattedyr,” Kristiania, 1912, 15 & 16 Hefte, p. 711,

® CETACEA—LILLIE. 121

standing. The floe was between the ship and the whales. This enabled us to estimate
the height ofthe highest dorsal fin in the school at about 5 feet. It was quite the
highest fin which we saw on a “ Killer,” and it was about the same height as the
fins in the school of “ High-finned” whales. We saw another school of “ Killers” with
variable dorsal fins off Coulman Island, on February 11,1911. The “ High-finned ”
whale seen on December 19, 1910, was among the floes of pack-ice, and Wilson
estimated the dorsal fin as being about 4 feet high. My own opinion is that the
“ High-finned” whale is probably only a variety of Oreinus orca, or possibly a new
species of that genus.

12. Globicephala melaena, Traill.

July SONS y- 3 : dS 20EN 2451312 We, travelling southward.
55 LEAL 3 : 42° 06’ S., 175° 13’ E.

August 8, ,, : : : 33° 37’ S., 171° 30' E.

October 2, ,, 3 : é 39° 40’ S., 178° 21’ E.

March 19,1913 . : : 55° 03’ S., 166° 36’ w

April 18; 5, 5 : ‘ 492 09'S.,, 55° U5!

We did not see this Cetacean to the south of Lat. 56° S.

13. ‘Delphinus delphis, Linnaeus.

June 20,1910. 3. 2. s,s 20 N., 13° 45 W.

mee CCB STEN, 149 43 W.
Gulyaesees = C Ct«‘(C‘ !SC*«éiD OPN, 99° BOW
LG hac: ee ye 1° 7"N., 21° 16’ W.

On June 20, 1910, a school of about 100 individuals of this species approached the
ship from the eastward and followed her all day. These dolphins agreed very well
with the figures of D. delphis with the exception of having rather more white on their
ventral surfaces, and their pectoral fins seemed to be entirely white.

14. Tursio peronii, Lacépede. (Text-fig. 14.)
October 20,1910 . . . . 42°51'S., 153° 56’ E.
March 30, 1912 = : : : 47° 04’ S., 171° 33’ E.

On both the above dates a pair of these whales were seen playing under the bows
of our ship. They seemed to roll over more than the other dolphins which we saw. On
October 20 the 7. peronii came with a herd of “Dusky Dolphins,” but they kept
separate. They were larger than Layenorhynchus obscurus. All the four specimens
were exactly alike and agreed with Gray's figure,* except that the tail-flukes were quite
white above and below. The demarcation between the black and white was very
pronounced (text-fig. 14).

* Gray, J. E., “Synopsis of the Whales and Dolphins,” 1868, Pl. 15, Fig. 1.

122 “TERRA NOVA” EXPEDITION.

15. Layenorhynchus obscurus, Gray.* (Pl. VIII, figs. 2-4.)

September 3,1910 . ; ; : 5 Bia ay Shy leche} 13.
October Tess ; : : 2 ALOIS 8 Sa el2oreeiebr
Ree 38° 52’ S., 146° 40’ E.

E 20, ,, 49° 51'S,, 153° 56! EB.

0 ue asa a ‘ : : 43° 41'S., 156° 56’ BE.
July Sees : : ; F : Bir ID TSS ilrteh? veh 1D
October Ue, Weil. 3 : : : 37° 49' S., 178° 39! BH.

f ee 41° 51'S., 175° 16’ E

oe Ne wea Off Kaikoura, N.Z.

é Gripes: > yen ee eee ‘i
December 15, ,, ‘ : : j : Off Banks Peninsula, N.Z.

- eh ele : 4 : ; 20 miles off Akaroa, N.Z.
February 7,1913 . : ; ; : 51° 56’ S., 168° 02’ EB.
March IB as : : : : 5 Off Lyttelton Heads, N.Z.

Schools of dolphins which we identified as belonging to this species were seen on
the above dates.

This dolphin does not seem to occur further south than about Lat. 58° 8. But
whenever we were approaching, or leaving, the coast of New Zealand we invariably met

°°?

Fia. 14.— Tursio peronii, Lacépede.

large schools of the ‘* Dusky Dolphin,” which used to follow us and play round the
bows of the ship, as though they were seeing us off or welcoming us back to temperate
lands.

These dolphins varied a good deal in colour. Three of the most common types of
pigmentation, in what appeared to be one species, are given in Plate VIL, figs. 2-4.

* True, F. W., Bull. U.S. National Mus., No. 36, ‘‘ Review of the Family Delphinidae,” 1889,
p. 104, Pl. XXIX. Racovitza, E., op. cit., pp. 59, 111-119. Wilson, E. A., op. cit., p. 8. Waite,
E. R., Records of the Canterbury Museum, N.Z., Vol. I., No. 2, 1909, Pl. VI., Fig. 2.

CETACEA—LILLIE. 123

The type shown in Fig. 2 was seen on September 3, 1910, and October 18, 1910.
The variety in Fig. 3 occurred on October 20, 1910, and on March 13, 1913.

The type in Fig. 4 was noticed on October 7 and 12, 1910, and on October 6, 1911.
The commonest form off the coast of New Zealand was type 3.

16. Lagenorhynchus wilsoni, sp. nov.

* An undescribed Dolphin,” Wilson, Nat. Antarct. Exp. 1901-1904, IT, Zool., ‘‘ Mammalia,”
Pe 9, tgs i.

December 5, 1910 . . 56° 41'S., 176° 23’ E. March 20,1912 . . 61° 49'S., 160° 04’ E.

March 8, 1911 . -. 64° 23’S., 161° 39’ E February 2,1913 . . 61°18'S,, 157° 53! E.
" One 62° 51'S.,160°55'E. | April 3, , . . 55° 11'S, 110° 21’ W.
i" lp eeSe: 2048: 161° 22°F. | ,, 7, » » . 55° 25'S., 88° 35’ W.
_ eee 56-28) Si, 169° 58" E. |

This dolphin would seem to be confined to a comparatively narrow band of the
Southern Ocean, just north of the pack-ice, between Lat. 65° S. and Lat. 54°S. It
was first described by the late Dr. E. A. Wilson when on board the “ Discovery” in
1901-4. He called it the ‘ Hourglass Dolphin.” The specimens seen by the “ Terra
Nova” were very like Wilson’s figures.*

From what we could see of this dolphin, as it swam round the ship, it appeared to
belong to the genus Lagenorhynchus, and | have named it after its discoverer.

Dr. Liouville ¢ has identified this dolphin as Lagenorhynchus jitzroyi, Waterhouse,
and it is possible that he may be correct, but until further evidence is forthcoming I
am inclined to regard it as a distinct species, on the grounds that it appears to be
slightly larger in size and has a more distinctly marked snout than L. jitzroyi. These
two points, together with the striking coloration shown by Dr. Wilson, appear to
justify the proposal to name this form in his honour. Wilson estimated the length of
this dolphin at 8 to 10 feet, which I think is rather too great. My estimate for a
number of specimens was about 6 to 7 feet. The leneth of L. jitzroyi is from 5 to 6 feet.

I am only too well aware of the unsatisfactory nature of this method of naming
species from external appearances alone, but until an expedition is organised to obtain
specimens of dolphins for a critical examination of their external and internal
characters, the nomenclature of many of the members of this family is likely to
remain uncertain.

* Wilson, E. A., op. cit., p. 9, Fig 7.
} Liouville, J., op. cit., pp, 165-177, Pls. VIII. and IX.

t

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LONDON:
PRINTED BY WILLIAM CLOWES AND SONS, LIMITED,
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. Cetacea, Plate I.

ae in i : A a the eva.

Lia a
PLATE 1.
The New Zealand Humpback whale, Megaptera nodosa, Bonnaterre (adult).

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* Cetacea, Plate II. :

Fic.

Fie.

Fic.
Fic.
Fie.
Fic.
Fic,

PLATE II.
Megaptera nodosa, Bonn.

1.—Dorso-lateral view of an individual of No. 3 type (sce text-fig. 2). Whangamumu, Bay of

Islands, New Zealand.
2.—No. 2 type text-fig. 2, showing the patch of pigmented skin, covered with barnacles, behind the

chin. Bay of Islands, N.Z.

3.—No. 3 type, text-fig. 2.

4.—No. 2 type, text-fig.
5.—No. | type, text-fig.
6.—No. 1 type, text-fig.

7,—_No, | type, text-fig. 2.

bo by be

Ventral view. Bay of Islands, N.Z.

Bay of Islands, N.Z.

Posterior ventral view, showing dapple-markings. Bay of Islands, N.Z,
Dorso-lateral view. Bay of Islands, N.Z.

Ventral view, showing barnacle-patch behind the chin,

Photographs by Mr, George Hutchinson, of Edinburgh,

grit. Antarctic (Terra Nova) Exped. rg1o.
Brit. Mus. (Nat. Hist.) Zoology, Vol. I. Cetacea, Pl. II.

Cambridge University Press

Cetacea, Plate III.

J PLATE III.

Megaptera nodosa, Bonn.

Fic. 1.—Exterior face of left periotic bone, with tympanic bulla attached. From a foetus, 13} feet
long. a, tympanic membrane. Bay of Islands, N.Z.

Fie. 2.—Interior face of the above.

Fie. 3.—Anterior view of the above. a, tympanic membrane.

Fic. 4.—Exterior face of right tympanic bulla of adult specimen. Bay of Islands, N.Z,

Fic. 5.—Interior face of the same. a, the attached malleus.

Brit. Antarctic (Terra Nova) Exped. 1910.
Brit. Mus. (Nat. Hist.) Zoology, Vol. I. Cetacea, PIL IT:

Cambridge University Press

Cetacea, Plate IV.

PLATE IV.

Fig. 1.—Physeter catodon, Linn. Thirty-six bulls and one female stranded at Perkins Island, Tasmania,
in February, 1911. Photograph by Spurling & Son, Tasmania.
Fic. 2.—Balaenoptera acutorostrata, Lacépéde. Head of calf 10 feet long. Dunedin, N.Z. Photograph
by Prof. W. B. Benham, F.R.S.
Fie. 3.—Mouth of the same. Photograph by Prof. W. B. Benham, F.R.S.
Fic. 4.—Megaptera nodosa, Bonn. Foetus, slightly enlarged. Bay of Islands, N.Z. Photograph by
Dr. W. G. Ridewood.

Brit. Antarctic (Terra Nova) Exped. 1910.
Brit. Mus. (Nat. Hist.) Zoology, Vol. I. Cetacea, Piy ly,

Fig. 1, Spurling & Son, Photographers, Tasmania Cambridge University Press

Cetacea, Plate V.

PLATE V.

Balaenoptera acutorostrata,. Lacépéde.

Fic. 1.—Calf 10 feet long. Lateral view of head. Dunedin, N.Z. Photograph by Prof. W. B. Benham, F.R.8.

Fic. 2.—Lateral view of the same specimen. x 59 y ” » »

Brit. Antarctic (Terra Nova) Exped. 1910.
Brit. Mus. (Nat. Hist.) Zoology, Vol. I. Cetacea, Pl, V2

Cambridge University Press

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Cetacea, Plate VI.

PLATE VI.
i. Balaenoptera acutorostrata, Lacépéde.

Fics. 1-6.—Various specimens “ blowing” in the pack-ice of the Ross Sea on March 3, 191
by Paymaster F. R. H. Drake, R.N.

Brit. Antarctic (Terra Nova) Exped. 1910.
Brit. Mus. (Nat. Hist.) Zoology, Vol. I. Cetacea, PIP VIE

a amen

6.

Cambridge University Press

Cetacea, Plate VII.

PLATE VII.

Fic. 1.—Balaenoptera musculus, Linn. Immature male, 65 feet in length. Bay of Islands, N.Z.
Fic. 2.—Balaenoptera borealis, Lesson. Female, 40 feet. Bay of Islands, N.Z.

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Cetacea, Plate VIII.

PLATE VIII.

Fic. 1.—Dorsal view of Balaenoptera acutorostrata. Lat. 75° 3'S., Long. 173° 41’ E., Jan
From a sketch by the late Dr. E. A. Wilson.
Figs. 2-5.—Lagenorhynchus obscurus, te Colour varieties from the Southern Ocean.

te

Brit

Brit. Mus. (Nat. Hist.)

