Natural History Reports / British Antarctic (“Terra Nova”) Expedition, 1910-13. Zoology

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Natural History Reports / British Antarctic (“Terra Nova”) Expedition, 1910-13. Zoology

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BRITISH MUSEUM (NATURAL HISTORY). 


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


NATURAL HISTORY REPORTS. 


ZOOLOGY. VOL. I: 


VER Piibiveebe: | 


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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; 
Dunzav & Co., Lrp., 34-36 Margaret Street, Cavendish Square, W.1; Oxrorp University Press, London, E.C. 4 ; 
Wuetpon & Westy, Lrp., 2, 3, & 4 Arthur Street, New Oxford Street, W.C. 2; 
Oxrver & Born, Tweeddale Court, Edinburgh ; 
AND AT THE 
Bririsn Musgum (Natura History), Cromwell Road, London, S.W. 7. 


[All rights reserved.| 


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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.  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 


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Trematomus hansoni, 2, 3, 27. 
3 


ap lepidorhinus, 3, 5, 6, 27. 
# loennbergii, 2, 3, 27. 
 
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Psammobatis cirrifer. 


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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» NATURAL HISTORY OF THE 
: ADELIE PENGUIN 


BY 
os Staf-Surgeon G. MURRAY LEVICK, R.N. 
ae (Member of the Expedition). 
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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. 

>  ~ 
- » > 
 ' 


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. 
age eat) Slave a a i SOR Bele 7 


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Do Avan (NarurAt History), Cromyell Road, London, 8,W, 
oe 1915. 

[AI rights reserved.) 


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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  : : : 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 

- = - 

LONDON: 
PRINTED BY WILLIAM CLOWES AND SONS, LIMITED, 
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: 
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= : 
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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). 


Ol6] 


I 


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CI19],) o140ue UY 


poaxy (CAOH B®. lk) 


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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 


i, 


A , ; pr 
! igs a) Ry ; ate 


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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. 


sroug Ayisteatuy obpraqgmeg xurd ‘apy Dg 


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IIA Id 2908399 ] 19, AS 0TOO7 
OIGl pedxy (SAON PAJ9],) O1}OURYUY Wag 


(ISI YEN) SNA Wag 


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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. 


Vga B G2. 


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Fishes, printed on Special paper. 


| nee 


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. 


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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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PLATE III. 


Fic. 1.—Pagetopsis macropterus, Bouleng. (p. 132) 
Fie. 2. 55 Ss ; 

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Fia. 4.—Gymnodraco acuticeps, Bouleng. (p. 132) 


Length of Specimen. 


14 mm. 
15 mm. 
19 mm. 
24 mm. 


Locality. 


McMurdo Sound. — 


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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. 


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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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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. 


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Larval Fishes,P1 VII. . 


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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, 


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Tripterysium and Heterosomata. 


Larval Fishes, Plate X. 


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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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