THE
VOYAGE OF H.M.S. CHALLENGER.
ZOOLOGY-VOL. XXVI.
REPORT
iW,
SCIENTIFIC RESULTS
OF THE
VOYAGE OF H.M.S. CHALLENGER
DURING THE YEARS i 8 7 3-7 6
UNDER THE COMMAND OF
Captain GEORGE S. NARES, R.N., F.R.S.
AND THE LATE
Captain FRANK TOURLE THOMSON, R.N.
PREPARED UNDER THE SUPERINTENDENCE OF
THE LATE
Sir C. WYVILLE THOMSON, Knt., F.R.S., &c.
REGIUS PROFESSOR OF NATURAL HISTORY IN THE UNIVERSITY OF EDINBURGH
DIRECTOR OF THE CIVILIAN SCIENTIFIC STAFF ON BOARD
AND NOW OF
JOHN MURRAY, LL.D., Ph.D., &c.
ONE OF THE NATURALISTS OF THE EXPEDITION
Zoology— Vol. XXVI.
IPublt'sfteti up ©ttier of ^)er jflajestp's ^ouernment
PRINTED FOR HER MAJESTY'S STATIONERY OFFICE
AND SOLD BY
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1 888
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CONTENTS.
I. — Report on the Crinoidea collected during the Voyage of H.M.S. Challenger,
during the years 1873-1876. Part II. — The Comatul^e.
By P. H. Carpenter, D.Sc, F.R.S., F.L.S., Assistant Master at Eton College.
(The Manuscript ivas received in Instalments between 20th December 1886
and 30th March 1888.)
II. — Report on the Seals collected during the Voyage of H.M.S. Challenger in
the years 1873-1876.
By Sir William Turner, Knt, M.B., LL.D., F.R.SS. L. & E., Professor of Anatomy
in the University of Edinburgh, Member of the General Medical Council.
(The Manuscript ivas received in Instalments between 29th January 1887 and 10 th
January 1888 ; the Appendix, 13th December 1887 and 25th February 1888.)
III. — Report on the Actiniaria dredged by H.M.S. Challenger during the years
1873-1876. Supplement.
By Professor Richard Hertwig.
(TJie Manuscript ivas received 21st January 1888.)
7i
EDITORIAL NOTES.
This Volume contains Parts LX., LXVIL, and LXXIII. of the Zoological
Series of Keports on the Scientific Results of the Expedition.
Part LX. — This Part comprises the second portion of the Report on
the Crinoidea, by Dr. P. H. Carpenter, F.R.S., and deals with the Comatul^e.
The First Part of the Report, dealing with the Stalked Crinoids, forms
Part XXXII. of the Zoological Series of Reports, and was published in 1884
in Volume XI., Zoology.
In the present Memoir Dr. Carpenter gives the results of his investi-
gations, which have extended over thirteen years, and the Report may be
regarded as practically a complete Monograph of the living species of
Comatulag.
The Report consists of 401 pages of letterpress, 70 lithographic plates,
and other illustrations.
Part LXVIII. — This Report on the Seals, by Professor Sir William
Turner, F.R.S., completes the Reports on Marine Mammals, the First Part
of which, on the Bones of the Cetacea, formed Part IV. of the Zoological
Series of Reports, and was published in Volume I., Zoology, in 1880.
The present Part consists of a description of the skeletons of the Seals
collected by the Expedition ; of a revised classification of the Pinnipedia ;
of a description of the brains of the Elephant Seal and Walrus, and a com-
parison with the brains of the Carnivora generally and of Apes and Man ;
also some observations on the viscera of the Elephant Seal. It is accom-
viii THE VOYAGE OF H.M.S. CHALLENGER.
panied by an Appendix on the Myology of the Pinnipedia, by Dr. W. C.
Strettell Miller.
The Report consists of 240 pages of letterpress, illustrated by 10 litho-
graphic plates and several woodcuts.
Part LXXIII. — The Report on the Actiniaria, by Professor Richard
Hertwig, was published in 1882, forming Part XV. of the Zoological Series
of Reports, in Volume VI., Zoology.
The present Memoir forms the Supplementary Report promised at that
time on a number of forms which reached Professor Hertwig too late for the
descriptions to be included in the original Report. It consists of 56 pages
of letterpress and 4 lithographic plates.
The Memoir was translated from the German by G. Herbert Fowler,
Esq., Ph.D., of University College, London.
John Murray.
Challenger Office, 32 Queen Street,
Edinburgh, 21s/ June 1888.
ERRATA FOR PART LX.
Page 34, Hue 22, for " sctosa," read " multispina."
Page 60, line 6 from bottom, for " Actinomctra conjungcns," read " Antcdon conjungcns."
Page 90, line 15, for " Antcdon fluctuans," read " Antedon elegans."
Page 90, line 23, for " the late Mr. Spedding," read " the Eev. T. E. E. Stebbing."
Page 93, line 11, delete the words "of Actinometra diffwilis (PI. LII. fig. 2)."
Page 94, line 5, for " costatus " read " costata."
Page 94, lines 12, 16, for " Antedon fluctuans, n. sp.," read "Antedon elegans, Bell."
Page 96, lines 1, 6, 12, 16, and page 97, line 6, for "fluctuans" read "elegans." (See p. 264.)
Page 97, line 21, for "Antcdon lidentata" read "Antcdon variipinna."
Page 110, line 8, for " Antedon dubia " read " Antedon variipinna."
Page 205, line 6, for " Antedon variipenna," read " Antcdon variipinna"
Page 252, line 5, for " Bidistichate," read " Tridistichate."
Page 320, line 12, for " a.3.2br.^-," read " a.3.2br.^-."
Page 322, line 8, for "figs. 1-3" read "figs. 1-3, 8."
Page 325, line 22, for " figs. 4-6 " read " figs. 4-7."
THE
VOYAGE OF H.M.S. CHALLENGER
ZOOLOGY.
REPORT upon the Crinoidea collected during the Voyage of H.M.S.
Challenger during the Years 1873-76. By P. Herbert Carpenter,
D.Sc, F.R.S., F.L.S., Assistant Master at Eton College.
PART II.-THE COMATULjE.
PREFACE.
The accompanying Report is the result of a study of the Comatulae which has been
carried on, with occasional breaks caused by bad health and by the pressure of other
work, since the latter part of the year 1875. Early in 1878 Sir Wyville Thomson was
good enough to place the Challenger collection in my hands for description ; and in the
following year a preliminary account of it was published in the Proceedings of the Royal
Society.
The group had been singularly neglected for many years previously. Midler's
classical memoir, Ueber die Gattungen und Arten der Comatulen, appeared in 1849, and
laid the foundation of all future systematic work, as well as of the descriptive
terminology now in use. Isolated species have been described by various authors during
the last forty years, but no serious revision of the group has ever been attempted,
though in 1862 Midler's classification was modified in one or two points by Dujardin
and Hupe.
Thirty-five species were described by Midler in 1849, the types of which are scattered
(ZOOL. CHALL. EXP, — PAKT LX.— 1888.) OoO a
11
THE VOYAGE OF H.M.S. CHALLENGER.
through the various museums of the Coutiueut ; and by the kindness of Sir Wyville
Thomson I was enabled to make a personal examination of almost all of these in the
autumn of 1880. Twenty of them belong to Antedon and fifteen to Actinometra, as
these genera are now understood. Four at least of Midler's species (1 of Antedon,
3 of Actinometra) appear to me to have no real value ; while 168 species belonging to
these two genera are considered in the present Eeport, viz., 120 of Antedon and 48 of
Actinometra. Of these there are only two which I have not personally examined, the
type of one having disappeared, while I have not as yet been able to visit the museum
which contains the other.
Of these 168 species 79 were discovered by the explorations of the Challenger
(Antedon 64, Actinometra 15), which also added two new genera, represented by 4
species, to the family Comatulidse. Professor Semper's dredgings in the Philippine
Islands had previously made known the existence of a third generic type, and a fourth
was obtained by the Gulf Stream explorations of the U.S. Coast Survey, though the fact
was not recognised at the time. Other species of each of these two genera were obtained
by the Challenger, making in all 180 species of this family, 88 of which were new to
science.
These numbers are considerably lower than those mentioned in the Preliminary
Report. At the time when that was published I had only seen three large Comatula-
collections besides that of the Challenger, viz., those of the British and Paris Museums,
and that made by Professor Semper in the Philippines. Since then, however, I have
examined many hundred Comatulse, including in many cases large numbers of individuals
belonging to the same specific type, and the experience thus gained has been of the
utmost value, by enabling me to unite under one specific name forms which at first had
seemed distinct to the less trained eye. In one or two cases the result has been that the
specific names appended to some of the plates which were first printed off have since
required alteration ; and the same is true of some of the earlier sheets of the text.
The preparation of this Report has been considerably delayed by the pressure of
other work and by interruptions of various kinds. The first four years after the
collection came into my hands were occupied pretty continuously by the framing of
specific diagnoses. These were nearly all completed when Sir Wyville Thomson died, in
March 1882 ; and my leisure time for the next three years was almost entirely devoted
to the completion of the Report on the Stalked Crinoids which he had left unfinished. I
hoped then that another year's work would enable me to revise my descriptions of the
Comatulse and suffice for the completion of this Report. But this object has been
seriously interfered with by a continual increase of professional duties, together with the
necessity of completing some long-delayed paleeontological work for the Trustees of
the British Museum. I have also been much hindered by a troublesome affection of
the eyes, which has frecmently entailed a prolonged cessation from work.
REPOET ON THE CRINOIDEA. iii
These, and other causes for delay which I need not mention, have not, however,
been altogether disadvantageous ; for various small and more or less local collections of
Comatulse have come into my hands during the past five years, and the experience gamed
by their examination has been of great value in causing me to modify some of my earlier
judgments respecting specific characters.
The seventy plates accompanying this Eeport have been drawn by the following
artists — Messrs. Berjeau and Highley, Parker and Coward, George West and Sons, and
Mr. W. S. Evans ; and I desire to express my thanks to all these gentlemen for the care
with which they have always endeavoured to carry out my wishes. I am also greatly
indebted to my friend Professor F. J. Bell, F.Z.S., of the British Museum, for the ready
way in which he has always given me the utmost facilities for examining the collection
of Comatula? under his care.
I have likewise to acknowledge the courteous kindness of Professor S. Loven at
Stockholm, Professor A. Schneider of Breslau, Dr. E. von Marenzeller of Vienna,
Dr. Otto Hamann of Gottingen, and Dr. F. Nansen of Bergen, who have been good enough
to send selected Comatulaa to me for examination from the collections in their charge.
It is proper also that I should record my indebtedness to the authorities of the numerous
Continental Museums which I visited in 1880, for the uniform courtesy with which
their collections were placed at my disposal.
It only remains, in conclusion, for me to express my sincere thanks to Mr. John
Murray for the kind patience with which he has borne the numerous delays in the
completion of this Report, to which I have referred above ; and also to Mr. W. E. Hoyle,
M.A., of the Challenger office, for the careful manner in which he has supervised its
passage through the press.
Eton College, April 1888.
TABLE OF CONTENTS.
MORPHOLOGY.
I. — Genekal Introduction,
II- — The Centeo-Dorsal and Calyx,
A. The Centro-dorsal,
Inferior Surface, .
Superior Surface and Interradial Symmetry
External Form,
Obliteration of Cirrus-Sockets,
B. The Chambered Organ,
C. The Rosette, ....
Basal Star,
D. TheRadials, ....
Radials of Antedon,
Radials of Actinometra,
E. Abnormal Conditions of the Rays,
III. — The Geographical and Bathymetrical Distribution of the Comatul^e,
Thaumatocrinus, Promachocrinus, and Eudiocrinus,
Antedon, Ten-armed Species,
Antedon, Multibrachiate Species, .
Actinometra, .....
IV. — The Geological History of the Comatul^;,
V. — Classification, .....
General Rules of Co?reatfu/a-strueture,
Methods of Formulation,
List of Antedon Species, ....
List of Actinometra Species,
VI. — Description of the Specimens,
Family Comatulidre, ....
Definition of Genus Tliaumatocrinus, .
Thaumatocrinus renovatus, P. II. Carpenter,
Definition of Genus Atelecrinus,
Synopsis of the Species,
Atelecrinus balanoides, n. sp.,
wy villii, n. sp., .
PAGE
1
6
6
7
10
13
14
16
19
22
23
24
25
27
29
31
32
34
35
37
41
44
46
53
57
63
63
66
66
68
70
70
72
VI
THE VOYAGE OF H.M.S. CHALLENGES.
Definition of Genus Eudiocrinus,
Characters of the Calyx,
The Cirri,
Geographical Eange, .
Synopsis of the Species,
Eudiocrinus varians, n. sp., .
semperi, n. sp., .
japonicus, n. sp., .
Genus Antedon, de Freniinville,
Definition and History of the Genus,
Oral Pinnules,
Sacculi,
Series I. Elegans-grou-p,
Antedon elegans, Bell,
multiradiata, n. sp.,
microdiscus, Bell,
Series II. Ten-armed Species,
Synopsis of the Groups,
1. The Ba$icurva-gco\\\>,
Synopsis of the Species, .
Antedon longicirra, n. sp.,
vcdida, n. sp.,
incerta, n. sp.,
gracilis, n. sp., .
lusitanica, n. sp.,
breviradia, n. sp.,
spinicirra, n. sp.,
acutiradia, n. sp.,
bispinosa, n. sp.,
laiipinna, n. sp.,
multispina, n. sp.,
echifiata, n. sp.,
basicurva, n. sp.,
incisa, n. sp.,
tuberosa, n. sp.,
parvipinna, n. sp.
flexilis, n. sp.,
aculeata, n. sp.,
denticulata, n. sp.,
pusilla, n. sp.,
2. The .4c«?a-group, .
Synopsis of the Species,
Antedon acoela, n. sp.,
discoklea, n. sp.,
3. The Eschrickti-groixp,
Synopsis of the Species,
Antedon eschrichti, Mull., sp.,
antarctica, n. sp.,
australis, n. sp.,
rhomboidea, n. sp.,
quadrata, n. sp.,
PAGE
73
75
76
79
81
81
82
84
85
88
91
92
94
94
93
97
99
99
100
102
103
104
106
107
109
110
112
113
115
116
117
119
120
124
126
127
128
128
130
131
131
132
132
134
136
138
138
144
146
148
149
REPORT ON THE CRINOIDEA.
Vll
PAGE
4. The Tenella-growp, . 156
Synopsis of the Species, .
157
Antedon phalangium, Miill., sp., .
158
hystrix, n. sp., .
165
tenella, Retzius, sp.,
169
exigua, n. sp., .
178
alternata, n. sp.,
179
C rosacea, Linck., sp., i
| petasus, Diiben and Koren, sp., )
181
dubeni, Bohlsche,
181
lineata, n. sp., .
183
remota, n. sp., .
184
longipinna, n. sp.,
185
tenuicirra, n. sp.,
186
Isevis, n. sp.,
187
hirsuta, n. sp., .
188
angustipinna, n. sp.,
189
abyssorum, n. sp.,
190
abyssicola, n. sp.,
191
5. The Millierti-gToxrp,
192
Synopsis of the Species, .
193
Antedon milberti, Miill., sp.,
194
anceps, n. sp., .
198
variipinna, Carpenter,
198
carinata, Lamarck, sp.,
199
parvichra, n. sp.,
204
informis, n. sp.,
205
Synopsis of Unclassified Species,
206
Antedon balanoides, n. sp.,
207
Series III. Bidistichate Species,
208
6. The S/nHifera-grou-p,
211
Synopsis of the Species, .
211
Antedon maeronema, Miill., sp.,
212
quinquecostata, n. sp.,
215
lusitanica, n. sp.,
217
flexilis, n. sp., .
217
patula, n. sp., .
219
robusta, n. sp., .
220
compressa, n. sp.,
222
7. The Palmata-gToup,
223
Synopsis of the Species, .
225
Antedon manca, n. sp., .
226
disciformis, n. sp.,
228
clemens, n. sp., .
229
marginata, n. sp.,
230
tuberculata, n. sp.,
232
conjungens, n. sp.,
233
similis, n. sp., .
235
occidta, n. sp., .
236
regalis, n. sp., .
237
Vlll
THE VOYAGE OF H.M.S. CHALLENGER.
PAGE
Series IV. Tridistichate Species, .
238
8. The Granulifera-group,
239
Synopsis of the Species,
241
Antedon angusticalyx, n. sp.,
242
inxqualis, n. sp.,
244
distincta, n. sp.,
247
multispina, n. sp.,
248
porrecta, n. sp.,
250
9. The Savignyi-gvourt,
252
Synopsis of the Species,
252
Antedon angustiradia, n. sp.,
253
anceps, n. sp., .
254
variipinna, Carpenter, .
256
quinduplicava, n. sp.,
-
262
Note on Antedon fiuduans (elegans), .
264
Genus Adinometra, Miiller,
266
Definition of the Genus, .
267
History,
268
Position of the Mouth,
273
Non-tentaculiferous Arms,
275
Ovoid Bodies,
275
Terminal Combs of Pinnules,
276
Centro-dorsal and Calyx,
276
Series I. Synopsis of the Groups, .
277
1. The Solaris-group,
278
Synopsis of the Species,
278
Adinometra pedinata, Retzius, sp.,
284
Solaris, Lamarck, sp.,
288
2. The Pcrac/aVra-group,
290
Adinometra paucidrra, Bell,
291
3. The Typica-gvoux),
294
Synopsis of the Species, .
295
Adinometra distincta, n. sp.,
295
typica, Loven, sp., .
296
multibradiiata, n. sp.,
299
Series II. Ten-armed Species,
300
4. The Echinopitera-gxowp,
301
Adinometra nieridionalis, Pourtales, sp.,
301
Series III. Bidistichate Species, .
302
5. The Stettir/era-gvour),
303
Synopsis of the Species, .
304
Adinometra pidchella, Pourtales, sp.,
304
mandata, n. sp.,
307
stelligera, n. sp.,
308
6. The Vidida-gxoux),
310
Synopsis of the Species, .
311
Adinometra elongata, n. sp.,
311
simplex, n. sp.,
312
rotalaria, Lamarck, sp.,
313
valkla, n. sp.,
314
REPORT ON THE CRINOIDEA.
IX
Series IV. Tridistichate Species, .
7. The Fimbriata-giQaj),
Synopsis of the Species, .
Aclinometra fimbriata, Lamarck, sp.,
cojppingeri, Bell,
mulfiradiata, Linn., sp.,
sentosa, n. sp.,
lineata, n. sp.,
8. The Parvicirra-group,
Synopsis of the Species, .
Actinometra parvicirra, Mull., sp
quadrata, n. sp.,
triclwptera, Mull., sp.,
divaricata, n. sp,
belli, n. sp.,
duplex, n. sp.,
nobilis, n. sp.,
parvicirra, Mull., sp.,
triclwptera, Miill., sp.,
littoralis, n. sp.
regalis, n. sp.,
Definition of the Genus Promachocrinus,
Synopsis of the Species,
Promaclwcrinus kerguelensis, n. sp,,
abyssorum, n. sp.,
naresi, n. sp.,
VII. — Bathymeteical Distribution and Station List,
H.M.S. " Lightning," 1868,
H.M.S. "Porcupine," 1S69,
H.M.S. "Porcupine," 1870,
H.M.S. "Valorous," 1875,
H.M.S. "Alert," 1875, .
H.M.S. " Knight Errant," 1880,
H.M.S. "Triton," 1882, .
H.M.S. Challenger, 1873-76,
PAGE
315
316
317
317
320
322
325
327
329
330
331
331
332
332
334
335
336
338
345
346
347
348
350
350
351
352
353
353
353
354
355
355
356
356
357
INDEX TO WOODCUTS.
Fig. 1 . Thaumatocrinus renovatus, P. H. Carpenter, ....
Fig. 2. Antedon basicurva, n. sp.; side view of the calyx and arm-bases after removal of three rays,
Fig. 3. The same, .........
Fig. 4. The fiftieth and next following brachials of Antedon quadrata and Antedon eschrichti, .
Fig. 5. Pinnules of Antedon inxqualis and Antedon angusticalyx, ....
Fig. 6. Diagrams showing the different positions of the mouth in Actinometra,
(ZOOL. CHALL. EXP. — PART LX. — 1888.) Ooo b
67
100
122
154
246
274
MORPHOLOGY.
I.— GENERAL INTRODUCTION.
The Comatulse constitute a group of Neocrinoids, which is so extensive, and differs
so much from the remaining members of the order, that a subordinal rank may not
improbably come to be assigned to it. The great variety and extensive distribution of
the species of Antedon and Actinometra at the present time recall similar facts about
Pentacrinus and Millericrinus in the Mesozoic rocks, and about Actinocrinus and
Platycrinus in the Palaeozoic series.
Although a few Palseocrinoids, such as Agassizocrinus and Edriocrinus, seem to
have been stemless and unattached in the adult condition, the enlargement of the top
joint of the larval stem into a cirrus-bearing centro-dorsal is not known to have
occurred in any Palaeozoic, or even in any Triassic Crinoid ; while the physiological
condition of the young Edriocrinus has been frequently reproduced in the Mesozoic
Holopidse and in the recent genus Holopus, which inhabits comparatively shallow
water in the Caribbean Sea, side by side with the free Crinoids or true Comatulae.
The real nature of the latter group was long misunderstood. Linck and Linnaeus
followed Llhuyd in regarding them as peculiar forms of the Sea-stars, to which the general
name Asterias was assigned by the great Swede. Early in the present century, however,
the free Crinoids were separated from the Asterids and Ophiurids by Lamarck. But he
entirely failed to recognise their relationship to Guettard's Pentacrinus, which he placed
among the Polypes, together with the various species of fossil Crinoids.
Five years before Lamarck wrote, the genus Antedon had been established by de
Freminville1 for a Feather-star from tropical seas ; while in the next year Leach2 united
all the known species of this type of the Echinodermata under the one genus Alecto. A
similar step was taken in 1816 by Lamarck,8 who proposed the genus Comatula and
assigned to it eight species, six of them being new. One of these had been previously
1 M«5moire sur un Nouveau Genre de Zoophites de l'Ordre des Radiaires, Bull. Soc. Philom. Paris, Bd. ii.
pp. 349, 350, 1811.
2 The Zoological Miscellany, London, 1815, vol. ii. p. 61.
3 Histoire Naturelle des Animaux sans Vertebres, ed. 2, Paris, 1816, torn. ii. p. 530.
(ZOOL. CHALL. EXP. — PART LX. 1887.) OoO 1
4*
2 THE VOYAGE OF H.M.S. CHALLENGER.
called Antedon by de Fremiuville, and Alecto by Leach ; but Lamarck's authority as a
zoologist, together with his description of six new species, was sufficient to make his
genus more widely known than either de Freminville's Antedon or Leach's Alecto. The
very appropriate name Comatula was afterwards used by Miller, Goldfuss, de Blainville,
Agassiz, and Midler ; while d'Orbigny1 gave it an increased importance by founding the
family Comatulidae. He referred to this family, however, not merely the various forms of
Feather-star, both recent and fossil, in which the base of the calyx is closed below by the
cirrus-bearing centro-dorsal piece, but also the remarkable genus Marsupites, which, in
the adult condition at any rate, was totally devoid both of stem and of cirri. Further
research has shown, however, that Marsupites represents a form of Crinoid which is
altogether different from that of the Feather-stars ; and it is now generally considered as
the type of another family altogether, the Marsupitidaa.
The limits of d'Orbigny's family Comatulidaa have varied considerably at different
times. Eugeniacrinus and its allies were referred to it by Dujardin and Hupe,2 whose
classification has not been adopted by their successors ; whilst a variety of generic
names have been proposed for the numerous fragments of fossil Coniatulse which occur
in considerable abundance at certain horizons in the Jurassic and Cretaceous formations,
viz., Glenotremites, Solanocrinus, Decacnemos, Decameros, Comaster, Hertlia, and
Geocoma. All of these, with one or two possible exceptions, find their place within
de Freminville's genus Antedon, as has been explained elsewhere.3 Some twenty-five
years ago this name was revived by Mr. Norman4 in a more restricted sense than that in
which it was proposed by de Freminville ; and this step has been generally followed, with
the great advantage of simplifying tbe nomenclature considerably.
In Midler's earlier writings upon the subject of the Feather-stars, the names Alecto
and Comatula seem to have been employed indifferently and as equivalent to one
another ; but he was subsequently led to distinguish two different types of Feather-star,
one with five ambulacral grooves converging upon a generally central mouth, as in
Pentacrinus, and the other with an excentric mouth and fewer than five disk-ambulacra.
He therefore considered these as subgenera of Lamarck's original genus Comatula, and
while distinguishing the first one by Leach's name Alecto, proposed to call the second
type by the new designation Actinometra.5 Neither of these two subgenera were ever
formally defined, and Midler only described three species of Actinometra. A fourth was
1 Cours elementaire de Paleontologie et de Geologie stratigraphique, Paris, 1852, vol. ii. fasc. 1, p. 138.
2 Histoire Naturelle des Zoophytes, l^chmodermes, Paris, 1862, p. 186.
3 See P. H. Carpenter, On the Genus Actinometra, Muller, with a Morphological Account of a new species (Actino-
metra polymorpha) from the Philippine Islands, Trans. Linn. Soc. Lond. (Zool.), 1879, ser. 2, vol. ii. pp. 13, 14 ; and also
On the Genus Solanocrinus, Goldfuss, and its relations to recent Comatula', Journ. Linn. Soc. Lond. (Zool.), 1880, vol.
xv. pp. 196-201.
4 On the Genera and Species of the British Echinodermata, pt. i , Ann. and Mag. Nat. Hist, 1865, ser. 3, vol. xv.
p. 98.
5 Ueber die Gattung Comatula, Lam., und ihre Arten, Abhandl. d. h. Akad. d. JViss. Berlin, 1849, p. 246.
REPORT ON THE CRINOIDEA. 3
subsequently added by Bohlsche1 after the type had been accorded generic rank by
Dujardin and Hupe,2 who improved considerably upon Midler's definition of it. The
German zoologist had not considered the position of the mouth as a point of any
systematic importance ; but he referred to Alecto both species like Antedon rosacea and
Antedon eschrichti, which have five symmetrically distributed ambulacra radiating from
a central mouth, and also species with an equally symmetrical grouping of the ambulacra,
but with an excentric mouth. Dujardin and Hupe, however, took no account of the
number of ambulacra diverging from the peristome, to which Mliller attached so much
importance ; but they pointed out that the distinctive character of Actinometra rather lay
in the excentric position of the mouth, which determined the course of the ambulacra
round the margin of the disk, instead of towards its centre. Nevertheless, they did
not transfer to Actinometra the various species of Alecto described by Muller with an
excentric mouth and symmetrically grouped ambulacra ; so that they did not make any
real addition to the genus, although they recognised its characters better than Muller
had previously done. Single species were subsequently added to it by various writers,
but it was never properly defined.
Having assigned a generic value to Actinometra, Dujardin and Hupe" did the same
for Midler's type Alecto, for which, however, they preferred Lamarck's name Comatula.
But three years later Mr. Norman replaced this by Antedon, a name which was originally
proposed earlier than either Alecto or Comatula; and at the same time he restricted it
to those species only in which the mouth is central or subcentral and the anus lateral.
Very nearly all subsequent writers have accepted this definition of Antedon ; but no
attempt was ever made to modify the Mullerian descriptions of Comatula in accordance
with it.
Towards the end of 1875, ten years after the publication of Mr. Norman's precise
definition of Antedon, I had the opportunity of studying a large collection of tropical
Comatulse which had been obtained by Professor Semper in the Philippine Islands ; and
it soon became evident that the number of ambulacra diverging from the peristome is so
variable as to be useless for the purposes of generic discrimination. At the same time
other characters seemed to be correlated with the central or excentric positions of the
mouth respectively ; and I came to the conclusion that the real distinction between
Antedon and Actinometra respectively is based upon this feature of their organisation,
the number of groove trunks connected with the peristome being a character of very
minor importance.8 I soon learnt that Professor Liitken had held this opinion for some
time past; and he also pointed out to me certain characters of the oral pinnules which are
always associated with the excentric position of the mouth. Since that time I have
1 Ueber Actinometra Bennettii unci eine neue Comatula Art (Autedou Dubenii), Archiv f. Naturgesch.,
1866, Jahrg. xxxii. Bd. i. p. 90.
2 Op. cit., p. 208.
3 Trans. Linn. Soc. Lond. (Zool.), 1879, ser. 2, vol. ii. pp. 17, 18.
4 THE VOYAGE OF H.M.S. CHALLENGER.
examined a very large number of Coniatuhe ; and I have almost always found a
terminal comb on the oral pinnules of those species which have an excentric mouth ;
while a variety of other characters are more or less constantly associated with these, as
wTill be explained in detail further on.
In the year 1866 a new Comatulid genus, Phanogenia, was established by Loven1 for
a remarkable tropical species with a stellate centro-dorsal bearing a few rudimentary
cirrus- stumps. The dredgings of the Challenger, however, have shown that this
condition is common to several species of Actinometra, with which the genus Phanogenia
corresponds in all essential respects. A third new genus of Comatulae was established in
1868 by Semper2 for a little five-armed type which he had discovered in the Philippines.
He called it Ophiocrinus, and for some years it was regarded merely as a subgenus of
Antedon. Eventually, however, after examination of the three species obtained by the
Challenger in the Pacific, together with Semper's original specimen, I satisfied myself
of its claim to generic rank, and I proposed to call it Eudiocrinus,5 instead of by Semper's
name Ophiocrinus which had been preoccupied by Salter. But about the same time that
this new generic name was proposed on account of all the known species being limited to
the Pacific Ocean, another specific type was discovered by the " Travailleur " in European
seas, and it was subsequently described by Perrier 4 as Eudiocrinus atlanticus.
One other genus of recent Crinoids, has been described, besides those just mentioned
{Antedon, Actinometra, Phanogenia and Eudiocrinus), viz., Comaster, Agassiz.5 The
leading character of this genus, according to its proposer, depended upon the number of
divisions in the arms, and was rightly disregarded by Goldfuss e who thought more of the
presence of basals on the exterior of the calyx as a generic distinction. Miiller 7 adopted
the genus in the sense in which it was understood by Goldfuss ; but he seems eventually
to have abandoned it altogether.8 This will doubtless prove to be its ultimate fate, as it
has not been seen by any naturalist since the time of Goldfuss, wdiose original specimen
of it was dissected and has since disappeared. If his account of it is correct, Comaster
must be a very remarkable type, differing in many respects from all other recent Comatulae,
as I have explained elsewhere ; 9 but I am strongly inclined to believe that its apparent
peculiarities are merely due to the wTant of knowledge respecting the internal structure of
1 Phanogenia, et hittills okiindt slagte af fria Crinoideer, Dfversigt. k. Vetensk. Akad. FbrhandL, 1866, p. 231.
2 Ophiocrinus, eine neue Cornatuliden Gattung, Archivf. Naturgesch., 1868, Jahrg. xxxiv., Bd. i. p. 68.
3 Descriptions of new or little known Comatuhe. I. On the species of Atelecrinus and Eudiocrinus, Journ. Linn.
Soc. Load. (Zool.), 1882, vol. xvi. p. 493.
4 Sur des Eudiocrinus de lAtlantique et sur la nature de la faune des grandes profondeurs, Comptes rendus, 1883,
t. xcvi. pp. 725-728.
5 Prodrome d'une Monographie des Radiaires ou Echinoderraes, Mem. Soc. Nat. Sci. Neuch., 1835, t. i., p. 193.
6 Beitrage zur Petrefactenkiinde, Nova Acta Acad. Oses. Leop., 1839, Bd. xix. A. p. 348.
7 Ueber die Gattungen und Arten der Comatulen, Monatsber. d. k. preuss. Akad. d. wiss. Berlin, 1841, p. 180.
8 Abhandl. d. k. Akad. d. Wiss. Berlin, 1849, p. 244.
9 Journ. Linn. Soc. Lond (Zool.), 1877, vol. xiii. pp. 454-456.
REPORT ON THE CRINOIDEA. 5
the calyx of Comatuke which was prevalent at the time of Goldfuss, and that Comaster is
in reality nothing but a large Antedon or Actinometra.
Apart from Phanogenia and Comaster, therefore — one, if not both, of which are
merely synonyms — no other Comatulid genera except Eudiocrinus, Antedon, and
Actinometra were known, to science before the collections of the Challenger and of the
United States Coast Survey ships came into my hands for examination. But one species
of Eudiocrinus was known, and only about twenty each of Antedon and of Actinometra
had been described, though many others were awaiting description in various museums.
Now, however, the number of recent species of Comatula is probably nearly four hundred,
and three new genera have been established, thus doubling the number known at the time
the Challenger returned. One of these generic types, Atelecrinus, was actually obtained so
long ago as 1868, during the earliest explorations of the Gulf Stream by Count Pourtales ;
but the single specimen dredged was so small and mutilated that its very striking
peculiarities escaped notice at the time. Equally imperfect and isolated examples of two
other species were dredged by the Challenger ; and it was not until several less mutilated
individuals were obtained by the " Blake " in the Caribbean Sea, that I was able to
realise that a new Comatula genus had been discovered.1 It presents so many larval
characters that I have called it Atelecrinus, as will be explained subsequently.
Atelecrinus can hardly be considered as a new genus discovered by the Challenger ;
but with P romachocrinus and Thaumatocrinus the case is altogether different. The
former genus 2 differs from all other Crinoids in the composition of the calyx, which has
ten primary radials instead of five only, as is normally the case ; and it is represented by
three distinct species, one from the North Pacific, one from Kerguelen, and one from a
depth of 1800 fathoms at Station 158 in the Southern Sea. At this Station too, there
was obtained a single specimen of another Comatula which I have no hesitation in
regarding as by far the most remarkable of all the Crinoids that have been dredged of
late years, viz., the extraordinarily archaic form TJiaumatocrinus, which presents certain
characters only to be found in some of the Palaeocrinoids. Its peculiarities were fully
described in the Report on the Stalked Crinoids,3 and I do not propose therefore to say
much about it here.
1 Report on the Results of Dredging under the Supervision of Alexander Agassiz, in the Gulf of Mexico, and in the
Caribbean Sea, 1877-79, and along the Atlantic Coast of the United States during the summer of 1880, by the
United States Coast Survey steamer " Blake," Lieutenant-Commander C. D. Sigsbee, U.S.N., and Commander J. R.
Bartlett, U.S.N., commanding. XVI. Preliminary Report on the Comatula?, Bull. Mus. Comp. ZooL, 1881, vol. ix.
No. 4, p. 16.
2 Preliminary Report upon the Comatuhe of the Challenger Expedition, Proc. Roy. Soc, 1879, vol. xxviii. p. 385.
3 Zool. Chall. Exp., part xxxii., 1884, p. 370.
THE VOYAGE OF H.M.S. CHALLENGER.
II.— THE CENTEO-DORSAL AND CALYX.
The principal morphological character which distinguishes the Comatulidse from the
remaining families of Crinoids is the development of cirri upon the top stem-joint, and its
separation from the remaining portion of the stem as the centro-dorsal plate. This
supports the ring of united radials, and, in the recent forms at any rate, closes up below
the dorsal extension of the body-cavity which is contained in their central funnel, as is
well shown in PI. III. figs. 3a, 3b, and PI. V. fig. 2c.
Most recent Comatulse are further distinguished from the Stalked Crinoids by the
metamorphosis of the embryonic basals into the structure known as the " rosette," which
is enclosed within the radial pentagon, and so is entirely invisible externally (PL I.
fig. 8c; PL II. figs. 3c, 5c; PL IV. fig. 3c; PL V. figs. 2c, bd). It will be well
to discuss these two structures separately, though they are naturally in very close relation
with one another.
A. The Centro-Dorsal.
The term " centro-dorsal plate " is a very old one, and was for a long time used in
various ways by different authors. In fact it was not till the remarkable developmental
history of the uppermost stem-joint had been made out by the late Sir Wyville Thomson
and Dr. Carpenter, that the term acquired any definite signification. Both these authors
used it to denote the enlarged and cirrus-bearing top stem-joint1 which is at first in no
way different from the remaining joints of the stem below it (PL XIV. figs. 1, 2, 8, 9).
Eventually, however, it enlarges, and five cirri, which are radially situated, are developed
upon it (PL XIV. figs. 3-6), so that it has very much the appearance of a nodal stem-
joint of Pentacrinus. A second series of cirri, alternating in position with the first,
subsequently appears (PL XIV. fig. 7), and others are afterwards developed in succession,
so that as was well said by Wyville Thomson,2 " the centro-dorsal plate in Antedon does
not belong to the cup. It represents a coalesced series of the nodal stem-joints in the
Stalked Crinoids."
At a certain period in the development of the young Comatula the centro-dorsal
1 The centro-dorsal plate of Comatula must not be confused with the dorsocentral plate of other Echinoderms.
This name is now generally restricted to the central plate of the abactinal system in Urchins and Stellerids. I believe
this to be represented in the Comatula? by the terminal plate at the bottom of the larval stem, as explained on p. 168 of
Part I. It is shown in PI. XIV. figs. 1, 9. Comatul* thus have both a centro-dorsal and a dorsocentral, while the
latter only is present in the remaining Echinoderms. Zittel has also given the name centro-dorsal to the enlarged
uppermost stem-joint of Apiocrinus; but this bears no cirri, and though undoubtedly homologous with the centro-dorsal
of Comatula;, should not, I think, receive a name which is now universally understood as denoting the presence of
cirri.
2 On the Embryogeny of Antedon rosaceus (Linck, Comatula rosacea of Lamarck), Phil. Trans., 1865, p. 536.
REPORT ON THE CRINOIDEA. 7
separates itself from the stem-joint below it. and the " head " of the Pentacrinoid larva
becomes a free-swimming Feather-star, the rest of the larval stem being left to waste
away. The precise epoch of growth at which this separation occurs varies greatly. Thus,
for example, the young Antedon tenella retains its stem until twenty or thirty cirri have
appeared on the centro-dorsal, which conceals the basals, and the pinnules are developed
upon all the lower arm-joints ; whereas in Antedon rosacea and in other species, the stem
is discarded when there are only ten cirri on the centro-dorsal, the basals are still visible,
and the lowest portions of the arms devoid of pinnules ; while the absolute size which is
reached by the mature larva before dropping off its stem varies considerably.
After the formation of the first two whorls of cirri no special regularity can be traced
in the manner of their development. The young ones normally appear between those
previously formed and the radial pentagon, so that their sockets are close to the margin
of the centro-dorsal (PI. I. fig. let; PL II. figs. 2a, 4a; PL IV. figs, la, 3a). But
as the centro-dorsal grows and new cirri appear round its margin, the older cirri which
are attached close to the dorsal pole drop away, and their sockets become gradually
obliterated by calcareous deposit. The earlier stages of this process are seen in PL I.
fig. 6a; PL II. figs, la, 3a, 5a; and PL III. figs. 6a7, 7a; and the result is that the
dorsal surface is usually left comparatively smooth, as seen in PL IV. figs, la, lb, 2a, 3a,
but in some species of Antedon the deposit of new material continues after the cirrus-
sockets are obliterated, and causes the dorsal pole to become rough and irregular (PL III.
figs. 4b, 5a ; PL XL fig. 3). On the other hand, the lower surface of the centro-dorsal in
most species of Actinometra is almost flat and extremely smooth (PL V. figs. 16, Id, 2b,
2d, 2e, 4b, 5b, 5c). This is owing to the very extensive and uniform manner in which
the new material is deposited, and it sometimes produces very singular results, as will
be explained subsequently.
During the Pentacrinoid stage of larval existence the young Comatula is provided
with a stem which encloses a neuro- vascular axis just as in an ordinary Stalked Crinoid.
This axis contains the downward extensions of the peripheral cavities of the chambered
organ within the centro-dorsal and of its central axis. When the centro-dorsal separates
itself from the lower part of the larval stem, a minute five-rayed perforation remains at
its dorsal pole, which corresponds to the central canal in the stem of a Pentacrinus, and
gave passage to the neuro-vascular axis above mentioned. In recent Coinatulse this
opening is closed up very soon after the entry upon the free stage of existence, by a
portion of the calcareous deposit already noticed ; though traces of it are sometimes
visible internally upon the floor of the centro-dorsal cavity (PL II. figs. 2b, 3b). There
are some fossil Comatulae, however, in which it seems to have remained permanently
open throughout life, so far as we can judge from the material at our disposal ; while
in other forms again it is extended into a large stellate impression which occupies a
considerable space on the lower surface of the centro-dorsal, and in the fossil condition is
8 THE VOYAGE OF H.M.S. CHALLENGER.
more or less obliterated. But there can, I think, be no question that in Antedon
■perforata, Antedon rugosa, Antedon striata, and other species from the English Chalk,
together with some foreign species like Antedon tourtiie, Antedon semiglobosa, and
Antedon retzii, the inferior surface of the centro-dorsal was marked during life by a
large stellate opening which was considerably more than would be necessary for the
simple downward passage of the neuro-vascular axis of the stem. It seems to me very
probable, as I have explained elsewhere,1 that the peripheral parts of this opening, which
are radially situated, may have given passage to tubular extensions of the body-cavity
into the stem, such as existed in Barycrinus, Cuprcssocrinus, and in other Palaeocrinoids.
An indirect confirmation of this view is afforded by the characters of the stem in the
Bourgueticrinidge, which resembles that of the young Comatula in all essential points.
The stem-joints of this family contain a set of five radial spaces which communicate with
one another from joint to joint, and probably also through the top of the stem with the
body-cavity within the calyx. The presence of these same radial spaces in the stems of
fossil Comatula? would account for the perforation of the lower surface of the centro-
dorsal, which would have effected the communication between the portions of the body-
cavity derived from the right peritoneal sac, that lie in the stem and in the calyx
respectively. In the ordinary species of Antedon the calycular portion of the ccelom is
much broken up by the rosette, and by the calcareous network which rests above it and
occupies the central funnel of the radial pentagon (PI. IV. fig. 36) ; but, as I have shown
elsewhere,3 there are five median grooves on the ventral surface of the radials which
extend outwards in a similar position over the skeleton of the rays and arms, and
lodge the lowest portions of their cceliac canals. They are more distinct in some species
than in others, but are well shown in Antedon carinata (PL III. figs, id, 3a), Antedon
disciformis (PI. IV. fig. 26), and in Actinometra lineata (PL V. figs. 2a, 2c). When
these grooves pass from the ventral to the inner faces of the radials and descend into the
central funnel, they become closed into canals by the union of their edges with those of
the spout-like radial processes of the rosette. These canals, which I have called the axial
radial canals, are therefore the proximal ends of the five cceliac canals of the arms and their
extensions into the pinnules. As a general rule they become closed up by calcareous
tissue, and so do not reach the dorsal surface of the radial pentagon, which presents no
real openings except the central one occupied by the rosette (PL I. fig. 8c). The five
radial and five interradial processes of this structure are separated by passages which lodge
the paired branches of the five primary cords proceeding from the nervous envelope of
the chambered organ. These ten openings are well seen in PL I. figs. 6c, 8c ; PL III.
figs. 4c, 56; and also in PL V. figs, lc, 2c, 2e, bd, 5e, but in Antedon quinquecostata
and Antedon disciformis there are five additional openings on the lower surface of the
1 On some New Cretaceous Coniatulre, Quart. Journ. Geol. Soc, 1880, vol. xxxvi. pp. 556, 557.
8 Trans. Linn. Soc. Land. (Zool.), 1879, ser. 2, vol. ii. pp. 77, 78.
REPORT ON THE CRINOIDEA. 9
radial pentagon, one at the inner end of each radial (PI. III. fig. 6?> ; PI. IV. fig. 2c).
These are the dorsal ends of the radial axial canals, which do not become obliterated as is
usually the case ; and in Antedon disciformis there is a small pit on the upper surface of
the centro-dorsal corresponding to each of these canals which terminate blindly in this
position (PL IV. figs. 2c, 2c/). Among recent Comatulse, however, the most striking
development in this respect is presented by Antedon quinduplicava ; for the radial axial
canals which pass over from the ventral to the inner faces of the radials turn outwards
ao-ain at the bottom of the calyx, and expand into relatively large bilobate cavities which
are formed by excavation in the apposed surfaces of the radials and the centro-dorsal
respectively, as is well seen in PI. IV. figs, lc, id.
Among the fossil Cornatulte there are several species in which the ventral surface of
the centro-dorsal is marked by five small radial pits of this kind, that receive the
ends of the radial axial canals. But in Antedon retzii they appear as actual perforations
in the ventral surface of the centro-dorsal which reach downward to the bottom of its
internal cavity, being in fact only separated from it by a narrow septum, and this is
occasionally absent, so that the centro-dorsal cavity which is naturally decagonal or
pentagonal in outline becomes stellate. This condition is very common in the stem-joints
of some Palseocrinoidea, such for example as Cupressocrinus, and I think there can be
no doubt that the radial openings or the extensions of the central canal in all such cases
served for the passage of canals containing water in communication with that in the
coelom above.
Messrs. Wachsmuth and Springer ' suggested long since that the complex stem of
many Palseocrinoids might have been " subservient to respiration " ; and the facts
mentioned above respecting the Bourgueticrinidse and the Comatulae certainly go far
towards supporting this view.
The ventral surface of the centro-dorsal is usually flat or slightly hollowed, rarely
very convex, except in species like Actinometra paucicirra, Actinometra typica, &c,
in which the greater part of the centro-dorsal is enclosed within the radial pentagon, as
will be explained shortly. The internal openings of the canals leading to the cirrus-
sockets are frequently visible on the floor of its cavity, as is well shown in Promachocrinus
kerguelensis and in Antedon antarctica (PI. I. figs, id, 6d). In both these species and
also in others the walls of the centro-dorsal cavity are marked by strong ribs, the lower
ends of which are more or less distinctly visible through the axial opening, projecting
beneath its bp, which their upper ends help to support. Five of them, those at the
interradial angles, are often considerably larger than the rest, and may be the only ones
visible. In other cases, however, both these and numerous smaller intermediate ribs are
visible through the axial opening, as is seen in PI. I. figs. Id, 6d. These ribs are much
more distinct in some individuals than in others of the same species. Thus, for example,
1 Revision of the Palseocrinoidea, part i. p 15, Proc. Acad. Nat. Sci. Philad., 1879.
(ZOOL. CHALL. EXP. — PART LX. — 1888.) *-*00 2
10 THE VOYAGE OF H.M.S. CHALLENGER.
they do not appear within the middle portion of the centro-dorsal in the specimen of
Anteclon eschrichti figured in PI. I. fig. Sd, though they are comparatively large in other
forms of this type, as I have noticed elsewhere.1
The peripheral part of the ventral surface of the centro-dorsal is divided by ridges or
grooves into the five trapezoidal areas in which the radial plates are lodged, and they
are occasionally marked by more or less definite pits which receive the ends of the radial
axial canals, as already explained (PI. IV. figs. Id, 2d). In most Comatulse every two
fossse are separated by one of the five basal grooves which lodged the rays of the basal
star, to be described subsequently. They are sometimes comparatively insignificant, as
in Antedon antarctica (PI. I. fig. 6d), while in the Pacific species they are usually very
strongly marked (PI. II. figs. 1-56). On the other hand, if no basal star is present, the
radial fossse on the centro-dorsal are usually separated by tolerably sharp ridges as in
Antedon eschrichti (PI. I. fig. 8d), Antedon quinduplicava (PI. IV. fig. id), and Antedon
disciformis (PI. IV. fig. 2d). The last-mentioned species, however, has indications of
basal grooves at the proximal ends of these ridges. The grooves are fairly distinct in
both the species of Promachocrinus which I have examined, but though the radials are
ten in number, there are only five fossse on the centro-dorsal, the ventral surface of
which is distinctly pentagonal in outline, with its angles interradial, just as in Antedon
(PI. I. figs. \c, Id, 5).
In fact, I know of no Comatula in which the general shape of the centro-dorsal is
not more or less distinctly pentagonal with its ventral ridges and angles interradial.
Wachsmuth and Springer regard this fact as indicating the probable presence of radially
situated under-basals in the Co matula -larva. Their extensive and important investiga-
tions into the structure of the calyx in the Palseocrinoids have led them to formulate the
following rule : 2 — " In species with under-basals, whenever the column is pentangular,
its longitudinal angles are directed interradially, the sides and columnar cirrhi radially."
They proceed to state3 that the centro-dorsal of Comatulse is interradial " and rests,
as in the Apiocrinidse, against the outer face of the basals, not within the basal ring " ;
while they continue — " upon this mainly we base the opinion that perhaps also the
Comatula? in their early larva had rudimentary under-basals. That these plates, if
present, were not observed, is not surprising, as they may have been very minute and
been covered entirely by the column."
Whether this be the case or not, the statement that the centro-dorsal of Comatulse
rests against the outer face of the basals is a somewhat misleading one. The " outer
face " can only mean that which appears on the outside of the calyx ; and this, from its
very nature, cannot rest against the centro-dorsal, for it would then be internal and con-
cealed.
1 Quart. Journ. Geol. Soc, 1880, vol. xxxvi. p. 47.
4 Revision of the Palaeocrinoidea, pt. iii. sect. 1, p. 7 (229); Proc. Acad. Nat. Sci. Philad., 1885.
3 Ibid., sect. 2, 1886, p. 298 (222).
REPORT ON THE CRINOIDEA. 11
The centro-dorsal is at first a simple ring, in no way different from the other stem-
joints ; but when the basals come to assume a definite shape and the calyx acquires the
doubly conical form of the Cystic! phase, the centro-dorsal becomes distinctly wider than
the annular stem-joints below it and takes on a pentagonal shape. The basals rest
against the sides of the pentagon, and its angles which fit in between them are therefore
radial in position, as seen in PI. XIV. figs. 1,8. At this early stage the basals are only
in contact with the centro-dorsal by their lower edges ; but it soon begins to increase in
diameter and extends itself over the bottom of the calyx in the manner described by
Dr. Carpenter.1 It increases at the same time in vertical depth, and the first cirri make
their appearance. These are radial in position, and the portion of the centro-dorsal
between every two sockets rapidly enlarges, so that it comes to project beneath each basal
plate, and the angles of the centro-dorsal thus become interradial instead of radial. This
change is very clearly seen in larvae which have only one or two cirri, so that one part of
the centro-dorsal shows the primitive radial symmetry, and another part the acquired
interradial symmetry.
Thus then the centro-dorsal of Comatula, when it first assumes definite form, has a
most distinct radial symmetry. Its angles occupy the same position with regard to the
basals as do those of the enlarged top stem-joint in Guettardicrinus and Apiocrinus,
which are also distinctly radial in situation. I desire to lay particular stress upon this
fact, because Wachsmuth and Springer, in support of their assertion that Neocrinoids are
built upon the plan of dicyclic Crinoids, have stated that the top stem-joint " is disposed
interradially in the Apiocrinidae, Pentacrinidae, and Comatulse, similar to dicyclic Palseo-
crinoids." 2 But the ridges and angles of the top stem-joint are radial in every species of
Apiocrinus, as is seen with especial clearness in Apiocrinus magnificus.3 Wachsmuth
and Springer 4 say, however, that " the plate in Apiocrinus magnificus is not, as should
be supposed from appearances, disposed radially, but interradially, as shown by comparison
with species having a pentangular stem. It attained its radial angles accidentally by
adapting its form to the basal concavity which is naturally angular." This is a form
of teleological argument which is very easily employed but is very difficult to refute.
Neither Wachsmuth nor Springer, nor any one else, is acquainted with the post-embryonic
development of Apiocrinus, and the changes which may or may not have taken place in
the symmetry of its top stem-joint ; though from the positive way in which the
American authors write one would imagine that they had watched the whole process of
the " accidental " change of symmetry which they describe. If the basal concavity
"naturally" has radial angles, it is surely a "natural" and not an "accidental"
circumstance that the top stem-joint which occupies this cavity should also have radial
angles. This is the case in every species of Apiocrinus, in the single species of
1 Researches on the Structure, Physiology, and Development of Antedon rosaceus, Phil. Trans., 1866, p. 742.
2 Revision, pt. iii. p. 299.
3 See de Loriol, Paleontologie Franchise, Terrain Jurassique, t. xi. pis. 46-49. * Revision, pt. iii. p. 297.
12 THE VOYAGE OF H.M.S. CHALLENGER.
Guettardicrinus, and in the majority of those of Millericrinus ; and yet it is considered by
Wachsmuth and Springer as a merely " accidental " occurrence, and the real symmetry of
the top stem-joint in the Apiocrinidse is described as interradial. It is actually and visibly
so in some twenty species of Millericrinus. But they belong to that aberrant section of the
genus which so closely approaches Pentacrinus in having a distinctly pentagonal stem with
interradial angles, and articular faces the sculpture of which is very different from that of
the typical Apiocrinidse and somewhat closely resembles that of the joint-faces in certain
Pentacrinidae. In all of these species the top stem-joint, like those below it, has interradial
angles, and the same is the case with the basal concavity into which it fits. But Wachs-
muth and Springer tell us that the " natural " shape of this concavity in the Apiocrinidse
is to have radial angles, and they have not attempted to explain its interradial symmetry
in these aberrant and Pentacrinus-l\ke forms of Millericrinus by reference to any causes
whatever, accidental or otherwise. Perhaps it has escaped their notice ; but whether
this be the case or not, it is somewhat surprising to students of the Neocrinoidea to be
told that the distinctive characters of the top stem-joint in the Apiocrinidse, presenting
themselves in each of the three genera, and in by far the greater number of the species of
this family, are due to " accidental " causes. Further discussion of this question, however,
would be impracticable at present. I merely wish to point out that as soon as the
centro-dorsal of the early larva of Comatula takes a definite shape its angles are distinctly
radial, just as is permanently the case in the top stem-joint of Apiocrinus, and this is
in itself an argument against the supposed change of symmetry in the latter type about
which Wachsmuth and Springer write so positively. But when the cirri appear on the
centro-dorsal and the basals begin to be transformed into the rosette, the outline of the
centro-dorsal changes. The basals are no longer the principal plates in the calyx, but
they undergo metamorphosis into the small rosette, and the centro-dorsal increases rapidly
in size, more so than any other part of the skeleton, " so that it soon comes to pass beyond
the circlet of basals, and to abut on the proximal edge of the first radials ; and instead of
stopping here it continues to increase in diameter until it conceals the whole inferior
surface of the first radials, and sometimes even encroaches somewhat on the second." l
Here then we see the reason for the interradial angles of the centro-dorsal in the
mature Comatula. It is an altogether secondary condition, and due to the fact that the
fossae on the ventral surface of the centro-dorsal lodge the radial plates, so that the ridges
separating them are interradial, just as in Apiocrinus the fossa? on the top stem -joint
lodge the basals and are interradial, so that the intervening ridges and the angles in which
they terminate are radial. Even in fossil Comatula? which have no rosette, but persistent
basals, these plates are usually quite small and do not form a closed ring on the exterior
of the calyx; so that the upper surface of the centro-dorsal is mainly occupied by the
radial fossa? and has interradial angles as in recent Comatula? (PI. I. figs. 5, 6a, 8d; PI. II.
1 W. B. Carpenter, Phil. Trans., 1866, p. 742.
REPORT ON THE CRINOIDEA. 13
figs. 1-5, 6). Thus then the interradial symmetry of the centro-dorsal is an altogether
secondary condition, and is due not to the possible presence of radially situated under-
basals, as supposed by Wachsmuth and Springer, but to the fact that the radials
themselves rest upon the plate, the primary radial symmetry of which becomes
altogether obscured when it begins to increase in diameter and to develop cirri,
coincidently with the retromorphosis of the basals.
The external form of the centro-dorsal varies very greatly among different species of
Comatulge. It is very distinctly conical in Atelecrinus (PI. VI. figs. 5, 7). In the
three chief of the remaining endocyclic genera (Antedon, Eudiocrinus, and Promacho-
crinus) it is occasionally somewhat hemispherical or subcorneal, with the cirrus-sockets
arranged rather irregularly (PI. I. figs, la, 6a, 8a; PI. II. figs. 1-3, a; PL III. figs. 46,
5a, la; PI. XXX. figs. 1, 2, 4); but in some cases, as in Antedon quinquecostata, it is
more distinctly pentagonal and columnar, with the sockets grouped in alternating rows
(PI. III. fig. 6d), while in Antedon balanoides it is distinctly conical (PI. XXXIII.
fig. 6). In other forms again the dorsal pole is flattened (PI. II. fig. 4a), and this is
especially the case in Antedon carinata and Antedon macronema (PI. III. figs, la, 36 ;
PI. IV. fig. 3a), which in this character, as in some others, exhibit a variation in the
direction of Actinometra. On the other hand, Antedon quinduplicava and Antedon
disciformis, which are still more like Actinometra in the small number of functional
cirrus-sockets and in the discoidal shape of the centro-dorsal, belong unmistakeably to the
genus Antedon in the relative height of the radials (PL IV. figs, la, 2a).
In most species of Actinometra the centro-dorsal is a thin flattened disc, often with
only one row of functional cirrus-sockets (PL IV. fig. 4a ; PL V. figs. 16, Id, 26, 2a",
2e, 46) ; though in Actinometra stelligera it is thicker and bears a comparatively large
number of sockets (PL V. figs. 56, 5c).
As a general rule the shape of the centro-dorsal is tolerably constant in any individual
species of Antedon, being hemispherical in Antedon eschrichti (PL I. fig. 8a; PI.
XXIV. figs. 10, 11), columnar in Antedon quinquecostata (PL III. fig. 6d), and more
discoidal in Antedon carinata (PL III. figs, la, 36). But in Antedon phalangium it
exhibits a very considerable amount of variation, being hemispherical in some forms, but
greatly elongated and conical in others (PL XXVIII. fig. 2).
In some species of Actinometra the obliteration of the cirrus-sockets on the centro-
dorsal is carried to a very much greater extent than in Antedon ; and the number of
functional sockets, which is at no time large, is often extremely small. In some types the
changes in the centro-dorsal do not stop here, but it is reduced to the condition of a flat
pentagonal plate within the ring of radials as in Actinometra paucicirra (PL LIV.
figs. 1-7) ; while in species like Actinometra typica (PL LVII. fig. 1), the sides of
this plate undergo resorption, so that clefts appear between it and the radials. This
gives the base of the calyx an appearance so different from that of the ordinary Comatulas
14 THE VOYAGE OF H.M.S. CHALLENGER.
that the genus Phanogenia was instituted by Loven1 for the reception of species
presenting these characters.
It was pointed out in my preliminary report,2 however, that the stellate appearance
of the centro-dorsal in Phanogenia (PI. LVII. fig. 1) "appears to be one of the concluding
stages of a long series of changes in the shape and relations of the centro-dorsal, which
do not commence until some time after the loss of the stem and the entry upon the
free state of existence." The earlier stages of these modifications are well shown in a
series of specimens of Actinometra 'paucicirra, which is very abundant at Cape York
(PL LIV.). In the youngest individual of the series the centro-dorsal is a thin
and slightly convex circular disc, about 2 mm. in diameter, which bears five pairs of cirri,
one pair opposite each interradius. They reach 6 mm. in length and consist of about
fifteen joints, which are tolerably mature in their general characters (PI. LIV. fig. 10);
the next stage is a slightly older individual in which all the cirri have fallen away from
the centro-dorsal and the obliteration of their sockets has commenced (fig. 9). This
process has been carried further in the larger and more distinctly pentagonal centro-
dorsal shown in fig. 8, though it has gone on rather unequally, some of the sockets
being much more obliterated than others.
Scarcely any trace of sockets can be made out in the original of fig. 5, but the centro-
dorsal is a thin pentagonal disc with the appearance of processes at some of its angles,
which are more probably, however, the ends of the basal rays. Its surface is much more
nearly flush with that of the radials in the full-grown specimen shown in fig. 2, still,
however, retaining its pentagonal shape. Fig. 1 shows another modification, each
angle of the pentagon being marked by a more or less deeply impressed pit in which the
basal ray is sometimes visible. The form represented in fig. 3 has a more rounded
centro-dorsal, which is flush with the radials at its edges, and shows the basal rays at its
angles; while there are indications of pits at the distal angles of the sutures between the
first radials. The sides of the centro-dorsal in this specimen are slightly concave, and
this character is much more distinct in figs. 6, 7, so that the shape becomes markedly
stellate. In the former the centro-dorsal (as viewed from the dorsal side) is above the
level of the radial pentagon; but in the latter it is relatively much lower, so that its
surface is flush with that of the radials, the proximal edges of which are convex in
correspondence with the stellate outline of the centro-dorsal. Fig. 4 shows a similar
case in which the centro-dorsal is pentagonal. The effect of its complete withdrawal
into the radial pentagon is to make it entirely invisible in a side view of the calyx, as
seen in PL V. fig. 3 b ; while the dorsal surface of the united radials becomes very
deeply hollowed for its reception (PL V. fig. 3c) instead of being slightly convex, as i3
more usually the case (PL V. figs, lc, 5d).
1 Phanogenia, ett hittils okandt slagte af fria Crinoideer, Ofvcrsigt k. Vetensk.-Akad. ForhandL, 1866, p. 231.
*Proc. Roy. Soc, 1879, vol. xxviti. p. 390.
REPORT ON THE CRINOIDEA. 15
The stellate condition of the centro-dorsal just described in Actinometra paucicirra
is sometimes reached by that of Actinometra parvicirra before the cirrus-sockets are
entirely obliterated. In one specimen of this variable type which was obtained by the
Challenger, the mature cirri have disappeared and are replaced by a few rudimentary
stumps, while the sides of the plate are so deeply hollowed by their sockets that its
outline is rather stellate than pentagonal. But it is still distinctly above the level of the
radials (PL LXI. fig. 3).
The six examples of the large Actinometra nobilis which were dredged in the
Philippines also exhibit a considerable amount of variation in the characters of the
centro-dorsal (PI. LXV. figs. 1-6). In the least modified form it is a rounded pentagonal
plate distinctly above the level of the calyx, with traces of about ten cirrus-sockets, one
of which contains a very rudimentary stump, and a well-marked process at each of its
angles (fig. 2). In another specimen it is distinctly sunk below the level of the radials,
with which it remains united externally by the interradial processes at its angles ; but
its sides are bevelled away, and most of them bear indistinct cirrus-sockets, in one of
which a small stump is visible (fig. 3). In the other four examples, however, the
centro-dorsal shows little or no trace of cirri, and is distinctly concave on its dorsal
surface ; while it is completely enclosed by the radial pentagon, united to it by the
interradial processes at its angles, but separated from it by very distinct clefts along its
sides. Its shape, however, is more pentagonal than stellate (PI. LXV. figs. 1, 4-G).
These clefts are rather deeper in Actinometra littoralis, though the centro-dorsal
retains its distinctly pentagonal form, and is about flush with the radials, with which it
is in contact by its lower angles (PL LXVII. fig. 1). On the other hand, in Actinometra
divaricata the centro-dorsal is very markedly stellate, and remains above the level of
the radials, the surface of which falls away considerably towards the sides of the centro-
dorsal, but not so much so as to give rise to definite clefts (PL LXIII. fig. 6).
In the Challenger specimen of Actinometra typica, however, in the original type of
Phanogenia, and in others which I have seen, the centro-dorsal is both stellate and sunk
below the radials, so that there are very distinct clefts between the latter and its incurved
sides ; and no one would think from its present appearance that it had ever been a
cirrus-bearing joint (PL LVII. fig. 1). But in Loven's specimen the metamorphosis
was less complete, for a few cirrus-stumps are figured as still attached to the stellate
centro-dorsal, which is slightly above the level of the radials. The facts stated above,
however, concerning Actinometra paucicirra, Actinometra nobilis, and other forms,
entitle us to assume that cirri were really present in the young Phanogenia, so that the
genus ceases to have the extremely anomalous character which Loven not unnaturally
attributed to it.
These clefts which occur at the sides of the centro-dorsal in Actinometra typica and
similar species must not be supposed to place the cavity of the calyx in communication
16 THE VOYAGE OF H.M.S. CHALLENGER.
with the external medium. They occur in several species of Actinometra in which the
centro-dorsal undergoes very little modification, as for example in Actinometra pectinata.
The small centro-dorsal of this species, as I have shown elsewhere,1 retains its cirrus-
sockets and its discoidal form, but has five minute openings round its margin ; and these
lead into spaces between its ventral surface and the lower surface of the radial pentagon,
which are formed by the apposition of depressions in each of these surfaces respectively.
But sections through the calyx of this type show that the radial spaces leading inwards
from these marginal openings terminate internally against the inner portion of the
ventral surface of the centro-dorsal, and are completely shut off from the radial axial
canals enclosed between the rosette and the inner faces of the radials. There is, therefore,
no such communication between the body-cavity and the exterior as the presence of these
radial spaces might be supposed to indicate. They are precisely homologous with the
interarticular pores in the stem of Pentacrinus, which lead inwards some little way, as
described in Part I., but are in no communication with the central canal of the
stem.