D. G. Lillie, pinx. Cambridge University Press

BRITISH MUSEUM (NATURAL HISTORY).

BRITISH ANTARCTIC ("TERRA NOVA”) EXPEDITION, 1910.

NATURAL HISTORY REPORT.

ZOOLOGY. VOL. |, No. 4. Pp. 125-156.

LARVAL AND POST-LARVAL FISHES.

BY

C. TATE. REGAN, M.A,
(Assistant inthe Department of Zoology, British Museum (Natural, History).

WITH FIVE FIGURES IN THE TEXT AND PLATES I-X.

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LARVAL AND POST-LARVAL FISHES.

BY C. TATE REGAN, M.A.

(Assistant in the Department of Zoology, British Museum (Natural History)).

WITH FIVE TEXT-FIGURES AND TEN PLATES.

].— Antarctic and Subantarctic Fishes. PAGE

1. Notolepis and Myctophun . : : : : 125
2. A new Paraliparis—. : : : : : : ; 129
3. The Nototheniiform Fishes : : : : ‘ : 129
II.—Fishes from the Tropical and South Temperate Zones ; : 134

-III.—Notes and Conclusions.

1. Pelagic Larvae in Relation to the Distribution of Species. 148
2. The Recapitulation Theory : : é ; : : 149
3. Development of the Fins . ; ‘ ; : : 150
4, Characteristic Features of Pelagic Larvae. ; ; : 151
5. Systematic Importance of Larval Characters ‘ ‘ eis?

I—ANTARCTIC AND SUBANTARCTIC FISHES.
1. NOTOLEPIS AND MYCTOPHUM.

THE pelagic fishes of the order Iniomi are represented by a very perfect post-larval
example of the strictly Antarctic Notolepis coatsii and by a series illustrating the
development of the more widely distributed Myctophum antarcticum. This ranges,
when adult, throughout the Antarctic, Subantarctic, and South Temperate Zones,
but the larval and post-larval specimens were all taken in the Subantarctic Zone, some
to the south of New Zealand, others to the west of Tierra del Fuego.

Notolepis coatsii, Dollo. (Pl. I, figs. 4, 5).

A post-larval example, 70 mm. long, is very similar to one figured by Roule
(Deux. Expéd. Antarct. Frangaise, Poissons, pl. III). It was taken on Dec. 28th, 1912,
at Station 269, 68° 37'S., 166° 14’ W., surface. The adipose fin is continuous with

VOL. I. T

—————_
Jragsomian Instig

Fa ©

126 “TERRA NOVA” EXPEDITION.

the procurrent part of the caudal, and extends forward to the dorsal. The small pelvic
fins are a little in advance of the vertical from the origin of the dorsal, and the anus is
a short distance in front of them; from the anus a membranous fringe runs backwards

to the anal fin.

y

SESS
_ iH WPVM PH

HMMM

Fig. 1.—Notolepis coatsii. Type, actual size.

Dr. W. 8. Bruce has very kindly lent me the type in order that the accompanying
figure might be drawn, but owing to the condition of the specimen (cf. Tr. R. Soe.
Edinburgh, XLIX, 1913, p. 233) this figure is largely a restoration. The “Terra Nova”
example agrees with the type in the number of myotomes (82) and fin-rays (8 dorsal and

Fig. 2.—Distribution of Notolepis coatsi?. All recorded captures are from the area bounded by the
extreme limit of pack-ice, ~ ~ ~ ~—~—~-—. Specimens taken by the “Terra Nova,” & ;
“Pourquoi Pas?” sf ; “ Scotia,” 4 ; “ Challenger,” @.

28 anal); but in the type, which is 105 mm. long, the anus is further back, below the
anterior part of the dorsal fin, and there is a separate adipose fin. On the other hand,
in a specimen of 50 mm., the anus is further forward, only a short distance behind

LARVAL AND POST-LARVAL FISHES—REGAN. 127

the head, so that the migration of the anus backwards during the development of
this species is established.

Moreover, it is evident that Prymnothonus is not a valid genus, but merely a
larval form of Notolepis, Paralepis, ete.

The capture of this example in the Ross Sea completes the evidence that Notolepis
coatsii is circumpolar, for it had previously been taken near Peter Island, at the
South Orkneys and in the Weddell Sea, and near Wilkes Land.

Myctophum, sp.

Some specimens, very much damaged, taken on March 27th, 1912, at Station
238; 52° 11'S., 167° 25’ E., 30 metres.

The largest, 13 mm. long, is very similar to the somewhat larger example of
M. punctatum, Rafin. figured by Holt and Byrne (Fisheries Ireland Sci. Invest., 1910,
VI, pl. I, fig. 1), but differs in that the anal papilla is separated by an interspace from
the anal fin. I count 40 myotomes and 20 anal rays; the dorsal fin appears to have
been bitten off.

Myctophum antarcticum, Ginth. (PI. I, figs. 1-3).

A number of examples of this circumpolar species, 10 to 18 mm. in length, were
taken in the Subantarctic Zone at Stations—

235 52° 41'8., 168° 15’ E., 10 metres, March 26th, 1912.

238) 9) B22 WS: 167° 25) Ey 30°, — March 27th; 192:
ZA0. = 5 1-57’ S., 167° 88’ BH, 4 «, © March 28th, 1912.
250,251 54° 2'8.,177° 0’ W., surface, Dec. 20th; 1912.
Doo. HAS 38) S., 176° 55! Wi, Dec. 21st, 1912.
808 . 55°29'S., 78°54' W., 4 metres, April 9th, 1913.

In adult examples, 60 to 100 mm. in length, I count 13 to 15 dorsal and 18 to
22 anal rays, and 40 to 42 myotomes. I find the same numbers in the larger
specimens (14 to 18 mm.) of the “Terra Nova” collection, which have the fin-rays
developed, but differ from the adult fish in the following points :—

(1) There are no scales or photophores, except a single photophore on each side
above the base of the pelvic fin.

(2) The dorsal and anal fins are lower, the caudal fin is less emarginate, the
adipose fin is longer, and the paired fins are shorter.

(3) The fish is more elongate, and the snout is proportionately longer, the eye
smaller and the maxillary shorter, not reaching the vertical from the
posterior edge of the eye.

(4) There is a prominent anal papilla, and from it a membranous fringe runs
forward to the base of the pelvie fins.

(5) There is a small sinus, subdivided by septa, persistent above the occipital

it 9

28 “TERRA NOVA” EXPEDITION.

region of the head; this is an expansion of the anterior part of the dorsal
fin-fold.

(6) Pigment is wanting, except for a median black spot on the parietal region and
some pigmented areas on the tail. In specimens of 14 mm. (and less) there
is a dark area above and another below on the caudal peduncle ; in larger
ones the dorsal area spreads forward on each side of the adipose fin, and

the ventral one may disappear, or may be replaced by a median series of

Distribution of Myctophum antarcticum. Recorded captures of the adult are marked + ; larval

Fie. 3.
and post-larval stages taken by the ‘Terra Nova,” ff. The map shows the extreme limit of
pack-ice ~ ~ ~~~, and the mean annual surface isotherms of 6° C.——-—-—-— and
12° C. —.—-—-—.

dark spots, probably the precursors of the infracaudal plates of the adult
female.

Examples of 10 to 12 mm. are often more elongate, and have no photophores
and no pelvic fins ; the adipose fin extends forward to the dorsal; the dorsal fin is low,
and its rays are undeveloped or just evident. The maxillary reaches only to below the
anterior part or middle of the eye, which is relatively larger than in the specimens of

14 to 18 mm., whilst the snout is correspondingly shorter.

ee ee eS

LARVAL AND POST-LARVAL FISHES—REGAN., 129

Myctophum antareticum is very similar in form, proportions, position of the fins
and number of rays, number of myotomes, ete., to the northern JL glaciale, Reinh.,
the principal differences between them residing in the number and arrangement of the
photophores. Holt (P.Z.S. 1898, pp. 552-560, pls. XLVI, XLVI) has described young
stages of the northern species, and it is interesting to note the resemblances and
differences between these and corresponding stages of M. antarcticum. There is a
general resemblance in the development of the two species, but MM. glaciale at 8 mm.
long corresponds to M. antarcticum of 10 to 12 mm. and at 11°5 mm. to MM. antareticum
of 16 to 18 mm., whilst at 14°5 mm. JL glaciale has the general characters of the adult
fish, except that the eye, although considerably larger than in smaller examples, is not
yet proportionately as large as in the adult. Post-larval examples of J glaciale differ
from corresponding stages of MV. antarcticum in that they are not noticeably elongate
in form as compared with the adult, the abdomen is prominent, the dorsal sinus extends
from head to dorsal fin, and the latter is much higher than in Mf. antarcticum.

Even if post-larval J. glaciale appear less elongate than the same stages of
M. antarcticum to a great extent because the depth is increased by the large dorsal
sinus and the protuberant abdomen, yet if these features be neglected, the southern
species is at this early age noticeably more slender than the northern one. Jn both, the
eyes at first grow at a slower rate than the fish as a whole, as is the rule in fishes
generally, but later on grow faster than the rest of the fish, so that they are propor-
tionately much larger in the adult than in the young. Holt’s explanation—that this
is due to the fact that the adults live at greater depths than the young—may be
unreservedly accepted.

2. A NEW PARALIPARIS.

Paraliparis terrae-novae, sp. n. (Pl. I, fig. 6).
A young fish, 35 mm. long, was taken in McMurdo Sound on Jan. 16th, 1912, at
Station 332, 77° 15'8., 166° 0’ E., 0-550 metres. It is very similar in most respects to
P. antareticus, Regan, but differs notably in the fewer fin-rays (dorsal 55; anal 43)
and in the form of the pectoral fin, which has no elongate lower rays. This is the

second Antarctic species of this genus.

3. THE NOTOTHENIJFORM FISHES.

The “ Terra Nova” collection includes young stages of Pleuragramma, Pagetopsis
and Chionodraco, from the Ross Sea and McMurdo Sound. As I have already made a
systematic revision of the Nototheniiformes and have given a general account of their
distribution, it seemed worth while to try and complete this work by describing the
egos and young of the whole group, so far as possible, from the material in the British

130 “TERRA NOVA” EXPEDITION.

Museum (Natural History), including some undescribed larvae from the ‘‘ Discovery ”
collection.

NoTOTHENIIDAE.

Notothenia (PI. II, fig. 2).

A mass of eggs and a number of newly hatched larvae taken on Sept. 21st,
1910, at Roy Cove, Falkland Islands, by Mr. Rupert Vallentin, evidently belong
to a species of Notothenia. The eggs have a diameter of about 1°5 mm., and the
capsules adhere by facets to form a loose mass, just as in our northern Cottus scorpius.
The newly hatched larvae have a length of about 6 mm., and in their general structure
are very similar to Cottus larvae, for they have a short abdomen and a long tail, the
mouth is well-developed, and the median fin extends from the head round the tail to
the yolk-sac ; the anus is placed a short distance behind the yolk-sac, instead of at its
posterior edge, as in Cottus, nor can I see an oil-globule in the yolk.

Loénnberg has noted that in August JV. tessel/ata and NV. sima had the ovaries well
developed, with eggs measuring 1 mm. or less; it is quite likely therefore that the
eggs and larvae described above belong to one of these species or to another species of
the ¢essellata group.

A second mass, also presented by Mr. Vallentin, is very similar to the first, but has
the eggs somewhat larger (diameter 1°7 mm.); it was found under a stone at the
Falklands on May 10th, 1910.

Of the Antarctic species of Notothenia Lonnberg has noted that N. larseni and
N. nudifrons were nearly ripe on April 19, and that in NV. rossi and N. gibberifrons the
ovaries were very small in May, and in J. coriiceps in August.

Late larval and early post-larval stages of Notothenia have not yet been found.
Two young examples of V. macrocephala, 40 mm. long, were taken by the “ Challenger”
in the tow-net on Jan. 8th, 1874, off Kerguelen. These are very different in appearance
from the adult fish, as they are bright silvery, with the back bluish. From this
coloration and from their method of capture it may be concluded that the young of this
species swim at the surface and that its wide distribution may be connected with this.
In XV. cyaneobrancha, which is restricted to Kerguelen, young examples of 40 mm. have
the mottled coloration of the adult.

Trematomus.

In examples of 7. bernacchii and T. hansoni taken by the “Southern Cross”
and “Discovery” I find that the genital glands are much larger in April than
in October. Females taken in April have well-developed ovaries with eggs 1 to
1°5 mm. in diameter. All the specimens that were preserved of those captured in traps
during the winter are males, with testes nearly ripe ; it may be that the females cease
feeding at the approach of the breeding season. It is probable that spawning takes
place about July, and that the eggs are not much, if any, larger than in Notothenia.

LARVAL AND POST-LARVAL FISHES—REGAN. 131

Pleuragramma antarcticum, Bouleng. (P1. II, figs. 3-6).
Larval and post-larval examples were taken by the “ Discovery ” at Ross Island,
at a depth of 6 to 10 fathoms.

Date of Capture. Length.
December 14th, 1902... ... 6-7 mm.
December 25th, 1902... ... 8-10 mm.
February 8th, 1904 La soo SEI Racaiiey,
February 21st, 1902... ... 15-19 mm.
April, 1903 Ags ... 15-25 mm.
May 23rd, 1902 aa ... 24 mm.
August 13th, 1903 be Lo —2o) Tam.
September 18th, 1903 ... ... 20-25 mm.

The “Terra Nova” also secured specimens from the Ross Sea and the coast of
Victoria Land :

Station. Date. Locality. Length.
Near 186 December 31st, 1910 Ross Sea, 190 fathoms 30-35 mm.
325 August 8th, 1911 Cape Evans, 10 metres 16-18 mm.
326 January 9th, 1912 Terra Nova Bay, 10 metres 10 mm.
337 January 22nd, 1912 Cape Bird, 80 metres 10 mm.
325 April 30th, 1912 Cape Evans, 10 metres 22-25 mm.

From these data it seems probable that Pleuragramma may breed in the early
Antarctic summer, that the eggs may hatch out about the beginning of December, that
the newly hatched larvae may be 6 mm. long or a little less, that by the winter they may
reach a length of 15 to 25 mm., and when a year old may be as much as 35 mm. long.
Larval and post-larval examples have been taken not far from the coast and at depths
not exceeding 80 metres. Young fish (30 to 35 mm.) in the Ross Sea at a depth of
190 fathoms, and adults from the Ross Sea, 158 fathoms, from near the Balleny
Islands, 254 fathoms, in addition to examples found frozen on the ice barrier and
others taken from seals’ stomachs.

Larvae of 6-7 mm. are very similar to those of Notothenia from the Falklands.
Figures (PI. Il) are given of examples 6, 11°5,13°5 and 25 mm. long. It will be noted
that the permanent caudal rays are making their appearance in the larva of 11 mm.,
and that in the larger ones they have assumed their final position. The differentiation
of the dorsal and anal rays proceeds from behind forwards, and the full number is not
developed until a length of 30 mm. is reached. Pectoral fins are present from the first,
but the pelvics are rudimentary in fish of 25 to 35 mm. and are absent in smaller ones.
The pigmentation at the bases of the vertical fins and on the dorsal surface of the
abdominal cavity appears to be characteristic.

In these larval and post-larval Pleuragramma the eye is proportionately smaller
than in the adult fish, no doubt because the latter descend to greater depths.

132 “TERRA NOVA” EXPEDITION.

Artedidraco (Pl. II, fig. 1).

Examples of A. loennbergii, Roule, and A. skottsbergii, Linnberg, taken by the
“Terra Nova” in McMurdo Sound, at a depth of 207 fathoms, on January 23rd, 1912,
include nearly ripe females with eggs 23 to 3 mm. in diameter. Lonnberg has recorded
nearly ripe females of A. mirus, with eggs of 25 mm., from South Georgia in May.

On January 28th, 1904, the ‘* Discovery ” obtained two larvae which I identify as
A. skottsbergii, at Hut Point, Ross Island, at a depth of 3 fathoms. These are 13 mm.
long, and judging by the development of the vertical fins and the size of the yolk-sac
they have not been hatched very long; one of them is figured (PL. I, fig. 1). The
short tail and large yolk-sac make this larva quite unlike that of Plewragramma in
appearance.

GYMNODRACONIDAE.

Gymnodraco acuticeps, Bouleng. (Pl. II, fig. 4).

A post-larval fish 24 mm. long was taken by the “ Discovery” on Jan. 28th,
1904, at Ross Island, at a depth of 3 fathoms. The caudal and pectoral fins are fully
developed, the pelvies are rudimentary, and the dorsal and anal rays are all present
but do not reach the edge of the fin. A large yolk-sac is still evident. The snout 1s
short and blunt as compared with that of the adult fish, and the oval nostril is
relatively large; the opercular spine is not yet developed.

CHAENICHTHYIDAE.

The type of Chaenichthys rugosus, Regan, from Kerguelen, taken between October
and February, is a ripe female with eggs 3 to 4 mm. in diameter. Lonnberg has
examined a ripe female of Champsocephalus guinari, taken in May at South Georgia ;
this had eggs 4 mm. in diameter.

Pagetopsis macropterus, Bouleng. (Pl. Ill, figs. 1-3).

Two larvae from the “Terra Nova” collection, taken at Cape Evans, McMurdo
Sound, may be referred to this species. The smaller, 14 mm. long, was captured on
May 13th; the larger, 15 mm. long, on June 28th, 1911, at a depth of 20 metres.
Two somewhat larger specimens, 19 and 20 mm. long, were obtained by the
“Discovery ” at Ross Island.

The wide mouth, cleft to below or beyond the posterior edge of the eye, at once
distinguishes these larvae from those of the Nototheniidae and Bathydraconidae ;
another distinctive feature is the precocious development of the pelvic fins. The
relatively short tail, the small number of myotomes, and the black colour of the pelvic
fin membrane are characteristic of this species.

The series illustrates well the growth of the produced snout of the adult fish from
the snub-nosed form of the larva.

LARVAL AND POST-LARVAL FISHES—REGAN. 133

Chionedraco kathleenae, Regan (P1. IV, figs. 2, 3).

A larva of 21 mm. was taken by the “Terra Nova” off Cape Evans at a depth
of 10 metres on October 19th, 1911. This is not quite so advanced as the young
Pagetopsis of 19-20 mm. From these it is readily distinguished by the longer tail
and the more numerous myotomes (about 60), whilst the pelvic fins are scarcely
pigmented except between the spine and the first soft ray. A fish of 32 mm. from the
“ Discovery ” collection is much more advanced, but the dorsal and anal rays are still
undeveloped.

Cryodraco (Pl. IV, fig. 1).

Three specimens, 16, 17 and 21 mm. in total length, from Ross Island
(“ Discovery ” collection) may belong to an undescribed species of this genus. The
myotomes number 55 or 56, fewer than in C. atkinsoni, in which I count 62. The
body is crossed by two dark bands, the anterior on myotomes 27 to 29, the posterior
on myotomes 40 to 42; there are indications of a third band at the base of the caudal
fin. The slender form, produced snout and elongate pelvic fins further distinguish
these fishes from larval or post-larval Pagetopsis and Chionodraco.

The data given above with regard to the breeding and development of the
Nototheniformes may be summarized as follows :—

Of the Bovichthyidae nothing is known, and examination of the material in the
Natural History Museum gives no results.

The breeding season varies considerably ; approximate dates are May and
September for species of Notothenia, July for Trematomus hansoni and T. bernacchii,
November for Pleuragramma antarcticum, January and May for species of Artedidraco,
May for Champsocephalus gunnari, etc., ete.

The eggs vary in diameter from 1 to 4 mm. when taken from ripe females
preserved in spirit ; the diameter would no doubt be greater if eggs freshly taken from
the sea were examined. The Nototheniinae seem to have the smallest eggs,
Artedidraco and the Chaenichthyidae the largest. In all probability the eggs are
demersal throughout the group, and in some species of Notothenia they are known to
adhere together in masses.

In some of the Notothenimae the newly hatched larvae are about 6 mm. long,
but in Artedidraco and the Chaenichthyidae they are probably twice as long or
even more.

The newly hatched larvae have a distinct mouth, the membranous median fin

_ extends forward above to the head and below to the yolk-sac; the anus is situated at
or a little behind the posterior end of the yolk-sac, and at the edge of the fin-
membrane ; pectoral fins are present, but the pelvics do not appear until much later,
except in the Chaenichthyidae. During growth the caudal is formed and assumes its

VOL. I. 15)

134 “TERRA NOVA” EXPEDITION.

terminal position before the dorsal and anal rays develop ; of these the posterior rays
appear first, at any rate in Pleuragramma.

Of the forms identified the larvae of the Nototheniinae differ from the rest in the
small size of the yolk-sac. Of those with a large yolk-sac Artedidraco is distinguished
from Gymnodraco by the short tail, whilst the Chaenichthyidae differ from both in the
wide mouth and the early development of the pelvic fins.*

Except Pleuragramma artarcticum larval and post-larval Notothenioids have only
been taken quite near the coast. There is some evidence that the young of Notothenia

macrocephala, a widely distributed species, may be pelagic.

II—FISHES FROM THE TROPICAL AND SOUTH
TEMPERATE ZONES.

Larval and post-larval fishes were taken, by means of plankton nets and the
young fish trawl, at or near the surface in the following areas :—
(1) North of New Zealand and round the Three Kings Islands. Stations 85-142.
July to September, 1911.
(2) Melbourne Harbour. Station 161. October, 1910.
(3) Temperate South Atlantic, about 200 miles from the coast of Uruguay.
Station 311. April, 1913.
(4) Off Rio de Janeiro. Stations 39-40. — April, 1913.
(5) Western Tropical Atlantic, south of the Equator. Stations 43-57.
May, 1913.
(6) Atlantic, south of the Canaries. Stations 16-17. June, 1910.
(7) Atlantic, south of the Azores and west of the Canaries. Stations 68-69.
May, 1913.

The nature of the collections made in these areas is shown by the following

2
a
2
3)

summary --—

New Zeanuanp AND TureEE Kines Isnanps.
Sardina neopilchardus Limnichthys fasciatus
Prymnothonus, sp. Cubiceps caeruleus
Myctophum coccot Centrolophus maoricus
Diaphus, sp. Thyrsites atun
Lampanyctus macropterus Lepidopus caudatus
L. longipinnis, sp. n. Tripterygium varium
Anguilla australis Monacanthus scaber
Scombresox forsteri Diodon, sp.
Scorpis violaceus Haplophryne mollis

* Onos, Brosmius, Molva, etc., resemble the Chaenichthyidae in the early development of the pelvic
fins, in this respect differing from Gadus (cf. Ehrenbaum, Nordisches Plankton, Hier und Larven yon
Fischen, 1905-1909).

ia

LARVAL AND POST-LARVAL FISHES—REGAN. 135

Mertpourne Harpour.

Oda balteatus Platycephalus, sp.
Pentaroge marmorata

TEMPERATE SoutH ATLANTIC.

Stylophthalmus paradoaus M. laternatum
Cyclothone microdon Ceratias, sp.
Myctophum benoiti

Rio DE JANEIRO.

Sardinella pseudohispanica Ancylopsetta quadrocellata
Cyclothone microdon Ancylopsetta, sp.
Glyphidodon, sp. Symphurus plagusia

Gobiosoma molestum

TropicaL SoutH ATLANTIC.

Stylophthalmus macrenteron, sp. 0. L. acuticeps, sp. n.
Cyclothone microdon L. hexastigma, sp. n.
Vinciquerria lucetia Hemirhamphus unifasciatus
Synodus synodus Cryptotomus ustus
Prymnothonus, spp. Scorpaena, sp.
Lampanyctus maderensis Bothus ocellatus

Leptocephalus muraenae unicoloris

ATLANTIC, S. OF THE CANARIES.

Lampadena chavesi

ATLANTIC, S. OF THE AZORES.