It is worth notice that in one fossil species, Actinometra loveni? from the Gault of
Folkestone, the centro-dorsal approaches the Phanoge ma-condition. It is an almost
pentagonal plate, scarcely above the level of the radials. from which it is separated by
narrow clefts, just as in Actinometra typica (PI. LVII. fig. 1), and in Actinometra
nobilis (PL LXV. figs. 3-5), and from the close resemblance of the calyx to that of these
and similar species which are nearly all inhabitants of quite shallow water (20 fathoms or
less), it would appear that the portion of the Gault Sea in which Actinometra loveni
lived cannot have reached any great depth.
B. The Chambered Organ.
Reference has been made above to the radial axial canals which are enclosed between
the rosette and the radials, and sometimes reach the ventral surface of the centro-dorsal.
Their character and relations were minutely described by myself in 1879, in my memoir
on Actinometra* They were shown both in longitudinal and in transverse sections, and
figures were also given illustrating their openings on the under surface of the radial
pentagon, together with the pits corresponding to these openings on the upper surface of
the centro-dorsal in Antedon rosacea. They were clearly distinguished from the five
cavities within the central capsule which were first discovered by Dr. Carpenter.4 He
1 Trans. Linn. Soc. Lond. (Zool.), 1879, ser. 2, vol. ii. pp. 89, 90, 102, 103.
2 See P. H. Carpenter, On some Undescribed Coruatulse from the British Secondary Rocks, Quart. Journ. Geol.
Soc, 1879, vol. xxxvi. p. 51.
3 Trans. Linn. Soc. Lond. (Zool.), 1879, ser. 2, vol. ii. pp. 77, 78.
4 Phil. Trans., 1866, p. 738 ; and On the Structure, Physiology, and Development of Antedon rosaceus, Proc. Roy.
Soc, 1876, vol. xxiv. pp. 218, 219.
REPORT ON THE CRINOIDEA. 17
gave the name "five-chambered organ" or " quinquelocular organ" to the structure
which had been described by Midler as a single-chambered heart. For he found it " to
contain five chambers clustered like the carpels of an orange round a central axis ; " and
he described these chambers as being surrounded by a fibrillar envelope which he
regarded as nervous in character. Marshall ' again spoke of the cavity of the centro-
dorsal as lodging " a sac divided by vertical septa into five radial compartments, and
hence called the chambered organ " ; and he went on to explain how this is " surrounded
by a thick fibrillar investment known as the central capsule." Ludwig had previously
adopted the same terminology,2 and, in fact, he was the first to speak of the " chambered
organ" without the numerical prefix, but he never used this expression to denote
anything else than the five chambers with their central axis inside the central capsule ;
while he further described and figured the radial axial canals,3 the relations of which to
the coeliac canals of the rays and arms were subsequently pointed out by myself.4 Their
connection with the body-cavity and their distinctness from the chambers of the so-called
heart were clearly recognised by Greeff,5 both in his figures and in his descriptions ;
while I am not aware that Teuscher,G the only other recent original writer ou the subject
up to the time of Perrier and Jickeli, ever used the expression "chambered organ" at
all, though he often referred to the " Kammern des Gefasscentrums," and he recognised
the connection of the radial axial canals with the cceliac canals of the rays.
Recently, however, Messrs. Vogt and Yung have figured not only the cavities with-
in the central capsule but also the radial axial canals, and the whole system of spaces
within the calcareous network occupying the centre of the radial pentagon, together with
some accidental cavities within the solid base of the centro-dorsal piece and in the radials
as " cavites dependantes de la cavite generale et constituant, dans leur ensemble, l'organe
dit cloisonne." 7 They say " Ce sont les espaces qu'on est convenu d'appeler, fort
improprement, l'organe cloisonne," and again " C'est la reunion de toutes ces excavations
internes, qui sont revetues de membranes, envoyant de cloisons transversales et dessinant
ainsi un systeme complique de lacunes cloisonnees, qui composent ce que les auteurs ont
appele l'organe cloisonne (Gekammertes Organ). C'est une denomination eminemment
impropre, vu que ce n'est pas un organe, mais une suite de cavites parcourues par l'organe
dorsal avec ses vaisseaux, et formant la continuation de la cavite generale du corps, du
coelome, qui entoure les intestins."8 The statements contained in the first passage
1 On the Nervous System of Antedon rosaceus, Quart. Joum. Micr. Sci., 1S84, vol. xxiv., N.S., p. 510.
2 Beitrage zur Anatomie der Crinoideen, Zeitschr.f. wiss. Zool, 1877, Bd. xxviiL pp. 315-326.
3 Ibid, p. 318.
4 Trans. Linn. Soc. Lond. (Zool.), 1879, ser. 2, vol. ii. p. 78.
6 Ueber das Herz der Crinoideen, Sitszungsb. d. Gesellsch. z. Beford. d. ges. Naturwiss. zu Marburg, 1876, No. 5, p. 93.
6 Beitrage zur Anatomie der EchinoJermen : I. Coniatula mediterranea, Jenaische Zeitschr., 1876, Bd. iii. pp. 244-
260.
7 Traite d' Anatomie comparee pratique, Livr. vii., 1886, p. 550, expl. of fig. 276.
8 Ibid., p. 530.
(zool. CHAX.L. EXP. PART LX. — 1888.) Ooo 3
18 THE VOYAGE OF H.M.S. CHALLENGER.
quoted and in the first paragraph of the second one are inaccurate, to say the least of it.
Messrs. Vogt and Yung do not name the authors who have used the term " chambered
organ" in this " very improper" sense ; but it is certainly neither Dr. Carpenter, Ludwig,
Greeff, Teuscher, Marshall, Jickeli, Perrier, nor myself ; and I know of no other original
writer on Crinoid morphology who has used the expression "chambered organ" at all.
The space represented in the figures to which the Swiss authors refer 1 is the radial portion
of the body-cavity within the calyx, which is clearly distinguished from the chambers
within the central capsule in all the figures given by Ludwig, Greeff, and myself ; and
not one of us has ever regarded this space as a part of the chambered organ, nor, so far as
I know, has any other writer on the subject. But from the mode of reference employed
by the Swiss authors it would apj^ear that Dr. Carpenter had made a great mistake, which
had escaped notice for twenty years until it was rectified by Messrs. Vogt and Yung ;
whereas in reality they are themselves in error, because they give a meaning to his name
which neither he nor any one else ever intended it to bear. The term " (five-) chambered
organ" as employed by him and by every one of his successors until now refers exclusively
to the cavities within the central capsule, which lie on the dorsal side of the rosette and
radial pentagon. But Messrs. Vogt and Yung erroneously interpret it as denoting the
entire system of cavities within the centro-dorsal plate and the ring of radials that rests
upon it; and this is certainly not a definite organ, but a part of the general ccelom, as
stated by the Swiss authors. These facts, however, were perfectly well known both to
Dr. Carpenter and to his successors, and I am entirely at a loss to know who the authors
can be who have used the term "chambered organ" in the "eminently improper" sense
described by Vogt and Yung. The Swiss authors seem to have entirely ignored or
misunderstood the writings of their predecessors, and have attributed to them a mistake
which never was made. But instead of rectifying this supposed mistake they have
converted it into a real one, and have perpetuated it both in their text and in the
explanations of their figures. Thus in fig. 276 the cavities within the central capsule on
the dorsal side of the rosette, and the portion of the body-cavity which is on the ventral
side of this structure and is enclosed by one of its radial processes, are marked alike " c,c,
cavites dependantes de la cavite generale et constituant dans leur ensemble, l'organe dit
cloisonne." No one but Vogt and Yung has used the term ' ' chambered organ" in this sense ;
and as they rightly speak of it as " eminently improper," one cannot but regret that it
should have been employed in a textbook of comparative anatomy for the use of students.
But Messrs. Vogt and Yung go even further than this. The space on the dorsal side
of the central capsule which is marked f in fig. 267 and c in fig. 276, and is described
as one of the cavities of the chambered organ, is nothing; but a rent in the organic basis
of the floor of the centro-dorsal piece. These rents often appear in the skeletal tissues
when very thin sections are cut, and I have been familiar with them for years. But I
1 Op. cit.J, fig. 264 ; c, fig. 276.
REPORT ON THE CRINOIDEA. 19
have many sections through the calyx, both of Antedon rosacea and of other species of
Cornatulae in which there is no trace of them. Three such undamaged sections are
figured in my Actinometra-memoir,1 and I certainly never expected to find an accidental
fracture in the skeletal tissue outside the central capsule described as a part of the
chambered organ, the cavities of which are entirely within this capsule, as explained above.
If there really be such a diverticulum of the body-cavity within the calcareous
substance of the centro-dorsal piece as is described by Vogt and Yung, i.e., between its
inner floor on which the central capsule rests and its external surface, its presence could
easily be demonstrated by rubbing away the outer surface of the centro-dorsal until this
cavity was reached ; and I would commend this method of proving the accuracy of their
anatomical descriptions to the attention of Messrs. Vogt and Yung. They have made a
precisely similar error in their description of the anatomy of the arms, figuring a large
rent in the skeletal tissue of an arm-joint as the " cavite de la syzygie." They will not
find this cavity if they will take the trouble to rub away the syzygial surface of an arm-
joint, which contains but one cavity, that of the axial canal.
Another extraordinary blunder which is committed by these authors in the
explanation of fig. 276 is their description of the fibres (b) which unite the first radials
to the centro-dorsal as the " muscles entre le premier et le second radial." Their mono-
graph contains many other errors of a similar kind, not only in their interpretation of
well-known anatomical facts, as in this last case, which they might have avoided by
consulting the works of their predecessors, but also misrepresentations of passages in
these writings. These, however, are more fitly dealt with elsewhere.2
C. The Rosette.
While the presence of a cirrus-bearing top stem-joint or centro-dorsal piece is common
to all Cornatulae, even including the aberrant Thaumatocrinus, this genus, together with
Atelecrinus (PI. VI. figs. 5, 7) and many fossil species, differs from the adult condition
of all other recent Comatulas in the presence of the basals on the exterior of the calyx.
It was for a long time supposed that the basals of other Crinoids were unrepresented
in recent Comatulse ; but their existence in the Pentacrinoid larva was eventually recog-
nised by Allman, Sir Wyville Thomson, and Dr. Carpenter ; and the last-mentioned
observer discovered the remarkable changes which they undergo during the later part
of Pentacrinoid life. These changes result in their transformation into the "rosette"
which lies close to the dorsal surface of the central funnel within the radials, and covers
in the upper opening of the centro-dorsal cavity that lodges the chambered organ (sensu
stricto). It is well seen in the figures of Antedon eschrichti, Antedon accela, Antedon
1 Trans. Linn. Soc. Lond. (Zool.), 1879, ser. 2, vol. ii. pi. viii. figs. 3, 4, 7.
2 The Morphology of Antedon rosacea, Ami. and Mag. Nat. Hist^ 1887, ser. 5, vol. xix. pp. 19-41.
20 THE VOYAGE OF H.M.S. CHALLENGER.
insequalis, and Antedon breviradia (PL I. fig. 8c ; PI. II. figs. 3c, 5c ; PI. III. figs. 4c,
56), and also in those of Actinometra maculata, Actinometra lineata, and Actinometra
stelligera (PI. V. figs, lc, 2c, 2c, 5d, be). Owing to its homologies with the basals of
other Crinoids, and through these forms with the corresponding plates in other Echino-
derms (e.g., the genitals of Echini), it is a very important structure, apart altogether
from its intimate relation to the great nerve centre lodged within the centro-dorsal plate
and to the axial cords proceeding from it.
One would have thought therefore that some account would have been given of it in
Vogt and Yung's work upon practical comparative anatomy in which Antedon rosacea,
the form studied by Dr. Carpenter, is taken as a type of all Crinoids. It is dismissed,
however, in less than half a dozen lines, and not a word is said of its morphological
relations. In fact the word "basals" does not once occur in the chapter on Crinoidea
in the treatise by Messrs. Vogt and Yung, who pay no attention to the comparative
anatomy of anything but the soft parts as revealed by thin microscopic sections.
Unfortunately, however, this too exclusive reliance upon one method of investigation
has led them into a serious but at the same time a somewhat ludicrous error. In the
figure given by Messrs. Vogt and Yung 1 " pour montrer la disposition du systeme
nerveux central et des organes dorsal et cloisonne " the chambered organ (as originally
described) is covered by a structure marked o. No explanation of this letter is given,
but I learn from Professor Carl Vogt that the missing explanation should be — o, tissu
conjonctif areolaire entourant 1' organ e dorsal et les cavities c de l'organe cloisonne'.
Now this structure which is marked o in Vogt and Yung's fig. 276 is in reality
nothing more or less than a part of the rosette of modified basals, which in the natural
position of the animal roofs in the internal cavity of the centro-dorsal that contains the
chambered organ, as is well shown in PL V. figs. 2c, 2e, and 5c. The relations of this
structure to the soft parts beneath it are entirely ignored by Messrs. Vogt and Yung,
though they were described at length by myself in 1879 2 and again in 1881. In the
latter year I published two sectional views3 showing the position of the rosette with
respect to the chambered organ, and another similar figure and description were given
later on by Marshall.4 But these have been altogether ignored by Messrs. Vogt and Yung,
who have also neglected to work out the point for themselves ; and the consequence is that
a structure which, though small and insignificant in Comatulse, is nevertheless homologous
with the five genital plates of Echini, is figured in a textbook of comparative anatomy as
" areolar connective tissue." In the same figure, too, a portion of the centro-dorsal piece,
which is in immediate contact with the central capsule, is lettered " e, mesentere."
There are many other points in the relations of the Crinoid skeleton which are
1 Op. tit., p. 550, fig. 276. 2 Trans. Linn. Soc. Lond. (ZooL), 1879, ser. 2, p. 78.
3 The Minute Anatomy of the Brachiate Eehinoderms, Quart. Journ. Micr. Sci., 1881, vol. xxi., N.S.,p. 186, pi. xii.
figs. 14, 15.
1 Lot. cit., pp. 508, 511, pi. xxv. fig. 1. See also Ludwig, lot. tit., Taf. xix. fig. 74.
REPORT ON THE CRINOIDEA. 21
altogether misunderstood by Vogt and Yung, who have apparently attempted to work
out the anatomy of the type by one method only, that of thin sections, and have almost
completely ignored its osteology. Had they devoted a little more attention to the
characters of a prepared skeleton of Comatula they would have avoided not a few errors
which are calculated to give the student an altogether erroneous conception, not only of
Crinoid morphology, but of that of Echinoderms in general. The basal plates are among
the earliest calcareous structures which appear in the larva of any Echinoderm, and their
relation to the great nerve centre of a Crinoid renders them additionally important
morphologically. But no student of Messrs. Vogt and Yung would ever learn of their
existence at all.
The gradual development of the rosette out of the original basal plates of the
Pentacrinoid larva was fully described by Dr. Carpenter,1 who showed that it is
" essentially formed at the expense of the secondary or ventral layer of the original
basals, the ends of the curved rays being the sole residue of their primary or dorsal
layer." Alternating with these spout-like processes, which are radial in position, are five
others of a more triangular form, which occupy a somewhat deeper situation within the
radial pentagon. The apex of each of them is attached to a suture between two con-
tiguous radials, just between the two adjacent apertures of their central canals. Each of
these canals receives a branch of the primary basal cord proceeding from the central
capsule, that lies on the dorsal side of the interradial process of the rosette ; and when
the rosette is in its natural position in the calyx, an opening for the passage of one of
these secondary basal cords is visible between every two of the processes of the rosette.
This is well seen in Antedon eschrichti (PI. I. fig. 8c). The example of this species
which is here represented, has a comparatively simple rosette, which is almost entirely
free from any trace of the accessory structures to which I have given the general name of
the "basal star," such for instance as is represented in figs. 1-5, c on PI. II. In all these
forms, and more especially in Antedon angusticalyx and Antedon insequalis (figs. 4c, 5c),
a larger proportion of the embryonic basal has been left unabsorbed than is usually the
case in . the European and Arctic Comatulae ; but the peripheral margins of each plate
remain, and form, by their union with the corresponding parts of the adjacent plates, the
structure which I have called the basal bridge. This is united to each radial along the
inner margin of its dorsal face, and partially covers in the two secondary basal cords
which are converging; on its single axial canal. It is well shown in Acti?iometra
maculata and Actinometra stelligera (PI. V. figs, lc, 5d) and also in the rosette of the
latter species disconnected from the radials as seen in fig. 5c ; and it appears, so far as I
am aware, to be of pretty constant occurrence in this genus, though absent or at any rate
undistinguishable in some species of Antedon (PI. III. fig. 6b).
United with each angle of the pentagon formed by the five basal bridges is one of the
1 Phil. Trans., 1866, p. 745.
22 THE VOYAGE OF H.M.S. CHALLENGER.
rays of the basal star, the relations of which are described at length in my memoir on
Actinometra.1 At the time this was written (1877) I had only been able to dissect the
calyx in a comparatively small number of Comatulse ; but a detailed examination of the
large amount of duplicate material obtained by the Challenger has shown that a basal
star is nearly always present both in Antedon and in Actinometra, so that species like
Anteclon tenella, Antedon hageni, Antedon phalangium, and Antedon rosacea, in which
it is not developed, are the exception rather than the rule ; while there may be traces of
it in some varieties of Antedon eschrichti, though not in others.
It occasionally happens that the rays of the basal star, or, more shortly, "the basal
rays," appear on the exterior of the calyx between the centro-dorsal and the first radials.
But there is no constancy about this character, even in individual species. Thus, for
example, a basal ray is visible in the dissected calyx of Antedon antarctica shown on
PI. I. fig. 6a, but there is no trace of it in either of the three specimens figured on
PL XXV. figs. 10-12. The basal rays sometimes appear externally beneath the alternate
radials of the ten-rayed Promachocrinus (PI. I. figs, la, lc), but this is not always the case.
I have also seen them in some individuals of Antedon carinata, though not in that shown
on PI. III. fig. la. They are generally to be seen in Antedon macronema (PI. IV. fig. 3a;
PI. XXXVIII. fig. 5), in Antedon longicirra (PL XVII.), and also in Actinometra
pulchella (PL IV. fig. 5c), and Actinometra stelligera (PL V. fig. 5b) ; while there are
other species, such as Actinometra rnaculata and Actinometra lineata, in which they are
only occasionally visible.
I have shown elsewhere that the basal rays have an entirely different origin from
either the primary or the secondary portions of the rosette. They are tertiary structures
formed by calcification in the synostosis between centro-dorsal and radials. Sometimes,
however, they are very substantial structures, and each of them becomes so firmly united
with an interradial portion of the rosette that it is often possible to get the entire complex
structure thus formed to break up into five separate parts, each representing one basal
plate. The results of this operation are seen in Antedon antarctica (PL I. fig. 7),
Antedon carinata (PL III. figs, lc, 2a, 2b, 3a, 3b), Actinometra nieridionalis, Actinometra
pulchella, and Actinometra paucicirra (PL IV. figs. 4b, 5a, 6b). Each of the compound
basals so isolated is a somewhat elaborate structure. The basal ray may be long and
narrow as in Actinometra nieridionalis (PL IV. fig. 4b), or short and stout as in Antedon
antarctica (PL I. fig. 7) and Actinometra paucicirra (PL IV. fig. 6b.)
At the proximal end of the basal ray are two openings, one on each side, which give
passage to the secondary basal cords ; and they are separated, when seen from the dorsal
side, by the interradial process of the rosette with portions of the basal bridge (PL IV.
figs. 4b, 6b, yS). The lateral boundaries of these openings are formed by the halves of two
of the radial spouts of the rosette which extend outwards from the base of the interradial
1 Trans. Linn. Soc. Lond. (Zool.), 1879, ser. 2, vol. ii. pp. 95-100.
REPORT ON THE CRINOIDEA. 23
process and represent the unabsorbed lateral portions of the primary layer forming
the embryonic basal plate. The ventral side of the basal ray in the three species
of Actinometra which are figured on PL IV. figs. 5a, 46, Gb, a, is marked by a
relatively large depression which forms the central end of the axial interradial canal.
This descends into the calyx over the apposed lateral edges of two radials, as is well
seen in PI. III. figs. Id, 7c, and PL V. fig. 2c. But in most cases it ends blindly
without reaching the dorsal surface of the radial pentagon at alL
D. The Radials.
The radials of Comatulse differ considerably from those of the Pentacrinidae, the
family of Stalked Crinoids to which the free forms are most closely allied. In Pentacrinus,
as in the Pentacrinoid larva of Antedon (PL XIV. figs. 2-9), the first radials appear
above the basals on the exterior of the calyx as relatively large convex plates. They
retain this character in Thaamatocrinus and to a less degree in Atelecrinus (PL VI. figs.
5, 7). both of which are permanent larval forms in other respects. But in the two
large genera Antedon and Actinometra there is a very considerable amount of variation
in the extent to which the first radials appear externally. Some forms, such as Antedon
eschrichti, show no indication of them at all (PL XXIV. fig. 11), or only traces of their
angles in the interradial portions of the calyx (PL I. fig. 8a) ; while in other cases, such
as Antedon elegans, Antedon longicirra (Pis. VIII. , XVII.), and Antedon macronema
(PL IV. fig. 3a; PL XXXVIII. fig. 5), they exhibit a relatively large outer surface between
the edge of the centro-dorsal and the second radials. Between these two extremes every
intermediate gradation may be traced. The former is due to the gradual enlargement of
the centro-dorsal, which spreads itself over the base of the calyx towards the end of
Pentacrinoid life and sometimes conceals the first radials altogether, as described by Dr.
Carpenter1 in Antedon rosacea. The second radials thus appear to spring directly from
the centro-dorsal (PL XIII. fig. 2); and this has sometimes led to species of Comatidse
being described as having only two radials. In fact d'Orbigny2 described a new genus
Comatulina, for a fossil species in which there are no external basals and " les bras
s'articulent immediatement sans intermediates a la piece centrale pourvue de ramules."
The full-grown Antedon accela presents this appearance, but the younger specimen
figured on the same plate (PL XVI.) shows comparatively large first radials; while a
more mature individual (PL II. figs. 3a, 3c) shows relatively less of them, owing to the
spread of the centro-dorsal. The figures of Antedon plialangivm on PL XXVIII.
show similar differences of growth, though in a less degree.
The various species of Actinometra exhibit among themselves essentially the same
1 Phil. Trans., 1866, p. 742.
2 Cours elementaire de Paleontologie et de Geologie Stratigraphique, 1850-52, vol. ii. p. 139.
24 THE VOYAGE OF H.M.S. CHALLENGER.
differences as do those of Antedon in the extent to which the radials appear on the out-
side of the calyx. In Actinometra maculata, Actinometra lineata, and Actinometra
stelligera the oentro-dorsal is so large that it actually supports the proximal ends of the
second radials, and nothing but the angles of the first radials can be seen externally
(PI. V. figs. 1, 2, 5, a, 6). Small portions of their sides can be seen in Actinometra Solaris
(PL V. figs. 4, 6, c) ; while in Actinometra paucicirra and in all the Phanogenia-\ike
forms the centro-dorsal only occupies a comparatively small space in the centre of the
radial pentagon, a considerable portion of which appears externally as seen in PL V. fig. 3c,
and in Pis. LIV, LXV.
Not only the centro-dorsal, but also the radials of Actinometra present very
considerable differences from the corresponding parts of the Antedon-c&lyx, though
these differences are less distinct in the fossil than in the recent forms of both genera.
The outer or articular faces of the radials in Antedon are always much inclined to
the vertical axis of the calyx, and are usually much wider at their dorsal than at their
ventral ends, so that their outline is trapezoidal (PL I. figs. 6a, 8a ; PL II. figs. 1-5, a ;
PL III. figs, la, 46, 5a, 6a7; PL IV. figs. 2a, 3a). Antedon carinata and Antedon
macronema are, however, somewhat exceptional in this respect, the width of their
articular faces being very much more uniform ; and they further differ from most species
of Antedon and resemble Actinometra (PL V. figs. 1-5, o) in the relatively great diameter
of the central funnel of the calyx (PL III. fig. id ; PL IV. fig. 3b). For as a general
rule the opening of the central funnel which is bounded by the upper edges of the radials
is very narrow, their ventral surfaces being quite small and having a steep inward slope.
Hence when the calyx is viewed from above the greater part or even the whole of these
inclined external faces is visible, always down to the opening of the central canal in the
transverse articular ridge (PL II. fig. Ad ; PL III. fig. 6c); while sometimes even the fossae
for the attachment of the dorsal ligament are visible in a superior view (PL I. figs. 6, 8, b;
PL II. figs. 1-3, 5, d; PL III. fig. 4a). This last is especially the case in Antedon macronema
(PL IV. fig. 3b), though the reverse is true of Antedon carinata (PL III. fig. id).
Most species of Antedon have large muscle-plates, which greatly increase the height of
the distal faces of the radials (PL I. figs. 6, 8, a ; PL II. figs. 1-5, a ; PL III. figs. 46, 5a,
6c/). They are fairly large in Antedon carinata (PL III. fig. la), but in Antedon
macronema they are quite small and linear and barely distinguishable in side view from the
pair of fossae immediately below them, though they are seen rnoi*e clearly when viewed from
above (PL IV. figs. 3o, b). This lower pair of fossae was described by Dr. Carpenter 1 as
serving for the attachment of the interarticular ligaments. The ridges which generally
separate them from the upper fossae mostly start from the raised rim round the opening
of the central canal and run more or less obliquely outwards to meet the sides of the
radials (PL II. figs. 2-5, a ; PL III. figs. 46, 5a, 6d). In Antedon antarctica, however,
1 Phil. Trans., 1866, p. 714.
REPORT ON THE CRINOIDEA. 25
they are almost perfectly horizontal (PI. I. fig. 6a), and so give the calyx a very different
appearance from that of the closely allied Arctic species Antedon eschrichti (PL I. fig. 8a) ;
though Promachocrinus kerguelensis, another southern form, resembles Antedon eschrichti
in this respect (PI. I. fig. la), and the same may be said of Eudiocrinus semperi (PI. III.
fig. 7a). In some forms of Antedon incisa the ridges do not, as is usually the case, start
from the rim of the central canal, but curve upwards slightly from the median vertical
ridge of the articular face, and the upper pair of fossae are therefore somewhat restricted
(PI. II. fig. la). In Antedon disciformis (PL IV. fig. 2a) the ridges run upwards from
the central canal for some little distance and then curve outwards, leaving a sort of
furrow between them, the bottom of which is sometimes slightly raised.1 But in most
species of Antedon there is a strong median ridge running down from the ventral edge of
the articular face towards the opening of the central canal (PL I. figs. 6a, 8a ; PL II.
figs. 1-5, a ; PL III. figs. 46, 5a, 6d). This is hardly traceable in Antedon carinata
(PL III. fis;. la), which rather resembles Antedon disciformis in having a tendency to the
intermuscular furrow that is so characteristic of Actinometra (PL V. figs. 1-5, b) ; while
in Antedon macronema the muscular fossae are so very slight, that the notch between
them reaches down to the upper margin of the raised rim of the central canal (PL IV.
figs. 3a, b), a character which rarely occurs in Actinometra.
Thus then the radials of Antedon carinata and Antedon macronema differ from
those of other species of Antedon and approach those of Actinometra. There is much
less difference than usual between the widths of the upper and lower ends of the distal
faces, which are comparatively low, so that their long axes are horizontal and not vertical
as is usually the case (PL I. figs. 6a, 8a). The centre of the upper surface is consequently
occupied by a wide funnel, the walls of which are formed by the ventral surfaces of the
radials (PL III. fig- 1^ ; PL IV. fig. 3b). Antedon carinata has fahiy large muscle-
plates ; but they are quite small in Antedon macronema, and the ridges separating the
muscle- and ligament-fossse are so slightly oblique as to be almost horizontal, though their
orio-iu from the prominent and large rim of the central canal is very marked. In each
case, however, the general appearance of the calyx is much more that of the Antedon
than of the Actinometra type. The calyx of Antedon macronema further presents many
resemblances to that of Pentacrinus, especially in the small size of the articular faces
and in the large portions of the radials which appear externally (PL IV. fig. 3a). Of all
recent Comatulse it is the one which most closely approaches the general type of the
Jurassic forms of Antedon; and as it is only known to occur on the Australian Coast,
this is a point of considerable interest.
The radials of Actinometra differ very largely from those of the typical Antedon.
Their distal faces are relatively low, and lie nearly or quite parallel to the vertical axis of
the calyx, while there is but little difference in width between their ujjper and their lower
1 This is less distinctly seen in an interradial view of the calyx than in a face view of a single radial.
(ZOOL. CHALL. EXP. — PART LX. 1888.) OoO 4
26 THE VOYAGE OF H.M.S. CHALLENGER.
ends. The vertical position of the articular faces is well seen in some forms of Actino-
metra lineata, which has an extremely "wall-sided" calyx (PI. V. fig. 2e) ; while in
Actinometra paucicirra their lower portions actually slope inwards as seen in PI. V.
fig. 3c. The ventral faces of the radials, which in Antedon have a steep inward slope
(PI. I. fig. 86), are almost horizontal in Actinometra, sloping very gently inwards towards
the central space. Hence the opening of the funnel becomes widely expanded, and
when the radial pentagon is viewed from above little or nothing is seen besides the proper
ventral faces of its component radials. All the species of Actinometra which I have
examined have smaller muscle-plates than those of any Antedon except Antedon
macronema (PI. IV. figs. 3a, b), so that the distal faces of the radials are very low and
the muscular fossae often quite inconspicuous (PL IV. figs. 4a, 5c ; PI. V. figs. 1-5, b, 5c).
They are separated from the lower pair of fossae by fairly prominent ridges which are
either horizontal or curved slightly upwards. These start from the sides of the radial,
run inwards towards the middle line, and then turn downwards so as to leave between
them a wide furrow, which gradually dies away below with the disappearance of its
bounding ridges. No recent Actinometra has the distinct rim on the ventral side of the
opening of the central canal that exists in every Antedon, even in Antedon carinata
(PI. III. fig. la) and in Antedon macronema (PL IV. fig. 3a), perhaps the nearest
approach to it being in Actinometra meridionalis and Actinometra pulchella (PL IV.
figs. 4a, 5c), where the lower edges of the ridges bounding the intermuscular furrow are
somewhat thicker than usual.
These differences in the structure of the calyx in the two chief genera of Comatulse
are of considerable importance. For it is only by means of an acquaintance with them
that the generic determination of the fossil Comatulae becomes at all possible. Every one
hitherto found in the Tertiary strata and in the Chalk, of which the entire calyx is known,
is an unmistakeable Antedon, both in the characters of the centro-dorsal and in those of the
radials. Antedon sequimarginata from the Gault is as clearly an Antedon as Actinometra
loveni from the same formation is an Actinometra. But some of the Neocomian and
many of the Jurassic Comatulae are less easily identified. The wide and low radials with
marked intermuscular furrows of Actinometra cheltonensis from the Inferior Oolite, and of
Actinometra ivurtembergica from the Corallian of Nattheim, indicate the generic position
of these types pretty clearly ; while Antedon scrobiculata with its high articular faces,
much narrower above than below, is an undoubted Antedon. But on the other hand,
the low and wide radials and thin centro-dorsal of Antedon picteti and Antedon
infracretacea are very suggestive of Actinometra; though in both types the articular
faces of the radials have a considerable slope and are altogether much like the corre-
sponding parts of other species which are unquestionably referred to Antedon. For the
present, therefore, the systematic position of these and of other somewhat generalised
types of early Comatulas must remain in doubt.
REPORT ON THE CRINOIDEA. 27
E. Some Abnormal Conditions of the Bays of Comatul^.
In the two large genera Antcdon and Actinometra, just as in Apiocrinus, Pentacrinus,
and Encrinus, there are normally five rays which divide upon the third joint above the
basals, i.e., the third radial is the axillary (Pis. VIII., LXVII.) ; and this is the general
rule among the Neocrinoids. The exceptions are Metacrinus and Plicatocrinus, the
former with four or six radials (primitively five or eight, as some of them are syzygial
joints), and the latter with only two as in one or two fossil Comatulse. It sometimes
happens, however, that an additional radial is inserted into the normally three-jointed
series, as for example in the Pentacrinus millleri mentioned in Part I.,1 and I have met
with a nearly similar case in Antedon alternate/, (PL XXXII. fig. 5); while Wagner2 has
noticed the same monstrosity in Encrinus gracilis. But the two outer radials of Antedon
alternata remain separate and are not united by syzygy, as in the Pentacrinus millleri
just mentioned. On the other hand, in one example of Antedon remota (PI. XXIX.
fig. 6) and in the only specimen of Antedon incerta (PI. XVIII. fig. 4) and in one of
Actinometra parvicirra (PI. LXI. fig. 1) the second radial is missing in one ray and
the axillary rests directly against the first radial as in Plicatocrinus and in many Palseo-
crinoids.
Another and more common variation is in the number of the rays themselves.
Excepting, of course, in Promachocrinus there are normally five rays in all Comatulidae ;
but forms with four and six rays are occasionally met with. I have a tetraradiate
specimen of Antedon rosacea, and one of a Japanese Antedon in Dr. Doderlein's
collection, and also one of Actinometra paucicirra from Cape York. In all these three
individuals the anterior ray (A) is missing, so that the mouth, instead of being radial
in position, is placed interradially between the rays E and B.
On the other hand the " Blake" collection contains a six-rayed form of Actinometra
pulchella. The disc is unfortunately concealed, so that the symmetry of the ambulacra
cannot be made out. But I am rather inclined to think from the appearance of the
centro-dorsal that it has the usual pentamerous symmetry, one of the radials being rather
larger than its fellows and also axillary, so that it bears two small rays, as sometimes
happens in Allagecrinus.3 Another variation characteristic of this genus occurs in
Actinometra midtibrachiata (PL LVI. fig. 3), one of the radials being considerably smaller
than the other four.
The only other six-rayed Comatula that I know is a small and dry Antedon in the
1 Zool. Chall. Exp., part xxxii. p. 311, pi. xv. fig. 2.
2 Die Encriniten des unteren Wellenkalkes von Jena, Jenaische Zdtschr., 1886, Bd. xx. (N.F. xiii.), p. 20, Taf. ii.
fig. 13.
3 See Carpenter and Etheridge, Contributions to the Study of the British Paleozoic Crinoids, — No. I. On Allagecrinus,
the Representative of a New Family from the Carboniferous Limestone Series of Scotland, Ann. and Mag. Nat. Hint.,
1881, ser. 5, vol. vii. pp. 288, 292.
28 THE VOYAGE OF H.M.S. CHALLENGER.
British Museum. But the disc is sufficiently well preserved to show that the additional
ray is inserted between the two of the right side (D and E).
The facts above mentioned may be usefully compared with similar variations which
have been noticed in other Echinoderms. In the only six-rayed Blastoid that I have
seen1 there are but five ambulacra, though a pseudo-radial plate without a sinus is
intercalated between radials C and D, so that the dorsal surface of the calyx is very
regularly hexagonal.
On the other hand Blastoids with only four ambulacra are more common ; but the
dorsal part of the calyx is more or less distinctly pentagonal, the fifth radial not being
incised for an ambulacrum. The two postero-lateral and the right antero-lateral one
(C, D, E) are the rays in which this modification has been noticed, C showing it twice
and the other two once each.
Two tetraradiate examples of Encrinus liliiformis have recently been observed by
von Koenen ;2 but it is curious that variations from the normal pentamerous symmetry
are rare among the Pelmatozoa, except in the genus Rhizocrinus. Four- and six-rayed
Urchins are not uncommon; while Ludwig3 found half a dozen six-rayed individuals of
Cucumaria planci in a collection of one hundred and fifty. In all cases the sixth ray
was intercalated between the two forming the bivium, a fact which may be compared
with the absence of the middle ray of the trivium in the three Comatulaa with abnormally
interradial mouths mentioned above.
1 See Etheridge and Carpenter, Catalogue of the Blastoidea in the Geological Department of the British Museum
(Natural History), London, 1886, pp. 40, 41.
2 Beitrag zur Kenntniss der Crino'iden des Muscheikalks, Abhandl. d. k. Gesellsch. d. IViss. Gottingen, 1887, Bd. xxxiv.
p. 23 (of separate copy).
3 Ueber Sechsstrahlige Holothurien, Zool. Anzeiger, 1886, Jahrg. ix. p. 476.
REPORT ON THE CRINOIDEA. 29
III.— THE GEOGRAPHICAL AND BATHYMETRICAL DISTRIBUTION OF
THE COMATULSE.
Our knowledge of the existing species of the Comatulse is at present so imperfect
that it affords but a slight foundation for any generalisation respecting their geographical
distribution and the origin of specific types. For they occur in the most extraordinary
abundance over certain large areas, such as the Caribbean Sea, and more especially the
Eastern Archipelago and Australasia. Every large collection that I have examined, and
they are many, contains a number of forms from the latter district, the specific relations
of which will require months of detailed work before they can be properly elucidated.
Nearly all of these are littoral species, and it is chiefly with regard to them that any
generalisation would be premature at present. But the dredgings of the Challenger have
accumulated a large mass of information concerning the Comatulse of other seas than those
of Australasia. This relates more especially to the Comatula-fawaa of the continental and
abyssal regions, about which we cannot expect to gain very much additional knowledge
in future. The Comatulse of the Arctic and Sub-Arctic seas are also pretty completely
known ; while the Strait of Magellan and the Southern Indian Ocean between Marion
Island and Melbourne have yielded some dozen species for comparison with those of the
northern circumpolar fauna.
The following conclusions, then, embody the condition of such knowledge of the
Comatulse as I have been able to gain from the study of the Challenger collection and
preliminary work upon the material dredged by the U.S. Coast Survey steamer "Blake";
together with my notes upon the Comatulse in the museums of London, Paris, Berlin,
Vienna, Copenhagen, Lund, Stockholm, Amsterdam, Leyden, Hamburg, Dresden, Kiel,
Munich, Stuttgart, and also upon the collections made by Professor Semper in the
Philippines, Dr. Doderlein in Japan, Dr. Anderson in the Mergui Archipelago, and Dr.
Hickson in North Celebes.
The Comatuke range in latitude from 81° 41' N. to 52° 5' S., being represented in
each locality by a ten-armed Antedon, a point which will be considered later.
Although abundant near the coasts in the Arctic Ocean and on both sides of the
North Atlantic, no Comatulse have been dredged there at a greater depth than 800 fathoms,
nor were any met with in either of the Challenger's two traverses of the North Atlantic;
while, though one species has been obtained at the Canaries and Madeira, there is no record
of any from the Azores, Bermuda, or the Cape Verde Islands. The two Mediterranean
species range as far north as Scotland, but I do not know of their passing the meridian
of 20° E., either in the Mediterranean or in the Baltic. In the Florida Channel and in
the Caribbean Sea, however, Comatulse have been dredged in abundance. But none arc
known from the African Coast between Cape Verde (Goree) and the Cape of Good Hope,
30 THE VOYAGE OF H.M.S. CHALLENGER.
except for one species of Anted 'on at the equatorial island Rolas. The only Actinometra
common to both sides of the Atlantic occurs at St. Paul's Rocks, and a few Caribbean
species both of this genus and of Antedon are common along the South American coast
as far south as Cape Frio (lat. 23° 1' S.); while in mid-Atlantic Antedon was dredged
in moderate depths near Tristan da Cunha and Ascension respectively.
Closely allied to the North Atlantic species are those occurring at Kerguelen and
Heard Island, together with a couple of forms inhabiting the Strait of Magellan. This
Southern Ocean has also yielded Promachocrinus. the unique Tliaumatocrinus, and at 2600
fathoms a minute Antedon which was also found at 2900 fathoms in the North Pacific.
Various Comatulae have been obtained at Simon's Bay, Natal, Madagascar, Zanzibar,
Mauritius, St. Helena, Rodriguez, the Red Sea, the Seychelles and Ceylon, with a
solitary species at Kurrachee, and in the Bay of Bengal. It is curious, however, that
none were found by Mr G. C. Bourne on the coral reefs of the Chagos group. But the
region in which Comatulae are most abundant is the great Eastern Archipelago, which
may be roughly described as a triangular area reaching 100° from east to west and 65°
from north to south, with its angles at Ceylon, Japan, and the Kermadec Islands.
Within this large area, which includes the Challenger Stations 170 to 236, Comatulae occur
in the most bewildering profusion. But, so far as I know, not one has been found on the
coasts of New Zealand, although Eudiocrinus and a ten-armed Antedon were obtained by
the Challenger at Station 169, within a comparatively small distance of the East Cape of
the North Island. The Challenger's dredgings between the Admiralty Islands and Japan
were among the deepest of the whole cruise, ranging between 1100 and 4475 fathoms,
and no Comatulae were met with between the equator and lat. 35° N. Three species were
obtained on the green mud off the Japanese coast between 345 and 775 fathoms, and one
in 2900 fathoms at Station 244 in the North Pacific. This form, Antedon abyssicola,
is the deepest Comatida known.
From Station 244 until the Straits of Magellan were entered, the dredgings of the
Challenger yielded no Comatida at all, a fact which is the more interesting because
almost the same statement holds good for the Ophiurids.1
Single species of Antedon are known from the Sandwich Islands and Chile, and of
Actinometra from Tahiti and Peru ; but except for these and for the two in the Strait
of Magellan, I know of no Comatida in the Pacific east of long. 150° E., not even on the
western shores of North America. Antedon rhomboidea and Antedon magellanica, if they
can be called Pacific species at all, are the only ones in that ocean south of lat. 40° S.
None occur in New Zealand nor in Tasmanian waters. These two Magellan species are
therefore somewhat isolated, as on entering the Atlantic the Challenger dredged no
Comatulae until reaching Station 320, in 600 fathoms, where three ten-armed species of
Antedon were obtained. The Falkland Islands, however, seem to have yielded nothing.
1 Zool. Chall. Exp., 1882, part xiv. p. 309.
REPORT ON THE CRINOIDEA. 31
The distribution of the two leading Comatula genera, Antedon and Actinometra,
cannot yet be fully worked out, owing to the large number of species which are still
undescribed ; but that of the other generic types is easily stated.
The archaic Thaumatocrinus has only been found at 1800 fathoms at Station 158
in the Southern Ocean, where it was associated with Promachoerinus abyssorum, which
also occurred at Station 147 (1G00 fathoms), together with three species of Antedon.
Another species of Promachoerinus is common at Kerguehm, and a third was obtained at
500 fathoms off the Meangis Islands. Three species of Atelecrinus are known, two from
the Atlantic and one from the Pacific. The unique specimen of the latter was found at
Station 174c in the South, Pacific, at 610 fathoms ; while one of the Atlantic species
is only known from Pourtales' dredgings in the Gulf Stream off Havana (450 fathoms).
The other, found by the Challenger in 350 fathoms at Station 122 off Pernambuco, was
subsequently met with by the " Blake " off Nevis, St. Lucia, and Granada, at depths
of 291 to 422 fathoms.
Eudiocrinus, first obtained in quite shallow water among the Philippines by Semper,
was dredged by the Challenger both in the North and in the South Pacific, at depths
varying from 565 to 1050 fathoms; while the " Travailleur " found Eudiocrinus
atlanticus at 896 metres in the Bay of Biscay.
In discussing the distribution of Antedon and Actinometra, the two principal genera
of Comatulse, it must be remembered that each of them, but especially Antedon, contains
a very large number of species, and they should be considered for this purpose to
represent subfamilies rather than genera. Thus, for example, the name Antedon is now
given to all recent endocyclic Comatulse with the basals metamorphosed into a rosette,
and five rays bearing ten or more arms, just in the same way as the name Echinus was
originally used for a variety of regular Urchins, which have now received different generic
names. The difference between the tiny ten-armed Antedon abyssicola inhabiting depths
of three miles and upwards in the Pacific (PI. XXXIII. figs. 1, 2), and the littoral Antedon
clegans, Antedon multiradiata, or Antedon regalis (Pis. VIIL, IX., XLVL), is no doubt
very considerable at first sight ; but there are so many intermediate links between the
simple and the complex forms, that no hard and fast generic lines can be drawn. At the
same time, a glance at the tabular keys to the species which are given in the following
pages will show that they fall into certain very well defined groups ; and the range of
each of these groups, both in depth and in space, may be profitably studied.
In the first place, all the species of Antedon which have the two outer radials united
by syzygy are limited to quite shallow water in the En stern Archipelago. They are
comparatively few in number, and have perhaps the most restricted geographical range of
any of the specific groups. On the other hand, the Antedon species of the simple ten-
armed type like Antedon rosacea, are most remarkably abundant, and also extremely
varied in their character, — Antedon abyssicola and Antedon tuberosa, or Antedon
32 THE VOYAGE OF H.M.S. CHALLENGER.
carinata, presenting several very striking points of difference (PI. XXIII. fig. 2 ; PL
XXXIII. figs. 1,2; PI. XXXIV.). They fall into several sets, each of which represents
a different type of Comat u/a-structure, and in several cases the distribution of these
sets is fairly well defined.
The ten-armed species of Antedon have a wider range both in depth and in space than
any other types of the genus. This is of course only to be expected ; for they represent
a somewhat early stage in the development of the Pentacrinoid larva, the radial axillaries
and the pairs of first brachials which they bear appearing soon after the opening of
the tentacular vestibule, when the whole number of tentacles does not exceed twenty-
five.
These ten-armed forms are the only species of Antedon which occur outside the
fortieth parallels of latitude, and at greater depths than 750 fathoms. There is one
possible exception to this last statement. Some examples of Antedon insequalis with
three distichals reached me, together with fragments of Pentacrinus naresianus and the
label of Station 175 (1350 fathoms). But there is no record in the Station Book of their
occurrence here, though two Comatulae are mentioned. But these [Antedon breviradia and
Antedon acutiradia, PI. XI. figs. 3, 5) have the general facies of deep-water forms ; and this
is not the case with Antedon insequalis and the arms of Antedon basicurva, which are
labelled as coming from this station. It may then, I think, be safely assumed that the only
Cornatulae dredged at Station 175 were the ten-armed Antedon acutiradia and Antedon
breviradia, the multibrachiate Antedon insequalis not really occurring at that station.
Disregarding this form, we find that out of twenty-nine stations where Antedon was
dredged by the Challenger, "Porcupine," and other British expeditions, at depths exceeding
200 fathoms, twenty-eight yielded ten-armed species. Multibrachiate species occurred at
six of these, and at one other station, this (Station 135g) being the only locality below
200 fathoms where the genus Antedon occurred, but was not represented by any ten-armed
form. Eleven of the twelve " Porcupine " stations ' and two of the seventeen Challenger
ones were beyond the parallels of 40°. But the remaining " Porcupine " station and six
of the fifteen Challenger ones within these limits yielded multibrachiate forms, though
never at a greater depth than 750 fathoms. The " Porcupine " species, however, Antedon
lusitanica, is a curious one. It is dimorphic, some individuals having ten arms only,
and some having one or more distichal series (PI. XXXIX. figs. 1, 3).
The nine dredgings of the Challenger at which Antedon occurred at depths between
700 and 2900 fathoms inclusive, yielded nine species of the genus, all of them small and
ten-armed, and half of them belonging to the group which contains the familiar Antedon
rosacea and Antedon tenella. Four of the fifteen dredgings between the fortieth parallels
at depths exceeding 200 fathoms were at 1000 fathoms and upwards, and they yielded
1 Under this general name I include all the dredgings of the " Porcupine," " Lightning," " Knight-Errant," and
"Triton."
REPORT ON THE CRINOIDEA. 33
four species of Antedon, three of which were each found at two or more different stations.1
Thus Antedon abyssicola, from 2900 fathoms (Station 244) in the North Pacific, also
occurs at 2600 fathoms (Station 160) in the Southern Sea; and the remaining abyssal
station south of lat. 40° S. (Station 147, 1600 fathoms), yielded three different species
of the ten-armed Antedon-type. The species dredged at Station 135e in 1000 fathoms
was only represented by Pentacrinoid larvae, but of the eight remaining abyssal forms
(found below 700 fathoms), one that occurred at four stations in the Pacific is closely
allied to Antedon tenella, which ranges down to 740 fathoms in the North Atlantic,
between 30° N. and 75° N.; while three others belong to the same group as this species
and Antedon rosacea, which ranges in shallow water from the Fseroe Banks to the
Canary Islands, and possibly even to the equator.
In like manner, the Magellan and Heard Island species from the furthest south are
the Antarctic representatives of Antedon eschrichti and Antedon quadrata, which are
widely distributed in the Arctic Ocean. In fact, the group to which these forms belong
has the greatest geographical range of any set of the ten-armed Antedon-type,.
Antedon eschrichti (PI. XXIV. fig. 11) and its close ally Antedon quadrata
(PI. XXVI. figs. 2, 3), are common in the Arctic Ocean between the meridians of 80° W.
and 70° E. They were found by the " Porcupine " in the Fseroe Channel, and by the
Challenger off Halifax, which is their furthest southern range (lat. 43° N.). No other
Comatulae but the dimorphic Antedon lusitanica were found in the North Atlantic below
650 fathoms, but this form does not at all approach the Eschrichti-gvowp. The Straits
of Magellan, however, contain two species belonging to it ; while Antedon australis, and
Antedon antarctica from the neighbourhood of Heard Island are also very closely
allied to, though not identical with Antedon quadrata and Antedon eschrichti, and
are the southernmost Comatuhe known (PI. XXV.; PI. XXVI. fig. 4). None of these
species, however, nor in fact any of the Eschrichti- group, extend down to any greater
depth than 650 fathoms; but some of the Comatulse from depths below this belong, as
we have seen, rather to the North Atlantic than to the Arctic fauna. Certain of them,
however, find their places in the group of ten-armed species which have the sides of the
rays flattened and more or less closely approximated. One of them {Antedon bispinosa,
PI. XX. fig. 3) was obtained at Station 147, together with two species of the Tenella-
group, and two others {Antedon acutiradia and Antedon breviradia, PI. XL figs. 3, 5)
were the only two dredged with certainty at Station 175.'2 All these three occurred
below 1300 fathoms.
With the exception of Antedon bisjnnosa from the Southern Sea and Antedon
lusitanica and Antedon multispina of the Atlantic, all the twenty forms with laterally
compressed rays {Basicurva-growp) inhabit the Western Pacific and Australasia; and only
1 The Antedon breviradia and Antedon alternata occurred both at 630 and 1070 and at 1350 fathoms respectively.
■ See ante, p. 32.
(ZOOL. CHALL. EXP. — rABT LX. — 1888.) Ooo 5
34 THE VOYAGE OF H.M.S. CHALLENGER.
one (Antedon denticulate), from 49 fathoms at Station 190 in the Arafura Sea, can be
called a littoral species. The remainder all belong to the continental or to the abyssal
zone. Most of them have covering plates and generally also side plates to the ambulacra ;
and the two ten-armed forms of Antedon from the Challenger dredgings which have
plated ambulacra but the rays not flattened laterally (Accela-growp) are even more
restricted in their distribution. One was found in 140 fathoms at Station 192 in the
Arafura Sea, and the other in 500 fathoms off the Meangis Islands (Station 214).
Not only are the ten-armed species of Antedon the most widely distributed as a
group, but they also have the most extensive individual range. Antedon eschrichti and
Antedon quadrata of the Arctic Ocean were dredged by the Challenger in lat. 43° N.
Antedon phcdangium ranges from the north of Scotland to Morocco and throughout the
western basin of the Mediterranean. The Protean Antedon rosacea also occurs in
the Mediterranean, extends from the Fseroe Banks to the Canaries, possibly even to
Cape Verde and the equator, and is perhaps also found on the American coast; whde
Antedon carinata is distributed between the parallels of 15° N. and 35° S., through
the Indian Ocean from Java1 to Zanzibar, along the Atlantic coast of South America from
St. Lucia to Eio Janeiro, and is also found at Valparaiso.
None of the multibrachiate forms of Antedon have anything like this geographical
range. In the western North Atlantic there is no species with more than ten arms north
of Florida, and the dimorphic Antedon lusitanica is the only one known on the eastern side.
This last and those from Japan are the most northerly multibrachiate forms, while Antedon
setosa from off Tristan da Cunha and the various species inhabiting Port Jackson and
near the Kermadecs are the most southern representatives of these many -armed types of
Antedon, which have almost exactly the same range in latitude as the genus Actinometra.
Examples of each of the two great groups, those with two and those with three distichal
joints, occur in the Caribbean Sea, and they are abundant between the Society Islands
and the Red Sea. But, as we have just seen, they have a very limited bathymetrical
range, only appearing at seven Challenger stations between 100 and 630 fathoms,
and at none where the depth exceeded this latter limit.
In some of the Antedon-syyecies dredged at all these seven stations the secondary arms
consist of three distichal joints, the axillary with a Syzygy, but at two of them bidistichate
forms also occurred, together with species of Actinometra ; and the single " Porcupine "
Antedon with more than ten arms is Antedon lusitanica from 740 fathoms, in the North
Atlantic, which sometimes has a distichal series of two joints. There are no tridistichate
species of Antedon in the North Atlantic, outside the Caribbean Sea ; though they occur
in the South Atlantic at Tristan da Cunha and Ascension, and at five stations below
100 fathoms in the Western Pacific and Australasia.
On the other hand, the bidistichate series represented by Antedon lusita?iica does
1 See the remarks on this subject on p. 202.
REPORT ON THE CRINOIDEA. 35
not range further south in the Atlantic than 10° S., though it has the same distribution
as the tridistichate series in the Pacific and is generally more fully represented, forms
like Antedon palmata, Antedon elongata, and Antedon indica being often met with in
considerable abundance and variety. This group is also much more common than the
tridistichate group in the Caribbean Sea, especially below 100 fathoms ; and it ranges
down to 270 fathoms, at least 120 fathoms deeper than any member of the tridistichate
group has yet been found in that locality.
The range of the genus Actinometra, both in depth and in space, is very much more
limited than that of Antedon. It corresponds very closely, however, with the geographi-
cal and bathymetrical ranges of the multibrachiate species of this genus, though both
alike are slightly more extensive than the range of Actinometra. Thus, for example, the
multibrachiate forms of Antedon almost reach the parallels of 40°; while the northernmost
Actinometra does not reach 36° N., either in the Atlantic or in the Pacific, and the
southernmost are those of the Cape of Good Hope (34° 24' S.) and Port Philip (37° 48'
S.). In like manner no Actinometra has been obtained with certainty at a greater depth
than 533 fathoms ; though it is possible that this should be extended to 610 fathoms in
the Pacific.1 But as we have just seen, the tri- and bidistichate groups of the multi-
brachiate species of Antedon extend down to 630 and 740 fathoms respectively.
Like these forms too, Actinometra is far more extensively developed in the eastern
than in the western hemisphere. Several species are known from Southern Japan, and
the genus is abundant all through the Eastern Archipelago and down the east coast of
Australia as far as Port Jackson ; whde a single species from the latter locality also occurs
at Port Philip and in King George's Sound (Actinometra trichoptera). A few more are
scattered at Ceylon, the Eed Sea, Madagascar, Port Natal, and the Cape of Good Hope ;
but they are not known at all from the West African coast, nor from South America
south of Cape Frio. From this region, however, a couple of species occur abundantly up
to the Caribbean Sea and the Gulf Stream, but they do not pass the parallels of 25° N.;
though in the East Atlantic one species has been dredged four times beyond the thirty-
fourth parallel and at much greater depths than in the Caribbean Sea, e.g., 1500 metres 2
( = 812 fathoms). This type (Actinometra pulcliella) is one of special interest, not only
from its singularly Protean character, but because it is the only Actinometra common to
the two sides of the Atlantic ; while it is also, with one exception, the only Actinometra
ranging below 300 fathoms. The genus has been dredged eleven times at depths below
200 fathoms, four times by English, once by French, and six times by American expedi-
1 There is no record of the particular dredging at the Station numbered 174 which yielded Comatulse, the depths
being 210, 255 and 610 fathoms, except that the last one yielded Atelecrinus wyvillii. Three species of Actinometra were
obtained, together with five of Antedon, and from their general facies I should be decidedly inclined to refer them to one
of the two lesser depths.
2 According to H. Filhol (La Nature, 1884, p. 330), an Actinometra, which I take to be Actinometra pulchella, was
obtained by the " Talisman " off Rochefort at this very unusual depth.
36 THE VOYAGE OF H.M.S. CHALLENGER.
tions. Actinometra pulchella occurs at every one of these eleven stations, excepting
No. 174 in the South Pacific; while it is the only Actinometra represented at six at
least of them, including three of the deepest ones. This may be partly explained by the
fact that only one of these stations was in the Pacific, all the remainder being in the
Atlantic and the Caribbean Sea, but it is especially noteworthy because Actinometra
jmlchella is a dimorphic species, some forms having only ten arms, and some having
bidistichate series on one or more rays. The three species obtained by the Challenger at
Station 174 were all multibrachiate forms expressing widely different types of the genus ;
but the "Blake" dredged a ten-armed species at 450 fathoms, off Havana, and the two
deepest stations in the Caribbean Sea where Actinometra pulchella occurred also yielded
ten-armed species.
These ten-armed forms of Actinometra which occur in the Caribbean Sea and along
the South American coast, represent an entirely different type of the genus from the ten-
armed species of the eastern hemisphere. The latter mostly belong to the type of Actino-
metra Solaris, with syzygies between the two outer radials, though a few forms occur in
which these joints are united by bifascial articulation, as in nearly every Antedon and
in the Actinometra meridionalis of the Caribbean Sea (PI. LVI. fig. 1). The Solaris-
type, however, has not yet been discovered in the Atlantic.
Of the multibrachiate species of Actinometra the tridistichate type seems to be the
more extensively distributed and not the bidistichate one as in the case of Antedon. Thus,
for example, Actinometra parvicirra (o.3.(3).(3)) occurs in South Africa, Timor, Ceram,
the Philippines, Japan, the Friendly Islands, and even on the coast of Peru, so that it
has a range in longitude of some 260°, occurring everywhere but in the Atlantic. This
is only approached by the ten-armed Antedon carinata, which occurs on both coasts
of South America and across the Indian Ocean from Java to Zanzibar.
REPORT ON THE CRINOIDEA. 37
IV.— THE GEOLOGICAL HISTORY OF THE COMATULtE.1
So far as our present information goes the family Coniatulidse first appeared in the
time of the Middle Lias and is therefore of somewhat less antiquity than the Penta-
crinidae which date back to the Trias. Comatulee were fairly abundant all through the
Jurassic and Cretaceous epochs and were especially so at certain periods, that of the
Corallian in Germany and Switzerland, for instance.
The geographical distribution of recent Comatulse is far more extensive than that of
their predecessors. The distribution of the former is practically world-wide ; but so far
as is yet known, with the exception of an Antedon from Algeria and another from Syria,
no fossil Comatulse have been discovered out of Europe, not even in the Indian
Tertiaries, which contain so many Echinoderm remains. None are known in America,
though stem-joints of the remarkable Pentacrinus asteriscus are very common at certain
horizons of the Jura-Trias over wide areas of the western territories; and this shows
that the conditions of that long-distant age were not altogether unfavourable to the
development of Crinoid life. On the other hand, the Middle Lias of France contains
two species of Antedon, the oldest yet known ; and the genus occurs, together with
Actinometra, in the lower Oolites of both France and England ; while if Bourgueticrinus
ooliticus, M'Coy, is a Thiolliericrinus, as supposed by de Loriol, it is the earliest known
species of this very singular genus.
Both Antedon and Actinometra, especially the former, are well represented in the
Corallian of the Jura, and there are several species of Antedon in the Neocomian of the
continent, together with a few in Britain. The Gault of Folkestone has yielded typical
forms of both genera, and there are several Cretaceous species of Antedon scattered
through Europe, the formerly obscure Glenotremitcs paradoxus being the best known.
We are only acquainted with one Eocene Comatida; though three species occur in the
French Miocene, and there are others in the Pliocene both of England and of Italy.
In the majority of cases only the centro-dorsal is preserved, though it is not
uncommon for the radials to remain attached to it. But individuals with any arm-
joints preserved beyond the calyx-radials are decidedly rare ; and in this respect the
Comatulse differ widely from the Pentacriims-type, isolated calyces of which are not
often met with, though the arms are frequently extraordinarily well preserved.
One singular instance of the retention of the arms or arm-bases is afforded by
Eudiocrinus hyselyi.2 But for this fact the existence of Eudiocrinus in the fossil state
x I am indebted to the kindness of M. P. de Loriol for much information respecting the fossil Comatulidae of
France and Switzerland, some of it being as yet unpublished.
2 See de Loriol, Monographic des Crinoides fossiles de la Suisse, Geneva, 1877-79, pi. xxi. fig. \4.
38 THE VOYAGE OF H.M.S. CHALLENGER.
would have remained unknown, as the characters of the calyx in the recent species have
not yet been sufficiently studied to give any satisfactory clue to the detection of their
fossil representatives.
Most of the fossil Comatulae have more or less well defined basals appearing
externally, which have not undergone metamorphosis into a rosette as is the case in very
nearly all the recent forms ; and it is probable that species like Antedon tessoni and
Antedon orhignyi, although they show no basals externally, will in reality prove to be
no exceptions to the rule. I feel some doubt, however, with regard to the Tertiary
species, only two of which arc represented by more than the centro-dorsal ; and this
affords but Uttle information respecting the presence or absence of a rosette. I am
inclined to think myself that in the matter of basals the Tertiary species resembled their
predecessors rather than their successors. But this view cannot be confirmed till the
discovery of a type which shows basals at the interradial angles of the calyx, or of one
in which these plates are visible on the under surface of the isolated radial pentagon.
But no Tertiary species of this kind are known, and neither Antedon alticeps nor
Antedon italica shows any traces of basals between the radials and the centro-dorsal.
The determination of the generic position of a Mesozoic Comatula is often a matter
of considerable difficulty ; and this is especially the case when only the centro-dorsal is
preserved. In most fossil Cornatulse this part bears a considerable number of cirri which
are distributed over the greater part of its surface ; and it reaches a fair degree of
thickness ; so that there can be no doubt that these types have been correctly referred
to Antedon. But there are a few forms in which the centro-dorsal is relatively much
thinner and the number of cirri, which are almost or entirely limited to its sides, is
reduced. This is the case, for example, in two species from the Great Oolite and Bradford
Clay respectively, which I take to belong to Actinometra, rather than to Antedon.
Specimens which have the radials preserved can in some cases be referred to Antedon
without any difficulty, owing to the large proportion of height to width on the articular
faces of the radials. Such are Antedon sequimarginata, Antedon incurva, and Antedon
scrobiculata, the calyces of which closely resemble those of the typical forms of
Antedon figured on PL II.
On the other hand, the generic identity of Actinometra loveni from the Gault is
equally indisputable. For there is no living Antedon yet known in which the centro-
dorsal loses all traces of its cirri and becomes separated from the flattened radial
pentagon by clefts at its sides ; while these changes are not uncommon in Actinometra
(PI. L VII. fig. 1 ; PI. LXV.). But in by far the greater number of Comatulas which have
the radials preserved, the height of these plates is quite small relatively to their width, as is
invariably the case in the living Actinometra (PL V.). When these radials rest on a thick
centro-dorsal which is marked by a number of cirrus-sockets [Antedon decameros, Antedon
greppini) there can be no question that the type in question belongs to Antedon. But
REPORT ON THE CRINOIDEA.
39
there are a few species with low and wide radials, the distal faces of which have a steep
slope, so that they do not enter largely into the ventral aspect of the calyx. Such are
the two which I have described as Actinometra cheltonensis and Actinometra imrtem-
bergica. Only the radials of the former species are known and the slope of the
articular faces is scarcely as steep as in most recent examples of the genus. It is steeper
in Actinometra wurtembergica, which seems to have had a thicker centro-dorsal and
more numerous cirri than is usually the case in recent species of the genus. One might
also be inclined to refer to this genus the Antedon picteti, de Loriol, and Antedon
infracretacea, Ooster, both of which occur in the Valangian and have low wide radials
with a thin centro-dorsal, bearing but few cirri. They retain, however, the sloping
articular faces which are so characteristic of Antedon; and I think therefore that, for the
present at any rate, they should be referred to that genus.
Table showing the Distribution of the Fossil Coniatxdee in Space and in Time.
A. = Antedon. a. = Actinometra. E. = Eudiocrinus. T. = Thiolliericrinus.
Lias, .
Middle Lias,
13
a
«
a
O
a
&
o
Ph
c
N
w
.2
CO
<
•A
p
V
m
OS
'E
<o
so
<
V2
A.
■
Bathonian, ....
Oxfordian, ....
A. a. T ?
A.