Vineiquerria lucetia Scombresox saurus
Paralepis speciosus

The majority of these were captured well out at sea, and are young stages of
oceanic fishes, or oceanic larvae (Leptocephalus, Bothus ocellatus) of coast fishes.
Exceptions to this are the larvae and young fishes taken in Melbourne Harbour and
others captured a few miles off Rio de Janeiro (Sardinella, Glyphidodon, Gobiosoma,
Ancylopsetta, Symphurus) ; also some of the New Zealand species were taken near the
coast and may never be oceanic.

Of some importance in its bearings on the geographical distribution of coast-fishes
is the capture of young Hemirhamphus unifasciatus and Cryptotomus ustus, Brazilian
species, far out in the Atlantic.

Of greater interest is the case of Limnichthys fasciatus, known previously as a
little fish of the rock-pools of New South Wales and Lord Howe Island ; now its range
is extended to New Zealand, and its occurrence in localities so wide apart is explained
by the capture of the young fish at or near the surface between the Three Kings Islands
and New Zealand.

136 “TERRA NOVA” EXPEDITION.

ISOSPONDYLI.
CLUPEIDAE.

Sardinella pseudohispanica, Poey. (PI. V, fig. 2).

Typical Clupeoid larvae, 7 to 8 mm. long,

with the anus far back, anal fin
undeveloped, dorsal fin posterior, and caudal rayed and terminal. There are stellate
chromatophores on the head, below the heart, and above and below the gut; they are
most distinct on the dorsal border of the hinder two-thirds of the gut. Vertebrae 46.

The number of vertebrae and the resemblance to the larval Pilchard (Sardina
pilchardus) lead me to identify these larvae as Sardinella pseudohispanica, which is the
Western Atlantic representative of the Mediterranean Allache (Sardinella aurita, Cuv.
and Val.); the genera Sardina and Sardinella are very closely related.

Stations 39, 40. Six miles off the mouth of Rio de Janeiro Harbour. 2 metres.
April 27th, 1913.

Sardina neopilchardus, Steind. (Pl. V, figs. 3, 4).

Three larval and post-larval fishes may be referred to this species. I count
16 dorsal and 16 anal rays and 52 or 53 myotomes. The smallest example, 12 mm.
long, is more advanced than the 11°5 mm. larva of Sardina pilchardus figured by
Cunningham, as the caudal fin is fully formed and terminal in position, and the anal
rays are appearing. The largest, 18 mm. long, appears to differ from Cunningham’s
24mm. S. pilchardus chiefly in details of pigmentation, but after examination of
Pilchard larvae from Plymouth, kindly lent by Dr. E. J. Allen, I am doubtful whether
there are any constant differences between the larvae of the European species and its
representative in the seas of Australia and New Zealand.

Station 135. Spirits Bay, near North Cape, New Zealand. 3 metres. Sept. Ist, 1911.

STOMIATIDAE.
Stylophthalmus macrenteron, sp. n. (Pl. V, fig. 1).

A post-larval fish, 33 mm. long, shows several resemblances to Stylophthalmus
paradoxus, Brauer, and may therefore be described as a Stylophthalmus, although it is
unlikely that it is congeneric with any of the species associated under that name. In
all probability it belongs to the family Stomiatidae ; the only other family that seems
possible is the Alepocephalidae. If this be a Stomiatid it may represent a young stage
of Eustomias obscurus, Vaillant, described from a single specimen taken near the
Azores.

Form elongate; head one-fifth of the length of the fish. Snout produced and
depressed ; lower jaw prominent ; maxillary toothed, not nearly reaching eye; inter-
orbital region broad and flat. Myotomes about 70. Dorsal 22, placed posteriorly.
Anal 40, extending forward in advance of dorsal. Protruding terminal portion of

—

LARVAL AND POST-LARVAL FISHES—REGAN. 137

intestine very long and supported behind by a cartilaginous rod. A series of seven
dark spots along the back.
Tropical Atlantic. Station 49. 18° 51’S., 33° 40’ W. Surface. May 6th, 1913.

Stylophthalmus paradoxus, Brauer.
A larva, 6 mm. long, of the type figured on pl. V, fig. 5, of the Valdivia report.
South Atlantic. Station 311. 35°29'S., 50° 26’ W. 2 metres. April 22nd, 1913.

GONOSTOMATIDAR.

Cyclothone microdon, Giinth. (Pl. V, fig. 5).

Several examples 6 to 8 mm. long have the fins fully developed, but the head is
intermediate between that of the Gonostomatid larva figured by Holt and Byrne
(Fisheries Ireland Sci. Invest. 1912, I, pl. II, fig. 5), and that of the adult fish, the
maxillary not extending back beyond the eye. I count 12 dorsal and 16 anal rays
and 30 myotomes.

Tropical Atlantic. Station 39. 6 miles off mouth of Rio de Janeiro Harbour.
2 metres. April 27th, 1913.

South Atlantic. Station 311. 35° 29'8.,50° 26’ W. 2 metres. April 22nd, 1913.

Vineiquerria lucetia, Garm. (PI. V, figs. 6, 7).

Seven examples, 8 to 10 mm. in total length, differ from the adult fish in that
they are more slender and the photophores are not developed; also the mouth is
smaller. A specimen of 15 mm. has the lower series of photophores developed, but
the upper series is incomplete. All are from the Atlantic, taken at the surface.

Station 45. 21°S., 37°50’ W. May 4th, 1913. Station 50. 18°S., 31° 45’ W.
May 7th, 1913. Station 68. 27° 22’ N., 33° 40' W. May 28th, 1913.

INIOMI.
SYNODONTIDAE.

Synodus synodus, Linn. (Pl. VU, fig. 4).

A larva of 14 mm. differs from those described by Max Weber (Siboga Fishes,
p. 82, fig. 28) in having 13 patches of dark pigment on each side of the gut instead of
‘11. Probably this is a specific difference distinguishing the Atlantie S. synodus from
the Indo-Pacifie S. varius. I count 60 myotomes and 9 anal rays.

Station 46. 20° 30'S., 36° 80’ W. Surface. May 4th, 1913.

SUDIDAE.
Prymnothonus, Richards.
The only larvae known to belong to this family are of the type described as

* Prymnothonus ” (Giinther, ‘‘ Challenger” Pelagic Fishes, p. 39, pl. V, 1889). These are

138 “TERRA NOVA” EXPEDITION.

larvae of Paralepis and related genera with produced snout. It has been shown above
(p. 126) for the Antarctic Notolepis coatsi that there is an extended backward migration
of the anus during the transition from the Prymnothonus stage to the adult fish.

A larva of 12 mm. has the snout moderately produced. There are 80 myotomes
and about 20 anal rays. The anus corresponds to the twenty-eighth myotome and the
origin of the anal fin to the fifty-fourth (PI. VII, fig. 1).

Tropical Atlantic. Station 50. 18° 8., 31° 45' W. Surface. May 7th, 1913.

A second larva of 16 mm. has the snout more produced than the preceding.
There are 116 vertebrae (52 + 64) and 30 or more anal rays (PI. VII, fig. 2).

Tropical Atlantic. Station 47. 20° 30’ &, 36° 30’ W. Surface. May 4th,
1915.

In the number of anal rays these examples agree well enough with known species
of Paralepis, and in the number of myotomes the first agrees with the Mediterranean
species that I have examined. Possibly P. borealis, a species that I have not seen,
may have the larger number of myotomes found in the second specimen.

A third ‘* Prymnothonus” is probably generically distinct from these ; it is a post-
larval fish, 22 mm. long, evidently related to Paralepis, which it resembles in the
structure of the head, but it has only 60 myotomes and 11 anal rays. The adipose
fin is above the posterior end of the anal. Dorsal and pelvic fins are undeveloped
(PL VIL, fig. 3).

Station 85. 24 miles W.N.W. from Cape Maria van Diemen, New Zealand.
2 metres. July 24th, 1911.

Paralepis speciosus, Bellotti.

Omosudis elongatus, Brauer, Valdivia Tiefsee Fische, p. 140, fig. 68 (1906).

This species is represented in the British Museum collection by two examples of
65 and 75 mm. from Messina. Bellotti’s specimens were 75 and 90 mm., Brauer’s from
8 to 30 mm., the larger full grown. There is therefore reason to suppose that this
is a small species, and that it assumes the adult form at an early age.

A larva of 8 mm. that I refer, with some doubt, to this species has the fin-rays
not yet developed, but the form of the head is already as in the adult fish. The
patches of pigment on each side of the gut number only five, instead of eight, but this
may be a larval character.

Station 69. 29° 10’ N., 33° 36’ W. Surface. May 29th, 1913.

This species was described from the Mediterranean (Bellotti, Atti. Soe. Ital. XX,
1877, fase. 1, p. 2, fig.), and has been recorded by Brauer from the Gulf of Guinea
and the Indian Ocean.

MycToPHIDAE.

Larval and post-larval stages of Myctophum and related genera were taken to the
north of New Zealand and in the Atlantic. The species of this group are so numerous

LARVAL AND POST-LARVAL FISHES—REGAN. 139

and have such a wide range that the definite assignment of larvae to their species 1s
very dithcult. This difficulty is increased by the fact that the head, owing to the
relatively smaller size of the eye and mouth and the greater length of the snout, has a
physiognomy quite unlike that of the adult fish. However, by counting the myotomes
and fin-rays and taking into consideration the position of the fins, the size of the
mouth, etc., it is possible to make determinations which may, in some cases, approximate
to the truth.

The general character of the development has already been described in dealing
with Myctophum antarcticum.

Myctophum benoiti, Cocco (PI. VI, figs. 1, 2).

Several examples, from 4 to 7 mm. in total length, may belong to this species.
They have much in common with the larvae described and figured by Holt and Byrne
(Fisheries Ireland Sci. Invest. 1910, VI, p. 29, pl. I, fig. 8), from the Irish Atlantic
slope under the name “ Scopelid larva, R 2,” but seem to be specifically distinct. They
are distinguished by their rather deep form, conical snout, strong teeth, and by
the distribution of the stellate pigment spots ; four large ones are present on each side,
respectively at the origin of the dorsal and anal fins, below the adipose fin and above
the end of the anal fin; usually there is a fifth on the side between the two last-named.
There is also a spot on the back behind the head and a mid-ventral series of small spots
from the end of the lower jaw to the origin of the anal fin. I count 12 or 13 dorsal
and 17 or 18 anal rays and 37 myotomes.

South Atlantic. Station 311. 35° 29'S., 50° 26’ W. 2 metres. April 22nd, 1913.

Myctophum laternatum, Garm. (Pl. VI, fig. 7).

A specimen 8 mm. long is probably of this species. It is moderately elongate ; the
snout is short and the mouth is small, the maxillary extending to below the middle of
the eye. The anal fin has 14 rays and commences below the posterior part of the
dorsal, which has 11. The myotomes number 35.

South Atlantic. Station 311. 35° 29'S., 50° 26’ W. 2 metres. April 22nd, 1913.

Myctophum coccor, Cocco.

An example 20 mm. long has the characters of the adult fish, except that the eye
is relatively smaller, its diameter being less than the length of the snout.

Station 86. Off Three Kings Islands. 3 metres. July 25th, 1911. .

Diaphus sp. (Pl. V1, figs. 3, 4).

Form rather deep; snout obtuse and mouth oblique. Length 4 to 5 mm. Dorsal
and anal fins each with about 14 rays; about 35 myotomes. The distribution of the
pigment is shown in the figures.

Station 135. Spirits Bay, near North Cape, New Zealand. 3 metres. September
Misty LOI:

140 “TERRA NOVA” EXPEDITION.

Lampadena chavesi, Collett (Pl. VI, fig. 8).

Four specimens, 10 to 12 mm. long, may belong to this species. I count 12-13
dorsal, 13-14 anal, and 15 pectoral rays and 38 myotomes. The fins are placed as in
the adult fish. The maxillary extends to below the middle of the eye. On each side
there is a photophore in front of the eye, another at the base of the pectoral fin, and a
third above the base of the pelvic fin. Two examples have 2 mid-dorsal stellate
blackish spots behind the adipose fin and a mid-ventral series of 3 or 4 linear spots
behind the anal fin.

Station 17. South of the Canaries. 26° 17’ N., 20° 54’ W. 10 metres. June 30th,
1910.

Lampanyctus maderensis, Lowe (Pl. VI, fig. 6).

An example of 9 mm. shows the supraorbital ridge ending in an antrorse spine ;
the cleft of the mouth extends to below the posterior edge of the eye. The myotomes
number 36 and the fins are as in the adult fishes figured by Goode and Bean, and
Brauer. There is a bar of pigment at the base of the caudal fin. In specimens of
5 mm. the spine is not developed, the mouth is smaller, and the adipose fin is longer.

Station 50. Tropical Atlantic. 18°8., 31°45’ N. Surface. May 7th, 1913.

Lampanyctus macropterus, Brauer (Pl. V1, fig. 5).

A specimen of 10 mm. may belong to this species; it is very similar to the
example of ZL. crocodilus of the same size figured by Holt and Byrne (Fisheries Iveland
Sci. Invest., 1910, VI, pl. I, fig. 3). There are 12 dorsal and 20 anal rays and 40
myotomes. There is a black spot at the base of the caudal fin, another above the anal
papilla, and some smaller ones on the lower part of the head.

Station 135. Spirits Bay, near North Cape, New Zealand. 3 metres. September
Ist, 1911.

Lampanyctus longipinnis, sp. n. (Pl. VI, fig. 9).

Dorsal 22. Anal 25. Myotomes 38. Evidently related to L. procerus, Brauer,
differing in the more numerous dorsal and anal rays.

A young fish of 15 mm.

Station 113 (N.E. of Three Kings Islands). 33° 12’ S., 171° 05’ HE. 3 metres.
August 9th, 1911.

APODES.

Following the plan adopted by workers on this group, I use the name Leptocephalus
as a generic term for larval Hels, and give new specific names to those that cannot be
identified.

Leptocephalus acuticeps, sp. n. (Pl. VII, fig. 5).
Very similar to L. oxycephalus, Pappenh. (Deutsche Siidpolar Exped. XV, Zool. VI,

LARVAL AND POST-LARVAL FISHES—REGAN. 141

p- 190, pl. IX, figs. 3, 5), from the Indian Ocean, but with fewer myotomes, 207
(174 + 33) instead of 220-230 (180-190 + 40). Form elongate ; head small; snout
rather produced and acute. Teeth in 1 + 6 pairs in the upper jaw and 1 + 5 in the
lower. Gut with a swelling (liver) at about the thirtieth myomere. A series of dots
along the axis and along the dorsal border of the gut; a group of similar dots at the
end of the tail. Dorsal and anal fins rayless ; a caudal fin.

Total length 47 mm.

Tropical Atlantic. Station 45. 21° 8., 37° 50’ W. Surface. May 4th, 1913.

ANGUILLIDAR.
Anguilla australis, Richards.

An elver of 50 mm.

Station 129. Off Three Kings Islands. Surface. August 26th, 1911.

oOo
cu)

MURAENIDAE.

Leptocephalus muraenae unicoloris (Pl. VI, fig. 7).

A specimen of 60 mm. has the general characters of L. euryurus and L. similis of
Lea (Michael Sars Exped. IT, 1, No. 7), and of the larva of Jfuwaena helena (Schmidt,
Medd. Komm. Havunders. Fiskeri, IV, 2, 1913, p. 5, pl. I, fig. 3). The myotomes
number 138 (64 + 74 ), and as this is nearly the number of vertebrae (136) counted in
Muraena unicolor, De la Roche, this Leptocephalid may be referred to that species,
which is represented in the British Museum collection by specimens from Algiers,
Madeira, the Azores and St. Helena, but does not as yet appear to have been recorded
from Brazil.

The snout is blunt and the lower edge of the mandible is somewhat convex ; there
are 7 or 8 teeth on each side in both jaws. The vertical fins are rayed and the caudal
is much reduced. The pigment is in the form of small dots; some on the anterior
dorsal fold and one at the base of each dorsal and anal ray; a line of dots along
the dorsal surface of the gut, and anteriorly some below it.

Tropical Atlantic. Station 46. 20° 30'8., 36° 30’ W. Surface. May 4th, 1913.

OPHICHTHYIDAE.

Leptocephalus hexastigma, sp. un. (Pl. VII, fig. 6).

Myotomes 164 (72 + 92). Head small; snout acute and produced. Upper jaw
with 10 pairs of teeth, the three. following the anterior pair strong and spaced, the
rest closer together and decreasing. Lower jaw with 9 pairs of teeth, the anterior pair
followed on each side by 5 strong and spaced teeth and 3 small posterior teeth.
Dorsal rays commence in advance of anus, anal rays a little behind it. Gut with 6

VOL. I. : x

142 “TERRA NOVA” EXPEDITION.

pigmented swellings, the third largest, the last 3 small; these correspond respectively
to myotomes 15, 22-23, 29-30, 47, 57 and 67-68. Total length 60 mm.
Tropical Atlantic. Station 50. 18°8., 31° 45' W. Surface. May 7th, 1913.
The resemblances to the Ophichthys larvae described and figured by Schmidt
(Medd. Komm. Havunders, Fiskeri, IV, 2, Let) make it probable that this is a
member of the family Ophichthyidae.

SYNENTOGNATHI.

: ScOMBRESOCIDAE.

Scombresox saurus, Walb.

Post-larval fishes, 10 and 18 mm. in total length, are strongly pigmented. The

snout is short and the lower jaw is prominent, but not produced. The fins are fully
developed ; the smaller specimen has the fold in front of the anal fin still persistent.

Station 69. South of the Azores. 29°10’ N., 33° 36’ W. Surface. May 29th, 1913.

Scombresox forstert, Cuv. and Val.
Post-larval examples of this species are similar to those of its Atlantic

representative.
Station 89. Off Three Kings Islands. Surface. July 25th, 1911.

HeEMIRHAMPHIDAE.

Femirhamphus unifasciatus, Ranzani.
Dorsal 13-14. Anal 14-15. Myotomes about 50. Lower jaw not produced.

Total length 10 mm.
Station 53. Tropical Atlantic. 5°8., 27°15 W. 2 metres. May 12th, 1913.

PERCOMORPHI.
ScoRPIDIDAE.
Scorpis violaceus, Haast.

A young fish of 38 mm.
Station 129. Off Three Kings Islands. Surface. August 26th, 1911.

POMACENTRIDAE.
Glyphidodon, sp. (PIE V UN ie. 5):
Dorsal x1r—xmi, 10-11. Anal m,10-11. Vertebrae 26 (11 + 15). The specimen
measures 5 mm. in total length.
Station 39. Six miles off the mouth of Rio de demets Harker: 2 metres.
April 27th, 1913

EARVAL AND POST-LARVAL FISHES—REGAN. 145

SCARIDAE.

Cryptotomus ustus, Cuy. and Val. (Pl. VIII, fig. 6).

Dorsal 1x, 10. Anal 12. Vertebrae 25 (9-10 + 15-16). These little fishes,
9mm. in total length, evidently belong to the family Scaridae or to the sub-family
Julidinae ; the structure of the vertebral column and of the vertical fins leaves no
doubt as to this. The number of fin-rays is somewhat less than in the Julidinae,
except Doratonotus megalepis, which differs from them in its strong dorsal and anal
spines, long praemaxillary pedicels, etc. After comparison with numerous species of
Scaridae I am of opinion that these young fishes belong to Cryptotomus ustus, or to
some species nearly related to it.

Station 49. Tropical Atlantic. 18° 51'S., 33° 40’ W. Surface. May 6th, 1913.

ODACIDAE.
Odax balteatus, Cuv. and” Val. (Pl. VII, fig. 4).

In identifying a post-larval fish of 6 mm. with this species I rely on the number
of vertebrae, 36 (19 + 17) and of anal and caudal rays; the anterior dorsal rays are
undeveloped, and the pelvic fins have not yet appeared.

Station 161. Melbourne Harbour. 12 metres. October, 1910.

LIMNICHTHYIDAE.
Limnichthys fasciatus, Waite.

Several young fishes, 16 to 20mm. long; some of the larger ones show the
characteristic markings of the adult fish.

Fic. 4,—Limnichthys fasciatus (x 6).

This species was originally described from adult specimens in full roe, 43 mm.
long. These were found in rock-pools at Lord Howe Island (Waite, Rec. Austral.
Mus. V, 1904, p. 178, pl. XXIII, fig. 4), and afterwards similar examples were taken
from pools near Sydney (Waite, t.c. p. 243). As Lord Howe Island is 600 miles from
the coast of New South Wales, one might have inferred that the species was oceanic at
some period of its life. This is proved, and its known range extended by its capture at
or near the surface to the north of New Zealand, off the Three Kings Islands, and also
midway between these two localities.

Station 120. 34° 26’8., 172° 14’ BK. Surface. Aug. 18th, 1911.

- 130. Off Three Kings Islands. Surface. Aug. 27th, 1911.

, 133. - Spirits Bay, nr. North Cape, N.Z. 20 metres. Aug. 30th, 1911.

_ itera He Bs 3 metres. Sept. Ist, 1911.
xX 2

144 “TERRA NOVA” EXPEDITION.

STROMATEIDAE.

Cubiceps caeruleus, Regan.

A young fish 20 mm. long.

Station 125. Between North Cape and Doubtless Bay, New Zealand. Surface.
Aueust 23rd, 1911.

Centrolophus maoricus, Ogilby (Pl. X, fig. 7).

A young fish 19 mm. long has the body deeper and the cross-bars stronger than
the larger examples already described (‘‘ Terra Nova” Fishes, p. 19).

Station 142. 34° 45'S8., 170° 45’ E. 2 metres. September 8th, 1911.

, N ew
Q -

ae es
‘ ZEALAND

ee

Fic. 5.—Distribution of Limnichthys fasciatus.
&), Littoral, from rock-pools; +, planktonic.

GEMPYLIDAB.

Thyrsites atun, Kuphras. (Pl. VIII, figs. 1-8).

Specimens 5 to 10 mm. long are very similar to those of 7. prometheus described
and figured by Giinther (“‘ Challenger” Pelagic Fishes, p. 7, pl. I, figs. C, D). There
are about 35 vertebrae and 18 to 20 dorsal spines.

Station 133. Spirits Bay, near North Cape, New Zealand. 20 metres. Augus
30th, 1911.

TRICHIURIDAE.

Lepidopus caudatus, Kuphyas. (Pl. VIII, fig. 7).

A specimen of 11 mm. is distinguished from the preceding by the more elongate

a

LARVAL AND POST-LARVAL FISHES—REGAN. 145

form and the numerous vertebrae (about 100). The pelvic fins are represented by a
pair of serrated spines.
Station 135. Spirits Bay, near North Cape, New Zealand. 3 metres. September
Ist, 1911.
GOBIIDAE.
Gobiosoma molestum, Girard.
A post-larval fish of 5 mm. has the form of the adult. The number of fin-rays—
dorsal vir, 13; anal 11—point to this species.
Station 39. Six miles off the mouth of Rio de Janeiro Harbour. 2 metres.
April 27th, 1913.
CLINIDAE.
Tripterygium varium, Forst. (Pl. IX, figs. 1, 2).
Numerous larval and post-larval examples, 6 to 14 mm. in total length. One of
6 mm. is figured (Pl. IX, fig. 1); only the caudal fin has traces of the permanent rays.
At 9 or 10 mm. the caudal is fully formed and terminal and the anal and third dorsal
have their full complement of rays, but there is no trace of the first and second dorsals
nor of the pelvies in these, nor in the larger examples of 13 or 14 mm. (PI. IX, fig. 2).
The absence of these fins made it difficult to determine these fishes, but after preparing
the skeleton of one of the larger specimens I was led to place them in the Clinidae, the
form of the skull and the arrangement of the elements at the base of the pectoral fin
being especially characteristic. The number of vertebrae, about 45, and of fin-rays,
12-13 dorsal, 25-27 anal, 14 principal caudal and not less than 14 or 15 pectoral, lead
to the determination of the species as Tripterygium varium.
Spirits Bay, near North Cape, New Zealand.
Station 133. 20 metres. August 30th, 1911.
“ 135. 38 metres. September Ist, 1911.
136. Surface. September 2nd, 1911.