A. a.
A. a.
A.
Jurassic,
Corallian, ....
Portlandian,
Lower Neocomian (Valangian),
Upper Neocomian,
A.
A. T.
A.
A.
A.
A. T.
A. E.
A. a. T.
A.
A.
Cretaceous,
Gault
Cenouianian (Low. Ch. ),
A. a.
A.
A.
h
*
Senonian (Up. Ch.),
A.
A.
A.
A.
)
r
Eocene, ....
A.
i
Tertiary, . -j
I
Miocene (Middle),
A.
A.
Pliocer.e, ....
i
A.
A.
Two points may be noted about the fossil Comatulas generally. The calyces of many
of them reach a considerable relative size, the centro-dorsal being sometimes as much as
9 to 13 mm. in diameter, which is greater than that of nearly every living representative
of the family except Antedon eschrichti; while this type and Actinometra solan's
40 THE VOYAGE OF H.M.S. CHALLENGER.
are almost the only living Crinoids with arm-bases anything like so massive as in
the fossil species. The Miocene Antedon rhodanica has a very large centro-dorsal ; but
the three species from the Norwich Crag and the two from the Italian Tertiaries are all
craite small.
Another character which presents itself in a large number of the Jurassic and
Cretaceous species is the retention of the five-rayed perforation on the lower surface of the
centro-dorsal, the peculiarities of which have been discussed in Chapter II.
The geological distribution of the three fossil genera of Comatulse is shown in
the foregoing table.
REPORT ON THE CRINOIDEA. 41
V.— CLASSIFICATION.
Until the time of Johannes Miiller the number of recognised species of Comatulae was
extremely small, not more than a dozen, in fact. Retzius had described one, Linnaeus
two, Lamarck seven, and two more bore the names of Diiben and Koren ; but only
three of them had more than ten arms, viz., Comatula rotalaria with about twenty to
twenty-two, Comatula Jimbriata with twelve to thirty, and Comatula midtiradiata, with
forty to fifty.
Under these circumstances the classification of the Comatulae presented no difficulties.
But Midler's descriptive work1 raised the total number of species to nearly forty,
about half of them having more than ten arms. This very obvious character afforded
him the means of separating his species into two groups, which he further subdivided
according to the arrangement of the syzygies in the arms. Thus, for example, there
are two sets of ten-armed Comatulae, those like Actinometra pectinata (PI. LIII. fig. 15)
in which the two joints above the radial axillary are each traversed by a syzygy, and
those like Antedon eschrichti (PI. XXIV. figs. 10, 11) in which the first syzygy is in the
third brachial. In like manner the multibrachiate forms were separated by Midler into
two sets, those in which the brachial axillaries are syzygial joints (PI. LXVIII. fig. 2), and
those in which the axillaries are simple and not traversed by syzygies (PI. XLV. fig. 2).
All the Comatulae known to Midler could be placed in one or other of these four
sets, no matter to which of the two subgenera they belonged, Alecto or Actinometra.
He never made any definite attempt to separate the species of Alecto from those of
Actinometra, no apparent system being determinable, either in the order of his
specific descriptions or in his tabular arrangement of most of the species in the
form of a key, a species of Actinometra not unfrequently intervening between two
of Alecto. For more than a dozen years after the publication of Midler's memoir the
classification of the Comatulae remained practically as he left it. No one took up
the subject, and no new species were described. In the year 1862, however, a step in
advance was made by Messrs. Dujardin and Hupe.2 They divided the recent Feather-
stars into three genera, Comatula, Lamarck, Actinometra, Midler, and Comaster, Agassiz,
the last named being a type which Midler had been unable to recognise as generically
distinct from Comatula. Of his own subgenera, Alecto and Actinometra, the
latter was raised into a genus by Dujardin and Hupe, who referred to it three species,
while they limited Lamarck's name Comatula to the forms previously referred by Midler
to Alecto, and regarded them as constituting thirty-one species. These were divided into
groups having respectively ten, ten to twenty, twenty, twenty-six to forty, and more than
1 Abhandl. d. k. Akad. d. Wiss. Berlin, 1849, pp. 237-265. 2 Op. cit., pp. 192-213.
(zool. chall. exp. — rART lx. — 1888.) Ooo 6
42 THE VOYAGE OF H.M.S. CHALLENGER.
forty arms. But the French authors altogether gave up Midler's method of grouping
both the ten-armed and the multibrachiate Comatulse according to the arrangement
of the syzygies in the arms and their subdivisions, placing for example Comatula
brachiolata and Comatula Solaris with syzygies in the first and second brachials between
Comatula adeonse and Comatula echinoptera, both of which have the third brachial a
syzygy. In like manner Comatula fiagellata with no syzygies in the brachial axillaries
is placed between Comatula japonica and Comatula timorensis, in both of which the
axillaries are syzygial joints. While therefore Dujardin and Hupe made a distinct
advance on Miiller's classification in recognising two generic types of Comatulse,
their rejection of the characters on which he relied, and rightly so, as being of much
systematic value was a decidedly backward step. For all subsequent work has shown
that the position of the first syzygy in the free arms and the presence or absence
of syzygies in the brachial axillaries are characters of very considerable systematic value,
without the aid of which the classification of the hundred or more species comprised in
each of the genera Antedon and Actinometra would be even more chaotic than it is.
For some fifteen years after the appearance of Dujardin and Hupe's Histoire
Naturelle, systematic work on the " Comatulse progressed with extreme slowness, the
most important step being Norman's restoration of the generic name Antedon, owing to
its priority over both Comatula and Alecto.1 New species were described by Bohlsche,
Grube, and Pourtales ; but they were never figured, and no attempt was made to assign
them places in the system either of Midler or of Dujardin and Hupe. Dr. Liitken had
examined from time to time a considerable number of Comatulse which had been collected
among the Pacific Islands by the agents of the Godeffroy Museum ; and he arrived at
the conclusion that the real distinction between Antedon (or Alecto) and Actinometra lies
in the central or excentric position of the mouth, the number of groove trunks reaching the
peristome being a character almost entirely devoid of the systematic importance attributed
to it by Miiller. Liitken's views were never published, and I only learnt of his holding
them after myself arriving at the same conclusions ; but he was good enough to inform
me at the same time of a character then unknown to me, which I have since found to be
of almost invariable occurrence in Actinometra, viz., the presence of a terminal comb on
the lower pinnules (PL LVI. figs. 2, 4). These facts were published in my memoir on
Actinometra,2 where I also endeavoured to classify the species of the genus that I had
been able to identify, by an extension of the method employed by Miiller.
While recognising the systematic importance of the presence or absence of syzygies
in the arms of Comatulse, Miiller made no attempt to classify the multibrachiate forms
according to the number of joints between the successive brachial axillaries, though he
furnished the means for doing this in his descriptions of many species, a process which
1 Ann. and Mag. Nat. Hist, 1865, ser. 3, vol. xv. p. 98.
2 Trans. Linn. Soc. Lond. (Zool.), ser. 2, 1877 [1879], vol. ii. pp. 18-29.
REPORT ON THE CRINOIDEA. 43
would have enabled him to separate types that are placed very near to one another in
his scheme. Thus, for example, Comatula palmata and Comatula macronema are placed
respectively next to Comatula japonica and Comatula reynaudi, though the distichal
axillary is the second joint above the radials in the first pair, and the third (or, counting
the syzygy, the fourth) joint in the second pair. It soon appeared to me to be a very
general rule among Comatulse that " the first and second segments beyond every
axillary, whether radial or brachial, are nearly always united together in the same manner
as the second and third (axillary) radials."
These observations rendered the classification of the Comatulas which were then known
(1879) a comparatively easy task; and during the next three years I described several
species both o£ Antedon and of Actinometra, arranging the multibrachiate forms according
as there were one or two joints between the successive axillaries of the arms, and by the
presence or absence of syzygies in these axillaries. The most common arrangements of
the arm-divisions are the following — two joints, the second axillary without a syzygy,
and three joints, the second bearing a pinnule, but the third axillary with a syzygy.
These of course would have been equally well distinguished in Midler's classification
according to the presence or absence of syzygies in the axillaries. But in Midler's
scheme there is no separation from the second of these types of species like Actinometra
sentosa (PI. LX VI. fig. 4) in which all the outer arm-divisions consist of two joints only,
but the axillaries are syzygial joints just like those of Actinometra japonica, or Antedon
reynaudi. In like manner Midler's classification provides no place for forms like
Actinometra paucicirra, in which the axillary is not itself traversed by a syzygy, but is
united to the preceding joint by syzygy instead of by an articulation (PI. LIV. figs. 1, 2).
If these characters be taken into account, and especially the mode of union of the two
outer radials, whether by articulation or by syzygy, the numerous multibrachiate species
of Antedon and Actinometra may be readily separated into comparatively large groups,
for the further subdivision of which a more detailed examination of anatomical characters
becomes necessary.
In the year 1882 Professor F. J. Bell : attempted " to apply a method of formulation
to the species of the Comatulidee." He stated that the leading differences between the
radial, distichal, and palmar series in different species of Comatulse " are to be found in
the varying arrangement of that mode of union to which Johannes Midler applied the
term syzygial" ; and he therefore inserted the letter R, D, or P into his formula
" whenever the respective axillary is a syzygy," placing before this letter and the generic
symbol the figure 1, 2, or 3, according as the first, second, or third brachial is a syzygial
joint. Bell further devised a very convenient method of briefly indicating the number of
joints in the cirri and also that of these organs themselves. I have been glad to adopt
1 An attempt to apply a Method of Formulation to the Species of the Comatulidc-e ; with the Description of a New
Species, Proc. Zool. Soc. Lond., 1882, pp. 530-536.
44 THE VOYAGE OF H.M.S. CHALLENGER.
this method, which will be explained further on, but, on the other hand, the mode of
formulation suggested by Bell to express the characters of the arm-divisions in the
multibrachiate Comatulse left very much to be desired. For the regular forms which
have two or three joints in each arm-division and the axillary a syzygy, his notation is
probably as short a one as could be devised. But it gives no means of distinguishing one
of these types from the other, or from that of Actinometra multiradiata in which both
occur together ; and where the successive arm-divisions consist of two joints only, without
syzygies in the axillaries, it gives no information at all respecting the number of the arm-
divisions, Antedon palmate/, with three axillaries above the radials having the same
formula as Antedon macronema with only one.
Bell's method is totally inapplicable to irregular types like Actinometra multifida,
which have syzygies in the distichal axillaries but none in those of the subsequent
divisions ; and the consequence is that species with forty arms receive exactly the same
formulae (excepting of course for the cirrus-characters) as others with only ten to twenty.
I have referred elsewhere l to other difficulties connected with Bell's method of formula-
tion, which is neither elastic enough to indicate exactly on what joint the syzygy comes
in the distichal or palmar series, nor does it state the number of joints in each division
when there are no syzygies in the axillaries.
For some years before the publication of Bell's suggestions I had been in the habit of
employing for my own use a method of formulation which should briefly express the
characters of the rays and their subdivisions, and yet at the same time be elastic enough
to meet all the variations of Coma £w/a-structure with which I was acquainted, together
with any others that I could consider as possible. It was based upon a knowledge of the
structure of over two hundred species, which has enabled me to make the following
generalisations.
1. All ten-armed species of Actinometra which have the two outer radials united by
syzygy have the first two brachials united in the same way.
Examples. — Actinometra pectinata (PI. LIII. fig. 15); no Antedon known.
2. All many-armed species of Actinometra which have the two outer radials united
by syzygy, either have (a) all the arm-divisions of two joints also united by syzygy, and
the first two brachials similarly united ; or (/3) there may be three distichals of which
the first two are articulated and the axillary a syzygy, while the subsequent divisions
(if any) consist of but two joints united by syzygy.
Examples. — (a) Actinometra paucicirra (PI. LIV. figs. 1, 2, 10); (fi) Actinometra
typica (PI. LVII. fig. 1).
3. If the two outer radials are united by bifascial articulation, the two next joints are
similarly united, whether there be ten or many arms. In the former case the third
brachial is always a syzygy.
On the Classification of the Comatulas, Proc. Zoul. Soc. Lond., 1882, pp. 731-741.
REPORT ON THE CRINOIDEA. 45
Examples. — Antedon eschrichti (PL XXIV. fig. 11), Antedon tuberculata (PL XLV.
fig. 2), Antedon multispina (PI. L. fig. 3); Actinometra meridionalis (PI. LVI. fig. 1),
Actinometra stelligera (PI. LVIII. fig. 1), Actinometra regalis (PI. LXVIII. fig. 2).
4. In by far the greater number of Comatulse which have the two outer radials
united bifascially, and only one further division, the third brachial is the first syzygial
joint above the distichal axillary, whether this be a syzygy or not ; and the two lowest
brachials are also united bifascially.
Examples. — Antedon disciformis (PI. XXXIX. fig. 4), Antedon variipinna (PI.
XLVIII. fig. 5) ; Actinometra elongata (PI. LVII. fig. 4), Actinometra quadrata
(PI. LXII. fig. 1).
There are some exceptions to this rule. Actinometra pidchella and Actinometra
stelligera have two articulated distichals, but the first two brachials are united by syzygy
(PI. LII. fig. 2 ; PL LVIII. fig. 1). This is also the case in both Antedon angusticalyx
and Antedon insegualis (PI. L. fig. 1 ; PL LI. fig. 2) each of which has three distichals
of the usual character; while in the group of which Actinometra Jimbriata is the type
(PL LXII. fig. 3) there are also three distichals followed by a syzygy in the second
brachial. These exceptional syzygial unions do not occur, however, when there are no
distichals present on the ray and the arms spring directly from the radial axillaries.
Under these circumstances the first syzygy is always on the third brachial just as in
Antedon eschrichti and Actinometra meridionalis (PL XXIV. fig. 11; PL LVI. fig. 1).
This is very well shown in Actinometra pulchella (2-^-j, Actinometra coppingeri (S.2br),
and Antedon multispina (3.y), (PL LII. fig. 2 ; PL LX. fig. 2 ; PL LXIX. figs. 1,2);
and it may also be noticed even in species which usually have palmar series, such as
Antedon porrecta and Actinometra lineata (3.2[(p.)br] ), when both these and the
distichals are absent on any part of a ray (PL LX. fig. 3). These multibrachiate species,
therefore, are exceptions to Rule 4 in their state of fullest development. But when the
primary arms remain undivided the position of their first syzygy is invariably that of
the ordinary ten-armed Comatuke, i.e., on the third brachial, just as is stated in Rule 3.
5. If the two outer radials are articulated and there are two subsequent axillaries,
so that palmars are present, the first arm-syzygy above the palmar axillary is in the
third brachial in all cases but the following : —
a. Two palmars united by syzygy ; the first two joints beyond the palmar and all
subsequent axillaries are also united by syzygy.
Example. — Antedon distincta (PL LI. fig. 1).
ft. Two palmars, the axillary a syzygy ; the second joints beyond the palmar and all
subsequent axillaries also have a syzygy.
Examples. — Antedon porrecta, Actinometra sentosa (PL LXVI. fig. 4).
A very singular exception to this rule is afforded by Actinometra stelligera, which
48 THE VOYAGE OF H.M.S. CHALLENGER.
has articulated distichals and palmars, but the first two brachials united by syzygy
(PI. LVIII. fig. 1), so that its formula is— a.2.2.^-.
6. Whenever any arm-division, distichal, palmar, or any other consists of three
joints, the first two are articulated by ligaments, the second bearing a pinnule, and the
third (axillary) is a syzygy, just as in the first three brachials of Antedon eschrichti
(PL XXIV. fig. 11) and Actinometra meridionalis (PL LVI. fig. 1). When, however,
there are only two joints, and the second (axillary) is a syzygy, the first has a pinnule,
just as in the arm-bases of Actinometra fimbriata (PL LXII. fig. 3).
Examples. — Antedon variipinna (PL XL VIII. fig. 5) and Antedon porrecta
(PL LII. fig. 3) ; Actinometra parvicirra (PL LXI. fig. 1) and Actinometra sentosa
(PL LXVI. fig. 4).
7. The hypozygal of a syzygy is always united to the preceding joint by a muscular
articulation.
The method of formulation which I have devised in accordance with the above rules
is as follows : —
Like Professor Bell I use R to denote the syzygial union of the two outer radials ;
and I assume in accordance with Rules 3 to 5 that the first syzygy on the arm is in the
third brachial, unless otherwise stated. If it is in the second brachial I put 2br at the
br
end of the formula ; and if the first two brachials are united by syzygy -h- is used.
In like manner, and in accordance with Rule 5, 2d and 2p would indicate that there are
two distichals or two palmars, of which the axillary is a syzygy ; and -3- or 4v that the
two distichal or two palmar joints are themselves united by syzygy.
The figures 1 or 2 alone would indicate that there is either only a single axillary
joint, or that the axillary is the second joint and bifascially united to its predecessor; and
a 3 would denote three joints of which the axillary is a syzygy.1 If one figure occurs alone
in a formula it indicates the presence of distichals only ; two figures that palmars occur
as well ; and so on, an additional figure or letter (p\ p", p'") being added for each fresh
division, e.g., Actinometra alternans, 3, 2, 3, 2, Actinometra sentosa 3.2 (p. p'. br).
This may be tabulated as follows : —
Symbol used.
Character. Distichal. Palmar.
One axillary joint, ..... 1 1
Two joints united by syzygy,
Two articulated joints,
Two joints, the axillary a syzygy,
Three joints, the axillary a syzygy,
1 It would of course be more consistent to write 3d or 3p ; but the syzyj
d p
1 ~2
2 2
2d 2?
3
nature of the third (axillary) joint is such
a constant character (Rule 6) that until an exception is met with, I prefer to use the figure alone for the sake of brevity.
REPORT ON THE CRINOIDEA. 47
Bell's method of indicating the varying characters of the cirri is as follows : —
" If there are from 1-12 cirri, we may say there are few ; if from 12-30 a moderate
number; and if more than 30 a large number; if there are not more than 20 joints to
the cirri we may look upon them as being few, if from 20-40 moderate, and if more
than 40 numerous. I propose to use the letters a, b, and c to represent few, moderate,
and numerous respectively ; while the letter for the number of cirri will form the
numerator and that for the number of joints the denominator of a fraction ; and where
there is a difficulty of decision one might write ab, or be. Antedon and Actinomet ra
may be usefully, though not of necessity, distinguished by making A or A' part of
the formula."1 Bell prefers to use A' for Actinometra rather than "a" as I have
suggested, because the a is used in the formula for the cirri. I do not see the
force of this objection, as the two letters occur at opposite ends of the species formula
and only the later one is italicised ; while A' is much too like A to be readily distin-
guished at a glance, apart from the possibility of printer's errors. Bell's suggestion
that "br." should be used instead of "b" for the brachials to avoid confusion with
the b of the cirrus-formula is a good one, however, and I have adopted it accordingly.
In my former method of formulation I denoted the presence of ten arms only by
inserting a 10 into the formula of the type, thinking it more convenient to indicate this
character, which is generally a sharply defined one, in a positive, rather than in a negative
manner. Bell thinks, however, that "A. 10" compared with "A. 3 " is very apt to
mislead and to give rise to the impression that the Antedon in question has ten distichal
joints. In deference to his scruples therefore I shall omit the 10 in future and write, as
he does, the specific formula of ordinary ten-armed Coniatuke like Antedon eschrichti,
with no other characters than the generic letter and the cirrus-fraction. Thus Antedon
be
phalangium is represented by A.—.
It often happens that some individuals of a species are more fully developed
than others, i.e., they have additional axillaries in the arm-divisions. Thus for
example, one or two bidistichate series are occasionally present in Antedon lusitanica
which usually only has ten arms (PI. XXXIX. figs. 1, 3); while palmars are sometimes
found in some forms of Antedon quinquecostata and of Antedon variipinna, but not in
others (PL XXXVIII. fig. 1 ; PL XLIX. fig. 1). Under these circumstances I write
the figure or letter which denotes the character that is variable between brackets,
e.g., A.(2), lusitanica; A.2.(2), quinquecostata; A.[3.(2)], variipinna.
In Bell's system, however, " when a character frequently though not always obtains,
the corresponding letter is put within brackets." 2 If this were only meant to imply that
certain characters present themselves in some individuals of a species, but not in others,
Bell's method would be the same as mine. But though he goes much further than I
1 Loc. cit., p. 531. -' Loc. tit., p. 532.
48 THE VOYAGE OF H.M.S. CHALLENGER,
do in theory, he is by no means consistent in practice. Two of his new species, Antedon
reginee 1 and Antedon briareus,2 are represented by single specimens only ; the palmars in
the former and the post-palmars in the latter, as shown in his figures, are not com-
plete all round the calyx, so that the number of arms is thirty-eight and seventy-one
respectively instead of forty and eighty. The corresponding symbols are therefore
enclosed within brackets in his specific formulae.
From my experience with Actinometra parvicirra I can quite believe it possible that
examples of Antedon regime may eventually be found in which there are no palmars and
so not more than twenty arms ; but until this is the case I see no reason to enclose the
sign for the palmars between brackets in the specific formula. Since, too, there may be an
axillary beyond the post-palmar in Actinometra briareus with its seventy to eighty arms,
I do not think it probable that examples of the type will ever be found without
some post-palmar series, i.e., with forty arms or less ; and the use of the brackets in this
case would be extremely misleading, though it is no doubt correct for the subsequent
division, which Bell ignored altogether. But even in this case I should wait to use the
brackets till the obvious reasons for doing so presented themselves.
Bell does not always follow his own rule of employing brackets when the arm-
divisions are not equal all round the calyx. Thus he describes Antedon irregularis
as having eleven to twenty-two arms, and he figured an individual in which half
the primary arms do not bear distichal axillaries.3 He does not, however, put the sign
for the distichals within brackets, as he ought consistently to do ; for the presence of the
distichal axillary is a character which, as he expresses it, " frequently but not always
obtains " in this species. In like manner his figure and description of Antedon elegans *
show that half, i.e., five, if not more, of the primary arms may remain undivided. But he
does not put the distichal figure in brackets as his system demands. His most serious
lapses in this respect are indicated by his formula for Actinometra parvicirra.6 He gives
it as A'. 3. 3. No brackets being used at all, the reader is led to infer that the presence
of three distichals and three palmars is a character which "always obtains" in this
species. But I described some specimens in 1879 which had only twenty arms and no
palmars developed at all ; while I also figured one with only three of the ten distichal
1 Report on the Zoological Collections made in the Indo-Pacific Ocean during the Voyage of H.M.S. "Alert,"
1881-82, London, 1884, p. 115.
2 Bell's formula for this type is very incorrect. Not only has he referred it to Antedon when it is in reality an
Actinometra, but he twice stated that the post-palmar series consist of two joints without syzygies in the axillaries
(pp. 155, 163) ; whereas as a matter of fact his figure on pi. xiv. shows that this is only the case in nine out of the
twenty-seven post-palmar series, just one-third of the whole ! ! In all the remaining eighteen series there are three
joints like the distichal series, with a syzygy in each axillary ; and this arrangement, which occurs in two-thirds of the
divisions, should therefore be regarded as typical. Furthermore there are four cases of axillaries above the post-palmars,
three of which have syzygies like the distichal axillaries. Bell takes no notice of these, however, and so misses the
opportunity of contrasting this type, a.3.2.3.3 with Actinometra alternans, a.3.2.3.2 ; and from the formula which
he gives, A.3.2(2), one would be led to imagine that the specific relations of the type were rather with Actinometra
multifida (a.3.2.2).
3 " Alert" Report, p. 161, pi. xiii. fig. A. 4 Ibid., p. 162, pi. xiii. fig. B. 5 Ibid., p. 155.
REPORT ON THE CRINOIDEA. 49
series present, so that the number of arms is reduced to thirteen,1 and Bell has himself
examined individuals with less than twenty.2 According to his rule, therefore, the
formula should be — A'. (3). (3). but in the formula which he actually gives the brackets
are altogether omitted. I should write it myself as — a. 3. (3), to indicate that whde some
distichal series are always present in every individual, palmar series may occasionally be
entirely absent. This appears to me to be the only possible way in which brackets can
be profitably employed. Bell, however, thinks otherwise, as is shown by the following
passage : 3—
" From the table of Antedon formulae some facts become at once apparent : —
" (a) There are six examples among the more than ten-rayed forms in which the arms
are not a regular multiple of ten — that is, not 20, 40, or 80 ; this is clear from the sign
for the palmar or post-palmar being in these cases placed within brackets."
The first line of this passage contains a repetition of an error in terminology which
was made by Bell in 1882,4 and was afterwards corrected by myself.5 He seems, how-
ever, to consider the point an unimportant one and continues to use the expression to
which I took exception. There are no ten-rayed forms of Antedon, though there are
plenty which are ten-armed. The arms were clearly distinguished from the rays by
Midler, who laid the foundation of the descriptive terminology now in use for the
Crinoids. But Bell persists in using the word rays when he only means arms. This is
unfortunate, as it leads to confusion between the five-rayed but ten-armed Antedon and
the truly ten-rayed Promachocrinus, a point to which I have before alluded.
Bell has evidently made the generalisation quoted above on the basis of his formulae,
without special reference to the individuals he examined. He describes his single
specimen of Antedon gyges as having forty-one arms, and I find this to be due to the
presence of one post-palmar series, of which Bell's formula gives no hint. He is thus
able to include this type among those forms in which the arms are a regular multiple of
ten, i.e., forty. Then again he gives the formula of Antedon articulata as A.2.2. But
the exact number of forty arms which this expression denotes does not occur in his
specimen, which also has one post-palmar series ; while I have seen individuals with less
than forty arms. According to Bell's own system the formula of this type and perhaps
also that of Antedon gyges should be A. 2. (2). (2). We find then that not only on the six,
but in all the eight multibrachiate forms of Antedon for which he gives formulae the arms
are not a regular multiple of ten. But this is in no way a specially remarkable fact. The
singularity would be if the number of arms always were a regular multiple of ten, as is
generally though not always the case in Aetinometra paucicirra (PL LIV. figs. 1, 2).
But this is a most exceptional species. No one can examine any large collection of multi-
brachiate Comatulae without becoming immediately aware of the extreme irregularity in
1 Trans. Linn. Soc. Land. (Zool.), 1879, ser. 2, vol. ii. pp. 51, 52, pi. ii. fig. 9. 2 " Alert " Report, p. 168.
3 Ibid., p. 155. 4 Proc. Zool. Soc. Lond., 1882, p. 532, note. 6 Ibid., p. 732, note.
(ZOOL. CHALL. EXP. — PAET LX. 1888.) OOO 7
50 THE VOYAGE OF H.M.S. CHALLENGER.
the extent of the arm-divisions. Individuals with a simdar distichal axillary on each
primary arm and no further division, so that the number of arms is exactly twenty, are
extremely rare, except in Actinometra paucicirra, and to a less degree also in Actino-
metra pulchella. Another Caribbean species {Antedon spinifera) not unfrequently has
exactly thirty arms, owing to the very regular presence of palmar axillaries upon the
inner pair of every four secondary arms. But I cannot call to mind any species of
Comatula among the many hundred forms which I have examined in which the total
number of arms is exactly forty, owing to the presence of ten distichal series and twenty
series of palmars. I have seen an Actinometra parvicirra with thirty-nine arms, an
Antedon articulata with forty, and Bell's unique specimen of Antedon gyges has forty-
one. But I do not remember any species which always has exactly forty, and I doubt
if there be one ; while I can say with tolerable confidence that no one wdl ever find a
specific type which always has ten distichal axillaries, twenty palmars, and forty post-
palmars, thus giving rise to exactly eighty arms. The logical result of Bell's use of
brackets therefore would be that every Comatula with eleven to nineteen arms should
have the symbol for the distichals placed between brackets ; for those with twenty-one
to thirty-nine arms there should be brackets round the palmar sign and generally also
round the distichal one as well ; while the formulae of types which have over forty and
less than eighty arms should have the last, if not the two last, symbols within brackets.
A reference to Bell's formulae 1 for Antedon articulata and Antedon gyges, and to
those for Actinometra alternans, Actinometra parvicirra, and Actinometra midtifida, will
show, however, that he has not written them out according to his own system, for none of
them have any brackets at all, although in each case he knows of individuals in which
the number of arms is not an exact multiple of ten.
There is another point too which he does not seem to have fully considered in the
construction of his formulae. The multibrachiate Comatuke, such as Antedon occulta
(PL XLVIII. fig. 1) and Actinometra stelligera (PL LVIII. fig. 1), in which the successive
arm-divisions typically consist of two joints each, the axillary without a syzygy, are as a
rule extremely regular in their characters. But the case is quite different in those forms
which typically have three distichals and three palmars with syzygies in both the axillaries.
It is extremely rare to meet with examples of these species in which one or more of the
three-jointed distichal and palmar series are not replaced by two jointed series without
syzygies in the axillaries. Thus, for instance, I have described specimens of Actinometra
parvicirra with eighteen and twenty arms respectively, in which half the distichal series
were two-jointed, and the other half three-jointed ; and a similar irregularity occurs
among the palmars. In twelve individuals, however, ninety-six out of one hundred and
eleven distichal series, and sixty-seven out of seventy-six palmar series, were three-jointed;2
and I was thus definitely enabled to make out the characters of the type and to write its
i "Alert" Report, p. 155. 2 Trans. Linn. Soc. Land. (Zool.), 1879, ser. 2, vol. ii. pp. 44, 45.
REPORT ON THE CRINOIDEA. 51
formula — a. 3. (3). This method, a determination of the characters present in the majority
of cases, is the only one which can be safely relied on for fixing the characters of a species ;
and it is therefore apparent that the formulae given by both Bell and myself for species of
which we have only seen single individuals are necessarily liable to subsequent correction.
Bell has encountered this difficulty of irregular arm-divisions, and has met it by
giving three formulae for one species which he names Actinometra variabilis.1 It seems
to me that two would have been sufficient, as the characters indicated by the first,
A'.3.2., are also expressed in the third, A'.3.(2).(2) ; while there must be a considerable
mistake somewhere ; for Bell's first and second formulas do not provide for more than
forty arms, though he gives the total number of arms as sixty to ninety. His second
formula is A'.3.3., which of course represents a very different type from A'.3.(2).(2).
So far as one may judge from his figured specimen, the last is much the most correct,
for out of thirteen palmar series only two consist of three joints. On some part of every
ray there are three divisions above the palmare, each, with but one exception, consisting
of two simple joints. I find that a similar arrangement presents itself upon each of the
other three specimens of this type, and I should therefore write its formula as — a.3.2.2.2.2,
not using brackets for the last figure because a fifth post-radial axillary occurs in each of
the four individuals examined. Neither of Bell's formulae, however, allow for more than
three post-radial axillaries, while his second one A'. 3.3. would indicate by the absence of
brackets a type with exactly forty arms, and regular distichal and palmar series of three
joints each all round the cup, i.e., such a form as Actinometra parvicirra, while in
reality Actinometra variabilis only resembles that species in the constant presence of
three distichals, its later arm-divisions being totally different from those of that type.
While therefore it is extremely desirable to be able to examine a good number of
individuals before attempting to describe and give a formula for any new specific type
of multibrachiate Comatulae, I do not think that there is any serious objection to describing
a species from one individual only. For so far as the characters of the arm-divisions are
concerned, I have found it to be an almost invariable rule that the characters which
present themselves most frequently in any one individual are those which distinguish the
species. Thus, for example, bidistichate series only presented themselves in five out of
twelve specimens of Actinometra parvicirra,2 in which the number of distichals is
typically three. Two of these individuals were certainly abnormal, the numbers of
bidistichate and tridistichate series being exactly equal. But in the other three specimens
the largest number of bidistichate series was three, and they never presented themselves at
all in seven individuals. The same may be said, though with a somewhat less degree of
certainty, respecting the palmar series, sixty-seven of the seventy-six present consisting
of three joints. Palmare only occurred in eight of the twelve specimens examined, and
were abnormal in but four of them, one species being unusual in having three two-jointed
1 "Alert" Report, p. 155. 2 Trans. Linn. Soc. Lond. (ZooL), 1879, ser. 2, vol. ii. p. 44.
52 THE VOYAGE OF H.M.S. CHALLENGER.
series, and only two of three joints. But, excepting in rare cases like this, the
predominant characters of the individual may be safely taken as those of the type, and
the formula constructed accordingly.
Bell's method of writing a formula for every slight variation, as he has done in the
case of Actinometra variabilis, would result in the following list of formulae for
Actinometra parvicirra.
(2) (2) (">)
a. (3) ; a.^; a.3.(2); a.3.7^ ;x a.3(3) ; a.2.7^ ; a.2.(3), and so on.
Such a collection of formula? would be worse than useless from its confusion, and
very far from being the shorthand system which Bell rightly wishes to see employed. It
would be much easier to refer to the specific diagnosis at once than to try and make out
the predominant characters of the arm-divisions from a supposed shorthand of this kind.
Two points must therefore be noted in determining the formula of a species.
1. What are the characters of the majority of the arm-divisions in a given individual, or
better still, in a number of individuals ? 2. Whether examples ever present themselves
in which a given character, such as the occurrence of distichal, palmar, or post-palmar
divisions, is sometimes entirely absent 1 In this case, but only in this, the corresponding
symbol should be put between brackets in the formula, e.g. —
Antedon lusitanica, A.(2).
Actinometra parvicirra, a.3.[3.(3)].
Actinometra multiradiata, a.3.2{p.(pf)br}.
But the fact that all the ten distichals or twenty palmars do not always occur in every
individual of a species is no reason for placing the corresponding symbol in brackets.
Were this done, I have no hesitation in saying that both symbols would have to be
enclosed in brackets in the formula of every species with less than forty-one arms and
no post-palmar divisions. This of course would be absurd, and render the use of
formulae altogether futile.
The principles of classification which have been explained above 2 enable us to divide
the numerous species of Antedon and Actinometra respectively into groups of very
variable size. These are arranged in the following lists, which contain the names of
all the species described by myself and my predecessors, Eetzius, Lamarck, Muller,
1 This is similar to the expression given by Bell for Antedon elegans, in which there are generally two palmars, but
sometimes three. His figured specimen presents one case of the latter to four of the former ; and it is therefore clear
that the formula should be written A.3.2.
2 I may just remark here that I cannot at all agree with the dictum of Walther that " Wer sich je mit Crinoiden
besehaftigt hat, der wird wissen, wie wenig specifischen Werth die Gabelungen der Arme besitzen" (Palreontographica,
1886, Bd. xxxii. p. 182). Walther's experience seems to have been limited to a comparatively small number of fossil
Crinoids, not always in the best state of preservation. But so far as concerns the recent Crinoids, both stalked and free,
the number and characters of the arm-divisions afford points of much importance in the discrimination of species. I am
convinced that the same may be said of the fossil Neocrinoids, if not of the Palaeocrinoids too, provided that a sufficient
range of specimens is brought under consideration.
REPORT ON THE CRINOIDEA. 53
Bohlsche, Pourtales, Grube, Bell and others, so far as my knowledge of them extends.
A few of my own undescribed species, from the " Blake " collection and elsewhere, are
also added, as they have received names somewhat prematurely, owing to their being the
hosts of Myzostomida, which have been described by Professor von Graff. One or
two of Professor Liitken's MS. names are included, as they belong to easily recognisable
types, e.g., Antedon protecta ; but others, such as Actinometra mutabilis and Actinometra
trachygaster, are omitted, as I have found a considerable variety of types under each of
these names in the different collections that I have examined.
I have added a few remarks respecting the absence of certain well-known names,
such as Antedon sarsii and Actinometra timorensis, and also with regard to the
presence of certain species in altogether different groups, both specific and generic, from
those to which they have been referred by their original describers — (See the numbers
before the names).
In order to exhibit more completely the range of structural variation which is to be
met with among the Comatuke, I have included the formulae for various species that I
have examined but have not j^et described. Some rather variable species appear in more
than one group, 1 and to draw attention to their peculiarities the name is followed in
each case by the formula of the other group in which the species also occurs. Thus
Actinometra pulchella (10),2 and (2.(2).^-
In one or two cases I have departed from the strict numerical sequence in order to
avoid separating too widely species which are really very closely allied, on the sole
ground that one has an axillary more than the other. Thus A. 3. ^-jp immediately
succeeds A^y, while a.3.2(p.br) and a.3.2(p,p',br) follow most naturally directly
after a.3.2.6r., instead of being separated from the latter group by several inter-
vening ones, as was the case in my preliminary list.3
List of Species.
Genus ANTEDON.
I. A.R.2.2.2.
A.R.3.2.(2). 1. elegans
A.R.3.3.3. midtiradiata
A. R. 3. 3. 3. 3. 2. microdiscus
'It has not been worth while to repeat the name in one or two cases, e.g., Antedon elegans (A.R.3.2.(2)), and I
have therefore used brackets for the last figure.
2 The 10 is not used here to indicate that there are ten joints in the distichal series, but as a short way of
denoting the presence of only ten arms. I trust that the abbreviation will not be misunderstood. See p. 47.
3Proc. Zool. Soc. Lond., 1882, pp. 746, 747.
54
THE VOYAGE OF H.M.S. CHALLENGER.
i. A. 10. [only ten arms].
abyssicola
discoidea
4. milberti
abyssorum
diiheni
milleri
accela
aculeata
duplex (2)
echinata
mvltispina ( 3— )
acutiradia
eschrichti
parvicirra
adeonse
exigua
parvipinna
alternata
jlexilis (2)
perspinosa
anceps (3)
gracilis
petasus
angustipinna
hageni
5. phalangium
antarctica
hirsuta
pinniformis
armata
hystrix
prolixa
australis
impinnata
pumila
bcdanoides
incerta
pusilla
bar en t si
incisa
6. quadrata
basicurva
informis
remota
bidens
Isevipinna
rhomboidea
bispinosa
Isevis
rosacea
brevipinna (2)
Isevissima
serripinna
breviradia
latipinna
spinicirra
carinata
lineata
7. tenella
carpenteri
longicirra
tenuicirra
columnaris
longipinna
tessellata
cubensis
3. loveni
tvberosa
dejecta
lusitanica (2)
valida
denticulata
magellanica
8. variipinna [3(2)].
. A. 2.
brevipinna (10)
Jlexilis (10)
patula
clemens
lusitanica (10)
pourtalesi (2.2)
compressa
macronema
quinquecostata (2.2)
disciformis
marginata
robusta
duplex (10)
A. 2. 2.
articulata (2.2.2)
imparipinna
palmata (2.2.2)
bimacidata
indica
pourtalesi (2)
brevicuneata
Isevicirra
protecta
elongata
manca
quinquecostata (2)
REPORT ON THE CEINOIDEA.
55
A. 2. 2.
regalis
similis
spinifera (2.2.2)
regime
spicata
tuberculata
A. 2. 2. 2.
sequipinna
Jiagellata
palmata (2.2.)
articulata (2.2.)
gyges
spinifera (2.2.)
conjungens
occulta
TV. A.S.%
a
angusticalyx 10. hmqualis (s.^^j
,.„ ( f.br. \
9. granulifera I 3. — t~ J
multispina (10)
A. 3.*f.
distincta
A.3.
anceps (10)
angustiradia
A.3.1.
A.3.2.
acuticirra
ludovici
A. 3. 2. 3.
A.3.2{(».)6r}
A.3.3.
A.3.3.3.
9. <jrra»jM&/«raf ^-"o")
reynaudi
savignyi (3.2)
quinduplicava,
savignyi (3)
porrecta
bipartipinna
10. insequalis (^--^j
variipinna [3.(2)]
variipinna [(3)]
■philiberti
REMARKS.
1. Antedon elegans, Bell. I place this species in the first group because I find
on examination that the two outer radials are united by syzygy. This important fact
escaped the notice of Bell ;x and his specific formula is incorrect in other points besides
the omission of the R. The species now appears therefore in an altogether different
group from that to which I at first assigned it on the basis of his description.
1 "Alert" Report, pp. 155, 162.
56 THE VOYAGE OF H.M.S. CHALLENGER.
2. Antedon microdiscus, Bell. This is another species like Antedon elegans which has
the two outer radials united by syzygy, although they were not so described by Bell, who
assigned to the type a formula so unusual for an Antedon,1 (A. 3. 3(3)), that I was led to
examine the spocies for myself, with the result mentioned above. His formula is defective
in another respect besides the all-important omission of the R; for it takes no account of
any arm-divisions beyond the third axillary above the radials, and could not therefore
apply to any species with more than eighty arms. He says, however, " probably as
many as 90 arms in an adult," and nine sets of quaternary arms are represented
in his figured specimen. They are absent, however, in one of the smaller examples
of his type, which for this and other reasons I am disposed to refer to Antedon multi-
radiata. But their presence is nowhere indicated in the formula given by Bell ; and he
also puts the 3 indicating the tertiary or post-palmar series in a bracket, which would
imply that the full number of eight series is not developed on every ray. I much doubt,
however, whether an example of this or of any other type will ever be found with
exactly eighty arms owing to the presence of forty post-palmar axillaries and none
beyond them ; and his formula only tells us that every individual of this species does
not conform to this very regular arrangement.
Bell not only omits all reference to the quaternary arms in his better developed
individuals of this type, but he says of the tertiary arms that " of the three joints the
axillary may or may not be a syzygy." His figured specimen has the full number
(forty) of tertiary arms, and the axillary is a syzygy in each case. But in the smaller
individuals there seem to be some exceptional series of two joints only, the axillary not
a syzygy. This is probably the condition alluded to by Bell, but it would have been
better if he had described it more precisely, for a series of three joints with the axillary
not a syzygy is an arrangement which I have not met with in any Comatula, though it
is to be found in the Pentacrinidse.
3. Antedon loveni, Bell. Bell has given the name loveni" to the form which
appeared as Antedon insignis in his first list f and as this is the host of Myzostoma
coriaceum, the name should be altered in the Report on the Myzostomida by Professor
von Graff.4 On the other hand Antedon loveni of Bell's first list has been since described
by him as Antedon pumila.5
4. Antedon milberti, Mull., sp. Midler's two species, Comatula milberti and
Comatula jacquinoti, appear to me to be identical ; and the second name thus becomes
a synonym of the first.
5. Antedon phalangium, Midler, sp. This Mediterranean species was for a long
time but very imperfectly known, and examples of it were described by Barrett from the
i " Alert " Report, p. 155. 2 " Alert " Report, p. 158.
sProc. Zool. Soc. Loud., 1882, p. 534. 4 Zool. Chall. Exp., part, xxvii., 1884, pp. 14, 18, 40.
6 " Alert " Report, p. 157.
REPORT ON THE CRINOIDEA. 57
Sound of Skye, first as Comatula woodwardii,1 and then as Comahda celtica,2 both of
which names must now lapse.
6. Antedon quadrata. This is the Arctic type which was referred to Antedon celtica,
Barrett, first by von Marenzeller 3 and afterwards by Duncan and Sladen,4 and has since
been rebaptized by myself.5
7. Antedon tenella, Eetzius, sp. This species, which dates back to 1783, was
described by Say in 1825 as Alectro dentata, and is better known in Europe as Alecto
sarsii, Duben and Koren.
8. Antedon variipinna, Carpenter. To this species I now refer my own Antedon
crenulata, together with the Antedon decipiens and Antedon irregularis of Bell. Some
forms of it seem to have but ten arms.
9. Antedon granulifera, Pourtales. There is a syzygy between the two palmar
joints of this species, which escaped the notice of Pourtales.6
10. Antedon insequalis, n. sp. Most examples of this species that I have seen
br
conform to the type A.3.^-, but in one individual there is a single series of two
palmars, the axillary with a syzygy, while the first brachial above it is also traversed by
a syzygy (PI. LI. fig. 2). I much doubt, however, whether this is anything more than an
abnormal development ; but I have recorded it in order to guard against the possibility
of similar forms being subsequently found and described as new species.
Genus AGTINOMETRA.
I. (i) a.p4r.
1. brachiolata 2. pectinata 3. Solaris.
d.(p.)br. . .
(n) a. K. — 2 — 4- paucicirra
(iii) a.R.3.^-^. distincta
a.R.3.^
2
br.
2
multibrachiata novse-guinex typica
T0n two species of Echinodermata, new to the Fauna of Great Britain, Ann. and Mag. Nat. Hist., 1857, ser. 2,
vol. xix. p. 33. 2Ibid., vol. xx. p. 44.
3 Die Coelenteraten, Echinodermen und Wiirmer der k. k. Osterreichisch-ungariscken Nordpol-Expedition, Denkschr.
d. k. Akad. d. Wiss. Wien, 1877 [1878], Bd. xxxv. p. 380.
4 A Memoir on tke Echinodermata of the Arctic Sea to the West of Greenland, London, 1881, p. 75, pi. vi.
tigs. 5, 6.
6 On the Crinoidea of the North Atlantic between Gibraltar and the Faeroe Islands, Proc. Roy. Soc. Edin., 1883-84,
voL xii. p. 375.
6 Bull Mus. Camp. Zobl, 1878, toI. v. No. 9, p. 215.
(ZOOL. CHALL. EXP. — -PART LX. 1888.) OoO 8
58
THE VOYAGE OF H.M.S. CHALLENGER.
II. a. 10 [only ten arms].
blakei
cumingi
III. (i) a. 2
br
2'
a. 2. 2.
br
a. 2. 2. 2.
br
T
echinoptera
meridionalis
maculata
pulchella ( 10 ; &2.^)
stelligera (2.2.2")
5. pulchella (2.(2) ^)
6. rubiginosa
pulchella ( 10 ; k2.2.-^\
stelligera (2.2.(2)^)
br
a.2.2.2.2.^.
nigra
(ii) a.2.2.
a. 2.
elongata
a. 2. 3.
rotalaria
a. 2. 3. 3.
valida
a.2.3.3.3.
2b:(o)
IV. (i) a. 3. 1.1. br.(i).
2
a.3.2&r.
borneensis
discoidea [3.2(p.6r)]
7 coppingeri
Jimbriata
&.3.2(p.br).
discoidea (3.2&r)
lineata (3.2br)
&.3.2(p. p'.br)
multiradiata [3.2(p.
(ii) a. 3.
japonica (3.3)
quadrata
8. parvicirra [3.(3.3)]
a. 3. 2.
a. 3. 2. 2.
multifida
a. 3. 2. 2. 2. 2. 9
. variabilis
a.3.2.3.
grandicalyx
a.3.2.3.2.
alternans
a.3.2.3.3.
10. briareus
divaricata
simplex
lineata \3.2(p.br)~]
multiradiata [3.2 (p.p'. br)]
trichoptera (3.3)
magnifica
REPORT ON THE CRINOIDEA.
59
a.3.-| (|)2.2.
belli
,3l(|),
duplex
,3.1(1)3.3.
11. nobilis
a. 3. 3.
japo7iica (3)
parvicirra (3)
robustipinna
trichoptera (3)
,3.3.2.
littoralis
,3.3.3.
,3.3.3.3.
bennetti (3.3.3.3)
parvicirra [3(3)]
bennetti (3.3.3)
regalis
schlegeli
peroni
1. Actinometra brachiolata, Lam., sp. I refer to this type the form described by
Muller ' under the MS. name Comatula rosea, Mus. Vienna. He noted its close relation-
ship to and possible identity with Lamarck's type, and an examination of both has con-
vinced me that they are really identical.
2. Actinometra pectinata, Linn., sp. I have come to the conclusion that Actinometra
affinis, Lutken, MS., which appeared in my former list,2 is identical with the form which
Retzius referred to the Asterias pectinata of Linnaaus. The Alecto purpurea of Muller3
is probably a young form of the same type.
3. Actinometra Solaris, Lam., sp. The type for which Bell gave a formula under
the name of Actinometra albonotata* is now regarded by him as a variety of Actino-
metra Solaris.6 The following specific names have also been applied to this type at
different times — hamata, imperialis, intermedia, robusta, and strota. (See p. 288.)
4. Actinometra paucicirra, Bell. The form described by Bell under this name8 turns
out to be only the premature stage of the type to which I referred in my Preliminary
Report as Actinometra jukesi? But it has not yet been formally described, and Bell's
name therefore must take precedence.
5. Actinometra pulchella, Pourt., sp. This is the Antedon ptdchella. of Pourtales,
another form of which was also described by him as an Antedon alata.s
1 Abhandl. d. k. Afcad. d. Wins. Berlin, 1847 [1849], p. 250. 2 Proc. Zool. Soc. Lond., 1882, p. 747.
3 Archivf. Naturgesch., 1843, Jahrg. ix. Bd. i. p. 132.
4 Proc. Zool. Soc. Lond., 1882, p. 535. 6 " Alert " Report, p. 165.
6 Ibid., p. 169. 7 Proc. Roy. Soc, 1879, p. 390.
6 Reports on the Results of Dredging under the Supervision of Alexander Agassis, in the Gulf of Mexico, by the
United States Coast Survey Steamer " Blake," Lieutenant Commander C. D. Sigsbee, U.S.N., Commanding, Bull Alus.
Comp. Zool., 1878, vol. v. No. 9, pp. 215, 216.
60 THE VOYAGE OF H.M.S. CHALLENGER.
6. Actinometra rubiginosa, Pourt., sp. This form was originally described as an
Antedon by Pourtales, before the two genera were distinctly separated.1
7. Actinometra coppingeri, Bell. The formula assigned to this type by Bell2 is
that of a ten-armed species, with a syzygy in the third brachial. But the number of
arms varies from twelve to twenty, and there is a syzygy in the second joint above the
distichal axillary.
8. Actinometra parvicirra, Mull., sp. This protean type has been variously
described under the following names — annulata, mertensi, rneyeri, polymorpha,
timorensis, wahlhcrgi.
9. Actinometra variabilis, Bell. This species appeared in my former list8 in the
group (A. 3.3.), this being the formula which I was led to assign to it on the basis of that
previously given by Bell.'
10. Actinometra briareus, Bell, sp. Bell has described this species5 as an Antedon
with the formula A.3.2.(2) ; though the majority of the palmar series are three-jointed
and some of them are followed by another series of the same character.
11. Actinometra nobilis, n. sp. Actinometra dissimilis of Part I.6 appears to be a
varietal form of this t)>p>e.
Some curious points of contrast may be noticed in the two lists given above. There
are three very distinct types of Actinometra in which the two outer radials are united
by syzygy. (l) The ten-armed (Actinometra Solaris) ; (2) those with two distichals
(Actinometra paucicirra) ; and (3) those with three (Actinometra typica). In the
latter case each subsequent division (if present) consists of only two joints united by
syzygy. On the other hand, all the recent species of Antedon yet described which have
the radials a syzygy have three distichals, while the palmars and subsequent divisions
either resemble the distichals, or consist of two articulated joints. Species of Antedon
like Actinometra paucicirra and Actinometra typica are, like those of the Solaris-ty\>Q,
yet to be described.
More than half the species of Antedon belong to the simple ten-armed type with
articulated radials like Antedon eschrichti (PI. XXIV. fig. 11) ; while half the remainder
have only two joints in each of the first three arm-divisions, as in Actinometra conjungens
(PI. XLV. fig. l). But there are not ten described ten-armed species of Actinometra
which have articulated radials, nor ten with two-jointed distichal series. Both these
types, which together include over three-quarters of the species of Antedon, thus
present themselves but rarely in Actinometra.
On the other hand, we find in this genus a much greater number and variety of the
1 List of the Crinoids obtained on the Coasts of Florida and Cuba by the United States Coast Survey Gulf Stream
Expeditions in 1867, 1868, 1869, Bull. Mus. Comp. Zool, 1869, vol. i. No. 11, p. 356.
2 Proc. Zool. Soc. Lond., 1882, p. 535. 3 Ibid., p. 747.
♦ Ibid., p. 535. 6 u Aiert » Report, pp. 155, 163.
6 Zool. Chall. Exp., part xxxii. pp. 110, 111.
REPORT ON THE CRINOIDEA. 01
tridisticnate species than occur in Antedon. I do not know any species of the latter
genus with articulated radials in which there is a fourth post-radial axillary, such as
occurs in Actinometra alternans, Actinometra variabilis, Actinometra magnified, and
Actinometra bennetti ; and it is decidedly rare to find a third axillary ; while the
singular variations presented by Actinometra belli and Actinometra nobilis are
altogether unknown in Antedon.
The above tables show that it is possible to make a preliminary classification of the
species of Comatuke by using the characters of their successive arm-divisions.
But how are we to deal with the seventy odd species of Antedon which have only
ten arms, or with the thirty more which have bidistichate primary arms ? The
characters of systematic value which may be employed for this further classification are
those of the cirri, arms, and pinnules. The number of the cirri themselves and also that
of their component joints are very useful characters within certain limits. Antedon
valida and Antedon parvipinna, which are both figured on PI. XV., are obviously cmite
distinct specific types; and the same maybe said of Antedon alternata and Antedon
incerta, represented on PI. XVIII. , not only as regards the cirrus-characters, but with
respect to the pinnules also.
The shape and the relative sizes of these latter organs, especially at the bases of the
arms, often afford characters of much systematic value, as will be seen by comparing the
flagellate lower pinnules of Antedon quadrata and Antedon australis (PI. XXVII.
figs. 8-16) with the stiffer ones of Antedon occulta and Antedon variipinna (PI. XLVIII.
figs. 2, 3) ; while those of Antedon valida, Antedon incerta, and Antedon macronema
(PI. XV. figs. 5, 6; PI. XVIII. fig. 5; PI. XXXVIII. fig. 4) are of an altogether
different type from either of the others just mentioned.
Another very useful character for systematic purposes is to be found in the shape of
the arm-joints. In one large group of Antedon-species the radial axillaries and the next
few joints beyond them have their apposed sides much flattened, as is well seen in
PI. XV. fig. 6. In the absence of this very striking peculiarity, the shape of the arm-
joints, as seen from the dorsal side, is often of much use in classifying species. Thus, for
example, the elongated joints of Antedon phalangium (PI. XXVIII. fig. l), the short
compressed triangular joints of Antedon patula (PI. XLIIL), and the rounded joints of
Antedon variipinna (PI. XLIX. fig. 1) all afford good specific characters ; while in the
genus Actinometra the contrast is strong between the short discoidal joints of Actino-
metra Jimbriata and the triangular ones of Actinometra elongata (PI. LVII. fig. 2;
PL LXII. fig. 3).
The condition of the ambulacra in the arms and pinnules is also of much use in
classification. Thus, for example, Antedon accela, Antedon incerta, Antedon insequalis,
and other forms have both side plates and covering plates on the pinnule-ambulacra,
which are often better defined than in the Pentacrinida) ; while in other species, such as
62 THE VOYAGE OF H.M.S. CHALLENGER.
Antedon eschrichti and Antedon carinata, the ambulacral plating is reduced to small and
irregular spicules without definite arrangement.
By means of these various characters, then, it is possible to subdivide the large
specific groups, both in Antedon and in Actinometra, and to make out a detailed
classification of the numerous species belonging to each genus. Except in a few cases,
however, it would be premature as yet to make any attempt at distinguishing the
structural and the adaptive characters respectively among those which we are at present
inclined to regard as of specific value.
VI.— DESCRIPTION OF THE SPECIMENS.
Class CRINOIDEA.
Order NEOCPJNOIDEA.
Family Comatulid^e, d'Orbigny, 1852; emend. P. H. Carpenter, 1888.
Crinoids with the calyx closed below by the enlarged top joint of the larval stem,
which develops cirri and generally separates from the stem joints below it, so that the
calyx is free. The basals may form a more or less complete ring on the exterior of the
calyx, or be only represented by an internal rosette. Five or ten rays, either simple or
more or less divided. The first axillary is the second, or (very rarely) the first joint
above the calyx-radials. Definite interradial plates usually absent.
The mouth central, except in one genus.
Remarks. — The family Comatulidse, which was established by d'Orbigny1 in 1852, is
practically equivalent to a group which was proposed more than twenty years previously
by de Blainville,2 under the name of the " Asterencrinides libres." So far as I am aware,
de Blainville was the first author to make any definite separation of the Feather-stars
from the remaining Stellerids.
He divided this order into three families, the Asteridea, the Asterophydea, and the
Asterencrinidca, which last Miller had previously called Crinoidea.
De Blainville further subdivided the Asterencrinidea into two sections, the first of
which was " les Asterencrinides libres." He defined it as having a " corps libre, et sans
tige qui servirait a le fixer " ; and he referred to it the single genus Comatula, Lamarck.
In the great work of Goldfuss,3 which was published a few years later, there is,
however, no special separation of the genus Comatula from the other Stellerids, and it
simply appears as the first genus in his order " Asterites liberi," altogether separate from
the Stalked Crinoids, which are classed as the Stilasteritae, though the resemblance between
them and Comatula did not escape the notice of Goldfuss. He gave an account of the
anatomy of two recent species, and referred to the genus some fossils from Solenhofen,
1 Cours elementaire de Geologie et de Paleontologie stratigraphique, 1852, t. ii. fasc. i. p. 138.
2 Diet. d. Sei. Nat., 1830, t. lx. p. 229.
3 Petrefacta Germanise, Diisseldorf, 1826-35, vol. i. p. 201.
64 THE VOYAGE OF H.M.S. CHALLENGER.
while he also gave the names Solanocrinus and Glenotremites to some other fossil forms
of which only parts of the calyx were preserved. Some of Goldfuss's species were made
the types of new genera by Agassiz,1 and Miiller referred to them as follows in his first
communication to the Berlin Academy on the subject of the Crinoidea :2 — ■
"Die ungestielten Crinoiden mit Armen bilden 3 Familien (l) Articulata, gen.
Comatula, Lam., und Comaster, Ag. (2) Costata mit schaligem gerippten Kelch und
entgegengesetzten Pinnulse, wovon sonst bei alien ubrigen Crinoiden kein Beispiel
vorkommt, gen. Saccocoma, Ag. (3) Tessellata, gen. Marsupites."
The above passage must not be understood as meaning that " Articulata," Miiller, is
a synonym of " Comatulidae," d'Orbigny, and should therefore take precedence of it. For
there were Stalked as well as Unstalked Crinoidea Articulata and Crinoidea Tessellata ;
and in the subsequent memoir on Pentacrinus Miiller made these the two primary
divisions of the Crinoidea, altogether apart from the question of the presence or absence
of a stalk. But in his second preliminary communication 3 he made a passing reference
to " die in der Familie cler Comatulinen enthaltenen Gattungen Comatula und
Comaster," the latter genus being regarded by him as identical with Solanocrinus,
Goldfuss.
Miiller never said anything more definite about the family Comatulinse, however,
though he recognised Alecto and Actinometra as two subgenera of Comatula, Lamarck.
The Stalked Crinoids remained in an equalhy chaotic condition for many years. But
about 1850 Bronn and d'Orbigny made separate attempts to class them into families.
The former author4 established the family Astylidse, though without defining it, and
referred to it the recent Comatula and three fossil genera. Among these were Marsupites
and Saccocoma, both of which, as we have seen above, had been made the types of
separate families by Miiller. This was also clone by d'Orbigny,6 who divided the Crinoidea
into ten families, one of which was the Comatulidse, and this name, or its shortened form
" Comatuke," has been in use for the family of the Feather-stars ever since, though the
number of genera referred to the family has varied enormously.
D'Orbigny included in it the recent Comatula, Lamarck, and three other genera which
were based on the characters of various fossil species. None of these, however, are now
recognised ; and the same is true of a number of genera established by other palaeonto-
logists ; for with one exception all the true Comatulids which have been as yet discovered
in the fossil state can be referred either to de Freminville's genus Antedon, which has
priority over Comatula, or to Midler's subgenus Actinometra, which has gradually acquired
generic rank. The exception is the five-armed species from the Valangien of Switzerland,
1 Mem. Soc. Sci. Nat. Neuchatel, 1835, t. i. p. 193.
2 Monatsber. d. k. preuss. Akad. d. Wiss. Berlin, 1840, p. 91.
3 Ibid., 1841, p. 179.
4 Lethasa Geognostica, 1851, Bd. i. Tb. 1, p. 23.
6 Cours elementaire de Paleontologie et de Geologie stratigraphique, 1852, vol. ii. fasc. i. p. 138.
REPORT ON THE CRINOIDEA. 65
which represents a recent generic type first discovered by Semper and named by him
Ophiocrinus. In consequence, however, of the preoccupation of this name it has been
since changed to Eudiocrinus. Thus then the great number of generic names which
have been given to the fossil Comatulse become reduced to three, Antedon, Actinometra,
and Eudiocrinus. Three new genera have been established by myself for new types of
recent Comatuke, viz., Atelecrinus, Promachocrinus, and Thaumatocrinus ; and these
six are all that could strictly be included in the family Comatulidas until quite recently.
Pictet1 has also referred to it both Marsupites and Saccocoma, but Dujardin and
Hupe2 removed Marsupites to the Cyathocrinidae, and added to the Comatulidae
the sessile Eugeniacrinus and its allies, which had been grouped under the Eugenia-
crinidae in Bronn's " Thierreich." Zittel a restored this family to its proper position
and restricted d'Orbigny's name to the Feather-stars proper; while Saccocoma was
replaced in Midler's group, the Costata, which had been established for- its: reception
in 1840.
Quite recently, however, it has become necessary to add a seventh genus to the
family, viz., the fossil Thiolliericrinus, which represents a permanent form of a late
stage in the development of the Antedon-laYva. It has been well described by
de Loriol4 as an Antedon with a Bourgueticmius-stera. The stem-joints of the
larval Antedon are closely similar to those which are characteristic of the family
Bourgueticrinidae, their faces bearing strong transverse ridges with a deep fossa on
each side.
In ordinary Comatulae the centro-dorsal, after separating from the stem beneath it,
soon loses all trace of its previous connections, owing to a more or less extensive deposi-
tion of limestone at its dorsal pole ; whereas in Thiolliericrinus the connection between
the lower stem-joints and the cirrus-bearing centro-dorsal seems to have been maintained
much longer, if not throughout life. For the under surface of the centro-dorsal bears a
well-developed articular facet like that on an ordinary stem-joint of Bourgueticrinus or
Rhizocrinus. It would appear therefore that the centro-dorsal with the few cirri which
were developed upon it remained permanently attached to the stem below, so that
Thiolliericrinus would represent the permanent condition of an Antedon-larva during the
development of its second whorl of cirri. We cannot be absolutely certain about its
characters, however, until an entire example of the genus has been discovered. But
the presence of an articular facet on the under surface of its centro-dorsal is a feature
which is sufficient to distinguish it very markedly from the six genera of recent
Comatulae.
1 Traite de Paleontologie, Paris, 1857, vol. iv. p. 287.
2 Op. cit., p. 186.
3 Handbueh der Palaeontologie, Bd. i., Abth. 1, p. 395.
4 Description de quatre Echinodermes Nouveaux, Mem. Soc. Pal. Suisse, 1880, vol. vii. p. 10.
(ZOOL. CHALL. EXP. — PART LX. — 1887.) 000 0
66 THE VOYAGE OF H.M.S. CHALLENGER.
The general relations of these seven genera of Comatulidse are expressed in the
following table : —
I. Centro-dorsal has no articular facet on its lower surface.
A. Five rays.
i. Mouth central or subcentral. Oral pinnules have no comb.
a. Kadials separated by interradials, . . . . 1. Thaumatocrinus, n. gen.
b. Radials united laterally.
(1) Basals persist as a closed ring. No pinnules on
lower brachials, . . . .2. Atelecrinus, n. gen.
(2) Basal ring incomplete or invisible externally.
a. Five arms only, . . . .3. Eudiocrinus, Carpenter.
j3. Ten arms, . . . . .4. Antedon, de Freminville.
ii. Mouth excentric or marginal. Oral pinnules have a terminal
comb, . . . . . . .5. Actinometra, Miiller.
B. Ten rays, . . . . . . . .6. Promarhocrinus, n. gen.
II. Centro-dorsal has an articular facet below, . . . . .7. Tldolliericrinus, fitallon.
Genus 1. Thaumatocrinus, P. H. Carpenter, 1883.
1883. Thaumatocrinus, P. H. Carpenter, Phil. Trans., 1883, pt. iii. p. 919, pi. lxxi.
Definition. — Calyx composed of a centro-dorsal, basals, radials, and primary inter-
radials, the latter resting on the basals and so separating the radials laterally. That on
the anal side bears a short jointed appendage. Mouth central and protected by five
large oral plates which occupy the greater part of the disk, and are separated from the
calyx-intei'radials by two or three rows of small irregular plates. Five arms only.
Remarks. — Thaumatocrinus has already been described and its peculiarities discussed
in Part I. of this Eeport (pp. 370-372), and it is not necessary therefore to refer again to
the reappearance of certain Palseocrinoidal characters in this remarkable genus. As
compared with the more typical Comatulse it is peculiar in having persistent basal and
oral plates, the latter occurring in no other Comatulid, and in the simplicity of the rays,
which remain undivided, so that there are only five arms, as in Eudiocrinus (PI. VII.).
Thaumatocrinus renovatus, P. H. Carpenter, 1883 (fig. 1; Part I. pi. lvi. figs. 1-5).
Description of an Individual. — The total width of the calyx across the disk is barely
2 mm.; and the height of the centro-dorsal and radials together is about the same. The
former is rounded below, with its central canal completely closed up, so that it must
have been detached for some little time from the remainder of the stem. The bases of
half a dozen cirri are attached to it, and there are pits for the reception of two or three
more. In the largest stump which is preserved the first two joints are quite short, as is
usually the case in all cirri ; but the third reaches a length of 1"5 mm., so that the cirri
must have been very like those of some species of Eudiocrinus, which have a succession
of very long joints followiug the short basal ones (PI. VII. figs. 2, 7).
REPORT ON THE CRINOIDEA.
67
The basals are almost trapezoidal, much wider below than above, and in contact with
one another by their truncated lower angles. The middle of the lower edge of each is
slightly tubercular. On their narrow upper edges rest the interradials. which are oblong
and a little higher than wide. Four of them terminate in a free edge at the margin of
the disk, where they are in contact with the lowest anambulacral plates. But that on the
anal side bears a small tapering appendage of four or five joints, the last of which seems
to end freely. The radials are larger than the interradials and somewhat strongly arched.
There is a muscular articulation between them and the first brachials ; but the articulation
between these and the next joints appears to be only bifascial. The arm-joints are long,
slender, and cylindrical. One arm seems to be broken at a syzygy in the sixth brachial ;
while another has a syzygy in the fourth, and again in the eighth brachial. The second
brachial bears the first pinnule, which is on the right side in three arms, and on the left
in the other two. The pinnules are very delicate and composed of long slender joints.
Fig. \.—Thaumatocrinus rcnovatus, P. H. Carpenter. A, The calyx, anal side. B, The disk from above, aa, anal
appendage; an, anambulacral plates; at, anal tube; b, basal ; i2, second brachial ; al, centro-dorsal ; i, interradial;
o, oral ; r, radial, x 15.
The central portion of the disc is occupied by five relatively large oral plates which
stand up around the peristome ; while between them and the margin are two or three
irregular rows of small anambulacral plates, some of them extending up on to the lower
part of the long anal tube. The brachial ambulacra are not plated, however, and lie in
the arm-grooves, close down between the muscles but with no traces of sacculi.
Colour in spirit. — Dirty white.
Locality.— Station 158, March 7, 1874; kit, 50° 1' S., long. 123° 4' E.; 1800
fathoms; Globigerina ooze ; bottom temperature, 33°-5 F. One specimen, much muti-
lated and probably young.
68 THE VOYAGE OF H.M.S. CHALLENGER.
Genus 2. Atelecrinus, P. H. Carpenter, 1881.
1869. Antedon, Pourtales (pars), Bull. Mus. Comp. Zobl., 1869, vol. i. No. 11, p. 356.
1878. Antedon, Pourtales {pars), Bull. Mus. Comp. Zobl., 1878, vol. v. No. 9, p. 214.
1881. Atelecrinus, P. H. Carpenter, Bull. Mus. Cornp. Zobl., 1881, vol. ix. No. 4, p. 16.
1882. Atelecrinus, P. H. Carpenter, Journ. Linn. Soc. Lond. (Zool.), 1882, vol. xvi. p. 488.
Definition. — Centro-dorsal acorn-shaped, and bearing five vertical double rows of
cirrus-sockets, those of each row alternating with one another, and with those of adjoining
rows. They have horseshoe-shaped rims, the arches of which are directed upwards
while the two ends slant downwards and outwards. Radials separated from the centro-
dorsal by a complete circlet of basals. The first six or more brachials bear no pinnules.
Remarks. — The first example of this genus which was actually obtained was dredged
by Pourtales1 in 1869 off Cojima on the coast of Cuba. Two small ten-armed Comatulae
were brought up from a depth of 450 fathoms, and were briefly described by Pourtales
under the name of Antedon cubensis. But the description given by him only applies to
the larger and more perfect specimen, which differs considerably from the smaller and
much mutilated one. He seems to have recognised that the two were different, for in
his description2 of the Crinoids obtained by the "Blake" expedition of 1877-78 he wrote
as follows : — " To this species (i.e., Antedon cubensis) I refer j^rovisionally two specimens
very much mutilated, having lost the cirrhi and the arms, differing somewhat from my
type specimen, but possibly the differences may be due to age." He then described an
individual dredged at Station 43 ("Blake") in 339 fathoms, to which I shall refer directly,
and added that a smaller and equally mutilated one had been previously dredged by
himself in 450 fathoms near Havana.
These two specimens are quite different from the type of Antedon cubensis, and also,
though in a less degree, from one another. Not only are the first radials visible, and the
second but little shorter than broad, as was mentioned by Pourtales, but the first radials
are separated from the acorn-shaped centro-dorsal by a complete circlet of basals, and
there are no pinnules upon any of the first six arm-joints, which are the only ones
preserved. An equally mutdated specimen of Pourtales' second type was dredged by
the Challenger (1873) in 350 fathoms, off Barra Grande (PL VI. fig. 7); seven more
perfect ones, making nine in all, were obtained off Nevis, St. Lucia, and Granada, during
the cruise of the "Blake" in 1878-79, between 291 and 422 fathoms; while a single
example of a third species (PI. VI. fig. 5) was dredged by the Challenger in the
neighbourhood of Fiji, in the year 1874.
These eleven individuals, representing three different species, are distinguished from
all other living Coniatulse by certain very definite morphological peculiarities, which
impart an interest to this type second only to that of the archaic Thaumatocrinus. Its
1 Bidl. Mm. Comp. Zobl, 1869, vol. i. No. 11, p. 356. * Ibid., 1878, vol. v. No. 9, p. 214.
REPORT ON THE CRTNOIDEA. GO
two leading characters are — (1) the persistence of the embryonic basals which do not
undergo transformation into a rosette, but remain on the exterior of the calyx between
the centro-dorsal and the radials ; and (2) the absence of pinnules from the lowest joints
of the arms (PI. VI. figs. 5, 7). A third character, of no great morphological value, but
important from its apparent constancy, is the acorn-like shape of the centro-dorsal, and
the arrangement of the cirrus-sockets upon it in alternating double rows, with the ends
of their horseshoe-like rims projecting somewhat outwards.
The extent of development of the basals of Atelecrinus varies with the size of the
individual, apparently diminishing with age as in the Pentacrinoid larvse of ordinary
Comatulse (PI. XIV. figs. 5-7). In the smallest specimen of Atelecrinus balanoides they
are wide but low pentagons which fall away very rapidly from their interradial apices to
the points where they meet one another beneath the radials. The middle of each basal
rests on the top of one of the interradial ridges at the upper end of the centro-dorsal,
just as the basals of Pentacrinus rest on the upper ends of the interradial ridges of the
stem. In older individuals, however, just as in the Antedon-larva (PI. XIV. figs. 5-7),
the amount of the first radials which is visible on the exterior of the calyx becomes
relatively less and less, and the same is the case with the basals. These are best
described as triangular, with their lower angles extended so as to meet those of their
fellows and separate the radials from the centro-dorsal by what is practically little more
than a line, only visible at all under specially favourable conditions of light. Each of
the basals, when isolated, has the form of a short triangular prism with a flattened plate-
like extension on each side. They are in complete lateral contact, so as to form an
unbroken ring on the under surface of the radial pentagon, very much as in Pentacrinus
alternicirrus or in Pentacrinus wyville-thomsoni. Atelecrinus cubensis has comparatively
large basals which are of nearly uniform height (0-5 mm.) all round the calyx, rising
very slightly at the interradial angles; while in Atelecrinus ivyvillii each basal is slightly
arched, with its apex interradial, and it is only in contact with the outer edge of the
centro-dorsal at the interbasal sutures (PI. VI. fig. 5).
All three species agree, however, in the absence of any rosette and in the persistence
of the basals upon the exterior of the calyx, a feature which appears in no other recent
Comatula except Thaumatocrinus and the very doubtful Comaster ; while a further
peculiarity lies in the complete closure of the basal ring so as to separate the radials
altogether from the centro-dorsal. Several, if not all, fossil Comatulae have persistent
primary basals in the form of prismatic rods, which meet one another in the centre of the
under surface of the radial pentagon, and extend outwards towards its interradial angles.
But they do not always reach the periphery so as to appear externally between the
radials and the centro-dorsal, as they gradually thin out ; and there is only one described
form in which there is a complete ring of united basals on the exterior of the calyx.
As regards the characters of its calyx, therefore, Atelecnnus is certainly to be
70 THE VOYAGE OF H.M.S. CHALLENGER.
regarded as a permanent larval form. The absence of the pinnules from the lower parts
of the arms points to the same conclusion, as has been explained elsewhere.1
I have only been able to examine the disk in the two Challenger specimens, and in
one of these it is not very well preserved. But they both agree in the slightly excentric
position of the mouth, and in the large size of the peristome, so that the anal tube is
pushed backwards behind the centre about as much as the mouth is in front of it
(PL VI. figs. 4, 6).
Unlike the two Endocyclic Comatulse with five rays and a rosette (Antedbn and
Eudiocrinus), Atelecrinus is not a littoral type at all, nor does it extend upwards above
200 fathoms. On the other hand it is not known to occur below 610 fathoms ; so that
bathymetrically it falls very far short of the archaic Thaumatocrinus (1800 fathoms).
Apart from this last type, however, the geographical range of Atelecrinus, although
fairly extensive, is the least so of the five-rayed Coniatulse. In the Caribbean Sea and
the East Atlantic it ranges from 24° N. to 9° S.; while it also occurs in the Pacific near
Fiji in 19° S. If the fossil calyx mentioned by Schliiter 2 as having persistent basals also
belong to this genus, it will date back to the Cretaceous period.
The three existing species of A telecrinus may be distinguished from one another as
follows : —
I. Second radials transversely oblong and but little incised. Basals not specially
prominent at the angles of the calyx, . . . . .1. balanuides, n. sp.
II. Second radials markedly incised and about as long as wide.
A. Basals separated from the centro-dorsal at its interradial angles, . 2. wyvillii, n. sp.
B. Basals produced outwards at the interradial angles, . . .3. eubensis, Pourtales, sp.
1. Atelecrinus balanoides, n. sp. (PL VI. figs. 6, 7).
1879. Antedon eubensis, Pourtales (pars), Bull. Mus. Comp. Zobl., 1879, vol. v. No. 9, p. 214.
1881. Atelecrinus balanoides, P. H. Carpenter, Bull. Mus. Comp. Zool., 1881, vol. ix. No. 4,
p. 16, pi. i. figs. 1-6.
1882. Atelecrinus balanoides, P. H. Carpenter, Journ. Linn. Soc. Lond. (Zool.), 1882, vol. xvi.
p. 489.
Centro-dorsal acorn-shaped, reaching 5 mm. high by nearly 35 mm. in diameter.
It bears five vertical double rows of cirrus sockets, the upper ends of which are separated
by more or less distinct interradial ridges. Four to six sockets in each row, the dorsal
pole, though rough, being free from functional sockets. The ends of their horseshoe-
shaped rims slant downwards and outwards, but are much more prominent in some
individuals than in others.
The cirri have three or four quite short, almost triangular basal joints. The next is
two or three times as long as wide, and its successors are much elongated, reaching
2"5 mm., with a slight tendency to overlap one another on the ventral side of the cirrus.
1 Bull. Mus. Comp. Zobl., 1882, vol. ix. No. 4, pp. 14, 15. 2 Zeitschr. d. deutsch. gcol. Gesellsch, 1878, p. 60.
REPORT ON THE CRINOIDEA. 71
There are probably about thirty-five joints, the length much exceeding the breadth till
the penultimate, which is followed by a very small terminal claw. The last six joints
taper rapidly.
The basal ring is a very thin plate, rising at the interradial angles into triangular
elevations, which are produced slightly outwards and rest upon the upper ends of the
interradial ridges of the centro-dorsal. First radials broad and tolerably flat, their size
varying with the age of the individual. Second radials more arched, oblong, and quite
free laterally, their breadth in the adult being one and a half times their length.
Axillaries pentagonal, sometimes twice the length of the second radials, into which they
have a slight backward projection. Their width is about equal to their length, but their
proportions and also those of the second radials vary slightly in different individuals.
First brachials well separated laterally, with their inner sides shorter than the more
rounded outer ones.
Second brachials irregularly quadrate, projecting slightly backwards into the first.
The following joints have oblique ends and markedly unequal sides. Except in the
syzygial joints, the length is at first less than the breadth, but gradually becomes more
equal, and exceeds it after the fifteenth joint. Terminal joints relatively longer and
more equal-sided. Arm-bases smooth, but the middle and later joints overlap slightly.
The first syzygium on the third brachial. The following syzygies at intervals of from
one to six, usually of two or three joints.
First pinnule nearly always on the twelfth brachial, and consisting of about a dozen
elongated joints. The following ones increase in size and in the number of joints,
decreasing again towards the arm-ends. The lower joints of the middle and later
pinnules bear irregular spinous processes on their dorsal edges.
Mouth somewhat excentric, and surrounded by a large peristome. A little way
behind this is the anal tube, which is also slightly excentric in position. Disk 6 mm. in
diameter. In the Challenger specimen a very few minute calcareous granules are visible
on its ventral surface, and also on its sides between the rays. The " Blake " specimens
are more naked. The brachial ambulacra lie close down upon and between the muscular
bundles, and have a few scattered sacculi at their sides. Colour of skeleton white or
brownish-white.
Locality. — Station 122, September 10, 1873; off Barra Grande ; lat. 9° 5' S., long.
34° 50' W.; 350 fathoms; red mud. One specimen. Also obtained by the U.S. Coast
Survey steamer "Blake" at five stations in the Caribbean sea, between 291 and 422
fathoms.
Remarks. — Although this species was not dredged till four years after Pourtales had
published his description of Antedon cubensis, I have preferred to regard it as the
type of the genus Atelecrinus for the following reasons. Pourtales' description of
72 THE VOYAGE OF H.M.S. CHALLENGER.
Antedon cubensis, which has pinnules on the second and following brachials, does not
correspond to the characters of the smaller specimen which he called by this name, as it
has a complete basal ring, and there are no pinnules on any of the arm -joints which are
preserved; while another reason is that the dredgings of the "Blake" in the years
1877-79 have led to the discovery of eight examples of Atelecrinus balanoides, all of
them better preserved than the single individual which was referred by Pourtales to
Antedon cubensis.
The first specimen of Atelecrinus balanoides known to science was dredged by the
Challenger in 1873 ; but its cirri had disappeared, together with the whole of the arms
above the fifth brachials (PL VI. fig. 7). The Pacific species {Atelecrinus wyvillii) was
not much better (PL VI. fig. 5), and it was not till I received the " Blake " collection in
1880 that I was able fully to reahse the singular peculiarities of the type represented by
the two Challenger specimens which are figured on PL VI.
The distinctive characters of Atelecrinus balanoides are (l) the transversely oblong-
shape of the second radials, wdiich are but slightly incised to receive the bluntly angular
proximal edges of the axillaries ; and (2) the outline of the lower part of the calyx, which
slopes uniformly downwards from the radials on to the centro-dorsal, without the basals
being specially prominent at the interradial angles as they are in Atelecrinus cubensis.
The difference is very much of the same kind as that between the basals of Pentacrinus
wyville-ihomsoni and Pentacrinus miilleri respectively.
The nine individuals of Atelecrinus balanoides which I have examined, all agree
very well in their general characters, but differ considerably in the relative proportions
of the two outer radials and of the lowest brachials respectively. In all of them which
have enough of the arms preserved, the first pinnule is on the twelfth brachial, except in
one arm of one individual, in which the tenth joint bears the first pinnule.
2. Atelecrinus wyvillii, n. sp. (PL VI. figs. 4, 5).
1882. Atelecrinus wyvillii, P. H. Carpenter, Journ. Linu. Soc. Lond. (Zool.), 1882, voL xvi. p. 492.
Description of an Individual. — Centro-dorsal acorn-shaped, 4 mm. high by 3 mm.
wide. The double rows of cirrus-sockets are well separated from one another by
intervening spaces, and do not reach the dorsal pole. Four, or rarely five, sockets in
each row, the ends of which stand out prominently and give a serrate appearance to the
lateral edge of the plate. The upper portion is uniformly smooth, without any interradial
ridges ; but the edge is marked by five slight incisions situated interradially.
The basals are nearly uniform in height throughout their whole width, but are some-
what arched in form. The apex of each arch is interradial, and the interval between it
and the notched edge of the centro-dorsal below is only occupied by perisome. Hence
the basal ring is really only in contact with the centro-dorsal at its five lowest points,
REPORT ON THE CRINOIDEA. 73
i.e., at the interbasal sutures, immediately beneath the middle points of the first radials.
The latter have exceedingly high muscle-plates projecting inwards ; but their dorsal
surface is barely half as long as that of the second radials. These are nearly square, but
deeply incised to receive the strong backward projections of the axillaries, which are
roughly rhombic and slightly wider than long.
First brachials well separated laterally, with the inner sides much shorter than the
outer ones, and the distal edge much incised to receive the strong backward projections
of the quadrate second brachials. The following joints have markedly unequal sides,
with a syzygy in the third or fourth, and again in the fifth, sixth, or seventh brachial.
Disk almost naked, 4 mm. in diameter. Mouth somewhat excentric and surrounded
by a large peristome, immediately behind which is the anal tube. Brachial ambulacra
close down upon and between the muscular bundles. Skeleton light brownish-white.
Locality.— Station 174c, August 3, 1874; lat. 19° 7' 50" S., long. 178° 19' 35" E.;
610 fathoms ; coral mud ; bottom temperature, 39° F. One mutilated specimen.
Remarks. — This type differs from the other two species of the genus in the greater
squareness of the second radials, and in the curious relation of the basals to the centro-
dorsal. They are of uniform height, as in Ateleerinus cubensis, but are not in contact with
the centro-dorsal at the interradial angles of the calyx, being separated from it on the
exterior by a gap which is filled up by perisome (PI. VI. fig. 5). Apart from its
purely morphological importance, this Pacific species is also interesting as showing the
wide distribution of the genus ; and it is the only one of the three which is known as yet
to extend below the limit of the continental line (500 fathoms), though each of the others
has been dredged below 400 fathoms.
*&v
Genus 3. Eudiocrinus, P. H. Carpenter, 1882.
1868. Ophiocrinus, C. Semper, Archiv f. Naturgesek., 1868, Jalirg. xxxiv. Bd. i. p. 68.
1869. Comatula (Ophiocrinus), P. de Loriol, Denksckr. d. allg. Schweiz. Gesellsch. f. d. ges. Nature".,
1869, Bd. xxiii. p. 57.
1879. Ophiocrinus, P. H. Carpenter, Proc. Roy. Soc, No. 194, 1879, p. 385.
1879. Ophiocrinus, P. de Loriol, Monographie des Crinoi'des fossiles de la Suisse, Geneva, 1877-79, p. 277.
1882. Eudiocrinus, P. H. Carpenter, Journ. Linn. Soc. Lond. (ZooL), 1882, voL xvi. p. 493.
1883. Eudiocrinus, E. Perrier, Comptes rendus, 1883, t. xcvi. No. 11, p. 725.
1886. Eudiocrinus, E. Perrier, Les Explorations sous-marines, Paris, 1886, p. 275.
Definition. — Centro-dorsal and calyx like those of Antedon; but the radials bear the
brachials directly without the intervention of axillaries, so that there are only five
undivided arms. Mouth central. Sacculi abundant, scanty, or absent altogether.
Remarks. — The genus Ophiocrinus was established by Semper in 1868 for an elegant
little Comatula with five undivided rays, which he had discovered in the Philippine
Islands ; and in the following year a fossil species was described by de Loriol from the
(ZOOL. CHALL. EXP. PART LX. 1887.) Ooo 10
74 THE VOYAGE OF H.M.S. CHALLENGER.
Neocomian of Switzerland. The generic value of the type was doubted by Schliiter j1
and I had formerly myself some hesitation in regarding it as equivalent to Antedon,
Actinometra, and Promachocrinus.2 For there is no definite character, except the
simplicity of the rays, which can separate Eudiocrinus from the ordinary ten-armed
Antedon ; and in one of the three species of the ten-rayed Promachocrinus the rays divide
so as to form twenty arms (PI. LXX.), while in the two others there are ten undivided
rays (PL LXIX. figs. 5, 9, 10). But this character alone would hardly justify the
separation of the simpler type of Promachocrinus from the twenty-armed form ; while
I have an abnormal specimen of an Antedon with only nine arms, owing to one of the
rays not dividing, which is the case with all the rays of Eudiocrinus.
Nevertheless, it sometimes happens that a character, which is only of specific value
in one type, may be of generic value in another. Five recent species of Eudiocrinus
are known, four of which range from Japan into the South Pacific Ocean (lat. 37° S.),
while one occurs in the East Atlantic, and another has been found fossil in the
Neocomian of Switzerland. The simplicity of the rays thus appears to be a character
of some morphological importance, and I am, therefore, disposed to admit the generic
position which was originally assigned to the type by Semper. Unfortunately, however,
it cannot continue to bear the name by which he described it. For Salter, fifteen years
before Semper's description of Ophiocrinus, had designated by the same generic name
an obscure Crinoid from the Devonian of South Africa ; and the confusion thus
existing was increased by the posthumous publication in the year 1878 of the late
Professor Angelin's monograph of the Swedish Silurian Criuoids, in which the name
Ophiocrinus is connected with a third and totally distinct type.
Professor Semper's genus being thus preoccupied, I proposed in 1882 to call the type
Eudiocrinus (evStos, calm), in allusion to the fact that the four recent species of it, which
were then known, were limited to the Pacific Ocean. Curiously enough, however, a
few months before I suggested this name, several specimens of a new species of Eudio-
crinus were dredged by the French exploring vessel " Travailleur " in the Gulf of Gascony,
and, therefore, in European Seas. The type was naturally designated as Eudiocrinus
atlanticus by Professor Perrier,3 who gave a brief description of the characters which
distinguish it from the Pacific species.
Eudiocrinus, like Antedon, has a central mouth (PI. VI. fig. 2), and a more or less
hemispherical or conical centro-dorsal, an isolated specimen of which could not be
distinguished from the corresponding part of an Antedon (PI. III. fig. 7a; PL VI. fig. 1;
PL VII. figs. 1, 3, 4). The radials, however, in the only recent species which I have
been able to examine, differ slightly from those of the ordinary Antedon-type which is
illustrated on Pis. I. -IV. The articular faces are low relatively to their width (PL III.
1 Zeitschr. d. deutsch. geol. Gesellsch., 1878, p. 4(1. s Quart. Journ. Gcol. Soc, 1879, vol. xxxvi. p. 41.
3 Comptes rendus, 1883, t. xcvi. No. 11, p. 725.
REPORT ON THE CRINOIDEA. 75
fig. 7a), a character which presents itself in Antedon carinata (PI. III. fig. la) and in
Antedon macronema (PI. IV. fig. 3a), and is more especially distinctive of the genus
Actinometra, in which the muscle-plates, well marked in Eudiocrinus, are very much
reduced in size (PI. V. figs. 1-5, b).
The special peculiarity of the calyx in Eudiocrinus semperi, however, is the way in
which the muscle-plates stand up above the sides of the radials, owing to their edges
being strongly folded in towards the central articular ridge which separates them
(PI. III. figs. 7a, 7c). In many species of Antedon the articular facets of adjacent
radials are in close contact along the whole length of their sides, as for example in
Antedon eschrichti (PL I. fig. 8a), Antedon basicurva (PI. II. fig. 2a), and Antedon
breviradia (PL III. fig. 4b). But in other cases the ventral edges of the muscle-plates
are more or less folded outwards from the centre of the calyx, so that its interradial
angles are marked by five notches, which lie at the upper ends of the sutures between
the radials as in Antedon antarctica (PL I. figs. 6a, 6b), Antedon incisa and Antedon
angusticalyx (PL II. figs, la, Id, 4a, Ad), the young Antedon breviradia and Antedon
quinquecostata (PL III. figs. 5a, 5c, 6c, 6d). But in Eudiocrinus semperi this notch
is continued down to the dorsal surface of the radials as a wide groove between the
everted muscle-plates of every two adjacent radials (PL III. figs. 7a, 7c) ; so that in a
dorsal view of the calyx (PL III. fig. 7b) its interradial angles are not sharp but deejay
incised. An indication of the same character appears in Antedon quinquecostata (PL III.
fig. 6b) ; but on the other hand the young calyx of Antedon breviradia, which has the
ventral edges of its muscle-plates strongly folded outwards (PL III. figs. 5a, 5c), pre-
sents a very sharply pentagonal outline in dorsal view (PL III. fig. 56). The same is
the case in Antedon carinata, which has rather markedly everted muscle-plates (PL III.
figs, la, lc, id); while on the other hand Antedon incisa, in which this latter character
is less evident, has slight notches at the interradial angles of the dorsal surface of the
radials (PL II. figs. la, lc, Id). The Eudiocrinus-c&lyx, therefore, presents no characters
which do not occur in some one or other of the many species of Antedon; but they are
all considerably exaggerated, and are combined together in a somewhat unusual manner.
The interradial sutures on the dorsal surface of the radials are marked by slight
grooves, and there are corresponding grooves on the upper face of the centro-dorsal. But
they do not appear to have been occupied by any tertiary basals in the form of a star
(PL III. fig. 7b). The rosette is tolerably distinct, with a large central opening and well
marked radial spouts. But the interradial processes are scarcely visible, so that there
appear to be only five openings, one at the inner end of each interradial suture (PL III.
fig. 76).
The calyx of the fossil species of Eudiocrinus {Eudiocrinus hyselyi) like that of
nearly all the secondary species of Antedon and Actinometra, is of a very generalised
type ; and, but for the discovery of specimens with the arms attached, it would have
76 THE VOYAGE OF H.M.S. CHALLENGER.
been impossible, as de Loriol 1 remarks, to differentiate the species from the numerous
forms of Antedon which occur associated with it.
Neither arms, pinnules, nor cirri of Eudiocrinus present any characters which can be
said to distinguish them from the ordinary Antedon-tj-pe ; and the disc with its central
mouth might be readily taken for that of an Antedon, except for the fact that the
primary ambulacra do not divide, but proceed straight on to the five arms (PI. VI. fig. 2).
The sacculi which are usually so abundant at the sides of the ambulacra in Antedon, are,
however, far less constant in Eudiocrinus. Abundant in Eudiocrinus indivisus, and
Eudiocrinus atlanticus, they are scanty in Eudiocrinus varians, and altogether absent
in the two remaining species, so far as my knowledge of them extends.
The cirri of Eudiocrinus atlanticus are described by Perrier 2 in the following terms :
— "II n'existe egalement entre les longues pieces des cirrhes dorsaux que de tres faibles
coussinets charnus, et les cirrhes, dans le plupart des echantillons, se montrent etendus
en ligne droite et rassembles dans une attitude qui rappelle celle que certaines araignees
donnent frequemment a leurs pattes."
" UE. atlanticus est, au point de vue de la locomotion, une interessante modification
du type Comatule ; il ne peut en effet, se fixer solidement aux corps etrangers, comme
le font les autres animaux du meme groupe, et il est probable qu'il repose le plus
souvent les bras et les cirrhes etendus sur le limon de l'Oeean, n'ayant a craindre,
dans les profondeurs ou il vit, ni les vagues ni les courants ; mais les masses musculaires
de ses bras indiquent qu'il doit etre aussi un habile nageur. La plupart des Antedon,
et surtout les Actinometra, sont au contraire organises pour s'accrocher solidement
aux corps sous-marins et nagent peu."
It appears to me that Perrier has (as usual) drawn a somewhat hasty conclusion from
the majority of his fifteen specimens of Eudiocrinus atlanticus, with their cirri fully
extended. A large collection of Comatulaa at any particular locality is sure to contain a
number of individuals with the cirri stretched out in a straight line. Antedon phalangium,
for example, has cirri very like those of Eudiocrinus, composed of elongated joints with
small interarticular bundles (PI. XXVIII. figs. 1-3). Great numbers of this species,
with which Perrier is well acquainted, were dredged by the " Porcupine " off the coast of
Tunis. The cirri of some are spread out horizontally ; while in others they are turned
directly downwards, so as to form a sort of basket below the centro-dorsal, and in yet
others the cirri are mostly bent upwards, so as to lie alongside the arms, as in the
examples of Antedon gracilis, and Antedon valida, figured on PI. XV. Indeed all the
three positions may occur in the same individual. The same variations appear in the
long-jointed cirri of Antedon macronema from Sydney Harbour (PI. XXXVIII. fig. 5).
I have seen individuals of this type in which some cirri are horizontally extended, while
others make two or three coils round the stem of a sea-weed or other support. The same
1 Mono£. Crin. foss. Suisse, p. 279. 2 Comptes rendus, 1883, t. xcvi. No. 11, p. V26.
REPORT ON THE CRINOTDEA. 77
difference of position occurs in the short cirri of Antedon carinata, numbers of which
were found by the Challenger at Bahia ; while many instances of the same kind occur
among the Comatulse dredged by the Challenger in the Eastern Archipelago and by the
" Blake" in the Caribbean Sea. (See also PL XXXIII. fig. 6, and PI. LXX.)
I do not think therefore that Perrier is entitled to consider Eudiocrinus atlanticus as
a specially interesting modification of the Comatula-type with regard to its locomotive
powers, for it presents no peculiarities which do not occur in several species of Antedon.
It is true that like other species of the genus (PL VI. fig. 1 ; PL VII.) it has large and
powerful muscular bundles between the successive arm-joints ; and from this perhaps we
may draw the conclusion that it was " un habile nageur." But the last sentence of the
passage quoted above, wherein Eudiocrinus atlanticus is contrasted with Antedon and
Actinometra as regards its swimming powers and mode of life, entirely ignores all that
has been written upon the subject of late years.
It is true that the muscular bundles of Eudiocrinus, as also those of Atelecrinus
(PL VI. figs. 4, 7) and of many deep-sea Coinatulas, appear large by contrast with those
of other types in which they do not appear prominently on the ventral surface of the
arms, owing to their being covered by a thick and more or less opaque perisome. But
when this is removed the large muscular bundles become visible, as seen in Dr. Carpenter's
figure of Antedon rosacea.1 The same is the case in Antedon eschrichti, the muscular
bundles of which, when properly exposed, have at least as great a relative size as those of
any Eudiocrinus ; and if the size of the muscle-plates on the arm-joints be any criterion
of the strength of the muscular bundles attached to them, there is little to choose in this
respect between Antedon eschrichti, Actinometra paucicirra, and Actinometra nobilis.
The position assumed by the cirri, and the appearance of the muscular bundles on the
ventral surface of the arms of Eudiocrinus atlanticus, are not therefore characters of such
importance as Perrier seems to think, when he contrasts this type with " la plupart des
Antedon." I do not see that this species, with its cirri between 15 and 20 mm. in length,
is any less well adapted for fixing itself to submarine bodies, than Antedon phalangium
and many other species of the same genus which have cirri like those of Eudiocrinus
atlanticus (PL XXVIII. ; PL XXX. figs. 4. 8 ; PL XXXIII. fig. 6). Neither do I know
what authority Perrier has for his statement that most species of this genus swim but little,
while implying that Eudiocrinus atlanticus swims a good deal. It certainly cannot be
anything more than a somewhat hasty generalisation, which he could not possibly have
made had he stopped to consider why the muscular bundles of Eudiocrinus appear so
large in contrast to those of " la plupart des Antedon." But when he goes on to speak
of the species of Actinometra as being those which are specially adapted to fix themselves
and to swim but little, he falls into very considerable error. For, as will be shown
immediately in reference to another part of his description of Eudiocrinus, he has not
1 Phil. Trans., 1866, pi. xxxiv. fig. 2.
78 THE VOYAGE OF H.M.S. CHALLENGER.
taken the trouble to make himself sufficiently acquainted with the works of his pre-
decessors, and has therefore committed himself to various statements which will not bear
investigation.
I do not know what grounds of fact he has for his assertion that the species of
Actinometra swim but little. If, as I believe, it is merely an inference from the supposed
small size of the muscular bundles between the arm-joints, his premises are wrong, as I
have explained above ; while I do not know that he has ever been able to observe living
species of the genus, and to notice their abstention from the performance of swimming
movements. On the other hand, Professor Semper has kept various forms of Actinometra
in an aquarium for weeks together, and his observation of the regular alternating
movements of their arms while swimming was mentioned by myself as long ago as
1877. 1 I pointed out in the same memoir, and again five years later'" that the cirri of
Actinometra are few in number, and almost entirely limited to the margin of the
discoidal centro-dorsal ; while those of Antedon are numerous and more or less extensively
distributed over the under surface of the centro-dorsal. But yet Perrier tells us that
' ' surtout les Actinometra " as compared with Antedon are adapted to fixing themselves
by their cirri. The extreme inconsistency of this assertion with the real facts of the case
becomes still more apparent, when it is remembered that in many species of Actinometra
the cirri borne on the centro-dorsal duricg early life drop off, and their sockets
become gradually obliterated (PI. LTV. figs. 1-9 ; PL LXV. figs. 1-6). It was
mentioned in my preliminary Eeport3 that I had found the centro-dorsal of many
Actinometra-s\>ecies to be in the form of a simple flat plate, more or less stellate in form,
but entirely devoid of cirrus-sockets ; while in other individuals only a few imperfect
sockets are present, owing to their not having been completely obliterated. The
occurrence of a fossil Actinometra presenting these characters was also noticed ; 4 and
other references were made to this peculiarity as it was found in a successively increasing
number of species of the genus.5 Copies of the papers in which this character was
described were sent to Professor Perrier, who seems nevertheless to be altogether
unacquainted with its occurrence. For it is difficult to see how Actinometra paucicirra or
Actinometra divaricata (PL LIV. fig. 1 ; PL LXIII. fig. 8), with its perfectly flat centro-
dorsal entirely devoid of cirri, can be regarded as one of those Comatulae which are
especially "organises pour s'accrocher solidement aux corps sous-marins."
After making these somewhat ill-considered remarks, Perrier goes on to describe the
disc of Eudiocrinus atlanticus, which is not more than 5 mm. in diameter and is thus
very small in proportion to the size of the arms, which attain 120 mm., while the cirri
are from 15 to 20 mm. long.6 Perrier then adds " II resulte de ce que nous venons de
1 Journ. Linn. Soc. Lond. (Zool.), 1877, vol. xiii. p. 446. * Bull. Mus. Gomp. Zool., 1882, vol. ix. No. 4, p. 13.
3 Proc. Roy. Soc, 1879, pp. 389-391. 4 Quart. Journ. Zool. Soc, 1880, vol. xxxvi. p. 51.
6 The Comatulae of the Leyden Museum. Notes from the Leyden Museum, 1881, vol. iii. pp. 196, 208.
QComptes rendus, 1883, t. xcvi. No. 11, p. 727.
REPORT ON THE CRINOIDEA. 79
dire que, malgre la simplicite de leurs bras, les Eudiocrinus, loin d'etre un type primitif
de Comatules, representent au contraire un type notablement modifie."
I do not quite know what Perrier would regard as a primitive type of Comatula,
and I have not been able to arrive at any fixed ideas upon that subject myself. But if
his inference that Eudiocrinus is a much modified type has qo better foundation than
is given in his description of Eudiocrinus atlanticus, as would appear from his own
remarks just quoted, I do not think that much can be said for it. This species approaches
more nearly to Antedon than any of the other four comprising the genus ; for it has a
bifascial articulation between the first two joints above the radials like Eudiocrinus
semperi and Eudiocrinus japonicus; but it also possesses what these have not, viz.,
abundant sacculi ; and these organs are abundant in Eudiocrinus indivisus as in Antedon.
This latter form is, however, much further removed from the ordinary Antedon-type than
Eudiocrinus atlanticus, owing to the syzygial union of the two joints above the radials,
which only occurs in a very few species of Antedon. Perrier's inference as to the notably
modified character of Eudiocrinus appears, however, to be entirely founded upon his
knowledge of the single Atlantic species ; while he makes some considerable errors in
his comparison of it with the other Comatula genera, Antedon and Actinometra.
Eudiocrinus has a somewhat wider geographical range than Atelecrinus, extending
over more than 70° of latitude in the West Pacific, and occurring at about 45° N. in the
Atlantic. The type of the genus was found near Bohol in the Philippine Islands by
Professor Semper some twenty years ago. A second species (Eudiocrinus semperi), was
dredged by the Challenger shortly after leaving Sydney, and again off New Zealand. A
third (Eudiocrinus varians), was met with off the north-east part of the Philippine
Group, at the lowest bathymetrical limit of the genus ; while a fourth came up from
565 fathoms, to the south of the Bay of Yedo, and has also been collected at lesser
depths in Japanese waters. To these must now be added the Atlantic species dredged
by the " Travailleur " in 896 metres l (486 fathoms). The bathymetrical range of the type
is thus very considerable, and it has been dredged four times below 500 fathoms, on two
of which occasions the depth exceeded 900 fathoms. The only fossil species known occurs
in the Valanp-ien and Lower Urgonien of Switzerland.
The species of Eudiocrinus fall into two unequal groups. The first one comprises
Semper's type (Eudiocrinus indivisus), in which the first two joints beyond the radials
are united by syzygy ; while in the four remaining species there is a bifascial articulation
between these two joints. In describing the other three Pacific species, I spoke of the
fourth brachials as being traversed by a syzygy and bearing a pinnule in Eudiocrinus
1 This depth (896 metres) is that mentioned by Perrier in his first description of Eudiocrinus atlanticus (Comptes
rendus, 1S83, t. xcvi. p. 725). Recently, however, he has said :— " Les Eudiocrinus viventa environ 1200 metres de pro-
fondeur, dans les regions vaseuses " (Les Explorations Sous-marines, p. 275), and on the same page is figured a specimen
of Eudiocrinus atlanticus from 1000 metres. It may be well to remember that Eudiocrinus indivisus and Eudiocrinus
japonicus have both been dredged in less than 50 fathoms.
80 THE VOYAGE OF H.M.S. CHALLENGER.
semperi and Eudiocrinus japonicus, while in Eudiocrinus varians the first pinnule is on
the second brachial.1 But in his description of Eudiocrinus atlanticus, Perrier2 says
" La premiere syzygie se trouve entre la quatrieme et la cinquieme piece des bras ; c'est
la cinquieme qui porte la premiere pinnule ; la place de la premiere syzygie distingue
V Eudiocrinus atlanticus de YE. indivisus, Semper ; celle de la premiere pinnule la
distingue des trois autres especes."
In reality, however, the position of the first syzygy and that of the first pinnule in
Eudiocrinus atlanticus are exactly the same as in Eudiocrinus semperi and Eudiocrinus
japonicus. In describing the fourth brachial of these two species as a syzygy I was
using precisely the same terminology as was employed by Muller3 in his diagnoses of
Antedon rosacea, Antedon phalangium, and Antedon eschrichti, when he wrote " Das
erste Syzygium befindet sich am dritten Armglied." Perrier however employs a different
terminology, which, as I have explained in Part I. and elsewhere,4 has several disadvantages
from a morphological point of view. He describes the fourth and fifth brachials as
united by syzygy. It is perfectly true that these are primitively the fourth and fifth
joints of the arm, exactly in the same way as the composite third brachial of Antedon
rosacea consists of the united third (hypozygal) and fourth (epizygal) joints of the
growing arm, as described by Dr. Carpenter.5 But since the hypozygals of all the
brachial syzygies of Eudiocrinus atlanticus, Eudiocrinus semperi, or of Antedon
rosacea entirely lose their individuality as arm-joints, bearing no pinnules and taking
no part in the movements of the arm, I believe that it is more correct for descriptive
purposes to follow Muller and to consider the compound or syzygial joint as one arm-
segment only. In accordance with the Midlerian terminology, therefore, I described the
fourth brachial of Eudiocrinus semperi as being or having a syzygy, after going into the
subject rather fully in two memoirs which were published in 1882.6 Perrier, however,
in apparent ignorance of all that had been written on the subject by Muller, Dr. Carpenter,
and myself, not only introduces, though seemingly without knowing it, a new descriptive
terminology, but also imagines that I had used it before him. He has made a very
similar error in his description of Democrinus (Rhizocrinus) , and it is much to be desired
that for the sake of future workers he would take the trouble to acquaint himself with
the current nomenclature before writing his descriptions ; or at any rate that if he decides
to introduce a new descriptive method, he would make some statement to that effect.
The present result is that he describes a difference between Eudiocrinus atlanticus and
Eudiocrinus semperi or Eudiocrinus japonicus, which does not exist in reality. In all
three species alike there is a syzygy in the fourth brachial, as Midler would have described
it, with a pinnule on the epizygal.
1 Joum. Linn. Soc. Lond. (Zool.), 1882, vol. xvi. p. 495. 2 Comptes renins, 1883, t. xevi. No. 11, p. 725.
3 Abhamdl. d. k. Akad. d. Wiss. Berlin, 1849, p. 252. tProc. Zool. Soc. Lond., 1882, pp. 734, 735.
5 Phil. Trans., 1866, p. 721.
6 Joum. Linn. Soc. Lond. (Zool.), 1882, vol. xvi. p. 515 ; and Proc. Zool. Soc. Lond., 1882, pp. 734, 735.
REPORT ON THE CRINOIDEA. 81
Although the principal diagnostic character on which Perrier established Eudiocrinus
atlanticus thus turns out to be due to an erroneous method of nomenclature, he men-
tions a subsidiary one which merits more attention. For he finds that " L'JE.
atlanticus se distingue egalement de ces dernieres especes par le nombre et la grandeur
des organes, si repandus chez les Crinoides, nomnies corps sp>hcriques ou saccules.
Les saccules manquent aux E. japonicus, et E. Semperi ; ils sont petits et rares
chez YE. varians."1 In this respect, therefore, the Atlantic species is sharply
distinguished from both Eudiocrinus semperi and Eudiocrinus japonicus, which resemble
it most closely in the structure of the skeleton ; while they have over twenty-five cirrus-
joints, of which there are only fifteen in Eudiocrinus atlanticus.
The mutual relations of the five species of Eudiocrinus may, therefore, be expressed
as follows : —
I. First two brachials united by syzygy. First pinnule on the second brachial, . 1. indivisus, Semper, sp.
II. First two brachials united by a bifascial articulation.
A. First pinnule on the second brachial, . . . . .2. varians, n. sp.
B. First pinnule on the fourth brachial.
1. Sacculi absent. Twenty-five or more cirrus-joints.
a. Disk plated. First brachials nearly oblong, . . .3. semperi, n. sp.
/?. Disk naked. First brachials trapezoidal, . . .4. japonicus, n. sp.
2. Sacculi abundant. Fifteen cirrus-joints, . . . .5. atlanticus, Perrier.
Eudiocrinus varians, n. sp. (PI. VII. figs. 3-7).
1882. Eudiocrinus varians, P. H. Carpenter, Journ. Linn. Soc. Lond. (Zool.), 1882, voL xvi. p. 496.
Centro-dorsal low, nearly hemispherical, bearing about twenty cirri in two rows
which leave the dorsal pole free. Two forms of cirrus occur in the same individual. — (l)
With two or three short basal joints, the last of which is nearly square, while the following
joint is considerably longer, and the succeeding ones still more so, reaching 3 mm. in
length. Terminal joints unknown. (2) Eight at least of the lower joints are quite short,
few of them being longer than wide, and that but slightly so. Remainder unknown.
Radials partially visible. First brachials nearly oblong, inclined to be trapezoidal, with
small lateral processes which are the edges of the muscle-plates for articulation with the
radials. Second brachial also nearly oblong, with traces of a backward process into the
preceding joint, a pinnule on the right and a small process on the left side. The
following joints have somewhat unequal sides, with a pinnule on the shorter and a large
wing-shaped process on the longer side, which ceases on the sixth, or may go on to the
eighth joint. Succeeding brachials quadrate and unequal-sided, with the pinnule on the
longer side. The twelfth and following joints are distinctly longer than wide. Syzygia
in the fourth and eighth or ninth brachials ; then an interval of two to five joints
between successive syzygia.
1 Comptes rendus, 1883, t. xcvi. No. 11, p. 726.
(ZOOL, CHALL. EXP. — PART LX. — 1887.) OoO 11
82 THE VOYAGE OF H.M.S. CHALLENGER.
The first few pinnules have wide basal joints, the fourth and fifth of which are
sometimes expanded towards the dorsal side. This is most marked in the larger
specimen. The later pinnule-joints are elongated, but very much more slender in the
small specimen than in the larger one. The lower pinnules appear to be the longer,
containing more numerous, though shorter joints. That on the fourth brachial in the
larger specimen is almost 12 mm. long, and consists of twenty-five joints.
Disk 5 mm. wide. It bears numerous calcareous nodules, but the brachial ambulacra
only have delicate rods and networks of limestone at their sides. Saccidi are present,
though small, inconspicuous, and few in number Skeleton white.
The smaller specimen is 3 "2 mm., and the larger 4 "5 mm. across the centro-dorsal.
Locality— Station 205, November 13, 1874; lat. 16° 42' N., long. 119° 22' E.;
1050 fathoms; grey ooze; bottom temperature, 37° F. Two mutilated specimens.
Remarks. — This is a very singular species. The two mutilated individuals described
above resemble one another very closely in the characters of the calyx and arms, while
the cirri and pinnules vary considerably. In the smaller one I can find no certain trace
of any but the long-jointed cirri like those of Eudiocrinns semperi and Eudiocrinus
japonicus (PI. VI. fig. 1 ; PI. VII. figs. 2, 7).
But in the larger form, which retains the bases of two, if not more of these, the
majority of the remaining cirrus-stumps consist of numerous short joints but little longer
than wide (PI. VII. figs. 3, 5).
In the smaller form again, most of the pinnules are quite slender and delicate, with
somewhat glassy joints, which are twice, or more than twice, as long as wide (PI. VII.
fig. 4). But in the larger one they are usually considerably stouter and more massive,
though one or two of the lowest pinnules are much more slender than their fellows, and
somewhat resemble those of the smaller individual (PI. VII. fig. 3). This species is at
once distinguished from Eudiocrinus semperi and Eudiocrinus japonicus, which resemble
it in having an articulation between the first two brachials, by the presence of a pinnule
on the second one.
Eudiocrinus semperi, n. sp. (PL III. fig. 7 ; PL VI. figs. 1-3).
1882. Eudiocrinus semperi, P. H. Carpenter, Journ. Linn. Soc. Lond. (Zool.), 1882, vol. xvi. p. 497.
Centro-dorsal small, nearly hemispherical, or somewhat flattened, thickly covered
with cirrus-sockets, except at the dorsal pole. These have strongly marked articular
rims around the opening of the central canal, and are from twenty to thirty in number.
Cirri probably 30 mm. long, and tapering, of twenty-six joints ; the first three or four
quite short, the next more than twice as long as wide, and the four following ones the
longest, sometimes exceeding 2 mm. The remainder diminish slowly in size, but exhibit
no traces of any dorsal spines.
REPORT ON THE CRINOIDEA. 83
Eadials partially visible. First brachials nearly oblong, widening slightly and then
narrowing a little. Second brachials quadrate, and appearing in a side view of the
specimen to project strongly backward into the first brachials, as the surfaces of both
joints rise towards the middle of their line of junction. The following joints have
unequal sides, the fourth having a syzygy and bearing a pinnule on the shorter side,
usually the right. The seventh joint is more oblong, while the eighth and following
brachials become more distinctly unequal-sided, the breadth being about equal to the
length of the longer side which bears the pinnule. Further out on the arms the length
gradually increases in proportion to the breadth, and the joints become more and more
cylindrical. Second syzygy from the seventh to ninth brachial ; and the later syzygial
intervals vary from one to four joints.
The lower pinnules are all about equal in length, and consist of some twenty joints.
Except in the first four or five pinnules all but the lowest joints are twice as long as
broad, or slightly longer, and more transparent and glassy than the cirrus-joints.
Ovaries short, not extending over more than three or four joints. Towards the arm
ends the pinnules gradually decrease both in length and in the number of joints.
Mouth central. Disk and arm-bases rather closely plated, but the brachial ambulacra
merely have irregular rods and networks of limestone at their sides. They lie close
down between the muscles and show no traces of sacculi. Skeleton white.
Disk 5 mm. in diameter. Radial pentagon 4 mm. Spread probably about 150 mm.
Localities.— Station 164, June, 12, 1874; lat. 34° 8' S., long. 152° 0' E.; 950
fathoms ; green mud ; bottom temperature, 36°"5 F. One specimen.
Station 169, July 10, 1874; lat. 37° 34' S., long. 179° 22' E.; 700 fathoms; blue
mud ; bottom temperature, 40°'0 F. Two specimens.
Remarks. — I have named this species after Professor C. Semper of Wiirzburg, to
whom we owe the discovery during his residence in the Philippine Islands of the type
species of Eudiocrinus {Eudiocrinus indivisus). The absence of pinnules on the second
and third brachials distinguishes Eudiocrinus semperi both from the type and also from
Eudiocrinus varians. Furthermore, both these species have sacculi, which are abundant
in Eudiocrinus indivisus, but rare in Eudiocrinus varians ; while I have not been able to
find them even on the pinnules, either of Eudiocrinus semperi, or of the closely allied
Eudiocrinus japonicus, though they are abundant in the Atlantic species.
Eudiocrinus semperi, like other Coinatulse, exhibits a certain amount of local variation.
All three specimens were obtained in a very mutilated condition, hardly anything
remaining of one of them but the calyx and the bases of the arms. But sufficient
remains of the other two to indicate a considerable amount of flexibility in some of their
characters. That from the lesser depth (Station 169) is the larger of the two, and its
disk bears larger and more numerous plates ; while there are fewer cirri on the centro-
84 THE VOYAGE OF H.M.S. CHALLENGER.
dorsal, and the pinnule-joints are somewhat shorter and less glassy than those of the
individual from Station 164. In the former also both the antero-lateral rays have the
first pinnule on the left side ; while the latter presents a curious variation. The first
pinnule is on the right side in the two posterior rays, and on the left in the left anterior
one, the right anterior one being broken at the syzygy in the fourth brachial. The
anterior ray has been repaired at this syzygy, but no pinnule has been developed on the
epizygal. The fifth brachial, however, bears a pinnule as usual on the left side, but that
on the sixth is on the same side ; so that the first pinnule on the right of the ray does
not come till the seventh brachial.
Eudiocrinus japonicus, n. sp. (PI. VII. figs. 1, 2).
1882. Eudiocrinus japonicus, P. H. Carpenter, Journ. Linn. Soc. Lond. (Zool.), 1882, "vol. xvi. p. 499.
Centro-dorsal relatively large, conical, and covered except at the dorsal pole by from
forty to fifty cirrus-sockets, each with a well-marked articular rim around the opening of
the central canal. Cirri more than 35 mm. long, tapering, and consisting of twenty-seven
joints. The first three are quite short, the fourth a good deal longer than wide, and the
next four the longest, but scarcely reaching 2 mm.; the following ones diminish slowly
in size, but have no traces of any dorsal spine.
Radials just visible. First brachials trapezoidal, the sides commencing to slope
inwards almost immediately beyond the proximal edge. The second brachials, as seen
from below, are also trapezoidal, being narrower along their proximal edges, where they
project backwards into the preceding joints, both surfaces rising towards the line of
junction. The next four or five joints have unequal sides, the fourth being a syzygy,
and bearing a pinnule on its shorter side. In the only specimen with all the arm-bases
preserved, one of them has the first pinnule on the left side. The fifth and one or two
following joints also have the pinnule on the shorter side. The next is more oblong, and
its successor again a syzygy, with the pinnule on its longer side. The succeeding joints
have still more markedly unequal sides, the breadth being about equal to the length of
the longer side. After the second syzygy there is an interval of four or five joints
between successive syzygia.
The lowest pinnules are apparently tolerably equal, consisting of some twenty stout
joints, of which only a few middle ones are longer than wide. Beyond the eighth
brachial, the pinnule-joints become relatively longer and thinner and the pinnules more
slender. Ovaries short, not extending over more than three or four joints.
Mouth central or subcentral. Disk naked, 7 mm. in diameter ; the brachial ambulacra
close down between the muscles, with a few supporting rods and networks of limestone,
but no traces of sacculi. Skeleton white.
REPORT ON THE CRINOIDEA. 85
Locality. — Station 235, June 4, 1875; lat. 34° 7' N., long. 138° 0' E.; 565
fathoms; green mud; bottom temperature, 38°-l F. Three much mutilated specimens.
Remarks. — It is with some hesitation that I have separated this species from the
preceding one. It is altogether larger and more massive than Eudiocrinus semperi, with
a larger and more distinctly conical centro-dorsal and more numerous cirri. The first
brachials have larger muscle-plates for articulation with the radials, and are more
trapezoidal in outline ; and as the second brachials are relatively longer than those of
Eudiocrinus semperi, and at the same time more trapezoidal in form, the base of each
arm is considerably constricted at the junction of its first two joints (PI. VII. fig. 1).
The general proportions of the remaining arm-joints and of the pinnules appear to be
much the same in the two types, excepting that in the smaller Eudiocrinus semperi the
joints of the lower pinnules are rather longer relatively to their width than in
Eudiocrinus japonicus. Of the twelve arms which are preserved in three individuals of
the latter species, only one has the first pinnule on the left side ; while in Eudiocrinus
semperi this appears to be normally the case in the two antero-lateral rays.
There are some specimens of Eudiocrinus in the University Museum at Berlin, which
were kindly shown to me by Dr. Hilgendorf, who had collected them in Japan. I think
that they are probably identical with the type just described. They have rather fewer
cirrus-joints, and the junctions of the first eight brachials are distinctly tubercular. The
tubercle between the first two is in the middle line, and those between the following-
joints lie alternately on either side of the arm. The three Challenger examples, however,
show no traces of these tubercles, with the exception of the median one, which is far
less marked than in the Berlin specimens.
Genus 4. Antedon, de Freminville, 1811.
1733. AcKaw^os, Linck, De Stellis Marinis liber singularis, Lipsice, 1733, p. 53.
1733. Caput Medusx, Linck, Ibid., p. 57.
1758. Asterias, Linnfeus (pars), Systema Naturae, 10th ed., Holniise, 1758, t. ii. p. G63.
1777. Asterias, Pennant (pars), British. Zoology, 2nd ed., London, 1777, vol. iv. p. 55.
1783. Asterias, Retzius (pars), K. Svensk. Vetensk. Akad. Handl., Ar 1783, t. iv. p. 241.
1805. Asterias, Retzius (pars), Dissertatio, sistens Species Cognitas Asteriarum, Lundae, 1805, pp. 33-35.
1811. Antedon, de Freminville, Bull. Soc. Philom. Paris, 1811, t. ii. p. 349.
1813. Asteriatites, von Schlotheim (pars), Taschenbuch fur die Gesammte Mineralogie, 1813, Jahrg. vii.
Abth. 1. p. 68.
1815. Alecto, Leach, Zool. Miscellany, London, 1S15, vol. ii. p. 01.
1816. Comatula, Lamarck (pars), Histoire Naturelle des Animaux sans Vertebres, Paris, 1816, t. ii. p. 530.
1820. Ophiurites, von Schlotheim (pars), Die Petrefactenkunde, Gotha, 1820, p. 326.
1821. Comatula, Miller, A Natural History of the Crinoidea, Bristol, 1821, p. 128.
1823. Comatulithes, von Schlotheim, Nachtrage zur Petrefactenkunde, Gotha, 1823, Abth. ii. p. 17.
1825, Aledro, Say, Journ. Acad. Nat. Sci. Philad., 1825, vol. v. p. 153.
1827. Pentacrinus, Thompson, Memoir on the Pentacrinus Europaeus, Cork, 1827, p. 10.
86 THE VOYAGE OF H.M.S. CHALLENGER.
1828. ComatuJa, Fleming, History of British Animals, London, 1828, p. -190.
182S. Pentacrinus, Fleming, Ibid., p. 493.
1828. Hibernula, Fleming, Ibid., p. 494.
1830. Comatida (Astrocoma), de Blainville (pars), Diet. d. Sci. Nat., 1830, t. lx. p. 229.
1830. Alerto, Cuvier, Regne Animal, Paris, 1830, t. iii. p. 228.
1830. Phytocrinus, de Blainville, Ibid,, p. 235.
1832. Comatida, Goldfuss (pars), Petrefaeta Germanise, Dusseldorf, 1832, t. i. p. 201.
1832. Glenotremites, Goldfuss, Ibid., p. 159.
1832. Solanocrinites, Goldfuss, Ibid., p. 166.
1834. Comatula (Astrocoma), de Blainville (pars), Manuel dActinologie, Paris, 1834, p. 248.
1834. Phytocrinus, de Blainville, Ibid., p. 255.
1834. Ganymeda, Gray, Proc. Zool. Soc. Lond., 1834, pt. ii. No. 14, p. 15.
1835. Comatula, Agassiz, Mem. de la Soc. d. Sci. Nat. de Neuchatel, 1835, t. i. p. 193.
1835. Pterocoma, Agassiz, Ibid., p. 193.
1835. Glenotremites, Agassiz, Ibid., p. 194.
1835. Ganymeda, Agassiz, Ibid., p. 194.
1835. Phytocrinus, Agassiz, Ibid., p. 194.
1835. Solacrinus, Agassiz, Ibid., p. 196.
1836. Comatula, Thompson, Edin. New Phil. Journ., 1836, t xx. p. 295.
1837. Solanocrinites, Bronn, Lethaea Geognostica, 2nd ed., 1837, p. 272.
1837. Decac7iemos, Bronn, Ibid., p. 273.
1839. Comatula, Goldfuss (pars), Nova Acta Acad. Cass. Leop., 1839, Bd. xix. pars 1, p. 348.
1839. Solanocrinus, Goldfuss, Ibid,, p. 349.
1839. Comaturella, von Miinster, Beitriige zur Petrefactenkunde, 1839, p. 97.
1840. Hertha, von Hagenow, Neues Jahrb. f. Mineral., 1840, p. 664.
1840. Comatula, Miiller, Monatsber. d. k. preuss. Akad. d. Wiss. Berlin, 1840, p. 91.
1841. Comatula, Forbes, History of British Starfishes, London, 1841, p. 5.
1841. Comatula, Goldfuss (pars), Neues Jahrb. f. Mineral., 1841, p. 818.
1841. Solanocrinus, Goldfuss (pars), Ibid., p. 819.
1841. Alecto, Miiller (pars), Monatsber. d. k. preuss. Akad. d. Wiss. Berlin, 1841, p. 182.
1843. Alecto, Miiller (pars), Abhandl. d. k. Akad. d. Wiss. Berlin, Jahrg. 1841 [1843], p. 203.
1843. Alecto, Miiller (pars), Archiv f. Naturgesch., 1843, Jahrg. ix. Bd. i. p. 131.
1844. Alecto, Philippi, Neues Jahrb. f. Mineral., 1844, p. 540.
1846. Alecto, Diiben and Koren, K. Svensk. Vetensk. Akad. Handl. Stockholm, 1844 [1846], p. 229.
1846. Comatida, Miiller (pars), Monatsber. d. k preuss. Akad. d. Wiss. Berlin, 1846, p. 178.
1848. Antedon, Gray, List of the species of British Animals in the collection of the British Museum, pt. i.,
London, 1848, p. 28.
1849. Comatula (Alecto), Miiller (pars), Abhandl. d. k. Akad. d. Wiss. BerliD, Jahrg. 1847 [1849], p. 246.
1849. Ganymeda, Bronn, Index Palseontologicus, Stuttgart, 1849, p. 182.
1849. Glenotremites, Bronn, Ibid., p. 182.
1849. Solanocrinus, Bronn, Ibid., p. 182.
1849. Comaturella, Bronn, Ibid., p. 182.
1849. Alecto, Bronn, Ibid., p. 183.
1849. Comatida, Bronn, Ibid., p. 183.
1850. Decameros, d'Orbigny, Prodrome de Paleontologie stratigraphique universelle des Animanx Mollusques
et Rayonnes, Paris, 1850, torn. ii. fasc. 1, p. 121.
1850. Comatula, Ibid., pp. 180, 274.
1851. Decacnemus, Bronn, Lethrea Geognostica, 3rd ed., 1851, Bd. i. Th. iv. p. 133.
1851. Comatida, Broun, Ibid,, Th. v. p. 176.
1851. Glenotremites, Bronn, Ibid., p. 177.
REPORT ON THE CRINOIDEA. 87
1852. Comahda, Forbes, Monograph of the Echinodermata of the British Tertiaries, 1852, p. 19.
1852. Comatula, Quenstedt, Handbuch der Petrefactenkunde, Tubingen, 1852, p. 599.
1852. Solanocrinites, Quenstedt, Ibid., p. 600.
1852. Comatula, d'Orbigny, Cours element, de Paleontol. et de Geol. stratigr., Paris, 1852, torn. ii. fasc. 1,
p. 139.
1852. Comatulina, d'Orbigny, Ibid., p. 139.
1852. Decameros, d'Orbigny, Ibid., p. 139.
1852. Pterocoma, d'Orbigny, Ibid., p. 139.
1857. Comatula, Barrett, Ann. and Mag. Nat. Hist., 1857, ser. 2, vol. xix. p. 33..
1857. Comatula, Pictet, Traite de Paleontologie, 2me ed., Paris, 1857, t. iv. p. 288.
1857. Comaturella, Pictet, Ibid., p. 289.
1857. Decameros, Pictet, Ibid., p. 289.
1857. Pterocoma, Pictet, Ibid., p. 289.
1857. Glenotremites, Pictet, Ibid., p. 290.
1857. Alecto, Lutken, Vid. Meddel. Nat. Foren. Kj0benhavn, 1857, p. 55.
1858. Solanocrinites, Quenstedt, Der Jura, Tubingen, 1858, p. 657.
1859. Comahda, Sars, Nyt Mag. f. Naturvid., 1857 [1859], Bd. x. p. 17.
1860. Comatula, Bronn {pars), Klassen und Ordnungen des Thierreichs, 1860, Bd. it., p. 233.
1860. Comaturella, Bronn, Ibid., p. 233.
1860. Glenotremites, Bronn, Ibid., p. 233.
1861. Alecto, Sars, Oversigt af Norges Echinodermer, Christiania, 1861, p. 1.
1861. Allionia, Michelotti, Rev. et Mag. Zool., 1861, ser. 2, t. xiii. p. 353.
1862. Comatula, Dujardin and Hupe (pars), Hist. Nat. des Zoophytes, Echinodermes, Paris, 1862, p. 192.
1862. Comaster, Dujardin and Hup6 (pais), Paid., p. 211.
1864. Alecto, Lutken, Vid. Meddel. nat. Foren. Kjpbenhavn, 1864, p. 213.
1865. Antedon, Norman, Ann. and Mag. Nat. Hist., 1865, ser. 3, vol. xv. p. 101.
1866. Antedon, Bohlsche, Archiv f. Naturgesch., 1866, Jahrg. xxxii., Bd. i. p. 92.
1866. Antedon, W. B. Carpenter, Phil. Trans., 1866, p. 695.
1866. Antedon, Loven, Ofversigt k. Vetensk.-Akad. Forhandl., 1866, No. 9, p. 224.
1868. Hi/ponome, Loven, Forhandl. Skand. Naturf. Christiania, 1868, t. x. p. liv.
1868. Comatula (Alecto), Pourtales, Bull. Mus. Comp. Zool., 1868, vol. i. No. 6, p. 111.
1868. Antedon, Sars, Memoires pour servir h la connaissance des Crinoides vivants, Christiania, 186S, p. 47.
1868. Solanocrinus, de Loriol, Monographie des Couches de l'etage Valangien des carrieres d'Arzier (Yaud),
Geneve, 1868, p. 84.