SCLEROPAREL
SCORPAENIDAE.

Scorpaena, sp. (Pl. X, fig. 6).

Fin-rays.—Dorsal x1, 1 9; Anal m1 5; Pectoral 18. Vertebrae 24 (10 + 14).

The specimen is 10 mm. long ; the head is armed with serrated spines, two prae-
orbital, four praeopercular, one at the angle longest ; the interorbital ridges are
continued back to a pair of strong occipital spines ; the infraorbital ridges are serrated.
The lower part of the pectoral fin is pigmented.

Station 53. Tropical Atlantic. 5°S., 27° 15' W. Surface. May 12th, 1913.
Pentaroge marmorata, Cuv. and Val. (Pl. X, fig. 5).

Larvae 6 to 7 mm. long may be referred to this species, with which they agree in

146 “TERRA NOVA” EXPEDITION.

the number of vertebrae (28), pectoral (10) and caudal (14) rays. At this stage the
pectoral rays are simple and elongate, the pelvic fins are rudimentary, and the dorsal
and anal are rayless. The head is armed with paired occipital, otic and supraorbital
spines ; the praeoperculum has two strong spines alternating with two smaller ones.
Station 161. Melbourne Harbour. 12 metres. Young fish trawl. October, 1910.

PLATYCEPHALIDAE.

Platycephalus sp. (Pl. X, fig. 4).

Numerous specimens, 5 to 7 mm. long, evidently pertain to this genus. I count
27 vertebrae and 18 pectoral rays. In all the dorsal and anal are rayless; in the
larger ones the end of the notochord is upturned and the hypurals and caudal rays are
evident ; at 7 mm. rudimentary pelvic fins are present. The snout is depressed and
rounded and the mouth is wide. There is a double spine on each side of the occiput,
and there are two strong praeopercular spines and a smaller one below.

Station 161. Melbourne Harbour. 12 metres. Young fish trawl. October, 1910.

HETEROSOMATA.
BorHipae.
Paralichthinae.

Ancylopsetta quadrocellata, Gill (P1. IX, fig. 3).

A specimen of 5 mm. has the general characters of the post-larval fish described
and figured by Kyle as Ancylopsetta sp. (Danish Oceanographical Exped. Flat-fishes, —
p- 143.) The head is similarly armed, the anterior dorsal rays are produced, ete.
Notable differences from Kyle’s specimen are that the body is deeper and the fin-rays
are fewer. I count 73 dorsal rays, the anterior 13 somewhat spaced and at least the
first 8 of these produced; the anal rays number 60, and the vertebrae 34 or 35
(9-10 + 25).

A second example of less than 3 mm. shows the characteristic pignientation and
the prominent abdomen, but the fins are rayless and the end of the notochord is not
turned upwards. - :

Station 39. Six miles off the mouth of Rio de Janeiro Harbour. 2 metres.
April 27th, 1913.

Aneylopsetta sp. (Pl. IX, fig. 4).

Two larvae, each 4 mm. long, may be provisionally referred to this genus. They
agree with the preceding in the armature of the head and the number of vertebrae, but
are not so deep, and lack the spots on the body. The fins are membranous except an
anterior dorsal of 7 prolonged rays.

Station 39. Six miles off the mouth of Rio de Janeiro Harbour. 2 metres.
April 27th, 1913.

EARVAL AND POST-LARVAL FISHES—REGAN. 147

Bothinae (Platophrinae).
Bothus ocellatus, Agass.

A specimen of 26 mm. is still externally symmetrical and is extremely similar to
the example of B. podas figured by Kyle (t.c. p. 98, fig. 16). I count 86 dorsal and
59 anal rays and 38 (10 + 28) vertebrae. .

Station 53. Tropical Atlantic. 5°S., 27°15’ W. 2 metres. May 12th, 1913.

CYNOGLOSSIDAE.

Symphurus plagusia, Bl. Schn. (Pl. IX, figs. 5, 6).

Dorsal 90-95 ; Anal 70-75 ; Caudal 10 ; Vertebrae 49 (9 + 40).

Two examples of 6 to 7 mm. are rather similar to S. /actea of this size described
and figured by Kyle (t.c. p. 132, pl. 1v, figs. 41-42), but the abdominal appendix is
lacking, and the distribution of the pigment is somewhat different. A specimen of
11 mm. has already metamorphosed, showing that the change takes place at a smaller
size than in the European species.
Stations 39, 40. Six miles off the mouth of Rio de Janeiro Harbour. 2 metres.
April 27th, 1913.

PLECTOGNATHI.

BaALisTIDAE.

Monacanthus scaber, Forst. (Pl. X, fig. 3).

A post-larval specimen 5 mm. long. The head is large; the external bones are
spinate. The posterior dorsal and anal rays are not fully formed, but the total number
exceeds 30. The considerable length of the tail posterior to the permanent caudal fin
is worth notice.

Station 133. Spirits Bay, near North Cape, New Zealand. 20 metres. August
30th, 1911.

DIODONTIDAE.

Diodon sp.

A young fish, 5 mm. long.

Station 92. 24 milesS. by W. from Three Kings Islands. Surface. July 27th, 1911.

PEDICULATI.

CERATIIDAE.

Ceratias sp. (P]. X, fig. 1).

Two examples, 6 mm. in length, are in many respects similar to the adult fish.
There are 4 dorsal, 4 anal, and 9 caudal rays; an interorbital papilla represents the
- illicium. The mouth is nearly vertical and there are pointed teeth in the jaws.

Station 311. South Atlantic. 35° 29’S.,50°26’W. 2 metres. April 22nd, 1913.

148 “TERRA NOVA” EXPEDITION.

ACERATIIDAE.

Haplophryne mollis, Brauer (Pl. X, fig. 2).

A specimen of 10 mm. is very similar in outline to the one figured by Brauer
(Valdivia Tiefsee Fische, pl. XVI, fig. 10); but this outline is that of the loose skin in
which the fish is enclosed, and that of the body of the fish is quite different. The dorsal
and anal rays are long, but only their tips project, and a rudiment of the illicium
is present, but is hidden under the skin.

Station 127. Off Three Kings Islands. Surface. August 25th, 1911.

IIl—NOTES AND CONCLUSIONS.
1. PELAGIC LARVAE IN RELATION TO THE DISTRIBUTION OF SPECIES.

The larval, post-larval, and young fishes of the ‘“‘ Terra Nova” collection were all
taken at or near the surface. The majority of those captured far from land, whether
in the Antaretic (Notolepis coatsii), Subantarctic (Myctophum antarcticum), South
Temperate or Tropical Zones (cf. p. 134) belong to oceanic species, either pelagic or
bathypelagic. On the other hand, most of those taken near the coast are young
stages of coast fishes which, when adult, may either swim near the surface (e.g,
Clupeidae) or live at the bottom (e.g., Heterosomata). This is in agreement with the
fact that the same species of coast fishes rarely inhabits areas separated by a wide
expanse of sea; for example, the majority of the Brazilian species are not found on
the West African coast. It may be inferred that the distribution of a benthic species
along a coast may be helped by a pelagic larval phase, but that unless this be
prolonged it will not serve to establish the species in places separated from its
original habitat by a wide sea.

Bothus ocellatus (p. 147), one of the Flat-fishes, is a good example of a benthic
fish with a prolonged pelagic larval phase. The ‘“ Terra Nova” example, more than
an inch long, is still transparent and symmetrical; it was taken in the Atlantic
in 5° §., 27° 15’ W., quite 300 miles from the American coast. Jordan and Evermann
give the habitat of this species as from “Long Island to Rio Janeiro, on sandy
shores” ; but a specimen in the British Museum collection proves that it also occurs
at Ascension, about 1,000 miles from Brazil. There can be little doubt that it has
reached this island owing to the long duration of its life as a pelagic larva.

The Eels, Apodes (p. 140, Pl. VII, figs. 5-7) are benthic fishes with pelagic larvae
that attain a considerable size and age and may migrate for long distances ; but the
majority of the Atlantic species of this group are not common to the eastern and
western coasts.

There is considerable evidence that some coast fishes with a wide geographical

ee

LARVAL AND POST-LARVAL FISHES—REGAN. 149

range do not have a particularly long larval life, but that the young fishes may be
oceanic.

Of the species dealt with in this report three may be selected to illustrate this.

(1) Notothenia macrocephala (p. 130), a species of the shallow water benthos ;
this is known from the Magellan and New Zealand districts and from Kerguelen ;
off the last-named the ‘“‘ Challenger” obtained young fishes, 40 mm. long, swimming
at or near the surface.

(2) Hemirhamphus wnifasciatus (p. 142) ranges from Florida to Rio de Janeiro,
and*belongs to a genus of herbivorous fishes that swim at the surface near the coast.
The capture of a young fish, 10 mm. long, far out in the Atlantic, is of interest,
as //, picarti, Cuv. and Val., from Algeria is believed to be the same species.

(3) Limnichthys fasciatus (p. 148). Previously recorded from rock-pools near
Sydney and at Lord Howe Island; the pelagic young of this little fish were taken
by the “Terra Nova” to the north of New Zealand, near the*Three Kings Islands,
and midway between these localities.

Further examples are Scorpaena (p. 145) and Cryptotomus ustus (p. 143).

It may be regarded as nearly certain that when a species of coast fishes occurs
in areas separated from each other by wide expanses of the ocean, it has a pelagic
phase in its life history of sufficient duration to enable it to travel or to be carried from
one such area to another. In some fishes (Bothus, Apodes) this pelagic stage ends
when the larva acquires the structure and habits of the adult fish, but in other cases
it seems that the young fishes, essentially similar to the adults in structure, may
differ from them in habits and swim across the ocean.

2. THE RECAPITULATION THEORY.

It is well known that the truth of the theory that ontogeny repeats phylogeny
is shown by almost every Teleostean fish in the development of its caudal fin, which
is at first ventral and then becomes terminal.

Two rather puzzling developmental features, the migration forwards of the
dorsal fin in the Clupeidae and the migration backwards of the anus in Notolepis,
Paralepis, ete., may possibly be explained by the same theory.

In the Clupeidae, and the closely-related Elopidae and Albulidae, which are the
most generalised living Teleosts, the dorsal fin is completely formed on the posterior
part of the back and then travels forward to its permanent position. The earliest fishes
that can be regarded as ancestral to the Clupeoids are the palaeozoic Palaeoniscidae :
in this family the dorsal fin was placed posteriorly, usually above the anal. Next
come the Semionotidae, which are a big step nearer the Teleosteans, and then the
Pholidophoridae, which may be regarded as the immediate ancestors of the Clupeoids ;
in both these mesozoie families, and especially in the Semionotidae, the dorsal fin is
placed well backward and in a number of the genera is at least partly opposed to the

VOL, I. Y

150 “TERRA NOVA” EXPEDITION.

anal. It seems possible, therefore, that the backward position of the dorsal fin of
larval Clupeoids may be an ancestral feature.

In Notolepis (p. 125, Pl. I, figs. 4, 5), Paralepis (cf. Prymnothonus, p. 137, Pl. VII,
figs. 1-3), etc., the larva has a short gut with the anus not far behind the head ; as
the fish grows the gut becomes relatively longer, and the anus travels backward until
it reaches its final position a short distance in front of the anal fin. In a paper on
the classification of the Iniomi (Ann. Mag. N. H. (8) VII. 1911, pp. 120-133) I have
pointed out that the long-bodied and long-snouted Paralepis is closely related to,
but more specialized than, Chlorophthalmus, a short-snouted fish of normal piscine
form, but with the anus nearer to the head than to the anal fin. It is not impossible
that the anterior position of the anus in the larval Paralepis may be due to the
evolution of this genus from a Chlorophthalmus.

3. DEVELOPMENT OF THE FINS.

The pectoral fins, which are principally concerned with balancing, are -usually
present in the youngest larvae, and the permanent fin-rays appear at a very early
stage. As a rule the caudal, used for propulsion, is the next fin to develop. The
hypurals and fin-rays make their appearance below the notochord, and then by flexion
of the latter are brought into a terminal position. Afterwards the dorsal and anal -
rays develop in the embryonic fin-fold and the pelvic fins grow out. There are many
deviations from and exceptions to the general course of development of the fins
outlined above.