1869. Antedon, Pourtales (pars), Bull. Mus. Comp. Zool., 1869, voL i. No. 11, p. 355.
1871. Glenotremites, Geinitz, Palaxmtographica, Bd. xx. Abth. ], Cassel, 1871, p. 91.
1872. Antedon, Geinitz, Ibid., Abth. 2, 1872, p. 18.
1872. Antedon, Wyville Thomson, Proc. Roy. Soc. Edin., 1872, vol. vii. p. 764.
1874. Comaster, Lundgren, Ofversigt k. Vetensk.-Akad. Forhandl., 1874, p. 66.
1876. Comatula, Quenstedt, Petrefactenkunde Deutschlands, Bd. iv., 1876, Asteriden und Encriniden, p. 163.
1876. Solanocrinites, Quenstedt, Ibid., p. 171.
1877. Antedon, P. H. Carpenter, Journ. Linn. Soc. Lond. (Zool.), 1877, vol. xiii. p. 439.
1877. Eallispo7igia, Wright, Proc. Roy. Irish Acad., 1877, ser. 2, vol. ii. p. 114.
1878. Antedon, von Marenzeller, Denkschr. d. k. Akad. d. Wiss. Wien, 1877 [1878], Bd. xxxv. p. 380.
1878. Antedon, Schliiter, Zeitschr. d. deutsch. geol. Gesellsch., Jabrg. 1878, p. 40.
1878. Geocoma, Fraas, Aus dem Orient, Stuttgart, 1878, Th. ii. p. 89.
1878. Antedon, Pourtales (pars), Bull. Mus. Comp. Zool., 1878, vol. v. p. 214.
1879. Antedon, de Loriol, Monographie des Crinoides fossiles de la Suisse, Geneva, 1877-79, p. 253.
1879. Antedon, Fontannes, Ann. Soc. d' Agriculture, Hist. Nat., et Arts utiles de Lyon, 1879, p. 50.
1879. Antedon, P. H. Carpenter, Trans. Linn. Soc. Lond. (Zool.), ser. 2, 1879, vol. ii. p. 16.
88 THE VOYAGE OF H.M.S. CHALLENGER.
1879. Antedon, P. H. Carpenter, Proc. Roy. Soc, 1879, vol. xxviii. p. 385.
1879. Antedon, Rathbun (pars), Trans. Connect. Acad., 1879, vol. v. p. 156.
1879. Antedon, Ludwig, Mitth. Zool. Stat. Neapel, 1879, Bd. i. p. 536.
1879. Antedon, Zittel, Handbuch der Palaeontologie, Palseozoologie, Bd. i. Abth. 2, 1876-1880, p. 395.
1880. Antedon, P. H. Carpenter, Quart. Journ. Geol. Soc, 1880, vol. xxxvi. p. -10.
1880. Antedon, Claus, Grundztige der Zoologie, 4tb ed., 1880, Bd. i. p. 335.
1880. Antedon, P. H. Carpenter, Journ. Linn. Soc. Lond. (Zool.), 1880, voL xv. p. 191.
1881. Antedon, Duncan and Sladen, Memoir Arctic Ecbinodermata, London, 1881, p. 73.
1881. Antedon, P. H. Carpenter, Notes from tbe Leyden Museum, 1881, vol. iii. p. 178.
1882. Antedon, P. H. Carpenter, Journ. Linn. Soc. Lond. (Zool.), 1882, vol. xvi. p. 501.
1882. Antedon, P. H. Carpenter, Bull. Mus. Comp. Zool. 1882, vol. ix. No. 4, p. 13.
1882. Antedon, Bell, Proc. Zool. Soc. Lond., 1882, p. 532.
1882. Antedon, P. H. Carpenter, Ibid., p. 746.
1884. Antedon, Bell, Rep. Zool. Coll. H.M.S. "Alert," Lond., 1884, p. 155.
1884. Antedon, Cams, Prodromus Faunoe Mediterraneas, Pars I., Leipzig, 1884, p. 84.
1884. Antedon, P. H. Carpenter, Proc. Roy. Soc. Edin., 1884, vol. xii. p. 360.
1885. Antedon, P. H. Carpenter, Zool. Chall. Exp., part xxxii., vol. xi., 1884 [1885], p. 137.
1885. Comatula, Quenstedt (pars), Handbuch der Petrefactenkunde, Aufl. 3, Tubingen, 1885, p. 913.
1885. Antedon, Ludwig, Leunis, Synopsis der Thierkunde, Dritte Auflage, Hannover, 1885, Bd. ii. p. 947.
1886. Antedon, P. H. Carpenter, Bijdragen tot de Dierkunde, 1886, Aflevering 13, vi. p. 5.
1886. Solanocrinus, Walther, Palajontographica, Stuttgart, 1886, Bd. xxxii. p. 175.
1886. Antedon, Walther, Ibid., p. 177.
1886. Antedon, Levinsen, Dijniphna-Togtets zoologisk-botaniske Udbytte, Kj0benhavn, 1886, p. 410.
1886. Antedon, Stuxberg, Vega-Expeditionens Vetenskapliga Arbeten, Stockholm, 1886, Bd. v. p 162.
1887. Antedon, P. H. Carpenter, Ann. and Mag. Nat. Hist., 1887, ser. 5, vol. xix. p. 83.
Definition. — Centro-dorsal usually somewhat hemispherical or conical, rarely discoidal,
and generally bearing at least twenty cirri, often several more, which leave but little of
its under surface free. Outer faces of the radials relatively high, with large muscle-
plates, and much inclined to the vertical axis of the calyx.
Disk with a central or subcentral mouth and five equal ambulacra, which extend on
to all the arms. These are ten or more in number, all of the same length, and may have
an ambulacral skeleton which is most differentiated on the pinnules. Sacculi almost
always present on the pinnules, if not elsewhere.
History. — This genus was established in 1811 by de Frdminville,1 who was the
first to remove the Feather-stars from the confusion of the Linnean genus Asterias and to
give them a definite generic rank. A similar course was taken three years later by
Leach, who was probably unaware of de Freminville's work, and established the genus
Alecto for the Feather-stars ; but the same can hardly be said of Lamarck, who deliber-
ately rejected de Freminville's generic name, replacing it by one of his own making, viz.,
Comatula, and he doubtfully referred de Freminville's species Antedon gorgonia to his own
1 Memoire sur un nouveau genre de Zoophites de l'ordre des Radiaires, Bull. Soc. Philom. Paris, 1811, t. ii.
p. 349. In this note de Fr^minville referred to an illustration of Antedon in the Encyclop^die Methodique ; and
Perrier has consequently been led to mention the latter work as that in which the name Antedon was first proposed
Now. Archiv. Mus. Hist. Nat., t. ix., 1886, p. 79).
REPORT ON THE CRINOJDEA. 89
new species Comatula carinata,1 which he had established in apparent ignorance of the
fact that Leach had proposed Alecto carinata in the previous year.
The authority of the great French zoologist and the appositeness of the name which
he proposed both contributed to cause this somewhat ostentatious neglect of the work of
a fellow-countryman to be overlooked by naturalists in general ; and Lamarck's name
was used in succession by J. S. Miller, von Schlotkeim, de Blainville, Goldfuss, Agassiz,
and Minister. Among these authors Miller deserves especial mention, for he was the
first naturalist after Llhuyd and Linck who distinctly recognised the morphologic; il
resemblance between the Feather-stars and the Stalked Crinoids, a point which Lamarck
had entirely failed to notice ; and Miller accordingly drew up a new generic definition of
Comatula which was based upon this idea.2 He seems to have preferred this name
to Alecto, which genus he regarded as less precisely defined than Lamarck's
Comatula.
Johannes Midler also used Comatula in his first communication to the Berlin Academy
upon the subject of the Crinoidea; but in the next year (1841) he formally adopted
Alecto, Leach, as the generic name of several new species, while in the year 1843 he
applied it to the six Lamarckian species which he had not previously mentioned in this
relation, and subsequently also to the Asterias multiradiata of Linnasus.
When Midler first proposed the name Actinometra he regarded it as denoting a genus
equivalent to Alecto; but he eventually reduced both these names to subgeneric rank
and assigned a generic position to Comatula. Dujardin and Hupe, however, dropped
Alecto altogether and restored generic rank to Actinometra, making it equivalent to
Comatula. Soon afterwards, Norman very rightly restored de Freminville's name,
Antedon, which had been suffered to fall into disuse ; and it is now universally used for
the typical Endocyclic Comatulse with five dividing rays, both recent and fossil. There
are, however, a very large number of generic names which have been applied to the
centro-dorsals of fossil Comatula?, both with and without the radials attached, e.g.,
Glenotremites, Solanocrinus, Hertha, Decameros, Decacnemos, Allionia, Comaster, &c.
Pterocoma and Geocoma were the names given by Agassiz and Fraas to species from
the Solenhofen Slate and the Chalk of the Lebanon respectively. Ganymeda, Gray, is in
all probability the centro-dorsal of Antedon rosacea ; while on the other hand, Hyponome,
Loven, is the detached visceral mass of an Antedon common at Cape York.
The stalked larva of Antedon was first described as a dwarf species of Pentacrinus,
which name Fleming proposed to change into Hibernida, this genus being distinguished
from Pentacrinus as then known by the presence of two openings to the digestive
canal. De Blainville, on the other hand, noticed the differences between the characters
1 In his description of Antedon gorgonia de Fn'minville referred to the Encyclopt'die Mt'thodique, partie des Vera,
pi. cxxiv., fig. 6. But Lamarck quoted this figure as representing his Comalula mediterranean
s Op. cit., p. 128.
(ZOOL. CHALL. EXP. — PART LX.- 1887.) OoO 12
90 THE VOYAGE OF H.M.S. CHALLENGER.
of the larval stem and those of the West Indian Pentacrinus, and so proposed to call
the European form Phytocrinus. This was rendered unnecessary, however, by J. V.
Thompson's discovery that Pentacrinus europieus is the young stage of Antedon rosacea;
and it is now clear that Kallispongia, Wright, is a real Comatulid larva, and not a
mimetic Keratose sponge, as was at first supposed.
An attempt has recently been made by Walther1 to re-establish Solanocrinus,
Goldfuss, as a genus distinct from Antedon. I do not think, however, that there are any
really good reasons for this change. The fossil species which Walther refers to Solano-
crinus appear to him to present no syzygial unions in the arms, and this is the only
character of any real generic value which he can bring forward as separating Solanocrinus
from Antedon. I have explained elsewhere,2 however, that the absence of syzygies in
the arms of Solanocrinus costatus, Solanocrinus imperialis, and Solanocrinus gracilis,
is to my mind less certain than Walther believes ; while I strongly suspect from his
figures and descriptions that in all these three types the two outer radials are united by
syzygy, just as in Antedon Jluctuans and Antedon multiradiata (Pis. VIIL, IX.). I
cannot therefore yet acquiesce in Walther's restoration of Goldfuss's genus, though it is
quite possible that this course may become necessary at some future time.
De Freminville did not give any etymology for his new generic name Antedon, and no
clue as to its gender is to be obtained from the name of his single species Antedon
gorgonia. But when the genus was re-established by Mr. Norman in 1865 he used
Antedon as a masculine noun, and in this course he was followed by Sir Wyville Thomson,
Dr. Carpenter, M. Sars, Liitken, Marion, von Marenzeller, Greeff, Ludwig, and others.
In 1877, however, it was determined by the late Mr. Spedding3 that Antedon is really a
feminine name, and should be more correctly spelt Anthedon. Since that date it has
been used as a feminine noun by Pourtales, Ludwig, Duncan and Sladen, Bell, Verrill,
J. V. Carus, Greeff, Dr. Carpenter and myself; though Schliiter, Rathbun, Marshall,
Herdman, Dendy, Vogt and Yung, Walther, and, till lately, Perrier, have continued to
use it in the masculine gender. In Perrier's latest publication,4 however, the following
passage occurs about the name : — "Antedon rosaceus qui, selon la remarque de Victor Carus
doit etre remplace par celui d' Antedon rosacea, Antedon etant une nymphe. Ce savant
expose est suivi d'une etude des niceurs des Antedon qui contient plusieurs constatations
interessantes." As I was unable to find the authority for Perrier's statement in any of
the zoological works of Professor Carus, I wrote to him upon the subject, and he was
good enough to inform me that the facts referred to by Perrier had been contained in a
letter and not in any of his published works. He also kindly gave me a reference to the
1 Untersuchungen ttber den Bau der Crinoideen, Palseontographica, 1886, Bd. xxxii. p. 175.
2 The Generic Position of Solanocrinus, Ann. and Mag. Nat. Hist, 1887, ser. 5, vol. xix. pp. 81-88.
3 Nature, 1877, vol. xv. p. 366.
4 Memoire sur l'Organisation et le Developpement de la Comatule de la Mediterranee (Antedon rosacea, Linck)
Nouv. Archiv. Mus. Hist. Nat, Paris, 1S86, t. ix. fasc. 1, p. 79.
REPORT ON THE CRINOIDEA. 91
following passage in Pausanias where Anthedon occurs as the name of a nymph
(ix. 22, 5):—
" Trj? Se Boiwn'as r<x eV dpicrrepa tov 'Evpinov, MeacrdftLov opos Kakov^vov, koX vtrh
avTco Boianw eVi 6a\d<j(Trj<; 770X15 icrrlv 'AvOtJSwv. yeviadai Se rfj itoXcl to oVo/za ol /j.ev
airb ' AvdrjSovos vvfi,(f)7]<;, ol Se 'Avdcova BvuacrTevaai Xe'youcrii/ ivravOa, IloseiSawo? re 7rcuSa
Kal 'A\kv6vt]<; rrjs "ArXauTO^."
" That part of Bceotia which lies on the left of the Euripus is called the Messabian
Mountain, and below it on the coast is a town of the Boeotians called Anthedon. Some
say that the town was called after a nymph Anthedon, others that Anthon a son of
Poseidon and Alcyone daughter of Atlas reigned there."
It is clear, however, that although Antedon is etymologically incorrect, De Freminville's
spelling of the. name must be retained ; but at the same time the question of its gender
may be regarded as finally settled.
Remarks. — De Freminville's definition of Antedon, like those of Alecto by Leach
and of Comatula by Lamarck, would apply almost equally well to all the various forms
of Feather-stars. But that given by Mr. Norman * is of a much more limited character, as
it commences with the words " Mouth central. Anus lateral." This character alone was
sufficient to separate Antedon from Midler's genus Actinometra as defined by Dujardin
and Hupe three years before, but we now know four other genera of Endocyclic
Comatulse.
The essential characters of the calyx of Antedon have been fully explained already,
and there is therefore no need to refer to them again. It is distinguished from Pro-
machocrinus by the presence of five rays only, and from Eudiocrinus by the fact that
these rays divide so that there are ten primary arms, which may themselves divide again.
The presence of pinnules on the arm-bases and the lateral union of the radials distinguish
Antedon from the two remaining genera of recent Endocyclic Comatulse, Atelecrinus and
Thaumatocrinus respectively ; while the want of a comb on the oral jnnnules, the
presence of sacculi, and the central mouth distinguish it very clearly from Actinometra.
The oral pinnules of Antedon are extremely variable in their characters. In some
species, such as Antedon mxdtispina, and Antedon angvstipinna, they are comparatively
small and insignificant (PI. XIII. fig. 1 ; PI. XXIX. fig. 1). In others like Antedon
gracilis, Antedon valida, Antedon incerta and Antedon lusitanica they have a number
of short, but very wide basal joints, and are therefore somewhat massive in appearance
(PI. XII. fig. 3 ; PI. XV. fig. 6 ; PL XVIII. fig. 5 ; PI. XXXIX. fig. 2). In Antedon
occulta, and in a large number of similarly bidistichate species, they are stiff and styli-
form and stand up round the edge of the disk as if to shield it from danger, a character
which Liitken has expressed in the specific name Antedon protecta. They are more slender
and flexible and consist of much elongated joints in Antedon longipinna and Antedon
1 Ann. and Mag. Nat. Hist, 1865, ser. 3, vol. xv. p. 101.
92 THE VOYAGE OF H.M.S. CHALLENGER.
exigua (PL XXX. fig. 1 ; PL XXXII. fig. 4) ; while iu the group of species allied to
Antedon eschrichti they are more or less flagellate, consisting of a large number of
relatively short joints (PL XXIV. figs. 1-3, 7-9 ; PL XXV. figs. 1-3 ; PL XXVII.
figs. 8, 9). There is much less variation in this respect in the oral pinnules of Actino-
metra, which are always provided with a terminal comb (PL LIII. figs. 3-6), a character
which never occurs in Antedon.
On the other hand, the sacculi which are almost invariably present in this genus,
never occur in Actinometra, even when species of the two genera are living side by side
in the same locality ; and this fact is a very strong argument against the theory of Vogt
and Yung l that the sacculi are symbiotic Algae, as I have explained elsewhere.2 There
are a few species of Antedon, e.g., Antedon quinquecostata, in which they are small and
poorly developed, though they are abundant in others obtained at the same localities ;
and in some other instances the condition of the specimen has been such that I have not
been able to assure myself satisfactorily of the presence of sacculi. This is the case for
example with the two specimens of Antedon abyssicola from 2900 fathoms, the greatest
depth at which Comatulas have been obtained ; but they are fairly abundant in another
individual of the same species from 2600 fathoms in the Southern Sea. There are few
species of Antedon in which they are not present ; though they are more variable in their
occurrence among the species of Eudiocrinus, as has been already explained.
The very definite relation of the sacculi to the side plates of the ambulacra in those
species of Antedon which have a highly differentiated ainbulacral skeleton is a very
strong argument against the views of Vogt and Yung that they are symbiotic Algae. It
was pointed out on p. 127 of Part I. how the distal edges of the side plates are notched
for the reception of the sacculi, and figures were given on pi. liv. illustrating this
character in four species of Antedon. But it does not occur at all in species of Peata-
crinus and Metacrinus which live at the same localities as these Coniatuhe (Stations 170a,
175, 192, 214), and I cannot think, therefore, that the problem of the nature of the sacculi
has been solved by Vogt and Yung. These are not the only difficulties which suggest
themselves in connection with the details of their theory as I have explained elsewhere.3
Classification. — It has been shown on a previous page how the numerous recent
species of Antedon may be associated together into groups of variable size, according to
the characters of the rays and of their subdivisions. The first group to be considered
includes those species in which the two outer radials are united by a syzygy and not, as
is most frequently the case, by a bifascial articulation. Five of the eight recent species
of Pentacrinus are distinguished by this character, and it occurs in several species of
1 Op. cit., p. 570.
2 On the Supposed Presence of Symbiotic Algae in Antedon rosacea, Quart. Journ. Micr. Sci., 1S87, new sen,
vol. xxvii. p. 386.
3 Ibid., pp. 380-384.
REPORT ON THE CRINOIDEA. 93
Actinometra, both ten-armed and multibrackiate, e.g., Actinometra pectinata, Actinometra
paucicirra and Actinometra typica (PL LIII. fig. 15; PL LIV. figs. 1, 2; PL LVII.
fig. 1).
I do not know, however, of any ten-armed Antedon belonging to this group, and the
three species immediately to be described, in which the rays divide three or four times,
present one very exceptional feature in their organisation. It is a very general rule
among Neocrinoids that the mode of union of the first and second joints beyond the
radial and all subsequent axillaries is the same as that between the two outer radials.1
But this rule does not always hold good in the case of syzygial unions, though it is true
amongst other species, of Pentaerinus wyville-thomsoni and Pentacrinus alternicirrus,
of Actinometra difficilis and Actinometra paucicirra (PL L1I. fig. 2 ; PL LIV.
figs. 1, 2), in all of which the two outer radials, the two distichals and the first two
brachials are respectively united by syzygy.
In Actinometra multibrachiata and in Actinometra typica there are three joints in
the distichal series, the first two articulated and the third a syzygy. But in the numerous
remaining arm-divisions there are only two joints which are united by syzygy like the
two outer radials (PL LVI. fig. 2 ; PL LVII. fig. 1).
The three species of Antedon now to be described are, however, still more irregular ;
for in neither distichal, palmar, nor brachial series are the first two joints united by
syzygy, as is the case with the two outer radials. This latter character seems to have
presented itself in three Jurassic species of Antedon. Quenstedt 2 has described the two
outer radials of Solanocrinus (Antedon) costatus as united by syzygy, and his description
is borne out by his figures, one of which shows a first brachial of such a size that I feel
tolerably certain of its being really a syzygial joint as in Actinometra strata and Actino-
metra pectinata (PL LIII. figs. 2, 15). Walther's recent description of Solanocrinus
costatus9 contains the passage "Radiale II. mit Radiale IIT. versckmolzen, doch durch
eine Nahtlinie getrennt ;" and it is odd that he did not follow Quenstedt in describing
the union as a syzygial one. The large size and the pentagonal shape of the radial
axillaries in his Solanocrinus imperialis seem to me to indicate clearly that these are
syzygial joints ; and I am very strongly inclined to believe that the large joints which
he describes as " Axillaria" are really compound joints, consisting of the first and second
distichals united by syzygy, as in Actinometra paucicirra (PL LIV. figs. 1,2). These
pieces are more distinctly separate in the five remaining distichal series of his specimen,
while in some cases at any rate, the large first brachials would appear to be syzygial
joints. The same may be said of Walther's single specimen of Solanocrinus gracilis,4 of
which he remarks as a possibility that the apparently simple second or axillary radial
" als verschmolzenes Radiale II. + Radiale III. aufgefasst werden konnte."
» See Part I. p. 49. 2 Encriniden, p. 17:2, Tab. 96, figs. 26, 28.
3 Op. cit., p. 172. 4 Ibid., p. 174.
94 THE VOYAGE OF H.M.S. CHALLENGER.
If then there really were syzygial unions between the two outer radials and the first
two distichals and brachials respectively of Solanocrinus imperialis and Solanocrinus
gracilis, these two species would represent a type which is not as yet known to occur
in the recent Antedon at all, but only in Actinometra paucicirra and its allies ; while
the ten-armed Antedon costatus is represented at the present time by the various species
belonging to the type of Actinometra Solaris (PL LIII. figs. 2, 15).
The three species of Antedon belonging to this first series which were dredged by the
Challenger, may be classified as follows : —
Antedon, Series I.
The two outer radials united by syzygy.
A. Three distichals, the axillary a syzygy.
I. Subsequent divisions of two articulated joints, . . . .1. fluctuans, n. sp.
II. Subsequent divisions like the distichals.
a. Three axillaries above the radials, . . . . .2. multiradiata, n. sp.
b. Four axillaries above the radials, . . . . .3. microdiscus, Bell.
1. Antedon fluctuans, n. sp. (PL VIII.).
Specific formula, A.R.3.2.(2).— -.
Centro-dorsal a thick disk, bearing about twenty- five marginal cirri. These have
thirty to thirty -five joints, of which the fifth to eighth are much longer than broad. The
following ones diminish in length and gradually develop a sharp forward projecting
spine which decreases slightly in the short terminal joints, but increases again on the
penultimate as the opposing spine to a strong recurved claw.
Three radials visible, the second free laterally, but united to the third by syzygy.
The rays are quite free and may divide four times. Three distichals, the axillary with a
syzygy. Palmars and post-palmars (when present) usually of two articulated joints.
Arms from twenty-six to nearly forty in number, and composed of short, smooth, and
obliquely quadrate joints. A syzygy in the third brachial ; the next anywhere between
the twelfth and the sixtieth, with others at intervals of six to twenty joints.
The second distichal bears a long, tapering pinnule of about forty joints, the basal
ones tolerably stout and the terminal ones small. The second, and sometimes also the
third brachials have similar but smaller pinnules, and the following ones decrease slowly
in size, becoming long and slender again towards the arm-ends.
Disk much incised, and the interradial regions more or less covered with rather large
plates. The ambulacra as far as the last axillary are raised and strongly plated ridges.
But those of the arms and pinnules, including even that on the second distichal, are
unprotected.
REPORT ON THE ORINOIDEA. 95
Colour in spirit, — the skeleton a light brownish-white, with the perisome sometimes
darker. Sacculi abundant, especially on the pinnules, and sometimes appearing on the
outer ends of the plated disk-ambulacra.
Disk 8 mm.; spread probably 80 to 90 mm.
Locality.— Station 190, September 12, 1874 ; lat. 8" 56' S., long. 136° 5' E .; 49
fathoms ; green mud. Two mutilated individuals and one fragment ; one varietal form.
Other Localities. — H.M.S. "Alert," 1881 ; Torres Strait. One specimen.
Remarks. — Three of the Challenger specimens agree very closely in their general
characters, though the frequency of the arm-divisions, and therefore the number of arms,
varies considerably. All the distichal series consist of the usual three joints with a syzygy
in the axillary; nearly all the palmars have but two joints without a syzygy ; post-palmars
are present in every individual, and in the majority of cases resemble the type of the
palmars, so that the arm-formula becomes A.R.3.2.2. (PL VIII. fig. 1).
There was, however, a fourth specimen obtained besides these three, from which it
differs in many points, though not, I think, sufficiently so to entitle it to a distinct
specific rank (PI. VIII. fig. 2). The colour of the calyx and arms is the same brownish-
white as in the type, but the cirri have a strong reddish-brown tint (which was probably
purple during life) with white bands at the inter- articular lines, and the lower joints are
shorter relatively to their width than in the type-forms, though remaining longer than
wide. The difference from the type is most apparent, however, in the arm- divisions.
For four out of the ten distichal series have but two joints, the axillary without a syzygy;
and out of the twelve palmar series which remain, six have two and the other six three
joints, while there are no post-palmars at all, although they occur in each of the three
type-specimens. The arms too, are somewhat more massive than in these last, and their
component joints, instead of being smooth and obliquely quadrate, are relatively shorter
and more wedge-shaped, with a slight tendency to overlap.
The differences between this individual and the other three, which agree so closely in
their general characters, are certainly very marked ; but it is difficult to find in any one
of them an adequate reason for specific distinction. This conclusion is confirmed by the
fact that a third form which combines certain characters of each of the other two was
obtained in Torres Strait by Dr. Coppinger of H.M.S. "Alert." Being in a somewhat
mutilated condition it was not described by Professor F. J. Bell in his Report on the
" Alert " Echinoderms, but was put aside until the arrival of better preserved material ;
and I am indebted to him for the opportunity of referring to it here. It resembles the
type-form in the shape of its arm-joints but has no post-palmar series ; and it further
resembles the varietal Challenger specimen in having purplish cirri with white bands as
described above. It is tolerably clear, therefore, that as in so many other cases, we are
here dealing with a somewhat variable specific type, and I propose to designate it
96 THE VOYAGE OF H.M.S. CHALLENGER.
accordingly as Antedon fluctuans. It is curious, however, that while the two most
dissimilar forms were dredged at the same Station in the Arafura Sea, the intermediate one
was obtained at a much less depth in Torres Strait, and that no others have been met with
at any intervening locality. When examining the " Alert " specimen I found a small
Myzostoma upon it, which Professor von Graff has named Myzostoma quadricaudatum.
Antedon fluctuans is a type of considerable interest from a systematic point of view.
For the syzygial union of the two outer radials is in no case accompanied by a similar
union of the first two joints after each subsequent axillary. Whether there be three
distichals, as is normally the case, or two only, as in some exceptional rays, there is
always the bifascial articulation between the first two joints above the radials and above
every successive axillary, just as in the ordinary many-armed Antedons. In the case of
Antedon fluctuans, the palmar and post-palmar series (when present) normally consist of
two joints only, and this character distinguishes the type from Antedon midtiradiata
and Antedon microdiscus, in which there are three joints between the successive arm-
divisions (PI. IX.; PL XXXVII. fig. 3). I know of no other described species but
these with which Antedon fluctuans is likely to be confounded, provided, of course, that
the syzygial union of the radials be properly recognised.
2. Antedon midtiradiata, n. sp. (PI. IX.; Part I. pi. lv. figs. 3, 4).
Specific formula, A. R.3.3.3.— .
Centro-dorsal a thick, slightly convex disk, bearing from twenty to twenty-five
marginal cirri. These are rather long, consisting of forty or fifty, or occasionally more
joints, few or none of which are longer than wide. The last half have a small blunt
spine projecting slightly forwards, which forms a strong and sharp opposing spine on the
penultimate.
Three radials visible, the second short aud free laterally, but united to the third by
syzygy. The rays are quite free and may divide four times, each series of three joints
with the axillary a syzygy. About forty arms of smooth and short triangular joints,
which become blunter and more square towards the ends. A syzygy in the third
brachial, the next anywhere between the sixteenth and forty-fifth joints, with others at
intervals of seven to nineteen joints.
The distichal pinnules of moderate length, consisting of about twenty-five stout
joints ; the size gradually decreases to that on the second brachial, and the next pair are
considerably smaller, the following ones increasing slowly in size, but never becoming
very large.
Disk much incised and paved with large plates between the ambulacra, which are
elevated ridges with plated walls, but the plating scarcely extends beyond the level of the
last axillary.
REPORT ON THE CRINOIDEA. 97
Disk about 15 mm.; spread probably about 12 cm.
Colour in spirit, — a deep reddish purple with patches of whiter tint. Sacculi deeply
coloured, and abundant along the sides of the pinnule-ambulacra.
Locality. — Station 187, September 9, 1874; off Booby Island, Torres Strait;
lat. 10° 36' S., long. 141° 55' E.; 6 fathoms; coral mud. Two imperfect individuals.
Remarks. — This species differs from Antedon fiuctuans in the composition of the
later arm-divisions, which resemble the distichal series in consisting of three joints with
a syzygy in the axillary, in the sensible decrease in the length of the pinnules after that
on the second brachial, and in the greater number of the cirrus-joints. The grouping of
the arm-divisions is the same as that of Antedon microdiscus, in which, however, there
is a fourth post-radial axillary (PI. XXXVII. fig. 4). This is absent in Antedon multi-
radiata, which has fewer cirri and smaller basal pinnules than Antedon microdiscus.
Several isolated disks of Antedon multiradiata were obtained by the Challenger in
Torres Strait, and two of them were figured in Part I. (pi. lv. figs. 3, 4). Owing to the
freedom of the rays, which are not bound together by perisome, the disks are very deeply
incised and have a markedly stellate appearance. The so-called recent Cystidean Hypo-
nome sarsii of Loven1 is nothing but one of these Antedon-disks covered with a
well-developed calcareous plating, both at the sides of the ambulacra and in the
interambulacral regions. It is not unlikely to have been the disk of Antedon multi-
radiata which was found in this condition at Booby Island by the Challenger, as it has
a more extensive plating than Antedon microdiscus, while Antedon bidentata, the other
species dredged at that locality, has a quite naked disk.
3. Antedon microdiscus, Bell (PI. XXXVII. figs. 4-6).
Specific formula — A.R.3.3.3.3.y.
1884. Antedon microdiscus, Bell, Rep. Zool. Coll. H.M.S. "Alert," London, 1884, p. 163, pi. xv.
Description of an Individual.— Centro-dorsal relatively large, with about three rows
of cirri on its sides, but the dorsal surface slightly convex and free from them. Forty to
forty-five joints in the cirri, few or none of them being longer than wide ; the distal
ones quite short, with tolerably well-marked spines, that on the penultimate being
sharp and distinct.
First radials barely visible ; the next two united by syzygy. The rays divide four
and sometimes five times, each series of three joints with the axillary a syzygy.
Arms of smooth joints, the lower ones short and nearly triangular, but becoming more
oblong after about the sixtieth. A syzygy in the third and again in about the thirtieth
brachial, with others at intervals of twelve to fourteen joints.
1 On Hyponome sarsii, a recent Cystidean, Canadian Naturalist, N.S., 1869, vol. iv. pp. 265-268.
(ZOOL. CHALL. EXP. PAET LX. 1887.) OoO 13
98 THE VOYAGE OF H.M.S. CHALLENGER.
Distichal and palmar pinnules large and stout, composed of forty to fifty joints, the
second one rather the longer. Their lower joints are large but not specially marked ; the
following ones diminish in size, but gradually develop a projection of the dorsal edge
at their distal end, which disappears in the smaller terminal joints. The third and
following pinnules decrease rapidly both in length and in stoutness, after which the
length slowly increases again.
Disk-ambulacra protected by a well-developed calcareous plating which ceases at the arm
bases ; anal tube also considerably plated, but the other interpalmar areas are unprotected.
Colour in spirit, — skeleton brownish-white, and the perisome mottled with grey.
Sacculi very abundant on the pinnules.
Disk 17 mm., spread about 20 cm.
Locality. — Station 186, September 8, 1874; Torres Strait; lat. 10° 30' S., long.
142° 18' E.; 8 fathoms ; coral mud. One specimen.
Remarks. — Only one example of the species was obtained by the Challenger, but it
did not come into my hands with the rest of the collection, having been given by Sir
Wyville Thomson to the National Museum at Stockholm, where I found it in August of
last year (1886), and Professor Loven has since been kind enough to send it over to
England for my further examination.
A larger specimen of the same type was obtained in 1881 at Port Molle, Queensland,
by H.M.S. "Alert," and was described by Professor F. J. Bell,1 together with some
smaller individuals already in the National Collection from Nicol Bay, Australia.
There was a very important omission, however, in Bell's description ; for he entirely
overlooked the fact that the two outer radials are united by syzygy, a character which,
next to those of the genus, is of primary importance for systematic purposes. His
description and figure rather led me to suspect the presence of this character long before
I saw the Challenger specimen, and my suspicions were verified when I examined his
types for myself. He likewise makes no mention of any axillaries beyond the post-
palmars, although such must be present to bring the number of arms up to ninety, the
number which he describes in the adult, while several quaternary arms are represented
. c
in his figure. His specific formula must be altered therefore from A.3.3.(3).— . to
A.R.3.3.3.3.— .
c
It is the presence of this fourth axillary above the radials which is one of the
characters distinguishing this species from Antedon midtiradiata. I have not seen any
specimen without it, though it is much more frequent in the individual from Port Molle than
in those from Nicol Bay and Torres Strait. These last resemble one another in Laving a
smaller number of cirrus-joints and a better-developed penultimate spine than in the type.
1 "Alert" Report, p. 163, pi. xv.
EEPOET ON THE CEINOIDEA. 99
The joints of the cirrus figured by Bell ' are much wider than long ; whereas in the
Challenger specimen this is only the case in the outer part of the cirrus, some of the
lower joints being as long as or longer than wide, and in premature cirri the length is
distinctly greater than the width, while the penultimate spine is especially prominent.
Bell described the second or palmar pinnule of his type specimen as being a good deal
longer than the first or distichal one. This is not the case, however, in that dredged by
the Challenger; and the pinnules show no trace of the slightly keeled basal joints
described by Bell. But the distal edges of the basal joints are somewhat sharp, and
beyond the sixth joint they project slightly over the bases of the succeeding ones
(PL XXXVII. fig. 6). This feature gradually develops into a blunt slightly spinous
process, which is most marked about the fifteenth joint and disappears altogether after the
twenty-fifth ; but in the palmar pinnule figured by Bell it is not visible till the eighteenth
segment and continues till near the end of the pinnule. It is this feature apparently which
led Bell to say — " the more distal joints are provided with a spine or tuft of spines."
The anal tube of this individual contains a species of Anilocra, the Isopod which
was described and figured in Part I. in the same position on the disk of Actinometm
rpaucicirra (see Part I. p. 133, pi. lv. fig. 1).
Antedon, Series II.
The two outer radials articulated ; ten arms.
Remarks. — It has been pointed out already that the number of species of Antedon
which have articulated radials and only ten arms is very considerable ; and it therefore
becomes necessary for the purposes of specific discrimination to arrange them into groups
of comparatively small size. I have had considerable difficulty in effecting this object, as
the absence of any axillaries on the arms deprives us of an important aid to classification.
By using the characters of the arai-bases and of the lower pinnules, however, I have
found it possible to classify most of the ten-armed species of Antedon in five groups,
which I propose to call by the names of their more characteristic or best known species, —
thus, the Eschrichti-growp, the Tenetta-group, &c.
The radials and lower brachials have flattened sides. Pinnule-arnbulacra generally plated, . 1. BaMcurva.
The rays not flattened laterally. Pinnule-ambulacra well plated, . • • -2. Acoela.
The first two or three pairs of pinnules long and flagellate, with numerous short and wide
joints, . . . . . . . . . . .3. Eschricktii.
The joints of the lowest pinnules, which are often long and slender, are longer than wide,
frequently very much so, . . . . . . ■ • .4. Tenella.
The first pair of pinnules are comparatively small, and their joints but little longer than wide ;
one or more of the second, third, and fourth pairs are longer and more massive, with stouter
joints than their successors, . . . • • ■ • -5. Milherti.
1 "Alert" Report, pi. xv. fig. d.
100
THE VOYAGE OF H.M.S. CHALLENGER.
The first of these groups to be considered is one which has hitherto been entirely
unknown to me except in fossil Coruatulae, and therefore consists entirely of new species
discovered during the cruise of the Challenger. I will call it the Basicurva-grou]), for
it is in Antedon hasicurva that its principal distinctive characters are, on the whole,
most clearly visible.
In all the species of this group not only the two outer radials, but also the lowest
brachials of adjacent rays, come into very close mutual apposition, so that their sides are
flattened against one another.1
Fig. 2, A, represents a calyx of Antedon hasicurva from which three rays have been
entirely removed ; and it then appears that the two lateral faces of each pair of outer
Fig. 2. — Antedon hasicurva, x 3. A. Side view of the calyx and arm-bases after the removal of three rays, so as to show
the sides and inner faces of the other two. The two outer radials, two lower brachials, and in a less degree also the
third and fourth, have their outer sides flattened against one another. The genital pinnules have the third and fourth,
and sometimes the fifth joints greatly expanded, but the following ones are smaller. B. The lower part of an arm from
its inner side, to show the flattened inner faces of the first three brachials, including both the hypozygal and the
epizygal of the third.
radials are quite smooth and flat, like those of the first radials at the bottom of the
calyx, and they are in close apposition with those of the second and axillary radials in
adjacent rays. In like manner the outer faces of the first brachials on adjacent rays
come into close mutual contact and are very perceptibly flattened. The same is true of
the second, and in a less degree also of the third brachials, all these three joints being
somewhat compressed laterally, with flattened sides and sharp, straight, outer edges.
The inner edges and sides of the second and third brachials present the same feature, as
seen in fig. 2, B, so that the lower portions of the arms lose their usual rounded character
1 This character is more or less distinct in some forms of Antedon milberti, but appears to be a varietal rather than
a fundamental one (see p. 197).
EEPORT ON THE CEINOIDEA. 101
and come to have a somewhat " wall-sided " appearance. In some species, such as Antedon
valida, Antedon incerta, and allied forms, which have large first pinnules, the second
brachials that bear them have quite short outer sides which are scarcely flattened
at all. But the lower pinnule-joints are extremely modified. Their outer sides are
flattened where they meet the corresponding pinnules of adjacent rays, while their inner
sides seem to have been cut away so as to let the pinnules lie close against the arm
(PI. XV. figs. 5, 6 ; PL XVIII. fig. 5).
On the other hand, there are some species in which the wall-sided nature of the arm-
bases is so comparatively inconspicuous that it might readily escape notice by an un-
trained eye. It is a peculiarity which becomes more prominent with age, the radial
and brachial joints of young individuals being always longer relatively to their width
than in the adult condition, so that those of adjacent rays come less closely into
contact.
This flattening of the apposed sides of the radials and lower brachials is a character
which, so far as my knowledge goes, does not appear in any of the Pentacrinidse, though
it is very marked in the fossil Solanocrinus costatus, Goldfuss. Quenstedt l figures a ray
of this species in which the flattened sides of the syzygial axillaries and of the two
lowest brachials are very clearly shown. He says that " von der Seite zeigten nur das
erste und zweite Armglied eine breitere Flache, die mit dem dritten pltitzlich schmal
wird, und alsdann ganz verschwindet, zum Zeichen, dass mit dem vierten Gliede die Arme
schon ganz getrennt waren, und sich mit ihren Innenrandern nicht mehr beruhrteu."
In like manner Walther 2 says of the same type " dass, wie bei dem Stiick von Quenstedt
die drei untersten Armglieder mit seitlichen Gelenkflachen eng verbunden sind und
einen Brachialen Pseudo-heleh bildeten, so dass sich also die Arme erst vom vierten
Glied an bewegen konnten."
In Encrinus and in some species of Apiocrinus this character is considerably exag-
gerated, the radials and the lower parts of the arms fitting very closely against their
fellows on either side. Traces of it also appear in Holopus, as described on p. 206 of
Part I.
In all but two of the twenty species which belong to the Basicurva-gvoxvp, and have
the lower parts of the rays flattened laterally and wall-sided, there are definite covering
plates on the pinnule-ambulacra which rest on a limestone band more or less completely
segmented into side plates, as for example in Antedon breviradia (PI. XIX. fig. 4). But
in two species, Antedon denticulata and Antedon pusilla, there is no ambulacral skeleton
at all, a peculiarity which separates them very distinctly from the other members of
the group.
1 Encriniden, p. 174, tab. xcvi. fig. 26a. 2 Loc. cit., p. 171.
102
THE VOYAGE OF H.M.S. CHALLENGER.
These may be classified very readily according to the characters of the cirri and lower
pinnules, as shown in the following scheme : —
1. The Basic urva-gromp.
The radials and lower brachials have flattened sides.
A. Pinnule-ambulacra plated.
I. Later cirrus-joints have dorsal spines.
a. First pinnule smaller than the second. Eighty cirrus-joints,
b. First pinnule longer than the second.
1. Sixty cirrus-joints or more.
Basal joints of first pinnule much flattened on the outer side.
First two joints of the distal pinnules expanded and trapezoidal,
First pinnule but little flattened on the outer side. First two
joints of distal pinnules not specially marked,
2. Thirty to fifty cirrus-joints.
(a) First pinnule flattened on the outer side. Cirri irregularly
disposed,
(i.) Pinnule on third brachial carinate, like that on second,
(ii.) Pinnule on third brachial small, like that on fourth.
a. Calyx and arm-bases not spinous.
* Second and third radials rounded, but not
specially convex.
Second radials very short,
Second radials of moderate length. Later
arm-joints carinate, .
** Second and third radials sharply convex or
carinate.
Axillaries wider than long,
Axillaries longer than wide,
/?. Calyx and arm-bases very spinous. Arm-joints
with large curved spines,
(b) First pinnule not flattened on the outer side. Cirri in ten
vertical rows, ......
3. Less than thirty cirrus-joints.
(«) Pinnule on third brachial a good deal smaller than that on
second. Arms spinous. First radials invisible, .
(b) Pinnule on third brachial not much smaller than that on
second. Arms smooth. First radials visible,
II. Less than thirty cirrus-joints without dorsal spines.
a. Pinnules of eighth and following brachials have broad lower joints, and
strong plates covering the genital glands.
1. Third and fourth joints of genital pinnules broad and nearly flat on
the outer side, but the fifth joint smaller.
(a) First radials concealed by centro-dorsal ; lower arm-joints with
raised and crenulated distal edges, . . . <
(b) First radials distinctly visible. Arm-bases smooth, .
1. longiciira, n. sp.
2. valida, n. sp.
3. incerta, n. sp.
4. gracilis, n. sp.
5. lusitanica,1 n. sp.
6. breviradia, n. sp.
7. spinicirra, n. sp.
8. acutiradia, n. sp.
9. bispinosa, n. sp.
10. latipinna, n. sp.
11. multispina? n. sp.
12. echinata, n. sp.
13. basieurva, n. sp.
14. incisa, n. sp.
1 This is also a Lidistichate species. See pp. 110, 217, and PI. XXXIX. fig. 1.
2 This is also a tridistichate species. See pp. 117, 248, and PI. LXIX. figs. 1, 2.
REPORT ON THE CRINOIDEA. 103
2. Lower joints of genital pinnules uniformly expanded.
(a) First radials concealed; less than twenty cirrus-joints.
(1) Calyx and arm-bases bluntly spinous; First pinnules
almost flagellate with twenty joints or more, . 15. tuberosa, n. sp.
(2) Calyx and arm-bases smooth ; first pinnules more
styliform, of about fifteen joints, . . . 16. parvipinna, n. sp.
(b) First radials visible; over twenty cirrus-joints, . . 17. flexilis,1 n. sp.
b. Pinnules of tenth and following brachials have the lower joints as long as
or longer than wide, and no extensive plating over the genital glands, 18. aculeata, n. sp.
B. Pinnule-ambulacra not plated.
I. Two radials visible; arm-joints short. Stoutest pinnule on sixth brachial.
Cirrus-joints long and smooth, . . . . . .19. denticulata, n. sp.
II. Three radials visible; arm-joints long. Stoutest pinnule on second brachial.
Cirrus-joints short and carinate, . . . . . .20. pusilla, n. sp.
1. Antedon longicirra, n. sp. (PL XVII.).
$pec ific formula — A. — •
Description of an Individual. — Centro-dorsal somewhat conical, bearing about
twenty cirri. These are enormously long (80 mm. or more), and consist of about eighty
segments, the lower ones of which are longer than wide. The middle joints are slightly
compressed laterally, and gradually develop a dorsal keel, which becomes rather large
in the shorter terminal joints, but is much reduced in size towards the end, and the
terminal claw is very small.
The ends of the basal rays are just seen above the centro-dorsal and the three radials
are visible, all rather long. The first two are oblong and the third pentagonal. The
axillaries and the first two brachials have sharp lateral edges and flattened sides ; all the
radials and the oblong first brachials are very convex, the centre rising to form a sharp
tubercle. Ten arms, of over one hundred joints, the lower ones triangular, but wider
than long, and gradually becoming carinate so as to develop a forward projecting dorsal
spine. Beyond the thirtieth joint they become laterally compressed and begin to overlap.
This is reduced again in the last few joints, which diminish rather rapidly in size.
Syzygies in the third and eleventh brachials, and afterwards at intervals of four to
six joints.
The pinnules are all stiff and styliform, consisting of elongated cylindrical joints.
The second pair are considerably longer than the first, which are relatively small. The
disk and ambulacra are well plated ; but the side plates and covering plates are not fully
differentiated on the pinnules. Sacculi very rare, or absent altogether.
Colour in spirit, — light whitish-brown.
Disk about 7 mm.; spread about 20 cm.
Locality. — Station 192, September 26, 1874 ; near the Ki Islands ; lat. 5° 49' 15" S.,
long. 132° 14' 15" E.; 140 fathoms; blue mud. One specimen.
1 This is also a bidistichate species. See p. 217 and PL XLII.
104 THE VOYAGE OF H.M.S. CHALLENGER.
Remarks. — This fine species is sufficiently distinguished by the great size of its cirri,
which are considerably longer than those of any other Antedon that I have seen with
the exception of Antedon valida (PI. XV. fig. 5). The appearance of the tertiary basals
and of all three radials on the exterior of the calyx, together with the small size of its
lower pinnules, separate it very clearly from the two species which come nearest to it in
the characters of the cirri, viz., Antedon valida and Antedon incerta (PI. XV. fig. 5 ;
PI. XVIII. fig. 4). Both of these show little or nothing of the first radials externally,
and have large and peculiarly modified lower pinnules.
Antedon longicirra is one of the few species of the genus which appear to be
unprovided with sacculi. The careful examination of several pinnules (both decalcified
and otherwise) has altogether failed to reveal their presence except in one doubtful case;
though from my experience with Antedon valida it is not improbable that they may
occur on other pinnules which I have not examined. But there are no notches for their
reception in the imperfectly differentiated side plates. In fact the ambulacra of this
species are more like those of the Pentaerinidae than is the case in any Antedon I know.
For there are no definite side plates, the covering plates resting on a continuous limestone
band which ceases some little way from the end of the pinnule, so that the last few joints
have no ambulacral skeleton at all, as in so many Pentacriniche (see Part I. p. 55 ; pi. xv.
fig. 7).
2. Antedon valida, n. sp. (PI. XV. figs. 5-8).
Specific formula — A. — .
Centro-dorsal large and conical with the ventral angles produced, and bearing about
fifteen cirri. These may reach 80 mm. in length and consist of about sixty -five joints,
of which the seventh to the twelfth are considerably longer than wide. From the
twenty-fifth onwards the joints are short and wide, with a strong dorsal spine.
The first radials just visible ; second and third strongly convex in the middle of their
line of junction. The borders of all three radials and of the lowest brachials are fringed
with strong spines. The axillaries and first brachials have straight edges and flattened
sides. Second brachial and hypozygal of the third flattened on the inner side only.
The junction line of the first two brachials somewhat tubercular.
Ten arms, of triangular, slightly overlapping joints, the later ones somewhat
compressed laterally. Syzygies in the third and about the twelfth brachials, and others
at intervals of four to fifteen joints.
The second brachials have large stout pinnules, the first eight joints of which have
broad and flattened outer sides. The third, fourth, and fifth joints have their inner
edges bent upwards and somewhat thickened, but in the next following joints these are
REPORT ON THE CRINOIDEA. 105
sharpened and form a keel. The third brachial has a similar but smaller pinnule with
flattened outer side. The next two pairs of pinnules have broad and carinate lower
joints, and the later pinnules are more stylifomi, with the two basal joints expanded and
trapezoidal, and the following ones elongated. Disk thickly covered with plates which
extend out on to the arms at the sides of the ambulacra, and also over the genital glands.
Pinnule-ambulacra have well defined side plates and covering plates. Sacculi very rare.
Colour in spirit, — light whitish-brown.
Disk 11 mm.; spread probably 20 cm.
Locality. — Station 214, February 10, 1875; off the Meangis Islands; lat. 4° 33' N.,
long. 127° 6' E.; 500'fathoms; blue mud ; bottom temperature, 41 °'8 F. Two mutilated
individuals, and one younger.
Remarks. — The very stout cirri of this fine species are as long as those of Antedon
longicirra, though consisting of fewer joints. This is owing to the greater length of the
first twenty joints in those of Antedon valida, as will be evident from a comparison of
the figures on Pis. XV. and XVII. The two species also resemble one another in the
almost entire absence of sacculi. Although I have examined many pinnules of Antedon
valida, there is only one in which I have been able to distinguish the sacculi at all
clearly ; and even in this there are not more than about a dozen on the whole pinnule.
They are sufficient, however, to show that sacculi may be present in Antedon longicirra,
Antedon acutiradia, and other species, although I have not been able to find them
on those pinnules which I have examined for the purpose.
Antedon valida and Antedon longicirra are, however, very distinctly separated by
the characters of the pinnules borne on the second brachials. In the latter species this
pinnule is comparatively inconspicuous and smaller than its successor ; but in Antedon
valida, as in Antedon incerta, it consists of short and wide joints, the lowest of which are.
flattened on the outer side, where they meet those of adjacent rays, and cut away on the
inner, so as to give a very singular appearance to the basal part of the pinnule, which it
is not easy to describe. It is well shown, however, on PI. XV. figs. 5, 6, and PI. XVIII.
fig. 5, and it reappears in a modified form in Antedon lusitanica (PI. XXXIX. fig. 2).
This flattening on the outer side of the first pinnule is much better marked in Antedon
valida than in Antedon incerta. But in both species alike the second brachial itself is
not much flattened on its small outer face (PI. XV. fig. 6), though its inner side and that
of the hypozygal of the third brachial are distinctly flattened (PI. XV. fig. 5 ; PI. XVIII.
fig. 5), while the outer face of the hypozygal is in no way specially marked. The distal
pinnules of Antedon valida are remarkable for the expanded and trapezoidal shape of
their two basal joints (PI. XV figs. 7, 8), a feature which scarcely appears at all in
Antedon incerta, though it is characteristic of the group of European and Circumpolar
species of which Antedon eschrichti is the type (PI. XXIV. fig. 13).
(ZOOL. CHALL. EXP. — PART LX. — 1887.) OOO 14
106 THE VOYAGE OF H.M.S. CHALLENGER.
Both Antedon valida and Antedon incerta have a very well-developed anarnbulacral
plating on the disk, which extends out on to the arms above the muscular bundles at the
sides of the ambulacra, as in many Pentacrinidae, and also over the genital glands. The
side plates and covering plates are better differentiated on the pinnules, however, than is
the case in that family. In the immature example of Antedon valida, which was obtained
at the same station as the two individuals above described, the axillaries and lower
brachials are more widely separated and have their sides less flattened than in the adult
condition. The basal joints of the first pinnules, however, have their usual appearance,
and also those of the distal pinnules- The axillaries are rather hexagonal than triangular,
and a considerable portion of the first radials is visible externally, while the arm-
joints are relatively longer and more quadrate, as is always the case in young
individuals.
3. Antedon incerta, n. sp. (PL XVIII. figs. 4, 5 ; Part I., ph liv. figs. 6, 7).
Specific formula — A.—.
Description of an Individual, — Centro-dorsal bluntly conical, with the upper angles
slightly produced. About twenty stout and long cirri, sometimes reaching 50 mm., with
nearly seventy joints. The ninth to twelfth are longer than wide, and the following ones
gradually shorten and develop a dorsal keel, which is most marked in the middle third.
First radials barely visible ; the next two somewhat sharply carinate. The axillaries
and first brachials with sharp edges and flattened sides. The second and the hypozygal
of the third brachial flattened on the inner side only. The junction line of the first two
brachials somewhat tubercular.
Ten arms, of tolerably smooth subtriangular joints, which gradually become quadrate.
A syzygy in the third brachial ; the next between the fifteenth and twentieth, and others
at intervals of seven to fifteen joints.
The lower pinnules are stout, with broad carinate joints, diminishing from the second
to the sixth brachial and then increasing slowly. The later ones are styliform with the
two lower joints slightly expanded. The basal joints of the first pinnule have their
outer sides somewhat flattened, and the third, fourth and fifth joints have their inner
edges truncated, so as to be flattened against the arm.
Disk thickly covered with plates which extend out on to the arms at the sides of the
ambulacra and also over the genital glands. The pinnules have well-defined side plates
and covering plates, most of the former being notched for the presence of sacculi, which
are small, but pretty regularly distributed.
Colour in spirit, — dirty yellowish-white.
Disk about 10 mm.; spread probably about 18 cm.
REPORT ON THE CRINOIDEA. 107
Locality. — Station 170a, July 14, 1874 ; near the Kermadec Islands ; lat. 29° 45' S.,
long. 178° 11' W.; 630 fathoms; volcanic mud; bottom temperature, 39°-5 F. One
specimen.
Remarks. — This is a smaller species than Antedon valida (PI. XV. figs. 5, 6), the
cirri, though containing the same' number of joints, not reaching more than 50 mm. in
length, as compared with 80 mm. in that species. The flattening of the outer side of the
basal pinnules is not so evident, and the lower joints of the distal pinnules show but
little trace of the expanded trapezoidal form which is so characteristic of Antedon valida
(PI. XV. fig. 8).. The sacculi too are much more abundant than in the latter species, the
side plates being notched for their reception, as described on pp. 83, 127 of Part I. and
shown in pi. liv. figs. 6, 7 (tibid.).
The ambulacra extend on to the genital pinnules as is also the case in Antedon
valida. But the plates covering the glands are much less developed than in the species
like Antedon acozla, which have no ambulacra on these pinnules, See Part I., pi. liv.
figs. 1-3.
One of the rays in the single- specimen of Antedon incerta is remarkable for having
the second radial axillary (PI. XVIII. fig. 4). It is smaller and more triangular than the
normal axillary seen in fig. 5, so that the pair of first brachials which it bears are in close
lateral contact with the axillaries of the two adjacent rays.
4. Antedon gracilis, n. sp. (PI. XII. figs. 3-5 ; PL XV. figs. 1-4).
Specific formula — A.—.
Centro-dorsal a low hemisphere with a roughened dorsal pole. About twenty cirri
which reach 30 mm. in length, and consist of fifty to fifty-five joints, a few of which are
longer than wide. The remainder are shorter and begin to overlap dorsally so as
gradually to develop a sharp spinous keel.
First radials scarcely visible ; the second short and sharply convex and the
axillaries widely hexagonal. Both joints and also the first two brachials are slightly
carinate and more or less fringed with small spines. They are also somewhat wall-sided,
with straight lateral edges which extend on to the hypozygals of the third brachials.
Ten arms, of elongately quadrate joints, the outer ones overlapping a little. Syzygies
in the third and about the thirteenth brachials, and then at very irregular intervals.
The second brachial bears a relatively large pinnule of some twenty joints, the lowest
of which are broad, with strong dorsal keels and flattened outer sides. A similar but
rather smaller pinnule on the third brachial ; the next pair much smaller and but slightly
carinate. The following pinnules increase slowly in length, and about the twentieth
108 THE VOYAGE OF H.M.S. CHALLENGER.
brachial they become boat-shaped at the base owing to the width of the third and the
next few joints.
Disk rather incised and much plated, as are also the lower parts of the arms. The
pinnule-ambulacra have fairly well-developed side plates, but the sacculi are small and
rare.
Colour in spirit, — brownish-white.
Disk 6 mm.; spread about 12 cm.
Locality.— Station 214, February 10, 1875 ; off the Meangis Islands ; lat, 4° 33' N.,
long. 127° 6' E.; 500 fathoms ; blue mud ; bottom temperature, 41°"8 F. Five
specimens.
Remarks. — The only example of this species which came into my hands with the
rest of the collection was the much mutilated individual represented on PL XII.
figs. 3, 4. But after this plate had been drawn I received three other specimens in a
much better state of preservation, together with the young form shown on PL XV. fig. 1,
Portions of the arms and pinnules are represented on figs. 2-4 of the same plate.
Ante-don gracilis occupies a curiously intermediate position between Antedon valida
and Antedon incerta on the one hand, and Antedon lusitanica and Antedon breviradia
on the other. The cirri are slender as in these latter species, while the two outer radials
are carinate, as in some forms of Antedon breviradia (PL XL fig. 5 ; PL XII. fig. 4).
This feature, however, also shows itself in the larger forms, Antedon valida and
Antedon incerta; while Antedon gracilis further resembles these types and differs from
Antedon breviradia and its allied species in the fact that the third brachial bears a
pinnule of the same kind as that on the second, smaller in size, but with similarly
carinate lower joints which are flattened on the outer side. This is not the case in
Antedon lusitanica, Antedon breviradia, &c, in which the pinnule of the third brachial
resembles its successor more than it does the large pinnule on the preceding joint. The
fringe of small spines on the radials and lower brachials rather obscures their straight
edged and wall-sided character ; but it is very distinct in the smoother individuals, and
the lateral flattening of the basal joints in the first pinnules indicates the position of the
type very clearly.
The pinnules above the twentieth brachial till near the end of the arm are remarkable
for the characters of the third and following joints (PL XV. figs. 2, 4). These are widely
V-shaped in section, so that the lower part of the pinnule has a boat-shaped appearance
when seen from the dorsal side (PL XV. fig. 4). This expanded part of the pinnule
encloses the genital gland and recalls on a smaller scale a similar arrangement in the
pinnules of Hyocrinus (see Part I., pi. Vc. fig. 10 ; pi. vi. fig. 1).
The enlargement of the lower joints gradually disappears towards the ends of the arms
(PL XV. fig. 2), and there is no indication of it in the pinnules of the youngest
REPORT ON THE CRINOIDEA. 109
individual obtained. This shows much more of the first radials externally than is
visible in the adult (PL XII. fig. 4 ; PL XV. fig. 1); while the pinnule on the third
brachial shows hardly any trace of the enlarged and carinate basal joints which appear in
the adult, but is more like its successor as in the group of species next to be described.
5. Antedon lusitanica, n. sp. (PL XXXIX figs. 1-3).
Specific formula — A.(2).— .
1884. Antedon lusitanica, P. H. Carpenter, Proc. Roy. Soc. Edin., 1884, vol. xii. p. 368.
Centro-dorsal hemispherical, roughened at the dorsal pole, and bearing twelve or
fifteen slender cirri. These reach nearly 30 mm. and consist of about fifty joints, of
which the fifth to the fifteenth are longer than wide. The following ones have a sharp
dorsal spine which is smaller again in the terminal joints.
First radials scarcely visible ; the second relatively short and trapezoidal, with traces
of a median ridge which is continued on to the axillaries. These are short, wide, and
pentagonal, with a slight backward projection in the middle of the proximal edge. The
axillaries and the first two brachials have sharp straight edges and flattened sides.
The second and the hypozygals are also sometimes flattened on their inner sides.
First brachials not much incised, and the outer portions of their dorsal surface are usually
much less convex than the remainder.
Ten arms, of smooth elou gated joints ; but in one individual there are two series of
two distichals each, the axillaries not syzygies. The third and the fourteenth or fifteenth
brachials are syzygial joints.
First pinnule considerably larger than the second ; its lower joints stout and wide,
with the outer sides somewhat flattened. The second and the three or four next joints
have their inner edges produced into strong keels which are slightly folded upwards.
The following pinnules are quite small and increase but slowly in length.
Disk 5 mm. in diameter, thickly covered with numerous small plates, those at the
sides of the ambulacra being rather more regularly arranged than the rest.
Pinnule-ambulacra not well defined, but the sacculi moderately developed.
Colour in spirit, — brownish-white or greenish- white.
Disk 5 mm.; spread probably about 12 cm.
Locality.— H.M.S. "Porcupine," 1870, Station 17a; lat. 39° 39' N., long. 9° 39' W.;
730 fathoms : bottom temperature, 49°-3 F. Ten mutilated specimens.
Remarks. — This is a peculiar species in many ways, and it is very unfortunate that
the ten individuals obtained by the "Porcupine" should have all been so mutilated, the
arms, except in two specimens, having broken away at the syzygy in the third brachials.
These two individuals are shown in PL XXXIX. figs. 1, 3. One is a ten-armed form
110 THE VOYAGE OF H.M.S. CHALLENGER.
with the bases of some arms preserved as far as the second syzygy ; while the other is
peculiar in having two distichal series, each consisting of two articulated joints, so that
the number of arms is raised to twelve. No trace of this arrangement appears on any of
the other nine specimens, but on the other hand there is no indication whatever of its
being due to fracture and subsequent regeneration, as is sometimes the case in other
Comatulae. A similar variation from the ordinary ten-armed type towards the bidisti-
chate group has been described by Dendy 1 in Antedon rosacea, and another is
j)resented by Antedon fiexilis (PL XLIL); while Antedon anceps, Antedon dubia, and
Antedon multispina are ten-armed species which are occasionally varied by the inter-
calation of tridistichate series.
If the discovery of better preserved material should show that the bidistichate
condition of Antedon lusitanica is a natural one and not a mere accidental variation, the
type will be worthy of special notice as the only Antedon found in European Seas which
has normally more than ten arms. It is already distinguished as the only European
Antedon with a plated disk and brachial ambulacra. The condition of the specimens
which I have been able to examine is not such as to afford much information respecting
the character of the ambulacral plates on the pinnules ; but it is sufficient at any rate to
show that sacculi are present and fairly well developed, as is not always the case in
species which have an ambulacral skeleton.
Antedon lusitanica was dredged at 740 fathoms in the East Atlantic, and its nearest
ally is undoubtedly Antedon breviradia, from 630 and 1350 fathoms in the South
Pacific (PI. XIX). Both species have short and wide second and third radials, Antedon
lusitanica especially so, while in most examples of this type the margin of the axillaries
and first brachials is much less rounded than the rest of their dorsal surface, and seems
to stand off from it as lateral processes, a character which is scarcely perceptible in
Antedon breviradia. The first pinnules of the two species are also different. The keels
on the inner edge of their lower joints in Antedon lusitanica are less prominent than in
Antedon breviradia, but at the same time they are more distinctly separated from one
another than is the case in that type ; while the lower cirrus-joints are relatively longer
(PL XIX. figs. 1, 2 ; PL XXXIX. fig. 3).
5. Antedon breviradia, n. sp. (PL III. figs. 4, 5, a-c; PL XL fig. 5 ; PL XIX.; PL
XX. figs. 1, 2).
Specific formula — A.—.
Centro-dorsal hemispherical or bluntly concave, roughened at the dorsal pole, and
bearing fifteen or twenty cirri. These have forty to fifty joints, or a few more, of which
1 Description of a twelve-armed Comatula from the Firth of Clyde, Proc, Roy. Phys. Soc. Edin., 1886, vol. ix.
p. 180, pi. x.
REPORT ON THE CRINOIDEA. Ill
the seventh to tenth are longer than wide. The following ones are shorter and develop
a well-marked spine.
First radials scarcely visible ; the next two short and convex, with occasional traces
of a median ridge, especially in young individuals. Axillaries short and widely
hexagonal, projecting backwards into the second radials. Both joints as well as the first
brachials have straight edges and flattened sides. The inner faces of the second and the
hypozygals of the third brachials are also slightly flattened.
Ten arms ; the lower joints triangular or quadrate, rather longer than wide ; the
distal ones laterally compressed and overlapping so as to become carinate.
A syzygy in the third brachial ; the next between the twelfth and twenty-fifth
(usually about the fifteenth), and others at intervals of from one to fifteen, usually three
or four joints.
The first pinnule, which is much larger than the second, consists of about a dozen
joints. The first six are wide and thick, with their outer sides somewhat flattened, and
the third to the fifth have their inner edges produced into expanded processes which are
slightly folded upwards. The next three or four pinnules on either side are quite small
and the length gradually increases, the later pinnules becoming styliform, with elongated
joints. In some arms the first two joints of the distal pinnules are rather expanded and
trapezoidal, but in others they are not specially modified.
Disk and brachial ambulacra well plated. Side plates and covering plates of the
pinnule-ambulacra generally well differentiated. Sacculi largely developed in some
pinnules and altogether absent in others.
Colour in spirit, — light brownish-white.
Disk 6 mm.; spread probably 16 cm.
Localities. — Station 170a, July 14, 1874 ; near the Kermadec Islands ; lat. 29° 45' S.,
long. 178° 11' W.; 630 fathoms; volcanic mud; bottom temperature, 39°'5 F. Five
specimens, one much mutilated, and another with cysts of Myzostoma murrayi.