The dorsal and anal fins usually originate in their final position, but in the
Clupeidae, and the related Albulidae and Elopidae, they develop more posteriorly and
the dorsal especially migrates forwards for a considerable distance after it has been
formed (ef. Pl. V, figs. 2-4). In these families the dorsal fin develops earlier than the
anal, but in the Iniomi the reverse is the case (e.g., Synodus synodus, p. 137, Pl. VII,
fig. 4; Prymnothonus, p. 137, Pl. VII, figs. 1-3). Sometimes the formation of the
dorsal fin, or the anterior part of it, is so delayed that the embryonic fin-fold disappears
first ; this may often happen with the spinous dorsal fin of acanthopterous fishes (e.g.,
Oda balteatus, p. 143, Pl. VIII, fig. 4; Tripterygium varium, p. 145, Pl. IX, figs. 1, 2).

Notable exceptions to the general rule that the anterior or spinous portion of the
dorsal fin develops later than the posterior or soft-rayed portion are the Trichiuroids
and some Paralichthmae. In the former (e.g., Thyrsites atun, p. 144, Pl. VIII, figs. 1-3)
the early formation of the spinous dorsal may be for purposes of defence, and one may
well believe that it would be effective in warding off the attacks of the predaceous
young of such fishes as Pomatomus saltator (cf. Agassiz and Whitman, Mem. Mus.
Comp. Zool. XIV, No. 1, 1885, p. 16).

In some Flat-fishes of the sub-family Paralichthinae (e.g., Ancylopsetta, p. 146,
Pl. IX, figs. 3, 4) the anterior dorsal rays are formed at a very early stage and grow

LARVAL AND POST-LARVAL FISHES—REGAN. 151

out into filaments which may be used as feelers, perhaps to find food at night.
When the pelvic fins develop precociously they also may take the form of spines
(e.g., Thyrsites) or of long filaments (e.g., Trachypterus) ; but in other cases they grow
out into large fins with the rays fully connected by membrane (e.g., Pagetopsis
macropterus, p. 132, Pl. III, figs. 1-3); presumably such fins would be used to help the
pectorals in balancing and to prevent the fish from sinking.

It seems that in most fishes the fins develop in the same order and that the
development of one or more fins out of their turn, precociously, may be for purposes
either of flotation, balance, defence, or perhaps nocturnal feeding.

4. CHARACTERISTIC FEATURES OF PELAGIC LARVAE.

Absence of accessory organs of respiration and of adhesive organs.—Larval
structures that occur in more than one group of fresh-water fishes are external gills
and adhesive organs; the latter enable them to remain in the place selected by the
parents until the yolk is absorbed, and external gills are advantageous when the water
is limited in quantity or deficient in oxygen; that these structures are not found in
pelagic marine larvae is not surprising.

Tnvisibility.—All pelagic larvae are transparent, and in some groups the larval
stage is prolonged until a considerable size is reached; these large larvae remain
transparent owing to the strong compression of their bodies and the looseness of their
tissues, and the change into the less compressed and more compact young fish is
accompanied by a shrinkage. This type of development is characteristic of the Apodes
(p. 140, Pl. VII, figs. 5-7), but it occurs also in the Elopidae and Albulidae.

Buoyancy and balance—It has been suggested that the dorsal sinus of the
Myctophidae (pp. 127, 138, Pl. VI) may serve as a float, and it seems likely that the
large pelvic fins of the larval Chaenichthyidae, especially Pagetopsis (p. 132, Pl. III,
figs. 1-3), may be spread to prevent the fish sinking,

Generally the vertical fin acts as a keel, and balance is maintained by movements
of the pectorals; possibly the protrusion of the terminal part of the gut in many
larvae may be connected with balance; in those described as Stylophthalmus the
protruded portion may be quite long, but it is difficult to understand why this feature
should be so exaggerated as it is in Stylophthalmus macrenteron (p. 136, Pl. V, fig. 1).
The terminal part of the gut, with its basal support, forms a long appendage; this
would, presumably, keep the fish steady, and make it difficult for it to turn over on its
back, but would retard its progress if it attempted to swim; in fact, one may suppose
that the chief effect of flexions of the tail would be to rotate the fish, the anal
appendage serving as a fulcrum.

Sense organs.—Certain fin-rays may be formed precociously and grow out into
long filaments; these may be used as feelers, perhaps to find food at night. In all
larvae the auditory, optic and olfactory organs appear to be well developed, but it is

y 2

152 “TERRA NOVA” EXPEDITION.

difficult to assign any reason for the stalked eyes of some early larvae (Stomiatidae,
especially Stylophthalmus), unless it be conceded that they may enable the larva to
see in all directions and help it to maintain a balance, and, as it is so small and not yet
an active swimmer, are not likely to be injured or to impede its progress.

Defence.—Many pelagic larvae have the bones of the head, and especially the
praeoperculum, armed with strong spines, no doubt defensive. This is well exemplified
by Thyrsites (p. 144, Pl. VIII, figs. 1-3), Platycephalus (p. 146, Pl. X, fig. 4) and
Ancylopsetta (p. 146, Pl. IX, figs. 3, 4). The precocious development, probably for
defensive purposes, of the spinous dorsal and pelvic fins in Thyrsites has already been
referred to.

5. SYSTEMATIC IMPORTANCE OF LARVAL CHARACTERS.

Except in a few groups, of which the Apodes are the most notable example, there
are no features that characterize pelagic larval fishes as belonging to one order or
another, and it is not easy to determine their systematic position unless they are
sufficiently advanced towards the structure of the adult fish. In all cases the number
of myotomes and of fin-rays, if these be developed, are of the greatest help ; with these
as a guide one may, by a patient process of trials and eliminations, determine
specifically some most puzzling examples, as, for instance, the post-larval Oda
balteatus, described and figured above (p. 143, Pl. VIII, fig. 4). Although it does not
appear that the diagnostic characters of the different orders are likely to be strongly
reinforced by larval features, yet the study of a series of larval and young fishes, such
as those collected by the “Terra Nova,” confirms and in no degree modifies ideas as to
the relationship of the Teleostean orders and families derived from the study of the
morphology of the adult fish.

The pigmented patches on the gut of the Sudidae and Synodontidae (Pl. VII,
fies. 1, 2, 4), the armature of the head in the Platycephalidae (PI. X, fig. 4) and
Triglidae, are examples of similar larval characters in related families. The
development of the anal fin before the dorsal in the Iniomi, the migration of the dorsal
in the Clupeidae, Albulidae, etc., may be cited as examples of developmental features
common to a series of related families.

LARVAL AND POST-LARVAL FISHES—REGAN.

Aceratiidae, 148.
acuticeps, Gymnodraco, 132.

*; Leptocephalus, 135, 140.
Albulidae, 149, 150, 152.
Ancylopsetta, 135, 146, 150, 152.

"3 quadrocellata, 135, 146.
Anguilla australis, 154, 141.
Anguillidae, 141.
antarcticum, Myctophum, 125, 127, 139, 148.

i Pleuragramma, 131, 133, 134.
antarcticus, Paraliparis, 129.
Apodes, 140, 148, 149, 151.
Artedidraco, 132, 133, 134.

Pr loennbergii, 132.

rf skottsbergii, 132.
atkinsoni, Cryodraco, 133.
atun, Thyrsites, 134, 144, 150, 151.
australis, Anguilla, 134, 141.
Balistidae, 147.
balteatus, Odax, 135, 143, 150, 152.
benoiti, Myctophum, 135, 139.
bernacchii, Trematomus, 130, 133.
Bothidae, 146.

Bothinae, 147.
Bothus, 149.

» ocellatus, 135; 147, 148.

» podas, 147.
Bovichthyidae, 133.
caeruleus, Cubiceps, 134, 144.
caudatus, Lepidopus, 134, 144.
Centrolophus maoricus, 134, 144.
Ceratias, 135, 147.

Ceratiidae, 147.

Chaenichthyidae, 132, 133, 134.
Chaenichthys rugosus, 132.
Champsocephalus gunnari, 132, 133.
chavesi, Lampadena, 135, 139.
Chionodraco kathleenae, 133.
Chlorophthalmus, 150.

Clinidae, 145.

Clupeidae, 136, 148, 149, 150, 152.
coatsii, Notolepis, 125, 138, 148.
coccoi, Myctophum, 134, 139.
coriiceps, Notothenia, 130.

Cottus scorpius, 130.

INDEX.

crocodilus, Lampanyctus, 140.
Cryodraco, 133.

7 atkinsoni, 133.
Cryptotomus ustus, 135, 143, 149.
Cubiceps caeruleus, 134, 144.
cyaneobrancha, Notothenia, 130.
Cyclothone microdon, 135, 137.
Cynoglossidae, 147.

Diaphus, 134, 139.
Diodon, 134, 147.
Diodontidae, 147.
Doratonotus megalepis, 143.
elongatus, Omosudis, 138.
Elopidae, 149, 150.
euryurus, Leptocephalus, 141.
Eustomias obscurus, 136.
fasciatus, Limnichthys, 134, 143, 149.
forsteri, Scombresox, 134, 142.
Gempylidae, 144.
gibberifrons, Notothenia, 130.
glaciale, Myctophum, 129.
Glyphidodon, 135, 142.
Gobiidae, 145.
Gonostomatidae, 137.
gumnari, Champsocephalus, 132, 133.
Gymnodraco, 134,

a acuticeps, 132.
Gymnodraconidae, 132.
hansoni, Trematomus, 130, 133.
Haplophryne mollis, 134, 148.
helena, Muraena, 141.
Hemirhamphidae, 142.
Hemiramphus unifasciatus, 135, 142, 149.
Heterosomata, 146, 148.
hexastigma, Leptocephalus, 135, 141.
Iniomi, 125, 137, 150, 152.
Isospondyli, 136.
kathleenae, Chionodraco, 133.
Lampadena chavesi, 135, 139.
Lampanyctus crocodilus, 140.

longipinnis, 134, 140.

Be macropterus, 134, 140.
55 maderensis, 135, 140.
5 procerus, 140.

larseni, Notothenia, 130.

| elie onl

154 “TERRA NOVA” EXPEDITION.

laternatum, Myctophum, 135, 139.
Lepidopus caudatus, 134, 144.
Leptocephalus acuticeps, 135, 140.
5 euryurus, 141.
- hexastigma, 135, 141.
Pe muraenae unicoloris, 135, 141.
“3 oxycephalus, 140.
similis, 141.
crmeieeer ae 143.
Limnichthys fasciatus, 134, 143, 149.
loennbergii, Artedidraco, 132.
longipinnis, Lampanyctus, 134, 140.
lucetia, Vinciguerria, 135, 137.

macrenteron, Stylophthalmus, 135, 136, 137, 151.

macrocephala, Notothenia, 130, 134, 149.
macropterus, Lampanyctus, 134, 140.

53 Pagetopsis, 132, 151.
maderensis, Lampanyctus, 135, 140.
maoricus, Centrolophus, 134, 144.
marmorata, Pentaroge, 135, 145.
megalepis, Doratonotus, 143.
microdon, Cyclothone, 135, 137.
mollis, Haplophryne, 134, 148.
Monacanthus seaber, 134, 147.
Muraena helena, 141.

» unicolor, 141.

muraenae unicoloris, Leptocephalus, 135, 141.
Muraenidae, 141.
Myctophidae, 138.
Myctophum, 127.

BS antarcticum, 125, 127, 139, 148.

oy benoiti, 135, 139.

S coccoi, 134, 139.

- glaciale, 129.

+ laternatum, 135, 139.

punctatum, 127.
acca 125, 150.
33 coatsii, 125, 138, 148.
Nototheniidae, 130.
Notothenia, 130, 133.
a corliceps, 130.
FA eyaneobrancha, 130.
Pe gibberifrons, 130.
— larseni, 130.
- macrocephala, 130, 134, 149.
= nudifrons, 130.
m rossi, 130.
+ sima, 130.
tessellata, 130.
ase Notothenia, 130.
obscurus, Eustomias, 136.
ocellatus, Bothus, 135, 147, 148.
Odacidae, 143.
Odax balteatus, 135, 143, 150, 152.

|

Omosudis elongatus, 138.
Ophichthyidae, 141.