Station 175, August 12, 1874 ; near Kandavu, Fiji; lat. 19° 2' S., long. 177° 10' E.;
1350 fathoms; Globigerina ooze ; bottom temperature, 36° F. One imperfect specimen
with a cyst of Myzostoma murrayi.
Remarks. — This is a singular species which unites three forms that I was at first
inclined to consider as distinct. Like Antedon htsitanica it is an exclusively abyssal
type, ranging down from 630 to 1350 fathoms, and individuals from each depth were
infested with the cysts of Myzostoma murrayi, von Graff (PI. XIX. fig. 2).
The second radials are relatively longer than in Antedon lusitanica and more
distinctly incised by the axillaries, which are hexagonal rather than pentagonal as in that
species (PL XL fig. 5 ; PI. XIX. fig. 1 ; PI. XX. fig. 1); while in the younger individuals
both the second and the axillary radials show distinct indications of a median ridge like
112 THE VOYAGE OF H.M.S. CHALLENGER.
that which is so marked in Antedon spinicirra and Antedon acutiradia (PI. XL
figs. 1, 3, 5). The characters of the cirri and of the first pinnules also separate Antedon
breviradia from Antedon lusitanica, which was probably without such distinctly carinate
outer arm-joints as occur in Antedon breviradia (PI. XL fig. 5 ; PI. XIX ; PL XX.
fig. 1). Some of the later pinnules of this species have the lower joints flattened and
expanded as in Antedon valida, while in other arms there is but little trace of this
peculiarity. There is a similar variation as regards the sacculi. On some pinnules they
are abundant, alternating regularly with the side plates ; on others there are very few,
and some pinnules are altogether without them.
The characters of the centro-dorsal and calyx of Antedon breviradia undergo a
considerable amount of change during develojiment, as will be seen on comparison of
figs. 4 and 5 on PI. III., which I at first took to represent distinct species. The centro-
dorsal is deeper and more conical in the older form, while its more numerous cirrus-
sockets are arranged in tolerably regular vertical rows. There are two of these rows
under each interradial angle, each with three sockets, which alternate with those of
adjacent rows, and the dorsal pole is covered with a number of short stout processes of
which there is but little trace in the younger individual (PL III. figs. Ab, 5a). The two
forms also differ in the characters of the radial pentagon. In the younger one (fig. 5b)
its under face is tolerably flat and smooth, with the rosette rather near the surface and
little or no indication of a basal star ; while in the older form (fig. 4c) the rosette is
more deeply sunk within the axial opening, and is surrounded by a fairly definite basal
star. The surface of each radial is also very convex and rises to one or two sharp points
near the middle of its distal edge. These are well seen in the side view (fig. ib), which
is considerably different from that of the younger individual (fig. 5a). The lower part
of the muscle-fossaa is occupied by two or three strongly marked ridges with intervening
furrows, which are altogether absent in the less mature form. The latter, however, has
the upper end of the muscle-plates more everted than in the adult condition, so that the
central opening of the calyx is relatively larger and more pentagonal in appearance
(PL III. figs. 4a, 5c). I have now no doubt, however, that these differences are merely
those of growth, and that they are not of specific value, as I supposed them to be when
PL III. was lettered.
7. Antedon spinicirra, n. sp. (PL XL figs. 1, 2).
Specific formula — A.— .
Centro-dorsal hemispherical or bluntly conical, bearing twenty to twenty-five cirri
with forty to forty-five joints, a few of the lowest of which are longer than wide, while
all but the basal ones have a sharp dorsal spine.
REPORT ON THE CRINOIDEA. 113
First radials just visible ; the second radials and axillaries sharply convex and almost
carinate. The second are partly free laterally and but little incised by the hexagonal
axillaries, which are much wider than loug, with a more rounded dorsal surface, but only
slightly overlapping the distal angles of the second radials. Both joints, and also the
first brachials, have straight edges and flattened sides. The inner sides of the second and
the hypozygals of the third brachials are likewise slightly flattened.
Ten arms ; the second brachials relatively short and oblong, not projecting much
backwards into the first, but both joints are sharply convex at their line of junction.
The next few joints are nearly square and the following ones obliquely quadrate, longer than
wide. The later joints overlap slightly and become somewhat sharply carinate. Syzygies
in the third and about the thirteenth brachials ; others at intervals of two or three joints.
The first pinnule larger than the second. Its lower joints relatively stout, with some-
what flattened outer sides, and the inner edges of the third to the fifth joints slightly
carinate. The pinnules of the third and following brachials small and increasing slowly
in length, the later ones sometimes showing a faint expansion of the two basal joints.
Disk much incised and well plated, and the brachial ambulacra slightly so. Pinnule-
ambulacra tolerably well defined, the side plates with intervening sacculi.
Colour in spirit, — light brownish-white.
Disk 4 mm.; spread probably about 9 cm.
Locality. — Station 164, June 12, 1874; near Port Jackson; lat. 34° 8' S., long.
152° 0' E.; 950 fathoms; green mud; bottom temperature, 36Q,5 F. Five specimens,
two much mutilated.
Remarks. — There are several points of resemblance between this species and the
younger forms of Antedon hreviradia, which show a tendency to carination of the two
outer radials (PL XL fig. 5 ; PL XX. fig. 1). But the radials differ considerably in their
other characters, while Antedon spinicirra has fewer cirrus-joints than the larger Antedon
hreviradia, with the basal ones relatively shorter and more spinous than in the latter
type. Another point of difference is afforded by the first pinnules, the lower joints of.
which are much less expanded and carinate in Antedon spinicirra than in equally
developed forms of either Antedon hreviradia or Antedon acutiradia. This last is
distinguished from both the preceding species by the great relative length of the radial
axillaries (PL XL fig. 3).
8. Antedon acutiradia, n. sp. (PL XL figs. 3, 4).
Specific formida — A. (~?j •
Centro-dorsal hemispherical, bearing about fifteen cirri, which have the fourth and
some of the following joints much longer than wide, with traces of dorsal spines.
(ZOOL. CHALL. EXP. PART LX. — 1887.) 0°° 13
114 THE VOYAGE OF H.M.S. CHALLENGER.
First radials just visible ; the second partly free laterally and deeply incised by the
sharp proximal angles of the axillaries, which are longer than wide. Both joints are
very sharply convex and almost carinate, but the axillaries are wider and have a more
rounded surface than the second radials, which are partly hidden beneath their lateral
angles. Both joints and also the first brachials have straight edges and flattened
sides. The inner sides of the second and the hypozygals of the third brachials are also
flattened.
Ten arms; the first brachials somewhat incised for the sharp proximal angles of the
second,1 both joints rising to their line of junction. The fifth and following joints smooth
and obliquely triangular, much longer than wide, the later ones becoming obliquely
quadrate.
Syzygies in the third and sixteenth brachials, and then at intervals of three or four
joints.
First pinnule much larger than the second ; its lower joints wide and thick, with
somewhat flattened outer sides. The third to fifth have their inner edges produced into
expanded processes which are slightly folded upwards. The next pair of pinnules are
rather larger than their immediate successors, but the following ones are quite small and
increase very slowly in length.
Disk well plated and the brachial ambulacra slightly so ; pinnule-ambulacra without
very definite side plates ; the presence of sacculi uncertain.
Colour in spirit, — light brownish-white.
Disk 4 mm.; spread probably 10 cm.
Zoca^'ty.— Station 175, August 12, 1874; near Kandavu, Fiji; lat. 19° 2' S., long.
177° 10' E.; 1350 fathoms; Globigerina ooze; bottom temperature, 36° F. Two
mutilated specimens.
Remarks. — This type is unfortunately only represented by two calyces and half a
dozen arm-fragments with their pinnules mostly broken. No entire cirri are preserved,
and the position which I have assigned to this species among those with thirty to fifty
spiny cirrus-joints is therefore a somewhat conjectural one. But it has so many points
of resemblance with Antedon spinicirra and the two preceding species, that I have little
doubt respecting the character of its cirri.
It is most closely allied to Antedon spinicirra (PL XL fig. l), but differs in the
sharper carination and the greater relative length of the axillaries (PL XL fig. 3).
The second radials are much compressed laterally so that they appear, as it were, at
a lower level than the axillaries, the lateral angles of which overlap and partly conceal
them. Traces of this arrangement are visible both in Antedon spinicirra and in
Antedon bisptnosa. In the former species the enlargment and carination of the lower
1 There is a considerable amount of variation in this respect.
REPORT ON THE CRINOIDEA. 115
joints of the first pinnules is less marked than in Antedon acutiradia, which in this
respect rather resembles Antedon breviradia.1
The state of preservation of the pinnules in the two individuals under consideration
is unfortunately such that it is impossible to speak positively respecting the presence or
absence of sacculi. But there is no trace of them in any of the few pinnules that I have
been able to examine.
9. Antedon bispinosa, n. sp. (PI. XX. figs. 3, 4).
Specific formula — A. — .
Description of an Individual. — Centro-dorsal almost columnar, bearing about twenty-
five cirri on its sides. These have thirty to thirty-five joints, the three lowest of which
are almost saucer-shaped, and the next ones much longer than wide. The remainder are
shorter and acquire a marked keel which becomes reduced to a spine in the terminal
joints.
Three radials visible, the distal edges of the first fringed with blunt spines.
Axillaries pentagonal, with a curved base, overlapping the short second radials laterally.
Each joint has a rounded and spinous centre raised above the lateral portions, which
meet those of adjacent radials by flattened sides.
Ten arms ; the margins of the lowest brachials fringed with blunt spines. First
brachials rounded and short in the middle line, but with depressed lateral portions
which meet one another by flattened surfaces all round the calyx. Second brachials
more square and scarcely projecting backwards into the first. The eighth and following
brachials become quadrate and slightly overlapping, with two or three large curved spines
near the distal edge, which become very prominent in the outer portions of the arms.
Syzygies in the third and eleventh to fourteenth brachials, with others at intervals of
three or more joints.
The lower pinnules all very spiny ; the first much larger than its immediate
successors, with the three or four basal joints somewhat flattened on the outer side, and
the second to fifth with the inner edges slightly keeled and folded upwards. The
pinnule on the third brachial but little larger than that on the fourth, and the following
ones become gradually longer, with overlapping spinous joints.
Disk strongly plated, and the brachial ambulacra irregularly so. Pinnule-ambulacra
with large covering plates and ill-defined side plates. Sacculi rare.
Colour in spirit, — white, with dark brown patches on the calyx.
Disk 6 mm.; spread probably 10 cm.
Locality.— Station 147, December 30, 1873; lat. 46° 16' S., long. 48° 27' E.; 1600
fathoms ; Diatom ooze ; bottom temperature, 3 4° "2 F. One specimen.
1 This character is hardly visible in the view of the calyx which is represented in PI. XI. fig. 3.
116 THE VOYAGE OF H.M.S. CHALLENGER.
Remarks. — This species has such very definite characters that it is not likely to be
confounded with any other. The spiny calyx and the double row of long hook-like
spines along the arms distinguish it very clearly. The radial axillaries come into contact
above the depressed lateral portions of the second radials just as in Antedon acuticirra,
and there is much the same sort of relation between the first and the second brachials.
It is rather a robust species for such a considerable depth (1600 fathoms). But the
sacculi are poorly developed, as is so often the case in the abyssal Comatulse.
10. Antedon latipinna, n. sp. (PI. X. fig. 3).
Specific formula — A. .
Description of an Individual. — Centro-dorsal subcorneal and marked by twenty
cirrus-sockets disposed in ten vertical rows. About forty joints in the cirri, a few of
them longer than wide. The remainder' are shorter and begin to overlap dorsally so as
to develop a sharp spinous keel.
First radials partly visible ; the second rather convex, short and oblong ; axillaries
pentagonal, with slight backward projections, wider than the second, but barely twice as
long. Both joints, together with the first two brachials and the hypozygal of the third,
have straight lateral edges and small portions of the outer sides flattened.
Ten arms, of short and smooth quadrate joints. Syzygies in the third and twelfth
brachials, with others at intervals of seven to nine joints.
The second brachial has a short stout pinnule of about fifteen joints, the lowest of
which are short, wide, and slightly carinate, but not flattened laterally. The following
pinnules diminish to about the third pair and then gradually increase, their joints
becoming elongated. Disk much incised and well plated. Side plates fairly distinct on
the pinnule-ambulacra ; sacculi apparently absent.
Colour in spirit, — light brownish-white.
Disk about 4 mm.; spread probably about 8 cm.
Locality.— Station 232, May 12, 1875; lat. 35° 11' N., long. 139° 28' E.,
345 fathoms ; green mud; bottom temperature, 41°"1 F. One mutilated individual.
Remarks. — This species differs from all the preceding ones in the characters of the
first pinnule, the lowest joints of which, though wide and slightly carinate, have no
indication of the flattening on the outer side which is so characteristic of Antedon valida
(PI. XV. figs. 5, 6), Antedon breviradia and others. The first two brachials and the
hypozygal of the third have the usual wall-like sides and straight edges, but these features
are less marked on the two outer radials. The cirri are arranged in ten very regular rows
on the centro-dorsal, which is another character of separation from the species previously
REPORT ON THE CRINOIDEA. 117
described ; though traces of it are apparent in Antedon breviradia (PL III. fig. 4b).
Only a few fragments of the arms are preserved, and no traces of sacculi are visible in
the broken pinnules which I have been able to examine.
11. Antedon multispina, n. sp. (PI. XIII. figs. 1-3; PI. XIV. figs. 5-7; PI. LXIX.
figs. 1-4).
Specific formula — A. ( 3- 9 ) -y-
Centro-dorsal hemispherical, bearing about twenty cirri of twenty-five to thirty joints,
a few of which are longer than wide. The remainder are shorter and overlap slightly
so as to develop a dorsal spine.
First radials invisible in the adult ; second very short (in the adult) and axillaries
widely pentagonal. The axillaries and first brachials have flattened outer sides and
straight edges ; and the inner sides of the second and hypozygals of the third brachials
are also flattened. Numbers of small spines on the calyx and arm-bases.
Usually ten arms, but one individual has two tridistichate series. Arm-joints
elongately quadrate, with tufts of numerous small spines at one or both ends. The first
pair of brachials borne on the distichal axillary are united by syzygy ; but above the
radial axillaries the third brachial is a syzygy, the next between the ninth and fifteenth,
and others at intervals of three or four joints.
The second brachial bears a pinnule of about twenty-five joints, the lowest of which
are wide, with their inner edges cut away a little and the outer sides slightly flattened.
The next pair of pinnules are much smaller, and the following ones gradually increase in
length, with the lower joints at first broadly V-shaped, but afterwards more elongated.
Disk and arms well plated. Side plates fairly developed on the pinnule-ambulacra,
with moderately abundant sacculi.
Colour in spirit, — -light brownish-white.
Disk 4 mm.; spread probably about 10 cm.
Loccdity. — Station 344, April 3, 1876 ; near Ascension ; lat. 7° 54' 20" S., long.
14° 28' 20" W.; 420 fathoms; volcanic sand. Four broken individuals and three
Pentacrinoid larvae.
Remarks. — This species has perplexed me a good deal, on account of the mutilated
condition of the specimens, three of which are quite immature, while the fourth, which is
apparently full grown, has twelve arms owing to the presence of two tridistichate series
(PL LXIX. figs. 1,2). I was at first inclined to regard it as a small variety of Antedon
porrecta (PL LII. fig. 3) which occurs at the same station. But the tridistichate
character is the only resemblance between the two forms, their cirri, arms, and pinnules
being altogether different; and I am therefore forced to conclude, as with Antedon
118 THE VOYAGE OF H.M.S. CHALLENGER.
lusitanica, either that the twelve-armed condition is a monstrosity, or that Antedon
multispina is a dimorphic species. In the latter case it is rather a curious one. For in
all the four arms which are borne on the two distichal axillaries (PI. LXIX. figs. 1, 2)
the first pair of brachials are united by syzygy just as in Antedon angusticalyx,
Antedon distincta, and Antedon ineequalis (PI. L. fig. 1; PL LI. figs. 1, 2), which do
not conform- to the ordinary rule of a syzygy in the third brachial.
Antedon multispina differs from the species already described in the preceding pages
in the small number of its cirrus-joints, which does not seem to exceed thirty. It
resembles most of them, however, in having a relatively small pinnule on the third
brachial, which is more like its successors (PI. XIII. fig. 3) than its predecessor. The
two outer radials, especially the second which are very short, can hardly be described as
wall-sided ; but this feature is very marked on the outer side of the first brachials and
on the inner side of the second and the hypozygals of the third brachials (PI. LXX.
figs. 1-3), while the first pinnules of adjacent rays are flattened laterally against one
another and their inner sides are slightly cut away at the base, so as to recall the condition
of Antedon incerta and allied species (PI. XVIII. fig. 5).
From about the twelfth brachial onwards the third and next following pinnule-joints
are expanded for the protection of the genital glands, having a broadly V-shaped section,
though this is less marked than in Antedon gracilis (PL XV. fig. 4).
Besides the three young individuals of Antedon multispina, one of which is figured
on PL XIII. fig. l,the Challenger also dredged three Pentacrinoid larvae, which presumably
belong to this species, as it is the only ten-armed form met with at this station. They
are relatively much larger and more robust than the corresponding larval stages of
any other species which I have seen, with the exception of Antedon eschrichti.
Figs. 3-7 on PL XIV. represent five larvae, all equally magnified, which belong respect-
ively to Antedon hystrix ? (fig. 3), Antedon tenella (fig. 4), and Antedon multispina
(figs. 5-7). Figs. 3-5 illustrate almost the same developmental stage in the three
different species, that namely when the first cirri make their appearance and a fair
number of arm-joints have been formed. Of the three larvae, that of Antedon tenella
is the oldest, having pinnules on the outer parts of the young arms, but it is altogether
less robust than that belonging either to Antedon hystrix? or to Antedon multispina.
The latter is remarkable for the shortness of its stem, which has only thirteen joints
below the centro-dorsal, the two lowest being quite short and resting on a large and
expanded dorsocentral plate (PL XIV. fig. 5). The centro-dorsal is a thin plate, but
little larger than the joints below it, and the rudiments of three cirri have appeared upon
it, the positions of the other two being indicated by imperfect sockets. In the next
stage (PL XIV. fig. 6) the first pinnules have appeared on the arms, not at their bases,
but about the eleventh or twelfth brachial, and the five radial cirri which were* first
formed are well developed so far as can be judged from their basal joints, which is all
REPORT ON THE CRINOIDEA. 119
that remains of them, while there are one or two slight indications of the second whorl
of cirri, the positions of which alternate with those of the first. The centro-dorsal has
increased considerably in thickness, as has also the joint below it, which has similar re-
entering angles, just as is the case in the infra-nodal joint in the stem of the Penta-
crinidse. Both of these larvse have quite low basals as compared with those of the other
species figured on the same plate, and especially Antedon hystrix? In this respect
they approach the Pentacrinidaa and the typical Apiocrinidae rather than Rhizocrinus
and Bourgueticrinus, which they resemble in the characters of the middle and lower
stem-joints. In the oldest larva, however, the basals are entirely concealed by the
centro-dorsal, which has now reached a considerable size, with the second whorl of cirri
well developed and even traces of a third, while there is only one discoidal joint below it.
(PL XIV. fig. 7). In Antedon multi 'spina, therefore, the basals become entirely concealed
before the end of the Pentacrinoid stage, as in Antedon tenella, though it is not the case
in Antedon rosacea.
The youngest of the three immature forms of Antedon midtispina is considerably
more advanced than the oldest Pentacrinoid larva. Not only the basals, but also portions
of the first radials are concealed, and the first two pairs of pinnules have appeared, but
from the fifth to the twelfth brachials the arms are devoid of pinnules. In the still
older form, shown in PI. XIII. fig. 1, the first radials are only just visible externally,
though the second are relatively much longer than in the mature form. All the arm-
joints are provided with pinnules, though the lowest ones are quite small, that on the
second brachial being much more like its successor than is the case in the adult ; while
there is but little trace of any expansion in the lower joints of the genital pinnules.
The spines of the cirri are present from the first, but those on the calyx, arms, and
pinnules do not appear till after the Pentacrinoid stage, while the lateral flattening of
the radials and lower brachials is one of the last characters to make its appearance.
This, of course, is only to be expected, for it is only when the arms become tolerably
wide that their lower portions come into close lateral contact.
In the arms of the larvse, as in the pinnules of the adult, the covering plates are
supported upon imperfect side plates. These alternate very regularly with the sacculi,
which are relatively much more abundant than in the adult.
12. Antedon echinata, n. sp. (PI. XXI. figs. 4, 5).
Specific formida — A.-r-.
Description of an Individual. — Centro-dorsal a low hemisphere with about twenty
cirri on its sides. These have some twenty-five joints, of which the fifth is longest, with
a slight dorsal projection at its distal edge which becomes a spiny keel in the short later
120 THE VOYAGE OF H.M.S. CHALLENGER.
joints. First radials just visible ; the second nearly oblong, not very convex, and barely
united laterally. Axillaries about twice their length, broadly pentagonal, with slight
backward projections. First two brachials nearly oblong. All these pieces have sharp,
straight edges fringed with spines, and very slightly flattened sides.
Ten arms, of smooth, obliquely quadrate joints, as long or longer than wide.
Syzygies in the third and about the thirteenth brachials, with others at intervals of
two to four joints.
First pinnule not much larger than that on the third brachial, and consisting of about
twelve joints, of which the first five are rather expanded, with the inner edges a little
cut away, and all have tufts of small spines along the dorsal border. The next two pairs
of pinnules decrease slowly in length and become less spinous. The later ones are long,
slender, and tolerably smooth.
Disk much incised and well plated ; brachial ambulacra but slightly so. The pinnule-
ambulacra have fairly definite side plates, and large sacculi are occasionally present.
Colour in spirit, — light brownish-white.
Disk 3 mm.; spread probably 60 or 70 mm.
Locality. — Station 170a, July 14, 1874 ; near the Kermadec Islands; lat. 29° 45' S.,
long. 178° 11' W.; 630 fathoms; volcanic mud; bottom temperature, 39°'5 F. One
specimen.
Remarks. — This little form clearly belongs to the group of species with spiny cirri
and relatively large first pinnules ; but it is distinguished from its allies by a few well-
marked characters. The number of the cirrus-joints does not seem to exceed twenty-five ;
while the bases of the rays have but slightly flattened sides, and the accompanying-
peculiarities of the lower joints of the first pinnules are barely recognisable (PI. XXI.
fig. 5). It approaches Antedon multispina in the abundance of the spines on the radials,
lower brachials and their pinnules ; but it differs altogether from that species in the
smoothness of the distal arm-joints as well as in the appearance of the first radials
externally, and in the relatively larger size of the pinnule on the third brachial. Some
of the later pinnules have a few sacculi of unusually large size, but others are entirely
without them.
13. Antedon lasicurva, n. sp. (PI. II. figs. 2, a-d; PL XXI. fig. 3; PI. XXII.
figs. 3, 4 ; woodcut, fig. 3 ; also Part I., pi. liv. fig. 9 ; pi. lv. fig. 7).
Specific formula — A.-r.
Centro-dorsal hemispherical, with a very rough dorsal pole and small interradial
processes. About twenty cirri, of eighteen to twenty very stout joints, most of which
REPORT ON THE CRINOIDEA. 121
are longer than wide. The earlier joints overlap slightly, and the later ones more so,1
especially on the dorsal side, so as to produce a blunt spine at the distal edge which is
rather sharper on the penultimate.
First radials entirely concealed in the adult, and sometimes portions of the second
also. These are short and band-like, in close lateral contact, with raised edges which
are often somewhat crenated, and there is usually a slight tubercle in the middle of the
distal border, corresponding to one on the axillary. This is short and pentagonal with a
wide, open angle and more or less crenated edges. The dorsal surface is very convex,
with the margins more or less flattened, and wall-like sides. First brachials short, nearly
oblong and closely united ; the second more wedge-shaped. Both joints rise towards
their apposed edges to form a median elevation like that between the second and third
radials. The first three brachials wall-sided with flattened margins like the axillaries.
The following joints short till about the twelfth, after which they are longer and more
triangular, gradually becoming quadrate ; the terminal ones elongated and slightly
compressed laterally. Ten arms of about one hundred and twenty joints. In the lower
parts of the arms the distal edge of each joint stands up as a sharp crenulated ridge from
which the surface slopes backwards. As the joints become longer, further out on the
arms, this sudden rise disappears, and they overlap in the ordinary way.
Syzygia in the third brachials and then very variable in position. The next between
the ninth and sixteenth brachials, and others at intervals of one to sixteen (usually
three to seven) joints.
The second brachial has a short pinnule of about twenty -two joints, of which the six
lowest are trihedral and rather broad, and much flattened on the outer side, with a
marked dorsal keel which is lost in the smaller terminal joints. A similar but rather
smaller pinnule on the third brachial. The next pinnule has fewer joints, but the third
and fourth are relatively broader, and in the succeeding pinnules very much so, with
their outer faces greatly expanded towards the ventral side. This condition is most
marked about the twelfth brachial, and then gradually decreases, being traceable to the
twenty-fifth or thirtieth. After this it is lost and the pinnnles gradually diminish in
stoutness, but do not increase much in length.
Disk much incised and completely plated, as are also the arms, both along the
ambulacra and at their sides. The genital glands protected by stout anambulacral
plates.
The ambulacra of the distal pinnules have well-defined side plates alternating with
but often partly concealing the sacculi. These are abundant and very large, especially
on the genital pinnules.
Colour in spirit, — young individuals a yellowish-brown ; the older ones a dark grey-
brown.
1 This is not well shown in the only cirrus remaining on the figured specimen.
(ZOOL. CHALL. EXP. — PART LX. — 1887.) OoO 16
122
THE VOYAGE OF H.M.S. CHALLENGER.
Disk 7 mm.; spread about 20 cm.
Localities. — Station 170a, July 14, 1874; near the Kermadec Islands; lat. 29° 45'
S., long. 178° 11' W.; 630 fathoms; volcanic mud; bottom temperature, 39°-5 F.
Several specimens.
Doubtful. — Station 175, August 12, 1874 ; near Kandavu, Fiji; lat. 19° 2' S., long.
177° 10' E.; 1350 fathoms; Globigerina ooze ; bottom temperature, 36° F. Some arm-
fragments only.
Remarks. — This species and the following one (Antedon incisa) are sharply
distinguished from the preceding members of this group of ten-armed Antedons by their
smooth, stout cirri, and the peculiar expansion of the lower joints in the proximal and
middle pinnules (PL XXI. fig. 2). The only other Antedon in which the latter character
Fig. 2. — Antedon basicurva, x 3. A. Side view of the calyx and arm-bases after the removal of three rays, so as to show
the sides and inner faces of the other two. The two outer radials, two lower brachials, and in a less degree also the
third and fourth, have their outer sides flattened against one another. The genital pinnules have the third and fourth,
and sometimes the fifth joints greatly expanded, but the following ones are smaller. B. The lower part of an arm from
its inner side, to show the flattened inner faces of the first three brachials, including both the hypozygal and the
epizygal of the third.
is similarly develojaed is a tridistichate species, Antedon inzequalis (PI. LI. fig. 2), which
occurs at the same two stations (Stations 170a and 174) as Antedon incisa, Antedon basi-
curva having been found at the former only. It is possible too that Antedon basicurva
and Antedon insequalis were obtained at Station 175 j1 though there appears to be much
doubt upon this point. But whether this be the case or not, it is clear that the pecu-
liarity in question is a local one and limited to this particular region of the South Pacific.
1 Only two Comatulce are recorded in the Station Book as having been found at this locality ; and as the depth is
considerable (1350 fathoms), I have little doubt that they are the two small forms of Antedon breviradta and Antedon
acutiradia already described. The arms of the latter were all loose, however, and it is quite possible that the few arm-
fragments of Antedon basicurva may have been amongst them ; but no calyx of this species was obtained.
REPORT ON THE CRINOIDEA. 123
The first pair of pinnules in Antedon hasicurva are considerably different from their
successors. That on the second brachial is rather the larger of the two ; but their general
characters are identical (PI. XXII. fig. 4). All the joints are quite short, but the first
five or six are broad, carinate and trihedral, with their outer sides flattened, somewhat
as in Antedon valida and allied species (PI. XV. fig. 6). This is well seen in the
woodcut (fig. 3). Traces of this lateral flattening are apparent in the pinnule on the
fourth brachial, but those on the fifth and following brachials have the third and
fourth joints very broad and expanded (PI. XXII. fig. 3), though the fifth joint is smaller
again and its successors very much so. These lower joints, which are so broad and almost
flat on their outer side, afford support and protection to the genital glands which are
situated on their inner faces. The ventral surface of the glands is covered by a pavement
of anambulacral plates, often with large sacculi imbedded in them here and there as
shown in Antedon incisa (PL XXI. fig. 2, a). But there are no side plates and covering
plates as in the distal pinnules.
This expansion of the third and fourth pinnule-joints is best developed about the
tenth or twelfth brachial, after which it gradually becomes less and less marked and the
later joints more and more elongated. But the third and fourth joints are often
distinctly broader and flatter than their successors as far out as the thirtieth brachial,
after which they assume a more elongated form.
In one quite young specimen, only about one-third the size of that figured on
PL XXII. , there is comparatively little trace of this expansion of the third and fourth
joints, even on the lower pinnules (PL XXI. fig. 3). The arms too are much smoother
than in the adult, the edges of the lower brachials being but slightly raised, and showing
no trace of the crenulation which is so marked in the more mature forms (PL XXII.
fig. 3). The first radials are just visible as narrow curved bands immediately above the
centro-dorsal, which are not smooth and continuous as usual, but broken here and there
by pits. In a slightly older individual they are only represented by a row of irregular
processes between the centro-dorsal and the second radials ; while in the mature form
they are altogether invisible, though traces of these processes appear after the removal
of the second radials (PL II. fig. 2a).
The upper surface of the centro-dorsal is so much larger than the base of the radial
pentagon (PL II. figs. 2, a, c, d) that the second radials partly rest upon it and so
completely conceal the first, as in some forms of Antedon rosacea.
The cirrus-sockets are peculiar for having a very large articular facet in the centre,
from which radiating processes extend all round to the margin of the socket, as seen in
PL II. fig. 2a. The dorsal surface of the radial pentagon is marked by a well-defined
basal star, the angles of which do not, however, appear externally (PL II. fig. 2c). The
central funnel of the calyx (PL II. fig. 2d) is smaller than in Antedon breviradia
(PL III. fig. 4a), as the ventral ends of the muscle-plates are less everted than in that
124 THE VOYAGE OF H.M.S. CHALLENGER.
type. There is a further resemblance between Antedon basicurva and the mature
Antedon breviradia, in the presence of transverse ridges and furrows at the lower ends of
the muscle-fossee (PL II. fig. 2<x ; PL III. fig. 46). I have found two radials without
them, however, in one calyx of Antedon basicurva.
Some of the individuals of this species, including the youngest one above mentioned,
are deformed by the cysts of Myzostoma willemoesii, and Myzostoma tenuispinum, von
Graff.
14. Antedon incisa, n. sp. (PL II. figs. 1, a-d ; PL XXI. figs. 1, 2).
Specific formula — A.—.
Centro-dorsal hemispherical, with a rough dorsal pole and small interradial processes.
About fifteen cirri of some fifteen to eighteen rather stout, smooth joints, most of which
are longer than wide. In the younger cirri the later joints overlap very slightly on the
dorsal side so as to produce faint spines ; this is lost in the older cirri, except in the
penultimate, which bears a strong opposing spine.
Three radials visible ; the first short and band-like with curved borders, meeting one
another above the interradial processes of the centro-dorsal ; the second somewhat longer,
in close lateral contact, and rather convex in the centre, but little incised for their
junction with the axillaries, which are also sharply convex, short and pentagonal, with
very open angles. The axillaries and the first four or five brachials have the marginal
portions of their dorsal surface flattened vertically, with sharp edges and wall-like sides.
Ten arms ; the first brachials almost oblong, and very convex in the centre ; the second
shorter and more wedge-shaped. The following joints smooth and rather short till about
the tenth, then longer and obliquely quadrate, becoming blunter and more elongated
towards the end of the arm, but without any overlap.
A syzygy in the third brachial ; the next frequently in the tenth, but sometimes not
till the eighteenth ; and others at intervals of one to nine, generally four to six, joints.
The second brachial has a longish pinnule of about thirty short joints, the lower ones
rather flattened on the outer side with a sharp dorsal keel. A similar but shorter pinnule
on the third brachial. The next pinnule is longer, with fewer but larger joints, the
fourth and fifth of which are broad and expanded towards the ventral side. The
expansion increases in the following pinnules, which have the third and fourth joints
largest. This is most marked about the twelfth, but is traceable nearly to the thirtieth
brachial. The remaining pinnules diminish in stoutness without increasing much in length.
Disk much incised and pretty completely plated, as are also the brachial ambulacra.
The genital glands protected by stout anambulacral plates. The ambulacra of the distal
pinnules have well-marked side plates, which are generally notched for the sacculi.
These are large and abundant, especially on the genital pinnules.
REPORT ON THE CRINOIDEA. 125
Colour in spirit, — brownish-white, or light yellowish-brown.
Disk 7 mm.; spread about 18 cm.
Localities. — Station 170a, July 14, 1874 ; near the Kermadec Islands ; lat. 29° 45' S.,
long. 178° 11' W.; 630 fathoms; volcanic mud; bottom temperature, 39'5° F. Four
specimens and two fragments.
Station 174 (b, c, or d), August 3, 1874 ; near Kandavu, Fiji; lat. (about) 19° 6' S.,
long, (about) 178° 18' E.; 255, 610, or 210 fathoms '; coral mud; bottom temperature
(at 610 fathoms), 39° F. Two specimens.
Remarks. — This species is readily distinguished from Antedon basicurva by the
smoothness of the arms and the appearance of the first radials externally (PI. XXI. fig. 1 ).
Another point of difference is that the wall-sidedness of the arm-bases extends out to the
fourth, or even to the fifth, brachial, which is not the case in any of the preceding
species. On the other hand the basal joints of the first pair of pinnules in the type form
(from Station 170a) are less flattened and not so distinctly trihedral as in Antedon basicurva
which occurs at the same station, so that the proximal pinnules are more like their
successors than is the case in that species. It is curious, however, that in the two
individuals from Station 174 (b, c, or d), where Antedon basicurva did not occur, the
lower joints of the first pinnule show a distinct tendency towards the trihedral form
characteristic of that type (PL XXII. fig. 4).
The first radials are completely visible in all the specimens of Antedon incisa, though
they are shortest in those which are most mature (Pi. XXI. fig. 1), barely reaching half
the length of the second radials, to which they are nearly equal in some of the younger
individuals. A comparison of figs. 1 and 2 on PL II. will show other differences between
the calyx of this type and that of Antedon basicurva. The former has a relatively
smaller centro-dorsal, so that the radial pentagon covers it completely, and no part of it-
is exposed when the second radials are removed (PL II. figs, id, 2d). The under view
of the radial pentagon is much the same in the two types, except for the portion of the
first radial, which appears externally in Antedon incisa (PL II. fig. lc). But the lower
ends of the muscle-fossae in the latter species are almost entirely without the ridge and
furrow markings which are generally present in Antedon basicurva; the articular facet of
the cirrus-socket is relatively smaller in Antedon incisa than in Antedon basicurva,
and the radiating processes round the edge of the socket are less distinct (PL II. figs.
la, 2a).
One of the six individuals obtained at Station 170a bore two large cysts of Myzostoma
tenuispinum, von Graff (PL XXI. fig. 1), a species which also infests Antedon basicurv<<
at the same station ; but there is no trace of cysts on either of the two examples of
Antedon incisa obtained at Station 174.
1 The exact station, and consequently the exact depth, are not recorded.
126 THE VOYAGE OF H..M.S. CHALLENGER.
15. Antedon tubcrosa, n. sp. (PI. XIV. fig. 9 ; PL XXIII. fig. 2).
Specific formula — A. — .
Centro-dorsal a thick disk with a rough dorsal surface. Fifteen to twenty cirri of
thirteen to fifteen joints, of which the fifth is longest. The following ones gradually
acquire a dorsal keel which passes into the opposing spine of the penultimate.
First radials concealed ; second and third both short and wide, the former closely
united laterally and the latter pentagonal with very open angles. The radials and the first
three brachials are straight-edged and wall-sided, and more or less carinate in the middle
line. Ten arms, of nearly one hundred and fifty joints ; small blunt tubercles on the
surface of the radials and arm-bases. First brachials oblong and closely united, the
second more wedge-shaped ; the third to the tenth brachials saucer-shaped, their distal
edges being more or less raised and crenulated. This feature disappears in the following
joints, which are more triangular, and elongate considerably towards the end of the arm.
The lowest joints, especially in the younger arms, have a marked dorsal keel, which
gradually dies away in the middle third, the terminal joints being quite smooth.
A syzygy in the third brachial ; the next between the tenth and twenty-sixth brachials,
and others at intervals of four to twelve, usually six to eight joints.
The first pair of pinnules tolerably equal, consisting of twenty to twenty-five joints,
the lowest of which are broad and slightly keeled. The next pinnules slowly increase in
length and in size, the third and following joints being expanded to receive the genital
glands, which are protected by strong plates. The fourth and fifth are larger than the rest,
but not markedly so. This ceases rather beyond the first third of the arm, and the
pinnules then become more slender, with the basal joints square or longer than wide.
Disk much incised and paved with small plates ; the arms moderately so, and the pinnule-
ambulacra have distinct side plates with intervening sacculi, which are also abundant in
the plating over the genital glands.
Colour in spirit, — the young arms nearly white, the older ones a dark brownish-grey.
Disk 5 mm.; spread 25 cm.
Locality. — Station 210, January 25, 1875; off the Panglao and Siquijor Islands;
kit, 9° 26' N., long. 123° 45' E.; 375 fathoms; blue mud ; bottom temperature, 54°1 F.
One entire specimen and another much broken.
Remarks. — The absence of the first radials externally and the roughness of the arm-
bases give this type a certain amount of superficial resemblance to Antedon basicurva
(PL XXII. fig. 3). But it differs altogether from that species and from Antedon incisa in
the characters of the pinnules, though the genital glands are protected by strong anambu-
lacral plates, as in both these types. In these again the third and fourth joints of the
genital pinnules are specially distinguished from the rest by the great breadth of their
REPORT ON THE CRINOIDEA. 127
outer sides, which are enlarged at the expense of the inner sides (PI. XXI. fig. 2 ; fig. 3
on p. 122). But the expansion of the pinnule-joints is much more uniform in Antedon
tuberosa, as they increase and then decrease gradually from the base to the end of the
pinnule, while both sides of each joint are enlarged, though the outer is slightly more so
than the inner one. In these characters Antedon tuberosa resembles the smoother
Antedon parvipinna and the ten-armed variety of Antedon fiexilis ; but apart from its
occasional bidistichate character (PL XLIL), the latter form is readily distinguished by
its longer cirri, the appearance of the first radials externally, and by the smoothness of
the arm-bases.
All three species are remarkable for the great development of the sacculi, which are
found not only between the side plates of the distal pinnules, but also along the medio-
ventral line of the plated genital pinnules as in Antedon incisa (PL XXI. fig. 2a) ; but
they are often absent in this position in other species, as, for example, Antedon acosla
(see Part I. pi. liv. figs. 1-3).
A young larva was obtained at Station 210, which must belong either to Antedon
tuberosa or to Antedon distincta (PL LI. fig. 1), the only two species found at this station.
The latter, however, is a multibrachiate form with both distichal and palmar series, and
from the appearance of the larva I think that it should most probably be referred to
Antedon tuberosa. It is represented in PL XIV. fig. 9, and is so very similar to the
corresponding stage of the Pentacrinoid of Antedon rosacea, except perhaps for being a
trifle more robust, that little need be said about it. The centro-dorsal is still small and
without any indication of its subsequent enlargement for the development of cirri, and
but very few arm -joints have appeared above the radial axillaries.
16. Antedon parvipinna, n. sp. (PL XV. fig. 9).
Specific formula — A.—.
Description of an Individual. — Centro-dorsal discoidal, bearing some fifteen marginal
cirri. These have about fifteen stout joints, of which the fifth and sixth are slightly
longer than wide.
Minute plates, probably the basal rays, rest upon the interradial angles of the centro-
dorsal and separate the bases of the nearly oblong second radials, the outer parts of
which are in close lateral contact. Axillaries short and wide with very open angles, and
also in close apposition. Both these joints have traces of a median keel which is
continued on to the arm-bases. The first brachials nearly oblong, with their outer sides
flattened ; the inner sides of the second and hypozygals of the third brachials also flattened.
Ten arms ; the joints after the eighth triangular, at first considerably shorter than
wide, but gradually becoming more nearly equilateral and finally quadrate. A syzygy in
128 THE VOYAGE OF H.M.S. CHALLENGER.
the third brachial, the next between the eighth and twelfth, and others at intervals of
five to nine joints.
The first pair of pinnules are relatively small and of tolerably equal size, consisting
of little over a dozen short joints. The second pair have longer joints, and those of the
sixth and following brachials have the third and the two or three next joints con-
siderably expanded so as to enclose the genital glands, which are protected by plates.
The fourth joint is larger than the rest, but not markedly so.
Disk and arms distinctly but not extensively plated ; the pinnule-ambulacra have
fairly denned side plates with intervening sacculi, which are also abundant in the plating
over the genital glands.
Colour in spirit,— white.
Disk about 4 mm.; spread probably 12 cm.
Locality.— Station 192, September 26, 1874; near the Ki Islands ; lat. 5° 49' 15"
S., long. 132° 14' 15" E.; 140 fathoms ; blue mud. One specimen.
Remarks. — This species comes very near to Antedon tuberosa (PI. XXIII. fig. 2), but
is quite free from the numerous blunt spines which are so characteristic of that species,
its calyx and arms being almost completely smooth. Furthermore, the first pinnules are
relatively shorter and much less flagellate than in Antedon tuberosa, consisting of a
smaller number of joints and having an altogether stiffer appearance.
17. Antedon fiexilis, n. sp. (PL XLIL).
Specific formula — A.(2).^.
Locality.— Station 192, September 26, 1874 ; near the Ki Islands; lat. 5° 49' 15" N.,
long. 132° 14' 15" E.; 140 fathoms; blue mud.
Remarks. — This species will be described in the bidistichate group ;T but its ten-armed
variety finds a place here. It resembles Antedon tuberosa in the uniform expansion of
the lower joints in the genital pinnules ; but it differs altogether from that species in the
appearance of the first radials externally and in the absence of spines on the arm-bases,
while the cirri are stouter and have a larger number of joints than in either that type or
Antedon parvipinna.
18. Antedon acideata, n. sp. (PI. XXIII. fig. 3).
Specific formula — A.—.
Description of an Individual.— Centro-dorsal subcorneal, bearing about fifteen cirri
in five irregular rows which are radial in position. The cirri have about eighteen joints,
1 See p. 217.
REPORT ON THE CRINOIDEA. 129
most of which are longer than wide and sharply compressed along, the dorsal edge, the
penultimate with an inconspicuous spine.
Three radials visible ; the first short and band-like, marked by occasional grooves and
projections. The second longer, with flattened lateral borders but sharply convex in the
centre, where they rise to meet the backward projections of the axillaries. These and
the first three brachials have a high centre and depressed margins like the second radials,
with sharp lateral edges and flattened sides. Ten arms ; the basal joints rather short,
with a sharp medio-dorsal line ; the following joints obliquely quadrate and more
distinctly carinate, so as to overlap. Syzygies in the third and about the fourteenth
brachials ; others at intervals of five to eight joints.
The first pinnule rather larger than its immediate successors ; their basal joints short
and laterally compressed, with a sharp dorsal edge. In the pinnules of the tenth and
following brachials the third joint and its successors are not expanded, but gradually
become longer than wide, and in the terminal pinnules are much elongated.
Disk invisible ; covering plates of the pinnule-ambulacra supported on a limestone
band which is not distinctly segmented. Sacculi variable, but not very common.
Colour in spirit, — light brownish-white.
Spread probably 15 cm.
Locality. — Station 214, February 10, 1875; off the Meangis Islands; lat. 4° 33' N.,
long, 127° 6' E.; 500 fathoms; blue mud; bottom temperature, 41°"8 F. One specimen.
Remarks. — This species has straight-edged and wall-sided arm-bases, just as in
Anteclon basicurva and Antedon incisa. But it differs altogether from these types in
the characters of the pinnules on the proximal third of the arm. So far as I have been
able to make out, without mutilating the specimen, the proximal pinnules have some-
what of the trihedral form with flattened outer sides which is characteristic of Antedon
basicurva (PI. XXII. fig. 4). Their next successors are altogether different, however,
the third and following joints gradually becoming relatively longer until they attain the
usual elongated shape which is characteristic of the middle and terminal pinnules. But
they acquire this shape at about the tenth or twelfth brachial, so that they differ from
the broad and expanded pinnule-joints in the corresponding part of the arm of Antedon
tuberosa (PL XXIII. figs. 2, 3), and the genital glands are unprotected by plates. The
side plates of the pinnule-ambulacra are not well differentiated, and the sacculi are
variable in their distribution, being moderately abundant in some pinnules and rare in
others.
The arrangement of the cirri is peculiar. There are none upon the interradial
portions of the centro-dorsal ; but beneath each ray there is a somewhat irregular vertical
row of two, three, or occasionally four sockets, all the rows converging on the apex of
the subconical centro-dorsal.
(ZOOL. CHALL. EXP. — PAET LX. 18S7.) OOO 17
130 THE VOYAGE OF H.M.S. CHALLENGER.
19. Antedon denticulata, n. sp. (PL XXII. figs. 1, 2).
Specific formula — A.y,
Description of an Individual. — Centro-dorsal hemispherical, with a denticulate rim.
Twenty to twenty-five rather slender cirri, of twenty-five to thirty smooth joints, nearly
all of which are longer than wide, the fifth and sixth longest.
First radials not visible ; second short and rather convex in the centre ; axillaries
short and widely pentagonal with slight backward projections. Both the radials and the
first two brachials are wall-sided, with straight edges and the margins of the dorsal
surface flattened.
Ten arms ; the lower joints nearly oblong and the following ones smooth, short, and
bluntly wedge-shaped, gradually becoming more oblong about the middle of the arm.
Syzygies in the third and twelfth or thirteenth brachials ; others at intervals of four to
six joints.
The pinnule on the second brachial is rather longer than that on the third, and the
length increases to the third pair (on sixth and seventh brachials). These consist of
about a dozen joints, the lowest of which are broad and slightly carinate. The next pair
are smaller with relatively longer joints and the following ones increase slowly in length.
Pinnule-ambulacra not plated ; sacculi apparently absent.
Colour in spirit, — very light brown.
Spread, perhaps 14 cm.
Locality.— Station 190, September 12, 1874; lat. 8° 56' N., long. 136° 5' E.;
49 fathoms ; green mud. One specimen.
Remarks. — This species and the following one are readily distinguished from all
those previously described with wall-sided arm-bases by the entire absence of any
ambulacral skeleton on the pinnules ; but they differ altogether from one another,
especially in the characters of the pinnules and of the arm-joints. Antedon denticulata
has quite short arm-joints, the lower ones obliquely quadrate and their successors more
nearly oblong, but always much wider than long (PI. XXII. fig. 1), and the third pinnule
is the largest. But in Antedon pusilla the arm -joints are as long or longer than wide
(PL XXIII. fig. 1), and the first pinnule is the largest.
There appear to be no sacculi in Antedon denticulata, or at any rate I have been unable
to find them. Their absence is remarkable in a form with unplated ambulacra, especially
as they are abundant in four individuals of Antedon fluctuans, which were dredged at
the same station.
REPORT ON THE CRINOIDEA. 131
20. Antedon pusilla, n. sp. (PI. XXIII. fig. 1).
Spec ifc formula — A . y .
Description of an Individual. — Centro-dorsal a low hemisphere bearing about fifteen
cirri with some twenty-eight joints, few of whieh are longer than wide, the distal ones
with a slioht dorsal keel.
First radials partially visible ; the second short and oblong with the centre of the
distal edge raised to meet the proximal edge of the axillary and form a tubercle. A
similar but smaller tubercle at the junction of the first two brachials. All four joints are
wall-sided and straight-edged, with the margins of the dorsal surface flattened.
Ten arms of smooth and elongated obliquely quadrate joints ; syzygies in the third
and then generally in the eighth or ninth brachials, with others at intervals of one to five,
usually three joints.
The first pinnule consists of about a dozen elongated joints and is considerably longer
and stouter than its successors, which decrease to about the fourth pair and then gradually
increase. The two lowest joints of the later pinnules are expanded and trapezoidal, but
the following joints are slender.
Pinnule-ambulacra not plated ; sacculi abundant.
Colour in spirit, — light brownish-white.
Spread about 7 cm.
Locality.— Station 192, September 26, 1874 ; near the Ki Islands ; lat. 5° 49' 15" S.,
long. 132° 14' 15" E.; 140 fathoms; blue mud. One specimen.
Remarks. — This is an elegant little form with long arm-joints and the first radials
visible externally, both of which characters are frequently indicative of immaturity. But
the great development of the genital glands, which are often found as far out on the arms as
the sixtieth brachial, seems to negative this idea in the present case. The two characters
just mentioned distinguish Antedon pusilla, from Antedon denticulata, from which it also
differs in the presence of tubercles on the rays and arm-bases, in the much shorter cirrus-
joints, and in the fact that the first and not the third pinnule is the largest. The distal
pinnules are very delicate and their two basal joints altogether different from the rest,
being expanded and trapezoidal, with their apposed edges much curved, as in many of the
Circumpolar species, while there are large and abundant sacculi on both arms and pinnules.
2. The Acmla-gvo\\\).
This group includes, at present, only two species, which differ from one another in
nearly all the characters of the cirri, arms, and pinnules, but are allied to the Basicurra-
group in the presence of a plated disk and of a well-defined ambulacra] skeleton,
characters which appear in no other ten-armed Comatulse.
132 THE VOYAGE OF H.M.S. CHALLENGER.
Pinnule-ambulacra well plated.
Less than twenty smooth cirrus-joints; genital pinnules with broadly expanded
joints and strong protecting plates, . ■ . . . .1. acaela, n. sp.
Forty or more spiny cirrus-joints ; no enlargement of the genital pinnules ; first joint
of lower pinnules much expanded, . . . . . .2. discoidea, n. sp.
1. Antedon accela, n. sp. (PI. II. figs. 3, a-d; PI. XVI.; also Part I., pi. liv.
figs. 1-4 ; pi. lv. fig. 5).
Specific formula — A. — .
Centro-dorsal subcorneal or hemispherical, bearing twenty-five to thirty cirri. These
have fifteen to eighteen smooth joints, nearly all of which are longer than wide.
Terminal claw sharp, with but little trace of an opposing spine.
First radials only visible in young specimens ; the second somewhat flattened, with a
convex proximal and concave distal border. Axillaries more convex, broadly pentagonal
or almost rhombic, with a wide distal angle, and sometimes projecting deeply backwards
into the second radials. The dorsal surfaces of both radials, with the two lowest brachials
and the hypozygals of the third, project beyond their faces and sides, especially the
latter, which fall away rapidly from the dorsal towards the ventral border.
Ten arms. First brachials nearly oblong with rounded outer edges ; the second convex
and irregularly pentagonal. About one hundred smooth triangular joints, as long as or
longer than wide. Syzygies in the third and then about the eleventh or twelfth brachials,
with others at intervals of two to five, usually three or four joints.
The first pair of pinnules have about thirty short joints, the lowest of which is a good
deal wider than the rest. From the fourth to the twenty -fifth brachials about three to
five of the pinnule-joints are greatly expanded laterally to enclose the genital glands, and
the first joint is much wider than its successors, especially in the lower pinnules. The
later pinnules have longer and more trihedral joints, the lowest of which are flatter.
Disk rather incised and completely covered with irregular plates bearing short
and blunt rod-like spines. Brachial ambulacra and interarticular spaces well plated, and
the expanded parts of the genital pinnules are completely enclosed in an arched
] lavement of flat plates, very regularly arranged and devoid of ambulacra. The ambulacra
of the later pinnules have very well-defined side plates, sometimes covering the sacculi
and sometimes notched for them. These are very abundant, except on the non-tentacu-
liferous genital pinnules.
Colour in spirit, — young individuals straw-coloured ; the older ones a dense brownish-
grey, becoming a dark grey in the most mature.
Disk 5 mm.; spread probably 20. mm.
Locality.— Station 214, February 10, 1875 ; off the Meangis Islands; kit, 4° 33' N.,
long. 127° 6' E.; 500fathoms; blue mud; bottom temperature, 41°-8 F. Several specimens.
REPORT ON THE CRINOIDEA. 133
Remarks. — This is a peculiar species for many reasons. In its general characters
it has many points of resemblance with Antedon basicurva, Antedon incisa, and
Antedon tuberosa ; but the sides of the basals and lower radials are not in close apposi-
tion and flattened laterally against each other as in those types, for they fall away
very rapidly from the dorsal towards the ventral surface, so that there is a considerable
space between every two rays, especially at the level of the articulations (PL XVI.
fig. 1).
This is smaller on the surface than it is deeper down, for the lateral edges of the
joints are produced outwards like the projecting eaves of a roof. This condition is
extremely marked in the case of the second radials, which have the proximal edge
similarly produced so as to overlap the minute portion of the first radials which appears
externally. The relative shapes of the two outer radials vary extremely. Their usual
appearance is represented in PL XVI. fig. 1 ; but in some individuals the second radials
are rather more oblong and show hardly any indication of an incised distal edge, while
the axillaries are widely pentagonal. On the other hand the axillaries sometimes project
far backwards into the second radials, which thus have a deeply incised distal edge, while
the proximal edge is also much curved.
In young specimens, such as that shown in PL XVI. fig. 2, a considerable portion
of the first radials is visible, but the projection of the edges of the next following joints
is almost as marked as it is in the mature individual. In the youngest specimen, with a
spread of about 80 mm., the external surface of the first radials is rather less wide than
that of the second, and a trifle more than half its length. It does not, however, increase
in size along with the corresponding parts of the two outer radials, but remains
undeveloped and is sometimes marked by small tubercular elevations like those on the
centro-dorsal, from which it is with difficulty distinguishable. These are situated in the
gap between the ventral edge of the centro-dorsal and the proximal edges of the second
radials, which project backwards so as to overlap and conceal them.
The most striking character of Antedon accela, and the one which allies it most
closely to the Basicurva-grou]), is the great size, both of the pinnule-joints and of the
protecting plates on the genital pinnules. Even the second pair of pinnules are enlarged
for the reception of the genital glands, three of their middle joints being expanded ; and
a little further from the disk the fifth and the four or five following joints are flattened
and produced laterally, as shown in PL XVI. fig. 2, the proximal joint being often
much enlarged at the same time. This expansion is not almost entirely limited to the
outer side only as in Antedon incisa (PL XXI. fig. 2), but it is equal on both sides of the
medio-dorsal line ; and the ventral portion of these expanded joints is covered by an
arched pavement of strong plates, few in number but of large size, and often very
regularly arranged, as seen in figs. 1-3 of pi. liv. in Part I. These protecting plates
are much larger and better developed than in either Antedon incisa or Antedon tuberosa.
134 THE VOYAGE OF H.M.S. CHALLENGER.
and they often alternate more or less regularly on opposite sides of the medio-ventral
line of the pinnule, where there is an opening in one of them for the exit of the genital
products.
In the young individuals obtained, even in those with a spread of 12 cm., there
is no trace of the enlargement either of the pinnule-joints or of the protecting
plates (PI. XVI. fig. 5), although both are visible in the older forms, which still show a
considerable part of the first radials externally. In the regenerated arms too the lower
pinnules are for some time quite small and inconspicuous, and altogether different from
those of the uninjured mature individuals. This is the case even when the arm
has attained almost its full size, and is absolutely larger than those of other individuals
not yet quite mature, but with comparatively large genital glands.
All these greatly enlarged genital pinnules are devoid of ambulacra, like the non-
tentaculiferous posterior arms of Actinometra ; but at about the position of the twenty-
fifth brachial there is a sudden diminution in size both of the pinnule-joints and of
the protecting plates, more especially of the latter. They become much smaller
and relatively more numerous, wThile the sacculi which are absent in the large lower
pinnules begin to appear, just as they show themselves in the genital pinnules of
Antedon angusticalyx from the same station ;* while eventually the ambulacral skeleton
shows itself above the small protecting plates, as in Antedon incerta.2 A little further
out on the arms these protecting plates disappear, and the ambulacral skeleton comes to
rest directly upon the pinnule-joints, as shown in PL XVI. fig. 4. The side plates are
very well differentiated and are often notched for the recejttion of the sacculi or of
portions of them ; but in other cases, when the sacculi are large, they are altogether
covered and concealed by the side plates.
2. Antedon discoidea, n. sp. (PI. X. figs. 1, 2).
Specific formida — A. — .
Centro-dorsal a thick disk, bearing fifteen to eighteen cirri in a single or partially
double row, with the dorsal surface free. The cirri reach 27 mm. in length and consist
of forty or fifty joints, a few of which at the base are longer than wide, and the following
ones gradually develop a sharp dorsal keel.
Three radials visible ; the first short, except at the angles of the calyx, where the
ends of the basal rays sometimes appear. Second radials short, wide and oblong, and
the axillaries barely pentagonal.3 Both joints have large muscle-plates, and their dorsal
surfaces rise towards the middle of their apposed edges. Pays well separated.
1 See Part I., pi. liv. fig. 5. 2 See Part. I., pi. liv. fig. 6,
3 The above description applies to the joints as seen in Ml face view. They have a very different shape in the
figure of the entire animal, owing to the angle at which the rays are set on the calyx (PI. X. fig. 1).
REPORT ON THE CRINOIDEA. 135
Ten arms. First brachial almost oblong ; the second bluntly triangular, with a large
lateral process bearing the pinnule-facet. The next few segments each have a process of
the same kind, but gradually decreasing in size. Arm-joints after the tenth, triangular,
as long as wide, and slightly overlapping, but more quadrate towards the end. Syzygia
in the third and beween the tenth and fourteenth brachials ; others at intervals of two
to five, usually three, joints.
The first two pairs of pinnules have twenty or more short joints, the first of which is
much expanded dorsally and the next two slightly so. This expansion gradually dies
away in the following pinnules, which increase in size, becoming stiff and rod-like,
and composed of long cylindrical joints, after the first two, which are laterally compressed.
Disk and brachial ambulacra much plated. Covering plates of the pinnule-
ambulacra supported on a well-developed limestone band, which is not clearly divided
into side plates ; the sacculi concealed by it are very large and closely set.
Colour in spirit, — yellow or yellowish-white, with occasional brown or purplish bauds.
Disk 6 mm. ; spread about 1 6 cm.
Locality. — Station 192, September 26, 1874; near the Ki Islands; lat. 5° 49' 15"
S., long. 132° 14' 15" E. ; 140 fathoms; blue mud. Four specimens.
Remarks. — This is a singular species in many ways and is readily distinguished by
the characters of the lower arm-joints and of the pinnules which they bear. The broader
end of each joint projects considerably from the general lateral line of the arm, so as to
form a large pinnule-facet ; and the dorsal part of the first pinnule-joint is expanded into
a large curved plate which covers in this facet. This plate, which i3 well shown in PI. X.
fig. 2a, is sometimes so large that the whole arrangement looks as if it were a malformation
due to the action of an encysting Myzostoma which had taken up its abode in the pinnule-
socket. It is largest in the first two pairs of pinnules, the remaining joints of which are
relatively quite short, especially in the first pair (PL X. fig. 2a), but by the fourth pair
(PI. X. fig. 2b) the two basal joints are less expanded, though the third is slightly so,
while the following joints are much longer and somewhat carinate. In the middle and
distal pinnules this tendency disappears and the two lowTer joints have the usual somewhat
flattened appearance.
The lateral projection of the arm-joints to form large pinnule-sockets is a point of
some interest because it occurs in some forms of the Jurassic Antedon costata, as for
example that figured by Walther on taf. xxv. fig. 6 of his memoir. He describes the
arm-joints as follows " Das dickere Ende tragt einen dorsal en Knoten und einen seitlichen
Fortsatz zur Insertion der Pinnula." 1
The sacculi are fairly developed on the arms of Antedon discoidea, and in the pinnules
they are large and extraordinarily abundant. They are covered, however, by the
1 Loc. cit., p. 172.
136 THE VOYAGE OF H.M.S. CHALLENGER.
substantial limestone band which supports the covering plates but is not properly
segmented, into side plates. The ambulacra! skeleton does not extend to the end of the
pinnules, several joints of which are entirely without it, as in Antedon longicirra from the
same station, and in many Pentacrinida?.
3. The Eschrichti- gYouj).
This group is very well defined both in its zoological characters and in its
geographical and thermal distribution. Of the seven species which it at present contains,
three are Arctic and the remaining four Antarctic. Antedon eschrichti and Antedon
quadrata are widely distributed through the Arctic Ocean, reaching a latitude of 79° N,
and 81° N. respectively in Smith's Sound. They extend across the North Atlantic into
the Barents Sea, and Antedon eschrichti, at any rate, was dredged by Nordenskjold in the
" Vega " as far east as long. 92° 20' E.. In the East Atlantic both species are found as far
south as the parallel of 60° in the cold area of the Fseroe Channel ; but the Challenger
dredged them both off Halifax in lat. 43° 2' N. This is their furthest southern limit, and
the third Arctic species belonging to this group [Antedon barentsi) has as yet only been
found near Vardo in the Barents Sea.
Two of the four Antarctic species occur in the Straits of Magellan, while the other
two were obtained by the Challenger at Stations 150 and 151, between Kerguelen and
Heard Island. The only species found at depths exceeding 200 fathoms are Antedon
eschrichti and Antedon quadrata, both of which extend downwards from the littoral
fauna, the former descending to 632 and the latter to 466 fathoms. The temperature
limits of this group are very well defined. There is no record of any of them having
been found in water of a higher temperature than 36° F. Antedon quadrata was
obtained at 29° and Antedon eschrichti at 30° in the cold area of the Fseroe Channel ;
while Antedon australis was dredged from a temperature of 35c,2 at Station 150. I
do not imagine that this is likely to have been greatly exceeded at any of the shallow
water localities, either in the Arctic or in the Antarctic Seas, where species of this group
have been obtained. They are not, however, the only Comatulse which occur in the colder
seas of the globe. Antedon prolixa was obtained in 25 fathoms at Discovery Bay
(lat. 81° 41' N.) together with Antedon quadrata, and has since been dredged in
abundance near Spitzbergen by the Norwegian North-Atlantic Expedition, from a depth
of 743 fathoms ; while the " Varna " found it at 50 fathoms in the Kara Sea.
The " Porcupine " and " Triton " met with' Antedon hystrix in the cold area of the Fseroe
Channel where Antedon eschrichti and Antedon quadrata also occur ; while Antedon
tenella, which reaches a latitude of 79° near Franz Joseph Land, is common in the Barents
Sea and on both sides of the Atlantic down to 740 fathoms, with a thermal range of
from 30° to 50° F.
REPORT ON THE CRINOIDEA. 137
Apart from the absence of any lateral flattening of the rays and arm-bases and of an
ambulacral skeleton, the special character distinguishing the Eschrichti-gcoxnp is the
flagellate appearance of the proximal pinnules. The two first pairs (on second to fifth
brachials), and sometimes also the third pinnule on the outer side of the arm (on sixth
brachial), consist of a large number of short and wide joints, the later ones of which are
often somewhat serrate (PL XXIV. figs. 1, 2, 7, 8 ; PL XXV. figs. 1, 2 ; PL XXVII.
figs. 8, 9, II, 12, 14, 15). The cirri are always numerous and composed of thirty to
fifty joints ; while the long arms bear numerous closely set pinnules, so as to give a very
feathery appearance to the general plume; The regular arrangement of the syzygies too
is very striking as compared with the Basicurva-grou]), the members of which exhibit
hardly any regularity in the grouping of the syzygies, except for the presence of one in
the third brachial as in most Comatula3 with articulated radials. In the JEschrichti-groujp,
however, as also in Antedon jyhalangium, Antedon rosacea, and allied species, the
syzygies are situated with great uniformity in the third, eighth, and twelfth brachials,
and afterwards at intervals of two or three joints. The position of the third syzygy is
less constant than that of the second, but does not vary to any great extent (PL XXIV.
fig. 11 ; PL XXV. fig. 12 ; Pis. XXVL, XXVIII.).
It will be seen from the following table that the differences between the individual
species mostly turn upon the characters of the third pinnule, the relative shape of the
arm-joints, and the number of the cirrus-joints. Antedon rhomboidea and Antedon
barentsi are species based upon single individuals ; but I have seen seven examples of
Antedon australis, and a considerable number of each of the other four species, those of
Antedon esehrichti and of Antedon quadrata being from several different localities.
Neither Antedon barentsi nor Antedon magellanica were obtained by the Challenger
at all, the former living in the Barents Sea,1 wlule Antedon magellanica was obtained by
H.M.S. " Alert," and was described as a variety of Antedon esehrichti by Bell.2 I have
pointed out elsewhere,3 however, that it is altogether separated from Antedon esehrichti
by the characters of its arm-joints, a point to which Bell did not refer, and I have since
examined several individuals of it which were dredged by the Italian corvette " Vettor
Pisani," and have no doubt whatever as to its being a good species. Although not
Challenger species, these two are included in the following list for the sake of
completeness.
!The Comatulje of the "Willem Barents" Expeditions, 1880 and 1881, Bijdragm tot de THerkunde, 1886, 13
Aflevering, vi. pp. 1-12.
2 Note on a Crinoid from the Straits of Magellan, Proc. Zool. Soc. Lond., 1882, p. 651.
3 Bijdragen tot de Dierkunde, 1886, 13 Aflevering, vi. p. 4.
(ZOOL. CHALL. EXP. — PART LX. — 1887.) OoO 18
138 THE VOYAGE OF H.M.S. CHALLENGER.
The first two or three pairs of pinnules long and flagellate, with numerous short
and wide joints.
A. Third pinnule equal to, or not much shorter than the second.
I. Joints of the third pinnule mostly wider than long, as in the first and second.
Arm-joints triangular, but quite short.
a. Forty or more cirrus-joints. Axillaries as wide as long. Arms
smooth. Third pinnule most like the second, . . . 1. eschrichti, Mull., sp.
b. Less than forty cirrus-joints. Axillaries wider than long. Arm-
joints overlap. Third pinnule most like the fourth, . . 2. antarctica, n. sp.
II. Third pinnule has fewer hut much longer joints than the first and second.
a. Less than forty cirrus-joints. Arm-joints triangular.
1. Arm-joints short and much wider than long, . . .3. australis, n. sp.
2. Arm-joints as long as or longer than wide, . . .4. rhomioidea, n. sp.
b, Over forty cirrus-joints. Arm-joints quadrate, and as long as or longer
than wide, ....... mayellanica, Dell.
B. Third pinnule composed of a few elongated joints, and much shorter than the
second.
t I. About forty cirrus-joints. The middle arm-joints quadrate, . . 5. quadrata, n. sp.
II. Twenty-five to thirty cirrus-joints. The middle arm-joints triangular ;
genital pinnules plated, ...... barentd, Carpenter.
1. Antedon eschrichti, Mull., sp. (PL I. figs. 8, a-d; PI. XXIV. figs. 4-14, woodcut,
figs. 4, c, D, on p. 154).
Specific formula — A. — .
1841. Alecto Eschrichtii, Miiller, Monatsber. d. k. preuss. Akad. d. Wiss. Berlin, 1841, p. 183.
1849. Comatula (Alecto) Eschrichtii, Miiller, Abhandl. d. k. Akad. d. Wiss. Berlin, 1847 [1849],
p. 254.
1854. Alecto Eschrichtii, Stimpson (?), Mar. Invert. Grand Manan, 1S53 [1854], p. 12.
1857. Alecto Eschrichtii, Liitken, Vidensk. Meddel. nat. Foren. Kj0benhavn, 1857, p. 55.
1860. Alecto glacialis, Walker, Journ. Dublin Sob., 1860 [1862], vol. iii. p. 70.
1862. Comatula Eschrichtii, Dujardin and Hupe, Hist. Nat. des Zoophytes, Echinodermes Paris,
1862, p. 199.
1866. Antedon Eschrichti, Loven, Ofversigt k. Vetensk.-Akad. Forhandl., 1866, No. 9, pp. 224,
230, figs. i-m.
1866. Antedon Eschrichtii, Verrill, Proc. Boston Soc. Nat. Hist., 1866, vol. x. p. 343.
1872. Antedon escrichtii, Wyville Thomson, Proc. Roy. Soc. Edin., 1872, vol. vii. p. 764.
1876. Comatula Eschrichtii, Quenstedt, Petrefactenkunde Deutschlands, 1876, Bd. iv. Asteriden
und Encriniden, p. 165, Tab. 96, fig. 16.
1879. Antedon Eschrichtii, P. H. Carpenter, Trans. Linn. Soc. Lond. (Zool.), ser. 2, 1879,
vol. ii. p. 29.
1880. Antedon Eschrichtii, d'Urban, Ann. and Mag. Nat. Hist., 1880, ser. 5, vol. vi. p. 261.
1881. Antedon Eschrichtii, Duncan and Sladen, Memoir Arctic Ecliinodermata, London, 1881,
p. 73, pi. vi. figs. 1-4.
1882. Antedon Eschrichtii, Hoffmann, Niederland. Arch. Zool., 1S82, Supplement Bd. i., Lief
3, p. 1.
1882. Antedon eschrichti, Bell, Proc. Zool. Soc. Lond., 1882, p. 534.
1882. Antedon eschrichti, P. H. Carpenter, Ibid., p. 746.
REPORT ON THE CRINOIDEA. 139
1884. Antedon eschrichti, P. H. Carpenter, Proc. Roy. Soc. Edin., 1884, vol. xii. pp. 364, 374.
1886. Antedon Eschrichti, P. H. Carpenter, Bijdragen tot de Dierkunde, 1886, 13 Aflevering,
vi. p. 5, pi. i. figs. 7-10.
1886. Antedon Eschrichtii, Levinsen, Dijmphna-Togtets zoologisk-botaniske Udbytte, Kj0ben-
bavn, 1886, p. 410, Tab. xxxv. figs. 7, 8.
1886. Antedon Eschrichtii, Stuxberg, Vega-Expeditionens Vetenskapliga Arbeten, Stockholm,
1886, Ed. v. p. 162.
18S6. Antedon Eschrichtii, Fischer, Die Osterreichische Polarstation Jan Mayen, Ed. iii., Wien,
1886, Echinodermen, p. 3.
Centro-dorsal hemispherical, bearing a very large number of cirri, reaching a hundred
in old specimens. The dorsal pole, which is somewhat flattened, is free, but elsewhere
they are very closely set and may reach over 70 mm. in length, consisting of forty to
sixty joints, but few of which are longer than wide. The later joints project slightly,
but do not form definite spines.
First radials almost entirely invisible in the adult ; second quite short, oblong or
crescentic, according to the amount of incision by the axillaries, and almost free laterally,
with large muscle plates. Axillaries more than twice their length, triangular or rhombic,
with incurved sides. They are about as long as wide, and have a sharp distal angle.
Ten arms, with over three hundred joints in a large specimen. First brachial deeply
incised, with a short inner and much longer outer edge. The second irregularly quadrate,
and the succeeding joints to the eighth nearly triangular, with the pinnules on their
shorter sides and their apposed edges rising to tubercular prominences alternately on the
outer and inner sides of the arm. The following joints smooth and triangular, much wider
than long, becoming quadrate towards the end of the arms. Syzygies in the third,
eighth, and twelfth brachials, with others at intervals of two or three joints.
The lower pinnules long and flagellate, composed of numerous short joints, rather wide
at the base. The second (on the fourth brachial) is usually the longest, reaching nearly
40 millimetres in length and consisting of about seventy joints. The third pinnule is
of variable length, but its lower joints are larger than those of the second, though
distinctly wider than long. In the first three or four pairs of pinnules the dorsal edges
of the lower joints are sharp and cut away at the ends, so as not to meet their fellows,
and in the small terminal joints this sharpened edge is produced into a bluntly angular
process, making the end of the pinnule somewhat serrate. The following pinnules are
shorter and more massive, with large lower joints, which are nearly square in outline and
gradually become longer than wide. The middle pinnules reach 30 millimetres with
about forty joints, the two lowest of which are flattened and somewhat trapezoidal, with
their apposed edges incurved. Genital glands long and fusiform. Disk and arms not
plated ; sacculi extremely abundant.
Disk 25 mm.; spread 50 cm. (maximum).
Colour in spirit, — light reddish-brown.
Localities.— H.M.S. " Porcupine," 1869, Station 57 ; lat. 60° 14' N., long. 6° 17' W.;
HO THE VOYAGE OF H.M.S. CHALLENGER.
632 fathoms ; bottom temperature, 30°"5 F. Three (or more) specimens. Also at other
unrecorded localities in the " cold area."
H.M.S. Challenger, Station 48, May 8, 1873 ; on the Le Have Bank; lat. 43° 4' N.,
long. 64° 5' W.; 51 fathoms; rock. Several specimens.
H.M.S. "Valorous," Station 1, July 22, 1875; off Hare Island, in Davis Strait;
lat. 70° 30' N., long. 54° 41' W.; 85 fathoms; sand and mud. One specimen.
H.M.S. "Alert," 1875 ; Franklin-Pierce Bay in Smith's Sound ; lat. 79° 25' N.
Other Localities. — Melville Bay ; Jan Mayen ; Spitsbergen Sea : Barents Sea ; Kara
Sea ; Coast of Siberia to long. 92° 20' E. (Stuxberg) ; Bay of Fundy 1 (Stimpson).1
Remarks. — Although described by Miiller in 1841, this species was never figured till
187G, when Quenstedt gave a rough, but very characteristic sketch of it in the Atlas of
the Petrefactenkunde Deutschlands,2 Five years later it was again figured and minutely
described by Duncan and Sladen in their well known monograph of Arctic Echinoderms.
The numerous examples of it which were dredged by the Challenger off Halifax
(Station 48) are by no means so large and well developed as individuals which I have
examined from higher latitudes, and notably those obtained in the Barents Sea by the
Dutch Arctic Expeditions, which are the finest examples of the type that I have seen.
The spread of these Atlantic specimens does not exceed about 40 cm., and there are not
more than two hundred arm-joints. The cirri and the lower pinnules are also fewer-jointed
and shorter in proportion, while the arm-bases are much less tubercular than in the more
northern forms. In these last the junction of the first two brachials forms a somewhat
prominent knob in the middle line of the arm, and there is another at the outer end of
the hue of articulation between the second and third. The next is at the inner end of
the articulation between the third and fourth, the one joint projecting forwards and the
other backwards to form a knob-like elevation. This usually disappears at the second
syzygy (on the eighth brachial), but may be continued out for three or four joints further,
and the result of it is that the fourth to the seventh joints are altogether different from
their successors in bearing their pinnules on their shorter sides (PI. XXIV. figs. 10, 11).
Beyond the third syzygy the joints are very distinctly triangular, but they are consider-
ably wider than long, and this disproportion increases in the middle and outer parts of the
arms, so that the successive pinnules are very closely set (PI. XXIV. fig. 13) ; and it is
only cpiite at the extremities that the joints become at all quadrate (PI. XXIV. fig. 12).
This is one of the best characters for distinguishing Antedon eschrichti from Antedon
quadrata (PL XXVI. figs. 1-3 ; PI. XXVII. figs. 5-7 ; fig. 4 on p. 154), which is com-
monly found associated with it, though it is shared with Antedon antarctica, as seen in
PL XXV. fig. 12.
1 Stimpson had some hesitation in referring his single specimen to Antedon eschrichti, on account of its small size,
and it may not improbably belong to Antedon quadrata.
2 Encriniden, tab. 96, fig. 26.
REPORT ON THE CRINOIDEA. 141
The middle and outer pinnules of Antedon eschrichti exhibit a modification of the
first two joints of essentially the same character as that which has already been noticed
in Antedon valida (PI. XV. fig. 2). The first joint is irregularly trapezoidal, or in some
cases almost crescentic, its distal edge being more or less concave, while the proximal edge
of the larger and more trapezoidal second joint is similarly incurved and only meets that
of the first near its ventral end, so as to leave a large gap on the dorsal side, which is
occupied by ligament (PL XXIV. fig. 13). This feature is very characteristic of nearly
all the Comatulas from Arctic and temperate seas, and also of some abyssal forms
(PI. XXVII. figs. 26, 27 ; PI. XXVIII. fig. 4 ; PL XXXII. figs. 5, 7), while it likewise
presents itself in certain tropical species ; but it never appears in Actinometra.
The axillaries of Antedon eschrichti vary considerably in their shape from triangular
to rhombic, according to the extent of their backward projection into the second radials.
In a few instances I have found them to be longer than wide ; but in most cases
the width is equal to or a trifle greater than the length, more than half of which is on
•the distal side of the line joining the lateral angles. This is chiefly clue to the acuteness
of the distal angle (PL XXIV. figs. 10, 11). The axillaries of Antedon antarctica have
much the same shape, but they are usually considerably wider than long (PL XXV.
figs. 8-12).
There is much variation both in the relative and in the absolute size of the flagellate
lower pinnules of Antedon eschrichti. Those figured on PL XXIV. figs. 7-9, are
the three first pinnules on the outer side of the arm of a specimen from Station 48, i.e.,
those borne by the second, fourth, and sixth brachials. The same three pinnules
of Antedon rhomboidea, Antedon antarctica, Antedon australis, and Antedon quadrata
are figured on Pis. XXIV, XXV. and XXVIL, and in all but the last (PL XXVII.
figs. 8-13) the third pinnule is but little smaller than its predecessors. In Antedon
eschrichti it has fewer joints than the first and second pinnules, but the basal ones are
somewhat larger, though still wider than long, and a few of the outer joints become
longer than wide, which is not the case in the first two pinnules (PL XXIV. figs. 7-9).
The third and fourth pinnules are in fact the transitional stages between the flagellate
basal pinnules and the larger genital ones which follow them. In Antedon antarctica,
however, the change is much more sudden (PI. XXV. figs. 1-3).
Fig. 10 on PL XXIV. represents a small but very interesting example of Antedon
eschrichti which was dredged by the " Triton " in the Fasroe Channel. The cirri are small
and comparatively delicate, not exceeding 20 mm. in length, and the arm-bases are but
slightly tubercular. All the arms have been broken and regenerated either at the second
(eighth brachial) or third syzygy (twelfth or thirteenth brachial). In one arm there are
two distinct changes of diameter, showing that the first regenerated part had undergone a
subsecpient fracture which has been again made good. One can therefore study the
appearances presented by the new arm-joints in various stages of growth. The lowest
142 THE VOYAGE OF H.M.S. CHALLENGER.
and therefore oldest of these new joints are most like those of the corresponding part of
the arm in the adult, i.e., triangular or very slightly quadrate, but relatively wide
in proportion to their length. These characters, however, do not disappear as they do in
the adult, where the joints become gradually shorter and shorter, with a markedly
triangular outline. But throughout the remainder of the restored arm the joints are
quadrate and comparatively long ; while the two lowest pinnule-joints show but few traces
of the flattening and peculiarities of outline which are so characteristic of the adult. It
is just in these characters, besides the smaller size of the first pair of pinnules, that
Antedon quadrata differs from Antedon eschrichti, and it is therefore to be regarded as
a permanently immature form of the latter species. Levinsen, indeed, considers the
two species as identical, a point which I shall discuss when treating of Antedon quadrata.
Two Pentacrinoids, besides that of Antedon tenella, were dredged by the "Porcupine"
in the cold area of the Faeroe Channel ; but I doubt whether either of them can be the
one referred to by Sir Wyville Thomson1 in the following passage : — "A single example of
a pentacrinoid in an early stage was found associated with Antedon escrichti. It
resembles closeby the larva of Antedon sarsii, but the specimen was not sufficiently
preserved for a critical examination."
The larva mentioned in the above passage is possibly that which I have represented
on PL XIV. fig. 2. Its developmental condition is intermediate between the second and
third stages of the larva of Antedon tenella, though I do not think that it can be referred
to that type, on account of its greater robustness, and for other reasons. But, on
the other hand, I do not imagine it to be the larva of Antedon eschrichti, to which
species I referred it conjecturally in 1884,2 together with the larva represented on fig. 3
of the same plate.
The Danish exploring vessel "Dijmphna" dredged forty-five individuals of Antedon
eschrichti in the Kara Sea. They were of all sizes from a length of 25 mm. upwards.
One Pentacrinoid was also obtained and has been figured by Levinsen.3 It is considerably
younger than the smaller of two Pentacrinoids which were obtained by the Dutch ship
" Varna," in the Kara Sea, during the summer of 1883, and were sent to me for examina-
tion.4 I had been unable to definitely refer them to any specific type, but they are so
closely similar to that figured by Levinsen that I have now no doubt of their belonging
to Antedon eschrichti. It is equally clear to me, however, that the larva which is
represented on PL XIV. fig. 2 cannot belong to Antedon eschrichti, as seemed possible
in 1884 ; while it is too large for the corresponding stage of Antedon tenella, so far as I
can judge from the figures and dimensions of the latter which are given by Sars.
1 Proe. Roy. Soe. Edin., 1872, vol. vii. p. 764. 2 Proc. Roy. Soc. Edin., 1884, vol. sii. p. 364.
3 Kara-Havets Echinoderrnata, Dijmphna-Togtets zoologisk-botaniske Udbytte, Kjtfbenhavn, 1886, p. 34 (414) Tab.
xxxv. fig. 8.
4 The proofs of this Report were corrected early in 1885, but for some reasons with which I am not acquainted, it
has not yet been published.
REPORT ON THE CRINOIDEA. 143
It may perhaps be only a younger stage of the larva shown on PL XIV. fig. 3, which
I formerly referred to Antedon eschrichti on account of its extremely robust character.
I am now satisfied, however, from Levinsen's observations, that the latter supposition is
incorrect. The stem and the bases of the arms are nearly as well developed as in his
larva of Antedon eschrichti, and the cirri of the first whorl have rather fewer joints ; but
the basals are relatively much higher than in the Eschrichti-l&zva, the axillaries
of which are of an altogether different shape from those of the larva dredged by the
" Porcupine." The latter is not likely to belong to Antedon quadrata, which must have
a larva very like that of Antedon eschrichti, if indeed the two species are not identical ;
and I conclude therefore that the "Porcupine" larva should be referred to Antedon
hystrix, the only other Comatula found in the cold area of the Faeroe Channel.
The cirri of Antedon eschrichti resemble those of Antedon rosacea and Antedon
phalangium in the dimorphic characters of their younger stages. Fig. 6 on PL XXIV.
represents an immature cirrus of the normal developmental type. The lower joints are
relatively longer than in the full-grown cirrus shown in fig. 4, and its outer part consists
of a large number of short and wide joints with a strong terminal claw. On the other
hand, fig. 5 represents a " small mature " cirrus which is shorter and composed of fewer
joints than the immature one just mentioned ; but the small terminal joints, instead of
being short, wide and smooth, are much more like those of the adult cirrus and have
slight dorsal projections, though there is only a very small terminal claw. The cirri are
very numerous and the centro- dorsal proportionately large,' so that it hides the first
radials completely, only very small portions of them appearing on the exterior of the
isolated calyx (PL I. fig. 8a). The rosette of Antedon eschrichti is near the dorsal
surface of the radial pentagon and very well defined, with ten distinct spout-like
processes, of which the interradial ones are a trifle the smallest (PL I. fig. 8c), but there
is no indication of a basal star around it, the dorsal interradial furrows being simple and
not provided with lateral folds, so that the interradial markings on the upper surface
of the deeply hollowed centro-dorsal simply indicate the boundaries of the radial
fossae.
The articular faces of the radials of Antedon eschrichti are very characteristic. In a
full-grown calyx the muscle plates stand up nearly vertically, but the lower parts of the
faces are less steeply inclined (PL I. fig. 8«), so that the lower fossae are pretty completely
visible in a top view, while the muscular fossae are mostly concealed (PL I. fig. 8b).
They are separated from the pair of fossae below them by slanting ridges which run
upwards and outwards from the thickened lower end of the intermuscular ridge
immediately above the opening of the central canal. The lower pair of fossae above the
articular ridge are thus but little smaller than the upper pair which lodge the great
ventral muscles (PL I. fig. 8a). In smaller individuals, however, they are more unequal,
the upper fossa? being considerably larger than the lower ones.
144 THE VOYAGE OF H.M.S. CHALLENGER.
At the most of the localities where Anteclon eschrichti is known to occur it is
infested by the parasitic Myzostoma gigas, Liitken. Station 48 also yielded Myzostoma
jimbriatum, von Graff.
■_'. Anteclon antarctica, n. sp. (PI. I. figs. 6, a-cl, 7, a, b; PI. XXV.).
Specific formula — A. y .
Centro-dorsal hemispherical, thickly covered with cirrus-sockets. Eighty or more
cirri, reaching 35 mm. in length, and consisting of twenty-five to thirty-five joints,
several of which are longer than wide. The later joints project slightly beyond the
bases of their successors, and the penultimate has a well-developed terminal claw.
First radials invisible, except at the angles of the calyx, where they are sometimes
separated by the ends of the basal rays ; the second quite short and band-like, very
convex in the centre and deeply incised. Axillaries usually rather wider than long,
subtriangular, with a backward process of variable size in the middle of the base, some-
times so large as to give the plate an unequally rhombic appearance ; first brachial much
incised, with a short inner and lono; outer eds;e.
Ten arms, but slightly tubercular at the base, the joints after the third syzygy being
quite short, triangular and slightly overlapping. They become slowly quadrate towards
the ends of the arms, but always remain wider than long. Syzygies in the third, eighth,
and twelfth brachials, and then usually at intervals of three joints.
Lower pinnules long and flagellate, with a serrate rlorsal edge, reaching 25 mm., and
composed of about sixty short joints, the basal ones rather wide.
The first two pinnules are nearly equal ; but the third, though of about the same
length, consists of fewer and larger joints, some of the lower ones being as long or
longer than wide. The following pinnules shorter and more massive,with large lower
joints, which are nearly square in outline and overlap considerably. The middle and
outer pinnules of more elongated but still overlapping joints, the two lowest broader and
more flattened, with their apposed edges incurved.
Disk and ambulacra naked ; sacculi abundant.
Colour in spirit, — light brown.
Disk 17 mm.; spread about 25 cm.
Locality.— Station 151, February 7, 1874; near Heard Island; lat. 52° 59' 30" S.,
long. 73° 33' 30" E.; 75 fathoms ; volcanic mud. Several specimens.
Remarks. — This is no doubt the species to which Sir Wyville Thomson referred
when he stated that Anteclon eschrichti had been obtained in the Southern Ocean.
The two types are unquestionably very closely similar in their general appearance ; ,
but at the same time they differ considerably in points of detail. The cirri of Anteclon
REPORT ON THE CRINOIDEA. 145
antarctica are much smaller than those of Antedon eschrichti, even in individuals of
equal size, not having more than thirty-five joints, a considerable proportion of which
are longer than wide, while the later joints project considerably more on the dorsal side
than is the case in Antedon eschrichti (PI. XXIV. fig. 4 ; PI. XXY. fig. 6). This is
especially marked in the younger cirri which are of the " small mature " type (PI. XXV.
fig. 7), while those which develop in the usual way, though both relatively and
absolutely larger, are much more smooth-jointed (PL XXV. figs. 4, 5).
Eather more of the first radials is visible on the exterior of the calyx in Antedon
antarctica than in Antedon eschrichti (PL XXIV. figs. 10, 11; PL XXV figs. 10-12),
and in some instances the ends of the basal rays appear between their lower angles
(PI. I. fig. 6a). As in Antedon eschrichti the shape of the second radials depends con-
siderably upon that of the axillaries. These are always wider than long (PL XXV.
figs. 8-11), but vary considerably in shape, even in the same individual. They are
almost triangular in some cases, and widely rhombic in others, owing to the strong back-
ward projection, which forms a sort of tubercle together with the very convex centre of
the second radial. There is a similar variation in the shape of the first pair of brachials
(PL XXV. figs. 10-12), and the junctions of the following joints are by no means so
tubercular as in the largest variety of Antedon eschrichti, though more so than in the
smaller and smoother Atlantic specimens, which have about the same size as the largest
individuals of Antedon antarctica that were obtained. In both species alike, however,
as in all the members of this group, the fourth to the eighth brachials bear pinnules on
their shorter sides. Beyond the third syzygy the arm-joints of Antedon antarctica are
even shorter relatively to the width than they are in Antedon eschrichti (PL XXIV.
fig. 11 ; PL XXV. fig. 12), and they have a very decided tendency to overlap which is
absent in that species, the arms of which are unusually smooth (PL XXIV. figs. 13, 14).
The same may be said of the pinnule-joints, especially of the genital pinnules nearest
the calyx ; while the long flagellate pinnules on the arm-bases are serrate from end to end
in Antedon antarctica (PL XXV. fig. 1-3), whereas in Antedon eschrichti the middle
joints are smooth with sharp edges but nothing more. The third pinnule of Antedon
antarctica is much more like its successor than is the case in Antedon eschrichti. Its
lower joints are considerably stouter than those of the second pinnule, some of them
being as long as or longer than wide (PL XXV. figs. 2, 3), whereas in Antedon eschrichti
they are distinctly wider than long. In fact the third pinnule of Antedon antarctica
resembles the fourth pinnule of Antedon eschrichti rather than its fellow, the third pinnule.
In the middle and outer pinnules there is a good deal of variation in the extent of
modification in the two basal joints ; but they are never so much flattened and so nearly
trapezoidal in form as they are in the larger Antedon eschrichti.
The centro-dorsal of Antedon antarctica is somewhat more conical than that of
Antedon eschrichti (PL I. figs. 6a, 8a), and the axial opening on the ventral surface is
(ZOOL. CHALL. ESP. PART LX. 18S7.) OoO 19
146 THE VOYAGE OF H.M.S. CHALLENGER.
relatively much larger than in that type (PI. I. figs. 6d, Sd), so as to reveal a large
number of the internal ridges which separate the inner openings of the cirrus-canals.
Five of these, interraclial in position, are much more prominent than the rest, as is well
seen in the figure (PI. I. fig. 6d). The basal grooves on the ventral surface of the centro-
dorsal are scarcely more distinct than they are in Antedon eschrichti (PI. I. fig. 8d).
But on the other hand the dorsal surface of the radial pentagon bears a very well-defined
basal star, of which there is rarely any trace in that species (PI. I. figs. 6c, 8c). The
rosette lies deeper than it does in Antedon eschrichti, and the basal rays connected with
it are unusually stout. This is most noticeable in their isolated condition (PI. I.
figs. 7, a, b; compare PL III. fig. 2, and PI. IV. figs. 4-6, all equally magnified). In some
cases their distal ends appear on the exterior of the calyx, as seen in PI. I. fig. 6a. This
figure too shows the difference between the articular faces of the radials in Antedon
eschrichti and Antedon antarctica respectively. Their slope is more uniform in the
latter species, as there is much less of an angle between the upper and lower parts of each
face than in Antedon eschrichti (PI. I. fig. 8a), and the consequence is that more of the
large muscular fossag is visible in the ventral aspect of the calyx (PI. I. figs. 6b, 8b).
The ridges which separate them from the lower pair of fossae are much more horizontal
than in Antedon eschrichti, so that the two pairs of fossse are of very unequal size
(PI. I. fig. 6a, 8a).
A detailed comparison of the two outer radials and of the lower brachials in the two
species respectively reveals a number of similar points of difference between them ; and
though they are so very closely similar in habit and in general appearance, as also in the
conditions of their existence, there can, I think, be no question that they are distinct.
3. Antedon australis, n. sp. (PI. XXVI. figs. 4, 5 ; PL XXVII. figs. 14-20).
Specific formula — A. — .
Centro- dorsal hemisjiherical, thickly covered with about fifty cirri. These have
twenty-five to thirty joints, nearly all of which are longer than wide. The later joints
are laterally compressed, and their dorsal edges project considerably beyond the bases of
their successors, thus giving rise to a strong spine in the last few joints. The young
cirri round the dorsal pole resemble the mature form, but have fewer joints ; those round
the margin may have thirty smooth and elongated joints which ouly develop spines
quite late.
First radials just visible ; the second short and nearly oblong, but little incised for
the axillaries, which are broadly pentagonal or triangular with a slight backward pro-
jection in the middle of the base.
Ten arms, of somewhat overlapping joints, but not tubercular at the base. The arm-
joints after the second syzygy are shortly triangular, gradually becoming quadrate, but
REPORT ON THE CRINOIDEA. 147
always much wider than long. Syzygies in the third, eighth, and twelfth brachials, and
then at intervals of two or three joints.
The first pair of pinnules (on second and third brachials) flagellate, about 12 mm.
long, and consisting of forty -five short joints, the basal ones of which are broad, flattened,
and somewhat carinate. The next pair sometimes nearly equal to and sometimes shorter
than the first. The third pair also shorter, with stouter joints, most of which are
distinctly longer than wide, and they generally bear fusiform genital glands. The
following pinnules more massive, with squarer joints, which become elongated further out,
while the two basal ones become flattened and trapezoidal, with their apposed edges
incurved.
Disk and ambulacra naked. Sacculi abundant in some pinnules, but less so in others.
Colour in spirit, — white, with purplish or brownish patches.
Disk 1 0 mm. ; spread about 1 2 cm.
Locality.— Station 150, Febuary 2, 1874; lat. 52° 4' S., long, 71° 22' E.; 150
fathoms ; coarse gravel; bottom temperature, 35°"2 F. Seven mutilated specimens and
one very young.
Remarks. — This is a smaller and more delicate species than Antedon antarctica, which
it resembles in the shortness of the arm -joints ; but the arms generally are much smoother,
and there are fewer cirrus-joints, while the third pinnule is much less like the second
than is the case in that species (PL XXV. figs. 2, 3 ; PI. XXVII. figs. 15, 16). The three
lowest joints are by no means so wide as in Antedon antarctica, but more nearly
square, while the following joints till quite near the end are very distinctly longer
than broad, which is not the case in Antedon antarctica. Even the first two pinnules
of Antedon australis have a tendency in this direction, as compared with the much
longer ones of Antedon eschrichti and Antedon antarctica (PI. XXIV. figs. 7, 8 ; PI.
XXV. figs. 1. 2).
Like both these last mentioned types and the other Arctic species as^ell {Antedon
quadrata, Antedon hystrix, and Antedon prolixa), Antedon australis affords an excellent
illustration of the dimorphic mode of development of the cirri. Its full-grown cirri
consist of some twenty- five to thirty-five joints, the first half of which, except those
just at the base, are considerably longer than wide. As they get shorter their dorsal
edges come to project more and more definitely beyond the bases of the succeeding joints,
so that the compressed terminal segments are distinctly spinous (PI. XXVII. fig. 20).
There are numerous young cirri of this type round the dorsal pole. They consist of
twenty joints, all of which, except the two or three short ones at the base, have a
forward projecting dorsal edge (PL XXVII. fig. 18). But on the other hand, the margin
of the centro-dorsal bears several young cirri of an altogether different type in various
stages of development (PL XXVII. figs. 17, 19). One of them has thirty elongated
148 THE VOYAGE OF H.M.S. CHALLENGER.
joints, all of which are perfectly smooth and without any traces of the dorsal projections
which are so characteristic of the joints composing the shorter and more centrally placed
cirri. These long marginal cirri eventually develop spines and only differ in their
greater length from those nearer the dorsal pole.
Besides seven individuals of Antedon australis found at Station 150, which are all
pretty equally developed, the Challenger also obtained a mutilated calyx of much smaller
size, from which all the arms had broken away at the syzygies in the third brachials
(PL XXVI. fig. 5). It may belong to this same species, but if so, it is remarkable in not
showing more of the first radials externally than is visible in the more mature forms,
though, on the other hand, the axillaries are relatively longer, as would be expected in a
young individual. This may also account for the elongated shape of the joints of the
first pinnule, and for the present it will be safest to regard this form, which has
numerous cirrus-sockets on the centro-dorsal, though barely 4 mm. in diameter, as a
young example of Antedon australis.
4. Antedon rhomboidea, n. sp. (PL XII. figs. 1,2; PL XXIV. figs. 1-3).
Specific formula — A.y.
Description of an Individual. — Centro-dorsal hemispherical, with a flattened and
cirrus-free dorsal pole. Sixty or more cirri, of thirty to thirty-five segments, several of
which are longer than wide. The middle joints project beyond their successors on the
dorsal side, and in the shorter terminal joints this projection develops into a blunt keel.
First radials just visible ; the second trapezoidal, rather deeply incised and rising
to tubercles at the junction with the rhombic axillaries, which are much wider than the
distal ends of the second radials.
Ten arms, with a median knob at the junction of the first two brachials, and others
alternating on the outer and inner sides till the ninth or tenth joint. The first brachials
deeply incised, with very short inner ends, which barely meet above the sharp angles of
the axillaries. Middle arm -joints triangular, soon becoming quadrate, as long as or
longer than wide, and slightly overlapping. Syzygies in the third, eighth, and fourteenth
brachials, and then at intervals of two or three joints. The first two pinnules on each
side about 22 mm. long, flagellate, and composed of fifty short joints, the basal
ones broad, flattened, and slightly carinate, and the later joints serrate. The third
pinnule of about the same size but with fewer joints, the basal ones being stouter and
the middle ones longer than wide. The following pinnules are more massive, with square
joints, which soon become elongated, the two basal ones but little modified.
Mouth subcentral, with rather large calcareous concretions round the peristome and
at the sides of the ambulacra on the disk and arm-bases. Sacculi very abundant.
Colour in spirit, — light brownish-white.
EEPOET ON THE CEINOIDEA. 149
Disk 12 mm.; spread at least 25 cm.
Locality.— Station 308, January 5, 1876 ; lat. 50° 8' 30" S., long. 74° 41' 0" W.;
175 fathoms; blue mud. One specimen.
Remarks. — This species resembles Antedon australis in the length of the middle
joints of the third pinnule, but it is altogether a much larger form, with considerably
longer arm -joints than occur in that species. In this respect Antedon rhomboidea and
Antedon magellanica have the same relation to Antedon australis and Antedon
antarctica that the northern species Antedon quadrata has to Antedon eschrichti.
Antedon magellanica has never yet been properly described, the type specimen having
been regarded merely as a variety of Antedon eschrichti,1 although its arm-joints are
mostly quadrate and as long as or longer than wide. It has many points of resemblance
with Antedon rhomboidea, but there may be over fifty cirrus-joints, while some of the
middle joints of the second pinnule are longer than wide. It is nearly half as long again
as the corresponding pinnule in an equal sized specimen of Antedon rhomboidea, while
the third pinnule is considerably shorter than the second. Several examples of the type
were obtained by the Italian corvette " Vettor Pisani," and I hope soon to be able to
describe them at length.
5. Antedon quadrata, n. sp. (PI. XXVI. figs. 1-3 ; PI. XXVII. figs. 1-13 ; woodcut,
figs. 4, a, b, on p. 154).
C
Specific formula — A.y.
1878. Antedon celficus, von Marenzeller, Denkschr. d. k. Akad. d. Wiss. Wien, 1877 [1878],
Ed. xxxv. p. 380.
1880. Antedon celticus, d'Urban, Ann. and Mag. Nat. Hist., 1880, ser. 5, vol. vi. p. 380.
1881. Antedon celtica, Duncan and Sladen, Memoir Arctic Eclnnodermata, London, 1881,
pp. 75-77, pi. vi. figs. 5, 6.
1881. Antedon celtica, P. H. Carpenter, Zool. Anzeiger, 1881, Jahrg. iv. p. 521.
1882. Antedon celtica, Bell, Proc. Zool. Soc. Lond, 1882, p. 534.
1882. Antedon celtica, P. H. Carpenter, Ibid., p. 746.
1884. Antedon quadrata, P. H. Carpenter, Proc. Koy. Soc. Edin., 1884, pp. 375-377.
1886. Antedon quadrata, P. H. Carpenter, Bijdragen tot de Dierkunde, 1886, 13 Aflevering,
vi. p. 7, pi. i. fig. 6.
1886. Antedon Eschrichtii, Levinsen {jpars), Dijniphna-Togtets zoologisk-botaniske Udbytte,
Kj0benhavn, 1886, p. 410.
1886. Antedon quadrata, Fischer, Die Osterreichische Polarstation Jan Mayen, Bd. iii., "Wien,
1886, Echinodermen, p. 3.
Centro-dorsal hemispherical, bearing fifty to seventy cirri, with about thirty to
thirty -five joints, several of which are longer than wide ; the later joints sharpened
dorsally but not distinctly carinate.
1 See Proc. Zool. Soc. Lond., 1882, p. 650.
150 THE VOYAGE OF H.M.S. CHALLENGER.
First radial very short ; the second longer and trapezoidal, somewhat incised by the
rhombic axillary and forming with it a slight prominence. Axillary as wide as or wider
than long, with a fairly open distal angle.
Ten arms, of some two hundred smooth joints. First brachial rather deeply incised,
with a short inner and much longer outer edge ; the second irregularly quadrate, and
the six following joints more oblong or obliquely quadrate, with the pinnule on the
shorter side. The first three joints above the third syzygy are sometimes nearly
triangular and as wide as long, but the following ones are distinctly quadrate and
gradually become longer than wide ; the terminal joints somewhat elongated. Syzygies
in the third and eighth brachials and then at intervals of two to four joints.
The first two pinnules long and flagellate, composed of numerous short joints, the
basal ones wide and slightly carinate, sometimes with dorsal processes, and the later
joints serrate. The third pinnule little more than half as long as the second, with much
fewer joints, the basal ones stouter, and the remainder mostly much longer than wide.
The following pinnules more massive, with square joints which gradually become
elongated. The two lower joints of the distal pinnules flattened and trapezoidal in some
forms, but only slightly modified in others.
Ambulacra not plated, but the disk sometimes bears a number of small calcareous
granules.
Colour in spirit, — white or brownish-white.
Disk 15 mm.; spread may reach 30 cm.
Localities. — H.M.S. Challenger, Station 48, May 8, 1873; on the Le Have Bank;
lat. 43° 4' N., long. 64° 5' W.; 51 fathoms ; rock. Several specimens.
H.M.S. "Valorous," Station 6, August 10, 1875 ; Davis Strait ; lat. 64° 5' N., long.
56° 47' W.; 410 fathoms ; sand and mud ; bottom temperature, 34°-6 F. One specimen.
H.M.S. "Alert," 1875; Franklin-Pierce Bay (in Smith's Sound); lat. 79° 25' N.;
and Discovery Bay (in Eobeson Channel), lat. 81° 41' N.; 25 fathoms ; hard bottom.
H.M.S. "Triton," 1882, Station 4; lat, 60° 22' 40" N. and 60° 31' 15" N., long.
8° 21' W. and 8° 14' W.; 327 to 430 fathoms; stones, mud; bottom temperature, 31° "5
to 30° F.
Station 6; lat. 69° 8' 0" N., long. 7° 26' 30" W.; 466 fathoms; stones; bottom
temperature, 29c,5 F.
Other Localities. — Barents Sea ("Tegetthoff" and " Willem Barents"); Kara Sea
("Varna") ; Jan May en (Fischer).
Remarks. — This species has a curious history and has caused me no little trouble.
I have no doubt that it was dredged by the " Porcupine" in 1869 in the cold area of
the Fseroe Channel, together with Antedon eschrichti, which was met with in compara-
tive abundance ; but it is not now to be found among the remains of the " Porcupine "
REPORT ON THE CRINOIDEA. 151
collection which have come into my hands. Three years later (1872) it was obtained by
the ill-fated "Tegetthoff " 5° west of Nova Zembla, and was minutely described by von
Marenzeller 1 who referred it to Antedon celticus, Barrett, sp. Little was then known of
the latter form, except for the very incomplete description of it which had been given by
Barrett, and for Sir Wyville Thomson's incidental references to the numerous examples of
it which had been dredged off the north coast of Scotland by the "Lightning" and
" Porcupine."
Von Marenzeller, regarding his Arctic specimen as identical with Barrett's type,
gave a careful description of them which enabled Duncan and Sladen to recognise the
same form among the Comatulse dredged by Sir George Nares's Arctic Expedition of
1875-76. These were well and carefully described by Sladen,2 who was the first to
figure the type, though still under Barrett's specific name " celtica." He used this
designation with some hesitation, however, owing to the prevalent want of knowledge
respecting Barrett's species ; and after writing his description of the more northern form
he saw for the first time some examples of the true Antedon celtica, which he recognised
as altogether distinct from the Arctic type. He therefore inserted a note to this effect,
but did not alter the name under which the latter had been described by both von
Marenzeller and himself. It will be shown further on that Barrett's species has proved
to be identical with the long but little known Antedon phalangium, Midler, sp.. of the
Mediterranean, and the specific name celtica being therefore unoccupied, I thought at
first 3 that it might conveniently be retained for the type described under this name by
von Marenzeller and Sladen respectively. This course, however, has seemed undesirable
for many reasons ; and in compliance with the wishes of both the above named naturalists
I propose to give it a new name altogether. I have therefore chosen one indicative of the
character by which the species is most easdy distinguished from Antedon eschrichti, viz.,
the markedly quadrate shape of the middle and outer arm-joints, as seen in PI. XXVII.
figs. 5-7, and in the woodcut, fig. 4 on p. 154.
Thanks to the kindness of Dr. von Marenzeller I have been able to examine the
single individual dredged by the " Tegetthoff," and I am satisfied that Sladen was right
in identifying it with those which he described from Smith's Sound and Robeson Channel.
Another example was dredged by the " Valorous " in Davis Strait in 1875, and when the
Challenger Comatulse came into my hands I found the same type among a quantity of
specimens of Antedon eschrichti from Station 4S on the Le Have Bank (51 fathoms).
The species was twice taken by the "Triton" (1882) in the cold area of the Fseroe
Channel. The "Willem Barents" met with it in 1880 near the locality of the
"Tegetthoff" dredging; and it was twice obtained by the "Varna" in the Kara Sea
1 Die Coelenteraten, Echinodermen, unci Wurnier der k. k. Osterreieliisch-TTngarischen Nordpol. Expedition,
Berikschr. d. k. Alaul. d. Wiss. Wim, 1877 [1878], Bd. xxxv. p. 380.
2 Op. cit., p. 75.
3 Note on the European Comatnhe, Zool. Anzeiger, 1881, Jahrg. iv. p. 521.
152 THE VOYAGE OF H.M.S. CHALLENGER.
(1883). Not improbably too it may have been among the collections made by the
earlier " Willem Barents " Expeditions and by the " Vega "; though, as in the case of the
" Porcupine " specimens, it was not distinguished from immature individuals of Antedon
eschrichti.
A careful study of all this material has convinced me, however, that the two forms
are very different in reality ; though, as I have pointed out above, Antedon quad rata
may in some sense be regarded as a permanent larval form of Antedon eschrichti. Its
first radials are not entirely concealed by the centro-dorsal, but appear above it as short
band-like plates (PL XXVI. figs. 1-3). The second radials have more sloping sides than
in the smaller forms of Antedon eschrichti, so as to be trapezoidal in general outline ; and
the axillaries have a blunter distal angle than in that type (PI. XXIV. figs. 10, 11).
The arm-bases are not tubercular, though the joints between the first and second syzygies
have the same backward projections on the sides which do not bear the pinnules that
occur in Antedon eschrichti. The relatively long quadrate shape of the arm-joints
immediately after the third syzygy is less marked in the Challenger specimens of Antedon
quadrata, the southernmost ones known (PL XXIV. fig. 2), than it is in the two obtained
further north by the " Triton " (PL XXIV. fig. 3), and in those from the Arctic Ocean
which have been figured elsewhere by Sladen ' and myself.2 But the middle and outer
arm-joints of the two species are always distinguishable, those of Antedon eschrichti
being short, generally triangular, and much wider than long, till quite near the end of the
arm ; while the brachials of Antedon quadrata are obliquely quadrate and the length is
more nearly equal to the width. This is especially marked in the " Valorous " specimen,
and is no doubt partly due to its not being quite mature, as in the young forms obtained
by the Challenger (PL XXVI. fig. 1) ; but it is also very distinct in the larger examples
from the Barents and the Kara Seas.
The other special mark of Antedon quadrata is the disproportion between the second
and the third pinnule, which has already been noticed by Sladen as distinguishing the
type from Antedon eschrichti (PL XXVII. figs. 9, 10, 12, 13). In the individuals of
the latter species which were obtained by the Challenger at Station 48, the third pinnule
is relatively much shorter than in the more northern forms. In large examples of
Antedon eschrichti from the Arctic Ocean it is of almost exactly the same length as the
second pinnule, as described by Sladen ; but in the West Atlantic representatives of the
type it is distinctly shorter (PL XXIV. figs. 8, 9). The southern forms of the two species
therefore approach one another in the characters of the pinnules, just as in those of the
arm-joints ; although the more northern varieties are entirely distinct in both respects.
Not only is the third pinnule of Antedon quadrata altogether smaller than the
second, but its component joints, while fewer in number, are also different in their
1 Op. tit., pi. vi. figs. 5, 6.
2 Bijdragen tot de Dicrkunde, 1886, 13 Aflevering, vi., pi. i. fig. 6.
REPORT ON THE CRINOIDEA. 153
relative proportions. The basal joints are stouter as in the following genital pinnules,
and their successors are distinctly longer than wide, indications of which appear in the
second pinnule (PI. XXVII. figs. 9, 10, 12, 13). There is no sign of this, however, in
Antedon eschrichti, the joints of the third pinnule being as wide or wider than long
(PI. XXIV. figs. 8, 9). Furthermore there is generally less trace in Antedon quadrata
of the modification of the two lowest joints in the outer pinnules, which is usually so
marked in Antedon eschrichti (PI. XXIV. fig. 13), though it is extremely well developed
in an example brought by the " Varna" from the Kara Sea.
Levinsen1 has recently united Antedon quadrata with Antedon eschrichti, on the
ground that the characters of this latter species as stated by von Marenzeller, Sladen, and
myself, all present themselves in immature examples of Antedo?i eschrichti. The
possibility of this being the case had of course naturally occurred to me ; but I decided
against it for various reasons.
Levinsen is not personally acquainted with Antedon quadrata, but only knows it
from the descriptions of Sladen and von Marenzeller, and from my own preliminary
account of its special marks, characters which, as I am fully aware, do occur in young
individuals of Antedon eschrichti, though not, I think, to the same degree that they do in
Antedon quadrata. Had Levinsen been able to compare an example of Antedon
quadrata with an equal-sized but immature individual of Antedon eschrichti, I believe
that he would have found differences between them which he would recognise as of
specific value.
One of the " special marks " which I mentioned as distinctive of Antedon quadrata
when the type was rebaptised, was the very definite quadrate shape of the middle and
outer arm-joints (PL XXVII. figs. 5-7) as compared with those of Antedon eschrichti
(PI. XXIV. figs. 11, 14), which are much shorter than wide, with their sutures less oblique
than in Antedon quadrata. The young Antedon eschrichti also has relatively long and
quadrate arm -joints with oblique sutures, and Levinsen assigns this as one of his reasons
for uniting the two species. I was of course perfectly aware of this fact when I named
Antedon quadrata, and described it as a permanently immature form of Antedon
eschrichti.2 Since the publication of Levinsen's memoir, which only reached me after the
preceding pages were written, I have gone into the subject again in the only way which
can possibly give a satisfactory result, namely, the comparison of the corresponding arm-
joints in equal-sized individuals of the two species.
Figs. 4, A and B, on the next page, represent the portions of the arms between the
fiftieth and sixtieth brachials of two individuals of Antedon quadrata from different
localities. In both cases the joints are of an obliquely quadrate shape and nearly as long
as wide. But in the corresponding part of the arm of a young Antedon eschrichti of
equal size the joints are more nearly triangular and considerably wider than long (fig. 4, c),
1 hoc. cit., p 413. 2Proc. Boy. Soc. Edin., 1884, vol. xii. pp. 374-376.
(ZOOL. CHAIX. EXP. — PART LX. — 1S87.) OoO 20
154 THE VOYAGE OF H.M.S. CHALLENGES.
and in full grown individuals this character is still more marked. This is shown in
fig. 4, d, which represents a portion of the third quarter of an arm in the largest specimen
of Antedon eschrichti that I have seen. It is drawn of the same size as the three other
arm-fragments, so that their differences may be the more readily compared. The extreme
shortness of the arm-joints is one of the most striking characters of Antedon eschrichti,
though it occurs also in Antedon antarctica (PI. XXV. fig. 12) ; and it manifests itself in
individuals which have not attained half their full size, while it does not appear in
Fig. 4. — A, B, The fiftieth and next following brachials of Antedon quadrata; x 6. C, The same joints in a young
Antedon eschrichti of equal size ; x 6. D, Arm-joints of a mature individual ; x 4.
examples of Antedon quadrata which have the same dimensions, as shown in figs. 4, A,
B, c. The difference may seem but a slight one in small pieces of arms like those figured ;
but it produces a very decided effect on the general facies of the whole plume of arms,
on account of the greater or less separation of the successive pinnules from one another.
A second point of difference between the two species is the relatively small size
of the third pinnule in Antedon quadrata, as compared with that of Antedon eschrichti,
(PI. XXIV. figs. 8, 9 ; PI. XXVII. figs. 9, 10, 12, 13). Levinsen l has pointed out that
as the second pinnule appears before the third, there is necessarily a period of growth at
which the third pinnule will be only about half the size of its predecessors, as is the
case in Antedon quadrata.
This of course is perfectly true ; but as I have pointed out above, the relative propor-
tions of its component joints are not the same in the two types. The difference is
similar to that which shows itself in the arms, i.e., the joints are relatively longer in the
lower pinnules of Antedon quadrata than in those of Antedon eschrichti. The following
measurements of the second and third pinnules in Antedon quadrata and in a
young Antedon eschrichti of equal size will make this point clear.
Second pinnule. Third pinnule.
Length. Number of joints. Length. Number of joints.
Antedon quadrata, 14 31 8 17
Antedon eschrichti, . 15 39 12 28
1 Loc. cit, pp. 32, 33.
REPORT ON THE CRINOIDEA. 155
Thus then the third pinnule of Anteclon quadrata is only % as long as the second ;
whereas in Antedon eschrichti it reaches | of the size of the second, and a similar
difference appears in the relative proportions of their component joints.
There is another consideration which, taken by itself, would have no special probative
value ; but it is not without importance when combined with the other evidence
given above. Antedon quadrata has been dredged at eleven stations altogether, but at
only five of these was it found in association with Antedon eschrichti. The " Triton,"
"Alert," "Valorous," "Tegetthoff " and "Varna" (bis) obtained examples of this type at
localities where Antedon eschrichti did not occur ; and in the last four cases they were
only single individuals.
These facts would seem somewhat improbable if Antedon quadrata is merely
an immature stage of Antedon eschrichti as supposed by Levinsen. It is a common
experience of Arctic dredging to find individuals of Antedon eschrichti associated together
in considerable abundance, and at various stages of development ; and one would
therefore not expect to find isolated examples of young individuals, unaccompanied by
older ones, quite so frequently as is mentioned above.
Sladen is the only naturalist, besides myself, who has had the opportunity of directly
comparing examples of the two species which were obtained at the same locality ; and in
spite of Levinsen's remarks, I am still inclined to think that he was right in separating
the two forms. I find it difficult to believe that the fine example of Antedon quadrata
which I have figured in the " Varna" report is merely a young stage of the Antedon
eschrichti obtained at the same locality ; though I am by no means prepared to state
definitely that it is not the case.
My present impression is that we have to deal with two distinct species, the smaller
of which, as I have remarked before, represents a permanently immature form of the
larger one.
Antedon quadrata is another of the species in which the cirri are strikingly dimorphic
in their character. The mature cirrus of an Atlantic specimen is shown in PL XXVII.
fig. 1, while fig. 2 represents one that is still immature as shown by the relative length
of the sixth and following joints. This cirrus has developed upon the ordinary plan, a
much earlier stage of which is seen in fig. 4 ; but fig. 3 represents another young cirrus,
altogether different in appearance and belonging to the " small mature" type, just as has
been described in Antedon antarctica and Antedon australis (PL XXV. fig. 7 ;
PL XXVII. fig. 18). In the young individual figured on PL XXVI. fig. 1, the spread
of which cannot have been more than four or five centimetres, most of the cirri seem to
have developed upon the small mature plan ; but a few rudimentary cirri of the other
type are to be found round the margin of the centro-dorsal, and there are more in a con-
siderably older though still immature individual.
The youngest form obtained shows less of the first radials on the exterior of the
156 THE VOYAGE OF H.M.S. CHALLENGER.
calyx than might have been expected ; but its early age is indicated by the great relative
length of the arm-joints and the small size of the proximal pinnules (PI. XXVI.
fig. 1).
This species ranges slightly further north than Antedon eschrichti, having been
obtained at Discovery Bay (lat. 81° 41' N.) together with Antedon prolixa, and
at Franklin-Pierce Bay (lat. 79° 25' N.) together with Antedon eschrichti, which was not
met with at the higher latitude. The bathymetrical range is greater, however, in
the larger form, which extends down to 632 fathoms, while Antedon quadrata was
not found below 466 fathoms in the same region of the Feeroe Channel ; the nearest
approach to this depth being the "Valorous" station in Davis Strait (410 fathoms).
The three "Triton" specimens are all of them small, like those of the "Tegetthoff"
and "Valorous"; while they have a stiff er and less feathery appearance than the
larger ones obtained further north by the " Alert " and " Willem Barents."
In fact they more nearly resemble the small individual figured by Sladen1 in
their general characters. The dorsal processes on the lower joints of the basal pinnules
are less prominent than usual ; while the peculiar characters of the first two pinnule-joints
in the outer parts of the arms are by no means so marked as in larger individuals.
The only Arctic species that approaches Antedon quadrata in the great disproportion
between the second and third pinnules is Antedon barentsi ; but it has much smaller
cirri with fewer joints, triangular joints in the middle of the arms, and the genital
pinnules protected by plates as in Antedon incisa, Antedon acoela, and other tropical
forms, though on a less massive scale.
Like Antedon eschrichti, with which it is often associated, Antedon quadrata
officiates as host to Myzostoma gigas.
4. The Tenella-groviTp.
Long-jointed lower pinnules.
The first three species on the list of those which I have included in this group,
Antedon phalangium, Antedon hystrix, and Antedon prolixa, have many affinities with
Antedon eschrichti and its allies, both in their distribution and in the characters of
their arms and cirri.
Antedon hystrix and Antedon prolixa are exclusively cold-water species, not
having been obtained south of lat. 60° N.; though Antedon phalangium occurs in the
Mediterranean and in the East Atlantic as far south as the Seine Bank (lat. 33° 47' N.,
long. 14° 1' W.). These species differ from the Eschrichti-growp, however, in the
characters of the lower pinnules, which, though often long, slender, and more or less
flagellate, consist of joints which are much longer than wide, as is particularly evident
1 Op. cit,, pi. vi. fig. 5.
REPORT ON THE CRINOIDEA. 157
in Antedon longipinna, Antedon tenella, and Antedon exigua (PI. XXX. fig. 2 ; PI.
XXXI. fig. 4 ; PI. XXXII. fig. 4). On the other hand, in Antedon eschrichti and its
allies the first two or three pairs of pinnules consist of short and wide joints (PI. XXIV.
figs. 1, 2, 7-9; PI. XXV. figs. 1, 2 ; PI. XXVII. figs. 8, 9, 11, 12, 14). The species of
the Eschrichti-grou-p are exclusively confined to the Atlantic and Circumpolar Seas,
but do not extend downwards below 650 fathoms. The Basicnrva-growp, however, is
principally limited to the Pacific, ranging from 140 to 1350 fathoms. But there are
three Atlantic species which occur at depths of 420 to 1600 fathoms. On the other hand
the members of the TeneZZa-group are chiefly confined to the Arctic Ocean, the Atlantic,
and the Southern Sea. Antedon prolixa is one of the two northernmost Comatulse
known ; and two more {Antedon hirsutq and Antedon exigua) reach further south than
any others except Antedon australis and Antedon antarctica. Of the remaining fourteen
species only five occur in the Pacific ; four of them range between 150 and 775 fathoms ;
while the fifth, obtained at 2900 fathoms in the North Pacific (Station 244), was also
found at 2600 fathoms in the Southern Ocean (Station 160). These were the two
deepest stations at which Comatulse were met with. The next deepest (1600 fathoms)
in the Southern Ocean (Station 147) yielded two more species of the Tenella -group,
the remaining members of which are confined to the Atlantic. One or two of them
are littoral species, like Antedon rosacea itself, and Antedon diibeni, but this is not
the case with the Pacific members of the group, which are not known as yet to occur
above 150 fathoms.
With a few exceptions, then, the Tenella-growp may be regarded as especially
characteristic of the Atlantic and Circumpolar Seas, just as the Basicurva-gvowp is
chiefly confined to the Western Pacific ; and in each case the exceptional species belong
to the continental or abyssal, but never to the littoral fauna.
Long-jointed lower pinnules.
A. Forty or more long cirrus-joints.
I. Second pinnule as long as the first, . . . .1. phalangium, Mull, sp.
II. Second pinnule smaller than the first.
a. Axillaries longer than wide; second radials very deeply
incised, ....... 2. hi/strix, n. sp.
b. Axillaries as wide or wider than long; second radials incised,
but not very deeply so, . . . . . prolixa, Duncan and Sladen.
B. Fifteen to thirty cirrus-joints.
I. Second pinnule distinctly smaller than the first.
a. Cirrus-joints mostly longer than wide, the lower ones very
much so.
1. First pinnule nearly three times as long as the second ;
syzygial interval two joints, . . .3. tenella, Retzius, sp.
15S THE VOYAGE OF H.M.S. CHALLENGER.
2. First pinnule less than twice as long as the second.
a. Syzygial interval two joints ; later cirrus-joints
not specially long, . . . .4. etigua, n. sp.
/?. Syzygial interval one joint; later cirrus-joints
elongated, . . . . .5. alteniata, n. sp.
b. Cirrus-joints not specially elongated.
1. Lower joints of genital pinnules not expanded; cirrus-
joints not spiny.
..,,., n ( rosacea, Linck, sp.
a. Lower arni-ioints triangular, . . . o. < ' ' f
( petasus, Dub. and Ivor., sp.
/8. Arm-joints quadrate from the fifteenth, if not
before, . . . . .7. diibeni, Bohlsche.
2. Genital pinnules having expanded lower joints; cirrus-
joints spiny, . . . . .8. lineata, n. sp.
II. First and second pinnules tolerably equal.
a. Less than twenty cirrus-joints which are not specially
elongated; syzygial interval two joints, . . .9. remota, n. sp.
b. Over twenty cirrus-joints.
1. Elongated cirrus-joints; lower pinnules very long.
a. Second pinnule slender like the first ; syzygial
interval one joint, . . 10. longipinna, n. sp.
/}. Second pinnule stouter than the first ; syzygial
interval two joints or more, . . .11. tenuicirra, n. sp.
2. Cirrus-joints not specially elongated.
a. Arms smooth; axillaries long, . . .12. Ixvis, n. sp.
ft. Arms serrate and spiny ; axillaries wide, . 13. hirsuta, n. sp.
III. Second pinnule longer than the first.
a. Cirrus-joints short ; syzygial interval one joint, . .14. angustipinna, n. sp.
b. Cirrus-joints long; syzygial interval one to five joints, . 15. abyssorum, n. sp.
C. Less than twelve cirrus-joints, . . . . . .16. abyssicola, n. sp.
1. Antedon phalangium, Mull., sp. (PL XXVII. figs. 23-29; PI. XXVIII. figs. 1-3).
be
Specific formula, A.—.
1841. Aleeto phalangium, Miiller, Monatsber. d. k. preuss. Akad. d. Wiss. Berlin, 1841, p. 182.
1849. Comatula (Aleeto) phalangium, Miiller, Abhandl. d. k. Akad. d. Wiss. Berlin, 1849, p. 253.
1857. Comatula Woodwardii, Barrett, Ann. and Mag. Nat. Hist., 1857, ser. 2, vol. xix. p. 33,
pi. vii. fig. 1.
1857. Comatula celtica, M'Andrew and Barrett, Ann. and Mag. Nat. Hist., 1857, ser. 2, vol. xx.
p. 44.
1862. Comatula phalangium, Dujardin and Hupe, Hist. Nat. des Zoophytes, fichinodermes,
Paris, 1862, p. 198.
1865. Antedon cellicus, Norman, Ann. and Mag. Nat. Hist., 1865, ser. 3, vol. xv. p. 104.
1872. Antedon mediterraneus, Wyville Thomson, Proc. Roy. Soc. Edin., 1872, vol. vii. p. 765.
1872. Antedon celticus, Wyville Thomson, Ibid., p. 765.
1879. Antedon phalangium, P. H. Carpenter, Trans. Linn. Soc. Lond. (Zool.), ser. 2, 1879,
vol. ii. p. 29.
1879. Antedon celtica, P. H. Carpenter, Ibid., p. 29, pL iv. figs. 1-8.
REPORT ON THE CRINOIDEA. 159
1879. Antedon phalangium, Marion, Ann. d. Sci. Nat. (Zool.), 1879, ser. 6, t. viii. p. 40,
pi. xviii.
1879. Antedon phalangium, Ludwig, Mittheil. a. d. Zool. Stat. Neapel, 1879, Bd. i. p. 537.
1880. Antedon phalangium, Ludwig, Ibid., 1880, Bd. ii. p. 53, Taf. iv. fig. 1.
1881. Antedon phalangium, P. H. Carpenter, Zool. Anzeiger, 1881, Jahrg. iv. p. 521.
1882. Antedon phalangium, Bell, Proc. Zool. Soc. Lond., 1882, p. 534.
1883. Antedon phalangium, P. H. Carpenter, Proc. Zool. Soc. Lond., 1882 [1883], p. 746.
1884. Antedon phalangium, P. H. Carpenter, Proc. Eoy. Soc. Edin., 1884, vol. xii. p. 361.
1886. Antedon phalangium, P. H. Carpenter, Trans. Linn. Soc. Lond. (Zool.), ser. 2, 1886,
vol. ii. p. 476.
Centro-dorsal hemispherical, conical, or somewhat columnar, bearing from twenty-
five to thirty-five cirri. The longest of these have forty to fifty joints, of which the
fourth and its immediate successors are somewhat longer than wide.
The following joints may either remain longer than wide and end in a sharp claw,
usually without an opposing spine, or become gradually shorter, so as to be square or
even shorter than wide at the end of the cirrus, with one or two opposing spines and a
smaller terminal claw.
Three radials visible ; the second but little united laterally, nearly oblong, strongly
convex, and more or less incised for the axillaries, which are of variable shape, usually
pentagonal and wider than long, but sometimes rhombic and sometimes hexagonal.
Ten arms of two hundred smooth joints. First brachials also of variable shape, but
usually not very much incised by the second ; the fourth and three following brachials
nearly oblong, with alternating backward projections and pinnules on the shorter sides.
Above the second syzygy the joints are at first triangular, and as long or longer than
wide, but they gradually become quadrate and finally elongated. Syzygies in the
third, eighth, and twelfth brachials, and then at intervals of one to seven joints,
usually two or three.
The first two pairs of pinnules (on the second to fifth brachials) are tolerably equal,
long and flagellate. They consist of thirty to thirty-five joints, of which the six or
eight at the base are quite short, and their successors much longer than wide. The
following pinnules are shorter, with fewer joints, but the length of the lower ones
gradually increases, except in the two at the base, which become somewhat flattened.
Ovaries long and fusiform. Disk naked. Pinnule-ambulacra sometimes imperfectly
protected by small and delicate plates ; sacculi very abundant.
Colour, — green when alive ; brownish-white in spirit. Disk 7 mm. ; spread about
30 cm.
Localities.— H.M.S. "Lightning," 1868, Station 13; kit. 59° 5' N., long. 7° 29' W.;
189 fathoms.
H.M.S. "Porcupine," 1869; the Minch ; 60 to 80 fathoms. Several specimens,
with Myzostoma alatum and Myzostoma pulvinar. Also off Loch Scavaig, Skye.
Cruise of 1870; Station 13, off Cape Mondego ; lat. 40° 16' N. long. 9° 37' W.;
160 THE VOYAGE OF H.M.S. CHALLENGER.
220 fathoms ; bottom temperature, 52° F. Several specimens, with Myzostoma
alatum.
Off Cape Sagres (near Cape St. Vincent) ; 45 fathoms. Several specimens.
Off Carthagena ; 80 fathoms. Several specimens, with Myzostoma alatum.
Bay of Benzert ; 50 to 100 fathoms. Abundant.
Skerki Bank ; 30 to 120 fathoms. Abundant.
Other Localities. — (Mediterranean) Naples ; Nice ; Marseilles. The Atlantic — the
Seine Bank, 88 fathoms (S.S. "Dacia"); off Cadiz ("Talisman").