Ophichthys, 142.

oxycephalus, Leptocephalus, 140.
Pagetopsis macropterus, 132, 151.
Palaeoniscidae, 149.

paradoxus, Stylophthalmus, 135, 136, 137.

Paralepis, 127, 138, 150.

Bs speciosus, 135, 138.
Paralichthinae, 146.
Paraliparis antarcticus, 129.

oi terrae-noyae, 129.
Pediculati, 147. ;
Pentaroge marmorata, 135, 145.
Percomorphi, 142.
Pholidophoridae, 149.
Platophrinae, 147.
Platycephalidae, 146, 152.
Platycephalus, 135, 146, 152.
Plectognathi, 147.

Pleuragramma antarcticum, 131, 133, 134.

podas, Bothus, 135, 147.
Pomacentridae, 142.
Pomatomus saltator, 150.
procerus, Lampanyctus, 140.

Prymnothonus, 127, 134, 135, 137, 138, 150.

punctatum, Myctophum, 127.
quadrocellata, Ancylopsetta, 135, 146.
rossi, Notothenia, 130.
rugosus, Chaenichthys, 132.
saltator, Pomatomus, 150.
Sardina neopilchardus, 134, 136.
» pilchardus, 136.
Sardinella aurita, 136.

+ pseudohispanica, 135, 136.
saurus, Scombresox, 135, 142.
scaber, Monacanthus, 134, 147.
Scaridae, 143.

Scleroparei, 145.
Scombresocidae, 142.
Scombresox forsteri, 134, 142.

5 saurus, 135, 142.
Scorpaena, 135, 145.
Scorpaenidae, 145.

Scorpididae, 142.

Scorpis violaceus, 134, 142.
scorpius, Cottus, 130.
Semionotidae, 149.

sima, Notothenia, 130.

similis, Leptocephalus, 14]. __
skottsbergii, Artedidraco, 132.
speciosus, Paralepis, 135, 138.
Stomiatidae, 136.
Stromateidae, 144.

‘Sudidae, 137,152.
Synentognathi, 142.
_ Synodontidae, 137, 152.
_ Synodus synodus, 135, 137, 150.

4, varius, 137.
synodus, Synodus, 135, 137, 150.
_terrae-novae, Paraliparis, 129.
tessellata, Notothenia, 130.
Thyrsites atun, 134, 144, 150, 151, 152.
_ Trachypterus, 151.

‘Stylophthalmus macrenteron, 135, 136, 151.
paradoxus, 135, 136, 137.

i : LARVAL AND POST-LARVAL FISHES—REGAN.

Trematomus bernacchii, 130, 133.

A hansoni, 130, 133.
Trichiuridae, 144.
Triglidae, 152.
Tripterygium varium, 134, 145, 150.
unicolor, Muraena, 141.
unifasciatus, Hemirhamphus, 135, 142, 149.
ustus, Cryptotomus, 135, 142, 149.
varius, Synodus, 137.
varium, Tripterygium, 134, 145, 150.
Vinciguerria lucetia, 135, 137.
violaceus, Scorpis, 134, 142.

: PRINTED BY WILLIAM CLOWES AND SONS, LIMITED, DUKE STREET, STAMFORD STREET, S.E., AND GREAT WINDMILL STREET, W.

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PLATE I.

Length of Specimen. Locality.
Fic. 1.—Myetophum antarcticum, Giinth. (p. 127). F 11 mm. S. of New Zealand, Station 252.
RiGH2: # a3 : : : 15 mm. * pits
IMes By af * ; 5 : 18 mm. BS Fa
Fic. 4.—Notolepis coatsii, Dollo (p. 125). ; : 50 mm. Weddell Sea. ;
Fig. 5. FF 3 ; 5 : 70 mm. Antarctic, Station 269.
Fic. 6.—Paraliparis terrae-novae, sp. n. (p. 129) . 5 35 mm. MeMurdo Sound, Station 332. —

Brit.Antarctic (Terra Nova) Exped.1910.
Brit.Mus.(Nat.Hist) © Zoology, Vol. 1. Larval Fishes,PLI.

G.M.Wocdwand del, st lith. Huth imp.

, M
a | Iniomi and Paraliparis.

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Larval Fishes, Plate II.

Fic. 1.—Artedidraco skottsbergii, Loennb. (p. 132)
Fic. 2.-—Notothenia, sp. (p. 130). : F 3
Fig. 3.—Pleuragramma antarcticum, Bouleng. (p. 131)
’ Fie. 4. bas aA :
Fie. 5. 5 3
Fie. 6.

PLATE II.

Length of Specimen. Locality.
3 3 mm. Ross Island.
6 mm. Falklands.

Ross Island.

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Larval Fishes,PLIl.

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PLATE III.

Fic. 1.—Pagetopsis macropterus, Bouleng. (p. 132)
Fie. 2. 55 Ss ;

Hicso: = a

Fia. 4.—Gymnodraco acuticeps, Bouleng. (p. 132)

Length of Specimen.

14 mm.
15 mm.
19 mm.
24 mm.

Locality.

McMurdo Sound. —

»”
Ross Island.

Brit. Mus.(Nat.Hist,))

Brit.Antarctic (Terra Nova) Exped.1910.
Zoology, VoL.l.

Larve. Fishes,P1. II.

G.M.Woodward del, et lith

Pagetopsis and Gymnodraco.

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Length of Specimen.
21 mm.
21 mm.
32 mm.

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’ Larval Fishes, Plate V-
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PLATE V. x
Length of Specimen. Locality. :
Fie. 1.—Stylophthalmus macrenteron, sp. n. (p. 136) 3 : 33 mm. Atlantic, Station 49.
Fig. 2.—Sardinella pseudohispanica, Poey (p. 136) : : 7 mm. Rio de Janeiro, Station 39.
Fic. 3.—Sardina neopilchardus, Steind. (p. 136). ; : 12 mm. New Zealand, Station 135.
Fia. 4. 55 a8 : : 5 18 mm. a
Fic. 5.—Cyelothone microdon, Giinth. (p. 137) : ; : 8 mm. Atlantic, Station 39.
Fic. 6.—Vinciguerria lucetia, Garm. (p. 137) . : : 9mm. Atlantic, Station 45.

Fie, 7. ne ~ ; F ; 15 mm. . be

ee ee at

H Brit. Antarctic (Terra Nova) Exped.1910.
-it.Mus. (Nat.Hist.) Zoology, Vol I. Larval Fishes,PLV,

G.MWoodward del.etlith.

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Larval Fishes, Plate Wile

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PLATE VI.

Length of Specimen. Locality.
Fic. 1.—Myctophum benoiti, Cocco (p. 139) . : , : 4 mm. Atlantic, Station 311.
Fie. 2. . 50 3 : : 5 : 7 mm. s 5
Fic. 3.—Diaphus sp. (p. 139). : : : ; : 4 mm. New Zealand, Station 135.
Fie. 4. ) : : : : é 5 mm PA rege
Fie. 5.—Lampanyctus inacropterus, Brauer (p. 140) : 2 10 mm. % 09
Fig. 6.—Lampanyctus maderensis, Lowe. (p. 140) . : ; 9 mm. Atlantic, Station 50.
Fic. 7.—Myctophum laternatwn, Garm. (p. 139). é 5 8 mm. 5 Station 311.
Fic. 8.—Lampadena chavesi, Collett (p. 140) : ; . 11 mm. » Station 17.

Fie. 9.—Lampanyctus longipinnis, sp. n. (p. 140) . ‘ : 15 mm. New Zealand, Station 113.

Brit.Antarctic (Terra Nova) Exped.1910.
Brit.Mus.(Nat.Hist) © Zoology, Vol.1. Larval Fishes,P1.VI.

G.M.Woodward del. et lith. Huth imp,

Myctophidae.

:

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7. : Larval Fishes, Plate VII.

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PLATE VII.

Length of Specimen. Locality.
Fig. 1.—Prymnothonus sp. (p. 138) : ‘ ’ ; : 12 mn. Atlantic, Station 50.
Eig, 2: ) sp. (p. 138). F : : : 16 mm. >» Station 47.
Fic. 3. 3 sp. (p- 138) 5 : : : : 22 mm. New Zealand, Station 85.
Fie. 4.—Synodus synodus, Linn. (p. 137) 3 : : 5 14mm. Atlantic, Station 46.
Fic. 5.—Leptocephalus acuticeps, sp. n.(p.140) —. 5 : 47 mm. 3 Station 45.
Fic. 6. 3 hexastigma, sp. n. (p. 141) . : : 60 mm. < Station 50.
Fie. 7. muraenae unicoloris (p. 141) : : 60 mm. a Station 46.

Brit. Antarctic (Terra Nova) Exped.1910.
Ue, oology, VoL.I.

.

Larval Fishes,P1 VII. .

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Length of Specimen. Locality.

PLATE VIII.
1.—Thyrsites atun, Kuphras. (p. 144) . >. : F 5 mm.
2. 5 : ‘ : : 6 mm.
3. Pe 5 : : : . 10 mm.
. 4.—Odax balteatus, Cuy. and Val. (p. 143). : : 6 mm.
- 5.—Glyphidodon, sp. (p. 142). : ; 5 mm.
. 6.—Cryptotomus ustus, Cuy. and Val. é 143) : =) oem:

7.—Lepidopus caudatus, Euphras. (p. 144). : : 11 mm.

New Zealand, Station 133.

33 ”

Melbourne, Station 161.
Rio de Janeiro, Station 39.
Atlantic, Station 49.

New Zealand, Station 135.

Brit.Antarctic (Terra Nova) Exped.1910.
Brit.Mus.(Nat.Hist) Z oology,VoLL. Larval Fishes, P1.VII,

G.M Woodward del. et lith. Huthimp.

Trichiuroidea and Percoidea.

Larval Fishes, Plate IX.

PLATE IX.
Length of Specimen. Locality.

Fie. 1.—Tripterygium varium, Forst. (p. 145) : : : 6 mm. New Zealand, Station 135.
Richens - ; A 13 mm. . as

Fic. 3.—Ancylopsetta quadrocellata, Gill. (p. 146) . 6 : 5 mm. Rio de Janeiro, Station 39.
Fig. 4. eS Jos GIG) 5 3 « : F) 4mm. A Pe

Fic. 5.—Symphurus plagusia, Bl. Schn. (p. 147). 7 mm. an 4

Fie. 6. 55 Ft P ae aes 11 mm. 4 a

Brit.Antarctic (Terra Nova) Exped.1910.
Brit.Mus.(Nat.Hist,) Zoology,Vol.l. Larval Fishes,P1. IX.

———— EZ,

j GLE :

ase

(\, ;
G.M.Woodward del. etlith. Huth imp.

Tripterysium and Heterosomata.

Larval Fishes, Plate X.

‘Fia.
Fic.
Fic.
Fie. 4
TG
Fig.
Fie.

PLATE X.

1.—Ceratias sp. (p. 147).
2.—Haplophryne mollis, Brauer & 148)
3.—Monacanthus scaber, Forst. (p. 147)
4.—Platycephalus sp. (p. 146).
.—Pentaroge marmorata, Cuv. and Val. (p. Say
6.—Scorpaena sp. (p. 145) .
7.—Centrolophus maoricus, Ogilb. ite 144)

Length of Specimen. Locality.
6mm. Atlantic, Station 311.
10mm. New Zealand, Station 127.
5 mm. Station 133.
7 mm. Meliounne, Station 161..
7 mm.
10 mm. Atlantic, Station 53.
New Zealand, Station 142.

19 mm.

Brit.Antarctic (Terra Nova) Exped.1910.
Brit.Mus. (Nat.Hist) Z oology,Vol.l. Larval Fishes,P1.X.

G.M.Woodward del.etlith. Huth imp.
| Pediculati, Scleroparei, Monacanthus and Centrolophus.

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