Remarks. — This species was described by Muller so long ago as 1841, though for
a long time but little was known about it. The original specimens which Muller
examined had been obtained at Nice and at Naples, but for many years afterwards the
type was never recorded as having been found at either of these localities or anywhere
else. It was obtained off the coast of Tunis by the "Porcupine" Expedition of 1870,
though the fact was not recognised at the time ; and it was not till 1879 that much
attention was directed to it. Professor Marion had dredged it four years previously in
the harbour of Marseilles, and he gave a careful analysis of its peculiarities, which was
accompanied by some excellent figures.1
Meanwhile, in the year 1857, a Comatula, which had been dredged by Mr. MAndrew,
in the sound of Skye, was briefly described by Barrett 2 as new both to science and to
the British fauna. He at first called it Comatula woodwardii, but on finding that
this specific name had been previously employed by Edward Forbes for a fossil species
from the Crag, he proposed to call it Comatula celtica, under which name it is
recorded as having been dredged in the Minch by the " Lightning " and " Porcupine "
in the cruises of 1868-69. The original specimens to which Barrett gave the name
Comatula celtica disappeared for a considerable time, and it was not till they were
discovered in the collection of the British Museum by Professor F. J. Bell that the
true nature of his type was revealed. They are somewhat smaller than those which
had been obtained in the Minch by Mr. Gwjm Jeffreys, and by the " Lightning " and
" Porcupine," and had been generally referred to Antedon celtica. But during the
next twenty years neither Barrett nor any British zoologist seems to have thought of
comparing them with the second Mediterranean species of Antedon, the first of which
(Antedon rosacea) is abundant on the British coasts ; while the examples of Antedon
celtica, which were dredged in abundance on the Tunis coast in 1870, were noticed by
Sir Wyville Thomson 3 in the following passage : — " Many examples of the form
known to continental naturalists under the name A. mediterraneus, Lam., sp., were
1 Draguages au large de Marseille, Ann. d. Sci. Nat., 1879, ser. 6, t. viii. pp. 40-45, pi. xviii.
2 On two species of Echinodermata new to the Fauna of Great Britain, Ann. and Mag. Nat. Hist., 1857, ser. 2,
vol. xix. pp. 32, 33, pi. vii. fig. 1.
3 Proc. Roy. Soc. Edin., 1872, vol. vii. p. 765.
REPORT ON THE CR1NOIDEA. 161
dredged in the Mediterranean off the coast of Africa. I do not feel satisfied that this
is identical with Antedon rosaceus of the coast of Britain, though the two specific
names are usually regarded as synonyms. There is a great difference between them in
habit, a difference which it is difficult to define." Sir "Wyville was unfortunately
prevented by the state of his health from accompanying the " Porcupine " in this
cruise, and only made a cursory examination of the Coniatulas subsequently. Had
he been able to work them out at leisure, I cannot but think that the rediscovery of
Antedon phalangium would have taken place five years earlier than it did. Professor
Marion, to whom it was eventually due, has been kind enough to provide me with
some of his specimens from Marseilles, and I have not the smallest hesitation in
identifying them with the Antedon celtica of the Ross-shire coast, and also with the
Antedon which was found by the " Porcupine " in such abundance in the Bay of
Benzert, and on the Skerki Bank, off the coast of Tunis. During this same cruise of
1870 the type was also obtained by the "Porcupine" in 220 fathoms off Cape
Mondego on the Portugese coast, and likewise in 45 fathoms off Cape Sagres. Several
specimens were obtained too in about 80 fathoms a little to the south of Carthagena.
The " Dacia " dredged it in abundance on the Seine Bank in 88 fathoms, and the
" Talisman " took it off Cadiz. It inhabits somewhat deeper water than Antedon
rosacea, both in the Mediterranean and in the Atlantic, and this accounts to some
extent for its having so long escaped notice.
Besides making a careful comparison of the external characters in numerous Scotch
and Mediterranean specimens, I have also compared the dissected calyces of examples from
both localities. Were they fossils, and the only material at my disposal, I should un-
hesitatingly refer them to the same species. In each case there is the same great
variation in the shape of the centro-dorsal, which may be either a thick disk, columnar,
hemispherical, or conical. But whatever its shape, the functional cirrus-sockets are
limited to two or three irregular rows around the equator, all the inferior portions of
the piece having the sockets more or less completely obliterated (PI. XXVIII. figs. 1, 2).
The appearances presented by the first radials are nearly or quite identical in examples
from the two localities. The figures which I have given of the Scotch Antedon celtica
would do equally well as illustrations of the same parts in Antedon phalangium;
though in some of the Scotch forms the transverse ridges separating the muscle-fossse
from those below them are less oblique than in the calyces which I have figured,1 and
I have not found this to be the case in any examples of the Mediterranean variety that
I have examined.
The chief difference to be noticed between the Scotch and the Mediterranean
varieties of this species is in the characters of the cirri. The maximum number of
joints in both forms is from forty -five to fifty ; but while in the Mediterranean
1 Trans. Linn. Soc. Lond. (Zool.), 1879, ser. 2, vol. ii., pL iv. figs. 1-8.
(ZOOL. CHALL. EXP. PART LX. 1887.) 000 21
1G2 THE VOYAGE OF H.M.S. CHALLENGER.
examples all the joints except those just at the base are much wider thau long
(PI. XXVIII. figs. 1, 2), the later cirrus-joints of the Scotch variety are relatively shorter
and thicker, so that in the extreme forms they are actually wider than long (PI. XXVII.
figs. 23, 24).
The longest cirrus which I have found in specimens from the Tunis coast
measures 52 mm., and contains forty-seven joints, while in one from the Minch there
are fifty-one joints, though the length is only 47 mm., and in the most extreme forms
from this locality there are forty-eight joints in a length of but 35 mm. I have pointed
out elsewhere l that these two types of cirri, apparently so different, are linked together
by a complete series of intermediate gradations, in all of which there is a great amount
of variation in the characters of the terminal claw and of its opposing spine. Evidently,
therefore, the only character of the cirri of Antedon phalangium on which we can at
all rely as having a sufficient degree of stability for specific distinction is the great
number of their component joints. This is common to Antedon 2^'olixa and to
Antedon hystrix, and it serves as a convenient means of separating these three
species from the large group which embraces Antedon rosacea, Antedon tenella, and
similar forms with shorter and fewer jointed cirri.
Marion2 has described the second radial of Antedon phalangium as " profondement
enchancree pour recevoir l'axillaire, qui est tres-grande." The four figures which he
gives of the calyx certainly bear out his statements. But I have seen individuals
from Marseilles that I owe to his kindness, and others from the Tunis coast, which have
much less quadrate axillaries, and therefore also less deeply incised second radials.
Other examples from the Tunis coast correspond to Marion's figures ; but in most of
these and in all the Scotch specimens the second radials are oblong in their general
outline, and but little incised, while the axillaries are subtriangular, subquadrate, or
more usually pentagonal, with their bases curving slightly outwards. Not unfrequently
there are forward projecting lateral processes on the second radials which are much more
marked in some individuals than in others. The axillaries may have slight processes of
the same kind, and they are continued on to the first brachials as a sort of flattening of
their outer sides, thus affording an approach to the condition of the BasicuTva-groxvp.
The two first brachials, just like the two outer radials, vary considerably in their
shape and mutual relations. Thus, for example, in the four individuals figured by
Muiion the two joints borne by the axillary are well separated from one another above
its distal angle, and the second brachials have an irregularly quadrate shape. But in
the specimens dredged by the " Dacia " on the Seine Bank the second brachials are
almost triangular in outline, and the two joints below them are closely united above
1 On the Variations of the Form of the Cirri in certain Comatuloe, Trans. Linn. Soc. Land. (Zool.), 1886, ser. 2,
vol. ii. pp. 475-480, pi. lvii.
2 Ann. d. Sci. Nat. (Zool.), 1879, sir. 6, t. viii. p. 43, pi. xviii. fig. 11.
REPORT ON THE CRINOIDEA. 163
the axillary. This is also the case in some of the Tunis specimens ; but in others the
first brachials are quite free laterally, as in those figured by Marion, and in the
" Porcupine " examples from off Cape Mondego and from the Minch. A considerable
amount of local variation in the shape of the outer arm-joints is also to be noticed.
The triangular joints beyond the second syzygy are distinctly longer in the
Mediterranean forms than in those from the Seine Bank and from the Minch ; and
this is still more marked in the outer part of the arm where the joints become quadrate
(PL XXVII. figs. 28, 29).
The length of the lower pinnules is usually somewhat greater in the Mediterranean
variety than in the Scotch one. Marion gives an average length of 12 to 17 mm. for
the four lowest pinnules (on the second to fifth brachials). They reach 1 5 mm. in the
largest examples from the Seine Bank. I have never, however, seen any Scotch
specimens in which either of the four lower pinnules was more than 13 mm. long. The
next two pairs are usually distinctly smaller, though I have occasionally found the
pinnule on the sixth brachial to be almost as large as that on the preceding joint.
Beyond the first four pinnules their component joints diminish considerably in number,
but the basal ones increase in length, so that the inequality in the length of the pinnules
is less marked than it would otherwise be. On the whole the disparity between the first
four pinnules and their successors is somewhat greater in the Scotch specimens.
The peculiar modification of the two basal joints of the outer pinnules, which reaches
its maximum in Antedon eschrichti, is also, as might be expected, more distinct in the
northern than in the southern variety of Antedon phalangium. The shape of the first
joint is much the same in both forms ; but as a rule the second is relatively narrower
in the Mediterranean variety (PL XXVII. fig. 27), so that the distinction between it
and its successors is less marked than in the northern form (PL XXVII. fig. 26).
There is, however, a considerable amount of variation in this respect, even in individual
arms.
The "Porcupine" dredging in 220 fathoms off Cape Mondego in 1S70 yielded a
single larva of this species, in which the radial plates have not yet made their appearance
(PL XIV fig. 1). It does not differ in any important respect from the corresponding
stage in the larva of Antedon rosacea. But the stem is a trifle less robust, as compared
with the size of the head, and the five sacculi which are so constant in the rosacea-laxva,
one between the bases of every two oral plates, do not appear to be present at all.
Fig. 3 on PL XXVIII. represents the youngest condition of the free stage of Antedon
phalangium that I have met with. It was obtained on the Seine Bank, by the " Dacia,"
together with others somewhat older. The first radials are more exposed than in the
adult, and the pinnule of the third brachial is much smaller than that on the preceding
joint, the next two pinnules being smaller still, while some of the following brachials are
altogether without pinnules. A considerable number of cirri are developed, however,
164 THE VOYAGE OF H.M.S. CHALLENGER.
and they already show very distinct indications of the peculiarities which are charac-
teristic of the type.
Ludwig * has made some observations as to the comparative distribution of calcareous
plates and granules in the Neapolitan examples of Antedon pkdlangium and Antedon
rosacea. He finds that while calcareous deposits are more or less developed on the disk
of Antedon rosacea, that of Antedon pkdlangium is almost or entirely naked, which I
find to be the case also both in Tunis and in Marseilles specimens, and in the Atlantic
ones as well. But in the British variety of Antedon rosacea the perisome of the disk may
be either naked or bear scattered tubercles containing groups of radiating calcareous
spicules, and the perisomatic skeleton of the larval arms and pinnules disappears in
later life. I have found no trace of it in any specimens of Antedon rosacea, even in
those from the north of Scotland ; though examples of Antedon phalangium from this
neighbourhood have delicate plates on the pinnule-ambulacra. Like Ludwig, however,
I have found small rods in the marginal leaflets on the pinnules of Antedon rosacea from
Naples, and also in a Marseilles specimen ; while in the Tunis variety of Antedon
phalangium I find delicate perforated plates, the rudiments of the covering plates
which are so largely developed in many tropical Comatulse. They are less distinct in
the specimens dredged by the " Dacia," and in those from Marseilles they are reduced
to small Y-shaped rods, but little better developed than those of Antedon rosacea. In
some individuals of the Scotch variety the pinnule-ambulacra are in this condition, while
in others they have delicate, but still very definite plates, as in the examples from the
Tunis coast. In those from 220 fathoms off Cape Mondego, however, these plates reach
a considerable relative size and have a closer network of limestone rods. There are
about three to each pinnule -joint, and they alternate pretty regularly with the sacculi,
just as the side plates do in Antedon acoela and in other forms from the Eastern seas.
They are much better defined than the side plates of many tropical species, but they do
not support any covering plates above them. On the other hand, they are altogether
different from the large and oval covering plates of Rhizocrinus, Bathycrinus, and
Hyocrinus, which are unsupported by side plates, and rest directly on the pinnule-joints.
Their occurrence in Antedon phalangium in the East Atlantic is the more interesting, as
the locality is within a few miles of that which yielded Pentacrinus wyville-thomsoni
and Antedon lusitanica, both with plated ambulacra; while the latter is the only
European Antedon with both side plates and covering plates on the pinnules.
On the whole I am disposed to confirm Ludwig's observations respecting the greater
length of the anal tube in Antedon phalangium than in Antedon rosacea; but the
difference is not great, and is of no value as a specific character. The only two
species which have any great amount of resemblance to Antedon phalangium are
Antedon hystrix and Antedon prolixa. But it differs from both of them in the greater
1 IJber enige seltenere Echinodernien des Mittelmeeres, Mitth. d. zool. Stat. Neapel, 1880, Bd. ii. p. 54.
REPORT ON THE CRINOIDEA. 165
length of the second pair of pinnules, which are nearly equal to the first ; while the back-
ward projection of the axillaries into the second radials is more marked in both these
types than in Antedon phalangium, more especially in Antedon hystrix.
Antedon phalangium serves as host to two species of Myzostoma, viz., Myzostoma
pulvinar and Myzostoma alatum. The former is only known from the Minch ; but the
latter occurs both there and in the Atlantic, off Cape Mondego, and also in the
Mediterranean where it was dredged off Carthagena. It is curious, however, that no
Myzostoma was obtained with the great number of individuals which were met with by
the "Porcupine" on the Tunis coast.
2. Antedon hystrix, n. sp. (PI. XXVII. figs. 21, 22 ; PI. XXVIII. figs. 4, 5).
Specific formula. — A. t-.
1884. Antedon hystrix, P. H. Carpenter, Proc. Roy. Soc. Edin., 1884, vol. xii. p. 365.
Centro-dorsal hemispherical or subcorneal, bearing fifty or sixty dimorphic cirri. The
longest, which are round the margin, reach nearly 50 mm., and consist of about forty-
five smooth joints, most of which are longer than wide. The smaller cirri nearer the
dorsal pole have only about twenty-five joints, which are relatively shorter and overlap
slightly.
First radials partially visible at the angles of the calyx ; the second comparatively
short and often not visible at all in the middle line of the ray, owing to their being very
deeply incised to receive the strong backward projections of the axillaries. These are
quadrate in form, with their sides curved, especially the anterior pair, and they are
distinctly longer than wide, sometimes seeming to overlap the centro-dorsal ; but much
less than half the length is in front of the line joining their lateral angles. The first
brachials have long outer sides and very short inner ones, but like the second radials are
almost invisible in the middle line of the arm, owing to the very strong backward
projections of the irregularly triangular second brachials, which nearly reach the axillaries.
Both on these joints and on the rudely oblong third brachials, which are much wider than
long, the pinnule-socket is placed nearer the dorsal surface than usual. The next
following joints are short and quadrate, with curved proximal and distal edges ; and the
pinnule is on the shorter side, the longer being marked by a backward projection.
Syzygies in the third and eighth brachials, and afterwards at intervals of three or four
joints. Ten arms ; the lower brachials triangular and slightly wider than long, slowly
becoming quadrate, and somewhat elongated towards the arm-ends.
The first pair of pinnules (on 2nd and 3rd br.) are much longer and stouter than
the next pair. They reach nearly 15 mm. and consist of some thirty smooth joints, the
first six of which are short and nearly square. The second pair have but eighteen or
166 THE VOYAGE OP H.M.S. CHALLENGER.
twenty slender joints, and are only about 6 mm. long. The following pinnules increase
gradually, both in length and in stoutness, reaching 15 mm. in the outer parts of the
arms. The two basal joints are flattened and trapezoidal, with incurved edges. Ovaries
long and fusiform, extending over the greater part of the length of the lower pinnules.
Disk naked or nearly so ; sacculi abundant.
Colour in spirit, — light reddish-brown.
Diameter of centro-dorsal 5 mm.; spread about 17 cm.
Localities. — H.M.S. "Porcupine," 1869. Cold area? Two specimens, with Myzo-
stoma cirriferum.
H.M.S. "Triton," 1882. Station 4; Lit. 60° 22' 40" N. and 60° 31' 15" N., long.
8° 21' W., and 8° 14' W. 327 to 430 fathoms; stones, mud; bottom temperature, 31°-5
to 30o,0 F. One specimen.
Remarks. — The two individuals of this remarkable type which were obtained by the
"Porcupine" in 1869, seem to have been regarded as belonging to Antedon eschrichti,
and so remained unnoticed, while their locality was not recorded. They did not come
into my hands till 1883, when I also received the Comatulse dredged by the "Triton"
and " Knight Errant." The former collection included another example of the same type
from the " cold area," and as its nearest ally is the Arctic species Antedon prolixa, the
" Porcupine " specimens may be safely referred to one of the "cold area" stations. All
three individuals agree very closely in their general features and especially in the curious
dimorphism of the cirri, which is almost as marked as in Antedon phalangium. A good
example of the mature smooth and long-jointed cirrus is shown in PL XXVIII. fig. 4,
and some of its younger stages are seen in fig. 5 round the upper edge of the centro-
dorsal. But the cirri attached nearer the dorsal pole are somewhat different in appearance
(PL XXVII. fig. 21). Many of them are comparatively short, with only about twenty-
five joints, which are as wide as or wider than long and have slightly expanded ends so as
to overlap their successors. This is especially marked on the dorsal side, which is produced
into a sharp forward projecting spine. These characters seem to disappear, however, as
the cirri increase in age and develop additional joints (PL XXVII. fig. 22), so that
eventually they are not very markedly different from the smooth and long-jointed cirri of
the other type.
Antedon hystrix has a considerable amount of resemblance to Antedon prolixa,
which was obtained in Eobeson Channel by Nares's Arctic Expedition in 1875, and was
subsequently well described by Sladen.1 Before giving a new name to the " Porcupine"
and "Triton" specimens, I compared them carefully with Sladen's type, and came to the
conclusion that they should be regarded as distinct. Subsecment research has justified
this view. I have elsewhere described two specimens of Antedon prolixa which were
1 Op. tit., p. T7, pi. vi. figs. 7-10.
REPORT ON THE CRINOIDEA. 1G7
dredged by the " Varna " in the Kara Sea ; and by the kindness of Mr. F. Nansen,
Conservator of the Bergen Museum, I have been able to examine half-a-dozen individuals
of the same type which was met with in abundance by the Norwegian North Atlantic
Expedition near Spitzbergen. These last come nearest to Antedon hystrix in their
general robustness, the more northern forms of the type being generally smaller, except
as regards the cirri. These seem to be a little less smooth and to reach a greater length
in Antedon prolixa than in actually larger individuals of Antedon hystrix. Thus, for
example, an imperfect cirrus of Antedon prolixa, with the extremity missing, which was
measured by Sladen, reaches 58 mm.; while I have not found any cirrus exceeding 50 mm.
in Antedon hystrix. This, however, is a point of minor importance. The great difference
between the two types lies in the characters of the two outer radials and of the two lowest
brachials. The second radials of Antedon prolixa are but little more incised for the
axillaries than those of Antedon phalangium, with which species Sladen has well compared
it, though the two differ altogether in the proportions of the second pair of pinnules ;
while the axillaries of Antedon prolixa are very regularly quadrate and as wide as or wider
than long, the line joining their lateral angles dividing them into two nearly equal parts.
In Antedon hystrix, however, they are longer than wide, and project so deeply backwards
into the second radials that they sometimes seem to overlap the centro-dorsal
(PL XXVIII. figs. 4, 5). The second radials are therefore almost invisible in the middle
line of the ray, though when seen from the side they appear to have a considerable relative
length and to form a projecting tubercle together with the axillaries, as is well shown in
PI. XXVIII. fig. 4. The shape of the axillaries therefore is not " very regularly quadri-
form" as described by Sladen in Antedon prolixa, but more pear-shaped, with much less
than half the length of the plate in front of the hue joining the lateral angles, a condition
exactly the reverse of that which occurs in many forms of Antedon eschrichti. In like
manner the second brachials of Antedon hystrix are relatively larger, and project further
backwards into the first than in Antedon prolixa, so that there is a more distinct tubercle
on the line of junction.
The " Triton " specimen of Antedon hystrix was dredged in about lat. 60° 30' N., and
Antedon prolixa was obtained by the Norwegian North Atlantic Expedition in lat. 76° X.,
near Spitzbergen. It is of course possible that future explorations in the intervening area
of the Atlantic may discover a series of forms intermediate between those which now
appear distinct ; and I should not be very greatly surprised if this should turn out to be
the case. AVere they really identical, Antedon prolixa would present just the opposite
condition to Antedon eschrichti, its northern variety being less robust than that found in
lower latitudes. The small examples of Antedon prolixa from the Kara Sea (lat. 71° N.)
are, however, very different from Antedon hystrix. The "Triton" specimen of the
latter type presents a very curious malformation, which is shown in PI. XXVIII. fig. 5.
The two second brachials of one ray jointly support a single arm, so that there are only
168 THE VOYAGE OF H.M.S. CHALLENGER.
nine arms, not ten. The two third, brachials are replaced by a single syzygial joint,
which has the shape of an axillary reversed, i.e., with the angle downwards, and it bears
a pinnule on the left side. The following arm-joints are of the usual character. A
somewhat similar monstrosity was noticed in Part I. (p. 347) as occurring in Metacrinus
angulatus, and Levinsen 1 has figured one of much the same in kind in Antedon
eschrichti.
Two larvae which were dredged by the "Porcupine" in the "cold area" come to be
referred to Antedon hystrix by a sort of process of exclusion, as I cannot identify them
with the Pentacrinoids either of Antedon tenella or of Antedon eschrichti, with which
latter species Antedon quadrata is very closely allied.
No. 1 (PL XIV. fig. 2). In this larva there is no trace of cirri, the anal plate
separates two of the radials, and the arms are just beginning to sprout from the radial
axillaries. There are five discoidal joints at the top of the broken stem, which is much
more robust than that of the corresponding stage of Antedon rosacea, while the head,
which exceeds 1 mm. in length, is nearly twice as big as that of the rosacea-larva. The
orals, which rest directly on the radials, recall those of Hyocrinus, having a deep median
groove, only more marked than in that type, with the lateral edges folded over somewhat
strongly. This character is less marked in the rosacea-larva, and the orals of Antedon
tenella in the first and second stages of the Pentacrinoid were described by Sars2 as
convex. In its general appearance the Pentacrinoid now under consideration comes
between the second and third stages of the larvae described by Sars, but is of larger size
than both. Sars gives the measurements of the head as 0"5 and 0'75 mm. respectively, the
larger individual having six brachials above the axillaries ; but in the " Porcupine " larva
there are only two short brachials, and the head reaches 11 mm. It resembles the larva
of Antedon tenella in the great height of the basals, but differs from it altogether in the
unusual shortness and width of the radials, especially the first. These plates are
relatively wider than the corresponding plates in an older stage of Antedon tenella,
whereas they would be relatively longer did the larva belong to this species. A similar
difference between the radials of two other larvae in almost the same stage of development
will appear on comparison of figs. 8 and 9 on PI. XIV. I think it not improbable that
this "Porcupine" larva may be a younger stage of that represented in PI. XIV. fig. 3,
which was also dredged in the cold area during the cruise of 1869.
No. 2 (PI. XIV. fig. 3). The stem, which is broken some 20 mm. from the calyx,
forms an attachment to a hydroid-tube at about its thirtieth joint, and is continued
downwards half-a-dozen joints further. There are six discoidal joints below the rudi-
mentary centro-dorsal, which bears the sockets of five short cirri. Only one of them
remains, however, reaching up to the top of the basals, which make up about half the
1 Op. cit., p. 35, Tab. xxxv. fig. V. •
2 Memoires pour servir a la connaissance des Crinoides vivants, Ckristiania, 1868, p. 48.
REPORT ON THE CRINOIDEA. 169
height of the cup. The second radials and axillaries are well developed, as are also the
arms, which are unfortunately broken at about the tenth joint or earlier. But even
under these circumstances the head has a length of 4 mm. A slightly bifid plate, having
a somewhat worn appearance, stands up in one of the interradii of the disk. It may be
one of the orals, or, as I am more inclined to think, the anal plate. For I cannot make
out anything corresponding to it in the other interradii, which are, however, but
imperfectly visible. A striking feature of this very robust larva, and one in which it
resembles Antedon tenella rather than Antedon rosacea, is the large development of the
arms before the appearance of the cirri. The radials and brachials are larger than those
of a recently detached individual of Antedon rosacea. This is also the case in the Penta-
crinoid of Antedoneschrichti and in that of Antedon midtispina, from near Ascension
(PI. XIV. fig. 7), which has a very robust appearance, like the larva now under con-
sideration. The latter can hardly be a younger stage of the Pentacrinoid of Antedon
eschrichti than that figured by Levinsen.1 Their difference in relative age is not great,
while they are very unlike in many respects. The " Porcupine " larva has high basals
and relatively wide first radials, with short, wide, and well-formed axillaries (PI. XIV.
fig. 3) ; while in the Pentacrinoid of Antedon eschrichti the basals are low, the radials
relatively high, and the axillaries rhombic, about as wide as long. It would appear for
the same reason that this larva cannot belong to Antedon quadrata, which is most
closely allied to, if not identical with, Antedon eschrichti, while it is clearly not that of
Antedon tenella, and the only other Comatula known to occur in the cold area is Antedon
hystrix.
The brachial ambulacra of this larva are protected by relatively large plates, not
unlike those which occur in some varieties of Antedon phalangium, but the armature of
the ambulacra in the mature Antedon hystrix consists of quite simple rods of limestone.
This difference may perhaps be explained by the fact that an absorption of the perisomatic
skeleton of the Pentacrinoid seems to take place in some forms of Antedon rosacea, as
noticed by Dr. Carpenter.2
3. Antedon tenella, Retzius, sp. (PL XIV. fig. 4 ; PI. XXXI. figs. 1-4).
€
Specific formula — A. y.
1783. Asterias tenella, Retzius, K. Svensk. Vetensk. Akad. Handl, 1783, t. iv. p. 241.
1788. Asterias tenella, Linn., Systema Naturae, ed. xiii., cura, J. F. Gmelin, Lipsiaj, 1788, t. i.
pars vi. p. 3166,
1805. Asterias tenella, Retzius, Dissertatio, sistens Species Cognitas Asteriarum, Lundae, 1805,
p. 33.
1825. Alectro dentata, Say, Journ. Acad. Nat. Sci. Philad., 1825, vol. v. p. 153.
1835. Comatula mediterranea (?), Sars, Beskriv. og Jagtagels, Bergen, 1835, p. 40, pi. 8, fig. 19 a-j.
1 Loc. cit, tab. xxxv. fig. 8. 2 Phil. Trans., 1866, p. 741.
(ZOOL. CHALL. EXP. — PART LX. 1887.) OOO 22
170 THE VOYAGE OF H.M.S. CHALLENGER.
1844. Alecto Sarsii, Diiben and Koren, K. Svensk. Vetensk. Akad. Hand!., 1844 [1846],
p. 231, t. vi. fig. 2.
1819. Comatula (Alecto) Sarsii, Mull., Abhandl. d. k. Akad. d. Wiss. Berlin [1847], 1849, p. 254.
1857. Alecto Sarsii, Liitken, Vid. Meddel. nat. Foren Kjobenhavn, 1857, p. 107.
1860. Comatula Sarsii, Alder, Ann. and Mag. Nat. Hist., 1860, ser. 3, vol. v. p. 74.
1861. Alecto Sarsii, Sars, Oversigt af Norges Eehinodermer, Christiania, 1861, p. 1.
1862. Comatula Sarsii, Dujardin and Hupe, Hist. Nat. des Zoophytes, Echinodermes, Paris,
1862, p. 199.
1865. Antedon Sarsii, Norman, Ann. and Mag. Nat. Hist., 1865, ser. 3, vol. xv. p. 103.
1866. Antedon (Alecto) dentata, Verrill, Proc. Boston Soe. Nat. Hist., 1866, vol. x. p. 339.
1868. Antedon Sarsii, Sars, Memoires pour servir a, la Connaissance des Crinoides vivants,
Christiania, 1868, p. 47, tab. v., vi.
1872. Antedon sarsii, Wyville Thomson, Proc. Roy. Soc. Edin., 1872, vol. vii. p. 765.
1874. Antedon Sarsii, Verrill, Amer. Journ. Sci. and Arts, 1874, vol. vii. p. 500.
1S79. Antedon Sarsii, P. H. Carpenter, Trans. Linn. Soc. Lond. (ZooL), 1879, ser. 2, vol. ii.
p. 29.
1880. Comatula Sarsii (?), A. Agassiz, Bull. Mus. Comp. ZooL, 1880, vol. vi., No. 8, p. 150.
1880. Antedon Sarsii, Verrill, Amer. Journ. Sci. and Arts, 1880, vol. xx. p. 401.
1880. Antedon Sarsii, d'Urban, Ann. and Mag. Nat. Hist., 1880, ser. 5, vol. vi. p. 381.
1881. Antedon Sarsii, P. H. Carpenter, Bull. Mus. Comp. ZooL, 1881, vol. ix., No. 4, p. 5.
1882. Antedon Sarsii, Verrill, Amer. Journ. Sci. and Arts, 1882, vol. xxiii. p. 135.
1882. Antedon dentatum, Verrill, Ibid., p. 222.
1882. Antedon sarsi, BeU, Proc. ZooL Soc. Lond., 1882, p. 534.
1883. Antedon dentata, P. H. Carpenter, Proc. ZooL Soc. Lond., 1882 [1883], p. 746.
1884. Antedon dentata, P. H. Carpenter, Proc. Roy. Soc. Edin., 1884, vol. xii. p. 362.
1884. Antedon dentata, Verrill, Ann. Rep. Commissioner Fish and Fisheries for 1882, Washing-
ton, 1884, p. 661.
1886. Antedon dentata, P. H. Carpenter, Bijdragen tot de Dierkunde, 13 Afdeling, vi. p. 9.
N.B. — Not Antedon Sarsii, von Marenzeller (1877),
nor Antedon dentata, Fischer (1886).
Centro-dorsal hemispherical or conical, bearing a great number of cirri, seventy or
eighty in large specimens. They have fifteen to nearly thirty joints, most of which are
longer than wide, the lower ones greatly so, and somewhat dice-box-shaped ; the later
joints with sharp spines which project forwards over the bases of their successors.
First radials just visible ; the second short, deeply incised by the rhombic axillaries,
which are as wide, or usually wider than long, and extend laterally beyond them.
Ten arms ; the first brachials barely meeting above the sharp distal angles of the
axillaries, and deeply incised by the quadrate second brachials. From the first to the
second syzygy the joints have backward projections on alternate sides, and the next
following joints are triangular, as wide or wider than long, gradually becoming obliquely
quadrate. Syzygies in the third, eighth, and twelfth brachials, and then at intervals of
two joints.
The first pair of pinnules (on second and third brachials) slender and flagellate, reaching
15 mm. or more in length, and consisting of nearly forty joints, most of which are longer
than wide. The next pair usually only one-third their length, with about ten joints,
EEPOET ON THE CRINOIDEA. 171
the lowest of which are stouter than in the first pair. The following pinnules similar
in character and of gradually increasing length, with fusiform genital glands. The
basal joints of the distal pinnules are quite short, with their apposed edges incurved, and
the following joints are greatly elongated. Disk naked ; the brachial ambulacra some-
times provided with delicate calcareous rods, which alternate with the numerous sacculi.
Colour in spirit, — white.
Disk reaching 7 mm.; spread may be 8 cm.
Localities-. — H.M.S. "Porcupine," 1869, Station 51 ; lat. 60° 6' N., long. 8° 14' W.;
440 fathoms ; bottom temperature, 42° F. One specimen.
Station 54 ; lat. 59° 56' N., long. 6° 27' W. ; 363 fathoms ; bottom temperature
31°"4 F. One specimen.
Station 55; lat. 60° 4' N., long. 6° 19' W. ; 605 fathoms; bottom temperature,
290,8 F. Two specimens.
Station 74; lat. 60° 39' N, long. 3° 9' W. ; 203 fathoms; bottom temperature,
47°'6 F. Three specimens.
Cruise of 1870, Station 17a; lat 39° 39' N., long. 9° 39' W.; 740 fathoms; bottom
temperature, 49° "3 F. One specimen.
H.M.S. "Triton," 1882; Station 2; lat, 59° 37' 30" N., long. 6° 19' W. ; 530
fathoms ; mud ; bottom temperature, 46c,2 F. Five mutilated specimens.
Station 5; lat. 60° 11' 45" N. and 60° 20' 15" N., long. 8° 15' W. and 8° 8' W.;
433 to 285 fathoms ; hard ground, stones ; bottom temperature, 43°"5 to 40°*8 F. Two
specimens, with Myzostoma carpenteri.
Other Localities. — The Shetlands ; Scandinavia; Kara Sea; Barents Sea; West
Atlantic, off the American coast from Nova Scotia to New Jersey.
History. — This species was separated by Eetzius1 in 1783 from the Asterias
pectinata of Linnaeus, the type of which was an Actinometra from the Indian Ocean,
but Linnaeus also referred to it the AeK<xKvr]iJLo<; rosacea and Acca»c^/Ao? barbata of
Linck. These seem to be the British and Mediterranean varieties respectively of the
somewhat protean type which is now known as Antedon rosacea. In fact Linnaeus's
description of Asterias pectinata would apply equally well to almost every ten-armed
Comatula ; and it was remarked by Eetzius that " the definition of the species is such
that it includes two species, namely, pectinata and tenella." He described the latter
form as being more debcate than Asterias pectinata, and as having the " bases brachiorum
duplicatorum multo longiores." 2 This is a distinction of almost generic value between
Antedon and Actinometra, in which latter genus the relative length of the radials is quite
small ; and Retzius pointed out further differences in the shape of the arm -joints between
1 Anmarkningar vid Asteri:e Genus, K. Svensk. Vetensh. Akad. Handl., 1783, t. iv. p. 241.
" Dissertatio, sistens Species Cognitas Asteriaruin, Lundie, 1805, p. 33.
172 THE VOYAGE OF H.M.S. CHALLENGER.
the Linnean type of Asterias pectinata and the new form which he proposed to call
Asterias tenella. He described the habitat of the latter as " St Croix" ; and this is given as
the island of " Santa Cruz " in Gmelin's edition of the Systema Naturae, where Asterias
tenella is added to Asterias pectinata and Asterias multiradiata of the earlier editions.
Lamarck took but little notice of these three species when he established the genus
Comatula. Asterias pectinata was not noticed by him at all, though he proposed a new
name, Comatula mediterranea, for Linck's Decacnemos rosacea, which had been included
by Linnaeus in Asterias pectinata ; while he referred Asterias tenella with a % to his
new species Comatula brachiolata,1 which we now know to be an Actinometra closely
allied to the type of Asterias pectinata. Lamarck, however, was the only post-Linnean
zoologist who recognised that Asterias tenella was a Comatulid and not a Star-fish, a fact
which would seem sufficiently obvious when we remember that Retzius had pointed out
how it had been hitherto confused with Asterias pectinata. Goldfuss, indeed, gave the
name Comatula tenella to a fossil from Solenhofen, which was one of the four species
subsequently placed by Agassiz in his new genus Saccocoma.
Asterias tenella seems to have entirely escaped the notice of Johannes Muller when
he examined the Eetzian collection at Lund in 1841, and it has consequently altogether
dropped out of the literature. The original of the type, however, is still extant, together
with the examples of Asterias pectinata and Asterias multiradiata from the Indian Seas
which are the types of these two species respectively. I have been privileged to examine
all three, and find Asterias tenella to be very different from Asterias pectinata, for it is
identical with the well-known Scandinavian species which was described in 1844 by
Diiben and Koren as Alecto sarsii.' This specific name has been in use for nearly forty
years, and the range of the type was extended to lat. 70° N. by the " Willem Barents";
whde the " Porcupine " had previously dredged it at various localities in the Fseroe
Channel and also at 740 fathoms as far south in the Atlantic as lat. 39° N.
In the year 1880, however, the same species was obtained several times off the coast
of New England by the explorations of the United States Coast Survey and Fish Commis-
sion. Two years later Mr. Verrill 3 recognised that Alectro dentata, which was described
by Say in 1S25 from a specimen found at Great Egg Harbour, New Jersey, is identical
with Antedon sarsii, which occurs in abundance at various depths off the American coast
from New Jersey to Nova Scotia. A restoration of Say's specific name thus became
inevitable, and the association of the type with the familiar name of a deservedly
honoured Norwegian naturalist was no longer possible. Now, however, it appears that
forty-two years before the publication of Say's name Retzius had described the same
species from the American coast, and I have much pleasure therefore in restoring his name.
1 Op. cit, p. 535.
2 Ofversigt af Skandinaviens Echinodurmei', K. Svensk. Vetenst Akad. Handl, 1844 (1846), p. 231, t. vi. tig. 2.
3 Notice of the remarkable Marine Fauna occupying the outer hanks off the Southern Coast of New England,
No. 4, Amer. Journ. Sci. and Arts, 1882, vol. xxiii. p. 222.
REPORT ON THE CRINOIDEA. 173
The non-employment of it by Diiben and Koren in 1844 is not difficult to understand,
tor the occurrence of the same specific type on both sides of the Atlantic was not such a
familiar idea then as it is now ; and Midler had taken no notice of Asterias tenella,
Retzius, in his "Neue Beitriige." These were published in 1843 after his visit to Lund,
and contained amended descriptions of Asterias pectinata and Asterias multiradiata,
the other two Comatulas mentioned in Eetzius's dissertation, which he had personally
examined. Midler's omission to notice Asterias tenella seems to have caused its
relegation to the class of species ineertse sedis, from which I am glad to be able to rescue
it. I am in some doubt, however, as to whether the " St Croix " of Retzius can be the
island Santa Cruz, as mentioned by Gmelin. Retzius did not repeat it in his later
dissertation, but simply said " Habitat in oceano Americano." Santa Cruz being a Danish
island, one can readily understand that specimens collected there might come into the
hands of Swedish naturalists ; but on the other hand it is in latitude 18° N., considerably
(about 20°) further south than any locality at which Antedon tenella has been dredged
by American naturalists. I have tried, however, but in vain, to identify the Retzian
type with any Caribbean Antedon, though it has all the characters of the Scandinavian
Antedon sarsii and of Alectro dentata, Say.
While, therefore, I have no doubt as to its identity with these two types, I should
hesitate for the present to quote it as a Caribbean species.
Remarks. — The Scandinavian variety of Antedon tenella, which was described as
Alecto sarsii by Diiben and Koren, is considerably smaller and less robust than individuals
dredged from deep water in various parts of the Atlantic. The cirri do not seem to
have more than about twenty joints, while there may be six or eight more in
individuals from the New England coast, Fseroe Channel, and Kara Sea. The
projecting spines at the distal ends of the cirrus-joints are also less developed
in the Scandinavian examples. In the larger forms from the West Atlantic the distal
ends of the joints in the long oral pinnules are fringed with strong spines, so that they
appear to overlap the bases of their successors, and this character is much less developed
in the European variety. On the other hand, the delicate calcareous rods at the sides of
the ambulacra, which Sars described in the larva,1 are larger in the Norwegian form than
in examples from deeper water ; and they are sometimes entirely absent in the American
variety. The latter is also remarkable for the want of constancy in the proportions of
its second pair of pinnules. In some examples these have only a dozen joints and are
not more than 5 mm. long, but one-third the length of the first pair. But in others
they reach 7 or 8 mm. and have as many as twenty joints, a condition which I have
not noticed in any individuals from the East Atlantic.
Antedon tenella is closely allied to two other cold-water species, Antedon hystrix and
Antedon prolixa, with both of which it has been found associated. It is distinguished,
1 Crino'ides vivauts, p. 51, t;il>. vi. tig. 20.
174 THE VOYAGE OF H.M.S. CHALLENGER.
however, by its smaller size, lesser number of cirrus-joints, and by the different proportions
in the lengths of the first two pairs of pinnules. Like Antedon prolixa, too, it ranges from
shallow water down to 700 fathoms; but it has a much more extensive geographical range.
Fischer 1 has recently come to the conclusion, which I believe to be an erroneous one,
that the specimens which were described by Duncan and Sladen 2 under the name Antedon
prolixa are in reality but " ausgewachsene Exemplare " of Antedon sarsii, auct., i.e,
of Antedon tenella. Two of the four Comatulse which he obtained at Jan Mayen
clearly belong, as he himself states, to Antedon prolixa, as defined by Sladen. The
length of an incomplete arm in the larger one is 120 mm. The cirri, composed of
twenty-eight to forty-three joints, vary in length from 20 to 60 mm. ; the first pinnule
with twenty-eight joints is 14 mm. long, and the second with twelve joints reaches only
4"3 mm.3 It appears from these numbers that Fischer's larger specimen is somewhat
better developed than Sladen's type, with which it is evidently identical. Fischer has
further compared it with the two individuals which were obtained in the Barents Sea by
the " Tegetthoff," and were referred by Dr. E. von Marenzeller to Antedon sarsii.4
Fischer's conclusion is expressed in the following passage, — " Wenn man nun erwagt,
dass mit Ausnahme der durch die Grossenverhaltnisse bedingten Unterschiede (das
grossere der von Marenzeller beschriebenen Exemplare hatte Arme von nur 80 mm. Lange)
namlich die geringere Anzahl von Eanken-Gliedern, — sonst keine Abweichungen zu
verzeichnen sind, so muss man nothwendigerweise zu der Uberzeugung gelangen, was ich
ubrigens an der Hand der spater zu beschreibenden Jugendzustande des Weiteren ausfiihren
werde, dass unter Antedon prolixa, Sladen nur ausgewachsene Exemplare von A. Sarsii,
welche bislang noch nicht erschopfend beschrieben waren, zu verstehen sind."
The largest cirri of the "Tegetthoff" specimens have thirty-three joints and reach
37 mm. long. These dimensions are altogether exceptional for Antedon tenella, in the
Scandinavian examples of which there are usually not more than eighteen or twenty
cirrus-joints, while there may be about twenty-five in those from the Kara Sea and the
Fseroe Channel, and twenty-eight or thirty in the American variety, with a maximum
length of 24 mm.5 I cannot help suspecting therefore that the "Tegetthoff" specimens
may really be the young of Antedon prolixa. Dr. von Marenzeller was kind enough to
send them to me for examination in 1881, and I have hitherto regarded them as he
seems to have done, viz., as abnormal forms of Antedon sarsii (tenella). At the
time I examined them I was unacquainted with Antedon prolixa, and the possibility of
their being the young forms of this type never occurred to me. But in the six
years which have passed since then I have seen many examples both of Antedon
1 Eckinodermen von Jan Mayen, Die Usterreiche Polarstation Jan Mayen, Bd. iii., Wien, 1886, p. 30.
2 Op. cit, p. 77, pi. vi. figs. 7-10.
3 This is accidentally printed as 43 mm. in Fischer's paper.
4 Denkschr. d. k Alcad. d. JViss. Wien, 1877 (1878), Bd. xxxv. p. 381.
5 Figs. 3, 4 on PI. XXXI. represent the average cirri of the Faeroe Channel and American varieties respectively.
REPORT ON THE CRINOIDEA. 175
prolixa and of Antedon tenella in various stages of growth, and the largest cirri that
I have met with in the most robust examples of Antedon tenella from the American
coast are little more than half as long as those of the "Tegetthoff " specimens of the
same size; while the measurements of these last correspond very well with those of the
young Antedon prolixa from the Kara Sea and Spitzbergen.1 Apart from this possi-
bility, however, it appears to me that Fischer attributes to Antedon tenella a much greater
variability in the size of the cirri than is justified by our knowledge of other Comatulae.
He asserts that the small Scandinavian form with cirri 10 mm. long, and consisting of
eighteen or twenty joints, which he thinks he got at Jan Mayen,2 is identical with
Sladen's Antedon prolixa which reaches more than twice its size, and has cirri of forty to
forty-five joints which reach 60 mm. long. The Scandinavian form is sexually mature and
presents all the characters of an adult Comatida ; and if it is only a dwarf variety there
must be some reason for its existence. But Fischer believes it to occur at Jan Mayen,
side by side with the large form which belongs to the type of Antedon prolixa. This fact,
if true, would seem of itself to indicate that the two forms are different ; for if the dwarfing
conditions were in operation at Jan Mayen, the large prolixa-tj^e would not exist there.
Even if we suppose that the "Tegetthoff" specimens really are a local variety of
Antedon tenella with unusually developed cirri, 37 mm. long, of thirty-three joints,3 there
is a great difference between these cirri and those of the mature Antedon prolixa, which
may have forty to forty-five joints, and reach 60 mm. in length ; and the difference is
still greater if we remember the average size of the cirri in the Scandinavian type. If it
1 Since the above remarks were printed, Dr. von Marenzeller has been good enough to send me the two
"Tegetthoff" specimens for re-examination ; and I have no doubt whatever that they are immature forms of Antedon
prolixa, for they agree with this type in all the characters of the cirri, calyx, arms, and pinnules, much better than
with either the American or the European variety of Antedon tenella. They were dredged by the "Tegetthoff" in 1873,
two years before Sladen's types were obtained by the " Alert," and are therefore the earliest discovered examples of the
species.
2 I have left this discussion almost exactly as it was written originally ; but Dr. von Marenzeller's kindness has
recently enabled me to examine the two small specimens from Jan Mayen which Fischer identified with the Scandinavian
Antedon sarsii (tenella) ; and I can state without hesitation that they do not belong to this species. It is no doubt the
case, as remarked by Fischer, that "Diese zwei exemplaren tragen sammtliche von Diiben & Koren und den
spateren Autoren fur Antedon Sarsii angegebenen charakteristischen Merkmale." But Diiben and Koren's description
of the type is over forty years old ; and subsequent writers have added little of importance to it. Fischer does not
seem to have made a direct comparison of his two small specimens from Jan Mayen with actual examples of the
Scandinavian Antedon sarsii, though this would have been by far the most satisfactory way of determining their real
nature. Their cirri are considerably larger than those of a Scandinavian form of equal size which has well-developed
genital glands and all the other characters of maturity. Its first radials are almost entirely concealed, while in Fischer's
specimens a considerable portion of them is visible, very much as in the young Antedon phalangiwm shown on PI. XXVIII.
fig. 3. Similar differences appear in the characters of the lower arm-joints of the two forms. In those from Jan Mayen
the joints are longer than wide, with incompletely developed pinnules ; while in a Scandinavian Antedon sarsii (tenella)
of equal size, these joints are aa wide or wider than long, and present the shape characteristic of the adult individual
(PL XXXI. fig. 1). The difference is so marked that Fischer can hardly have overlooked it if he really did compare the
two types. But the shape of the arm-joints is not a character to which previous authors have paid much attention ; ami
if Fischer simply attempted to identify Antedon sarsii from the published descriptions of it, his reference of the two
small forms from Jan Mayen to this type may be readily understood.
3 See note 1.
176 THE VOYAGE OF H.M.S. CHALLENGER.
be admitted that the perfect cirri of sexually mature individuals of the same species may
vary in size from eighteen to forty -five joints, and in length from 10 to 60 mm., the
characters of the cirri become altogether valueless for systematic purposes, and Bell's
formula? for expressing them briefly are of no use whatever, while the various schemes of
classification of the different specific groups which are given in this Eeport must be to
a large extent rearranged.
Such an extensive range of variation in the characters of the cirri as is demanded
by Fischer's theory is one of which I have no experience whatever. Antedon eschrichti
ranges over nearly forty degrees of latitude ; but in the small Atlantic variety, as in the
large Arctic one, there are over forty joints in the cirri, which reach in the former to little
short of the length that they do in the latter type. The cirri of Antedon phalangium
have the same number of joints (about forty-five) in the Mediterranean as in the Minch,
though the joints are much shorter in the latter locality, so that the total length of the
cirri is reduced. Both these species thus resemble Antedon prolixa in having over forty
cirrus-joints in their southern, as well as in their northern variety ; and they thus afford no
support whatever to Fischer's theory of the great range of variation in Antedon tenella.
If Antedon tenella of Scandinavia, the Arctic Ocean, and the Atlantic is merely a
dwarf or undeveloped variety of Antedon prolixa, young examples of the latter species
should present all the characters of Antedon tenella ; but this is very far from being the
case. Two immature individuals of Antedon prolixa in different stages of growth were
obtained by the " Varna " in the Kara Sea, and others at about the same stage as the
larger of these were kindly sent to me by Mr. F. Nansen from the dredgings of the
Norwegian North Atlantic Expedition near Spitzbergen. I do not think that any one
could possibly refer them to Antedon tenella. I have compared the smaller form with
an absolutely larger example of Antedon tenella which was dredged by the " Willem
Barents " in the Barents Sea at no great distance from the locality of the " Tegetthoff's "
dredgings. The calyx of the latter is altogether more robust than that of the former, and
the first radials are concealed, while both the axillaries and the second brachials have
assumed the shape characteristic of the adult condition. The former (Antedon prolixa),
however, shows its immaturity by the appearance of a considerable portion of the first
radials externally, by the shape of the axillaries and of the second brachials, which is not
that of these respective joints in the adult, and by the greatly elongated arm -joints.
The cirri, on the other hand, are much better developed than those of the more mature
and absolutely larger Antedon tenella, as seen from the comparison of measurements A
in the following table : —
Antedon prolixa. Antedon tenella.
A. B. A. B.
Length of cirri, .... 14 '5 mm. 33-5 mm. 11 mm. 18 mm.
Number of joints, . 25 „ 34 „ 20 „ 26 „
REPORT ON THE CRINOIDEA. 177
Measurements B represent the results of a similar comparison of the yet more robust
Antedon tenella from the West Atlantic with other immature examples of Antedon
prolixa, larger than that already considered, but absolutely smaller than the individual
of Antedon tenella with which they are compared. In each case alike the cirri of
Antedon prolixa with an incompletely developed calyx and arms, are longer and have
more numerous joints than individually larger examples of Antedon tenella. The
difference in absolute size may be judged from the fact that in Antedon prolixa (B)
there is a length of but 4'5 mm. between the pinnule on the seventh and that on the
thirteenth brachial ; while in Antedon tenella this distance measures 7 mm., and yet the
cirri of this latter type are not much more than half the length of those of the young
Antedon 2^'olixa.
The reverse is the case with the pinnules, however ; the first pinnule of Antedon
tenella has nearly forty joints, and reaches 15 mm., while that of the young Antedon
prolixa, 10 mm. long, has but twenty-seven joints, though its cirri are so much better
developed than those of the other species.
It will, I think, be evident from the above-mentioned facts that Antedon tenella and
Antedon prolixa are different species, and not merely different stages of growth of one
and the same type, as supposed by Fischer. The most robust examples of Antedon tenella
with fully-developed arms and pinnules have very distinctly smaller cirri than immature
and absolutely smaller examples of Antedon prolixa. This demonstrates the fallacy of
Fischer's conclusion, which he further endeavours to support by the following passage : —
" Sollten noch irgend welche Zweifel entstehen, so werden dieselben wiederlegt durch die
Thatsache, class ich gleichzeitig mit den bereits beschriebenen Exemplaren zwei Penta-
cfinus-&ta,dieu auf einer Rhynconella aufsitzend fand, die volkommen mit den Beschrei-
bungen tibereinstimmen, die Sars in seinen ' Memoires des crinoides vivants ' gibt, und
auf Taf. v und vi abbildet."
I must confess that I cannot see the force of this reasoning. Fischer found two
examples of Antedon prolixa at Jan Mayen, and two smaller forms, together with two
Pentacrinoids, all four of which he referred to Antedon sarsii (tenella). But I do not
understand at all why the occurrence of these two Pentacrinoids ' should render it so
certain that Antedon tenella is only an immature Antedon prolixa. Precisely the same
reasoning would entitle me to assert that Antedon tenella is only an immature form of
Antedon eschrichti. For the latter species was dredged by the " Porcupine " in the
cold area together with Antedon tenella and its Pentacrinoid ; and in like manner
Antedon rosacea was obtained on the Skerki Bank together with Antedon phalancjium
and its Pentacrinoid. Antedon midtispina with three Pentacrinoids was dredged by
the Challenger at the same Station (No. 344) as Antedon porrecta ; while Antedon
1 I cannot help suspecting that there may be differences between these Pentacrinoids and the larva of Antedon
tenella which have escaped Fischer's notice.
(zoou chall. exp. — part lx. — 1887.) Ooo 23
178 THE VOYAGE OF H.M.S. CHALLENGER.
tuberosa and its Pentacrinoid were found associated at Station 210 with Antedon distincta.
But these facts afford us no grounds for asserting that Antedon rosacea and Antedon
phalangium, Antedon porrecta and Antedon midtispina, Antedon tuberosa and Antedon
distincta are respectively identical types.
Thus then it appears to me that neither of Fischer's reasons for believing in the
specific identity of Antedon tenella and Antedon prolixa has any probative value ; and
I see no necessity, therefore, for reducing Antedon prolixa, Sladen, to the rank of a
synonym as Fischer has done. (See the footnotes on p. 175.)
One example of the Pentacrinoid larva of Antedon tenella was dredged by the " Porcu-
pine " in the Faeroe Channel. It is a trifle more advanced than that represented by Sars
in figs. 9 and 11 on Tab. V. of his classical " Memoires." The arms are longer, with no
pinnule below the eighth joint. There is, however, but one cirrus, which seems to be the
only one as yet developed, though it is of considerable size, reaching up to the level of
the radial axillaries on the opposite side of the larva to that shown in the figure (PI. XIY.
fig. 4). The stem is attached by a slight calcareous expansion at about its thirty-fifth
joint to one of the rays of a Rhabdammina abyssorum ; and it then passes on to form
two other spreading attachments, with radicular branches sprouting from them over a
portion of a tubular Hydroid.
Two of the " Triton " specimens of Antedon tenella from the Fseroe Channel were
infested by Myzostoma carpenteri, the only species of Myzostoma yet found in association
with this Comatulid.
4. Antedon exigua, n. sp. (PI. XXXII. figs. 1-4).
Specific formula — A.-r.
Centro-dorsal hemispherical, almost covered by some fifty cirri of about twenty joints.
The lowest of these are much longer than wide, but the distal joints are short and
compressed.
First radials nearly or quite invisible ; the second short, almost concealed in the
middle line by the blunt hinder angles of the axillaries. These are broadly rhombic,
with a sharp clavicular process, and extend laterally beyond the second radials. The
surfaces of both joints fall away laterally from their medio-dorsal line. Ten arms ; the
first brachials with very long outer sides but a short centre and inner sides, the proximal
ends of which just meet their fellows above the clavicular. They are raised in the median
line to meet the sharp hinder angles of the large quadrate second brachials. Arm-joints
oblong till the second syzygy, with more or less distinct, alternating, backward processes ;
the following joints subtriangular, gradually becoming obliquely quadrate. Syzygies in
the third, eighth, and twelfth brachials, and then at intervals of two joints.
REPORT ON THE CRINOIDEA. 179
The second brachials bear greatly elongated pinnules of thirty or more cylindrical joints.
A similar one on the third brachial, sometimes with rather stouter joints. The next pair
are considerably shorter and stouter, and bear more or less developed genital glands. The
following pinnules all have relatively stout joints, with the basal pair but little modified.
Disk and ambulacra naked ; sacculi abundant.
Colour in spirit, — light reddish-brown, the skeleton somewhat whiter.
Disk 6 mm.; spread probably 17 cm.
Localities. — Off Marion Island ; 50 to 75 fathoms. Two specimens.
Station 145, December 27, 1873; off Marion Island; lat. 46° 43' 0" S., long.
38° 4' 30" E.; 140 fathoms ; volcanic sand. One specimen.
Remarks. — This species, which represents Antedon tenella in the Southern Sea,
differs from it in the shortness of the later cirrus-joints (PI. XXXII. fig. 3) and in the
characters of the lower pinnules. The second pair are relatively large and stout, with
more or less developed genital glands, which do not appear in Antedon tenella until the
fourth or even the fifth pair. They are especially large and well developed in the two
examples from the smaller depth, and the pinnule-joints are proportionately stout
(PL XXXII. fig. 2). Another point of difference from Antedon tenella is the greater
backward extension of the axillaries, so that the second radials are almost entirely con-
cealed in the middle line of the ray, while there is but little modification of the basal
joints in the distal pinnules.
5. Antedon alternata, n. sp. (PL XVIII. figs. 1-3 ; PL XXXII. figs. 5-9).
Specific formula — A.—.
Centro-dorsal more or less hemispherical, bearing some twenty-five to thirty-five cirri
of about fifteen smooth joints, most of which are longer than wide.
First radials just visible ; the second short and somewhat incised by the rhombic
axillaries, which are usually wider than long, with incurved distal edges. Ten arms ; the
first brachials barely meeting above the sharp angles of the axillaries and somewhat
incised by the quadrate second brachials. The next joints square or oblong till the
second syzygy, and the following ones elongately quadrate with very oblique ends.
Syzygies in the third, eighth, and twelfth brachials, and then at intervals of one oi
sometimes two joints, the latter being the more common at first.
The second brachial has a slender pinnule about 7 mm. long, and consisting of twenty
elongated joints ; the third has a similar but shorter one. The next pair are still shorter
but have stouter joints, one or both of them having well-developed genital glands, and
the following ones gradually increase in length, becoming slender and delicate, with the
two basal joints more or less flattened.
ISO THE VOYAGE OF H.M.S. CHALLENGER.
Disk and ambulacra naked ; sacculi fairly abundant, especially on the outer pinnules.
Colour in spirit, — the skeleton white, with the perisome brownish.
Disk 4 mm.; spread probably 5 cm.
Localities.— Station 169, July 10, 1874; lat. 37° 34' S., long. 179° 22' E.;
700 fathoms ; blue mud ; bottom temperature, 40° F. One specimen.
Station 170a, July 14, 1874; near the Kermadec Islands; lat. 29° 45' S., long.
178° 11' W.; 630 fathoms ; volcanic mud ; bottom temperature, 39°"5 F. Two specimens.
Station 218, March 1, 1875; lat. 2° 33' S., long. 144° 4' E.; 1070 fathoms; blue
mud ; bottom temperature, 36°"4 F. One specimen.
Station 236, June 5, 1875; lat. 34° 58' N., long. 139° 29' E.; 775 fathoms; green
mud ; bottom temperature, 37°'6 F. Four specimens (two of them young) with
Myzostoma cornutum.
Remarks. — This is another of those very interesting species which are widely dis-
tributed in the abyssal region. I was at first inclined to separate the two varieties from
the South and the North Pacific respectively (PI. XVIII. fig. 1 ; PI. XXXII. fig. S),
but the additional experience of variable specific characters gained between 1879 and
1887 has led me to abandon this idea. The axillaries and second brachials of the more
northern forms have sharper proximal angles than in the southern variety ; while the
joints both of the cirri (PI. XXXII. fig. 9) and of the pinnules (PL XXXII. figs. 5, 7)
are relatively longer. Sometimes also the first two or three syzygial intervals after the
twelfth brachial consist of two joints (PL XXXII. fig. 8), instead of one only as in the
southern variety (PL XVIII. fig. l) ; though in the outer parts of the arms syzygial and
simple joints alternate with great regularity (PL XXXII. figs. 5, 7).
The two young individuals obtained at Station 236 chiefly differ from the more
mature form in the relatively greater length of the joints composing the cirri, arms, and
pinnules, and in showing more of the first radials externally ; this is especially the case
in the youngest specimen, which has not yet developed its genital glands, and is only
about 30 mm. m diameter. The appearance of the first radials externally gives the
calyx a considerable amount of resemblance to that of Antedon abyssicola from Station
244 (2900 fathoms), but this, though absolutely larger, shows more of the radials than
appears in the young Antedon alternata, and has fewer joints in its cirri (PL XXXIII.
fio-. 1), while the syzygial interval in the outer parts of the arms usually consists of more
than one joint.
The sino-le individual of Antedon alternata which was dredged at Station 218 is
peculiar in having four radials on one ray (PL XXXII. fig. 6). So far as it is possible to judge
from the characters of the other rays, the third of these seems to be the intercalated joint.
Antedon alternata is readily distinguished from Antedon tenella and from Antedon
exigua by the presence of only one joint between every two successive syzygies in the
REPORT ON THE CRINOIDEA. 181
middle and outer portions of the arms (PI. XXXI. figs. 1,4; PI. XXXII. figs. 1, 5, 7).
It also differs from Antedon exigua in the greater length of the outer cirrus-joints
(PI. XXXII. figs. 3, 8,9).
Attached to one of the specimens obtained at Station 236 was a single individual
of Myzostoma cornutuni, von Graff.
c Antedon rosacea, Linck, sp.
' \ Antedon petasus, Dtiben and Koren, sp.
be
Specific formula — A.—.
No Coniatuke which can be referred to either of these closely allied species were
dredged by the Challenger, but a single individual which seems to belong to Antedon
rosacea was obtained by the " Porcupine " somewhere in the Fasroe Channel ; while five
young specimens were met with on the Tunis coast, together with large numbers of
Antedon phalangiu/m. Another example of the " Porcupine " variety was obtainedby
the "Knight Errant" in 1880, to the north of the Hebrides ; and the "Triton" dredged
Antedon petasus on the Fasroe Banks in 1882.
The mutual relations of these various forms, and of the numerous examples of what
is commonly called Antedon rosacea from different localities in Britain *and elsewhere,
constitute a problem of no little difficulty, and one which I prefer to leave undecided
for the present. I have been collecting the materials for its solution for some years past,
and shall hope in course of time to be able to arrive at a definite opinion on the subject.
Antedon rosacea has been described by G-reeff as occurring at the Canary Islands and
even at the Equatorial Island of Rolas,1 in the Gulf of Guinea. But the question
whether the forms mentioned by him are identical with the North British variety which
goes by the same name, is one which cannot be definitely decided without a careful
comparison of the individuals in question and of others from intermediate localities.
7. Antedon diibeni, Bohlsche (PI. XXXVII. figs. 1-3).
Specific formula — A.—.
1866. Antedon Dubenii, Bohlsche, Archiv f. Naturgeseh., 1866, Ed. i. p. 9.
1868. Antedon Dubenii (?), Verrill, Trans. Connect. Acad., 1868, vol. i. p. 365.
1879. Antedon Dubenii, P. H. Carpenter, Trans. Linn. Soc. Lond. (Zool.), ser. 2, 1879, vol. ii. p. 29.
1882. Antedon dubeni, Bell, Proc. Zool. Soc. Lond., 1882, p. 534.
1883. Antedon dubeni, P. H. Carpenter, Proc. Zool. Soc. Lond., 1882 [1883], p. 746.
Centro-dorsal a slightly convex pentagonal disk, bearing from thirty to forty cirri,
which have from twelve to fifteen smooth joints, the outer ones stouter than those at the
base, laterally compressed, and rather longer than wide.
1 Echinodermen, beobachtet auf einer Reise nach der Guinea-Insel Sao Thome, Zool. Anzeiger, 1882, Jahrg. 5,
pp. 116, 159.
182 THE VOYAGE OF H.M.S. CHALLENGER.
First radials almost entirely concealed ; the second oblong, and not united laterally ;
axillaries acutely triangular. There is a variable amount of calcareous plating on the
perisorne between the rays. Ten arms ; the first two brachials tolerably similar in shape,
oblong or subtriangular, the second being rather the longer. A few joints after the
second syzygy may be triangular, but they soon become quadrate, with the sutures but
little inclined, so as to be somewhat squarish in outline, becoming elongated towards the
ends. The lower and middle joints may overlap more or less, but the distal parts of the
arms are almost smooth. Syzygies in the third, eighth, and twelfth brachials, and then
at intervals of one to six joints.
The second brachial bears a tapering pinnule of some twenty-five or thirty elongated
and overlapping joints, and reaches over 10 mm. in length ; that of the third brachial is
about half its size, with twelve or fifteen joints. The next pair are of about the same
length, and the following pinnules gradually increase, becoming very long and slender
towards the arm-ends.
There are a few scattered granules on the ventral surface of the disk, especially in
the anal interradius. Sacculi very abundant on the pinnule-ambulacra.
Colour in spirit, — yellowish-brown or brownish-white.
Disk about 7 mm.; spread 8 or 9 cm.
Locality. — Bahia, 20 fathoms. One specimen. Also Rio Janeiro (Bohlsche), and
the Abrolhos (Verrill) ?
Remarks. — By the kindness of Dr. Otto Hamann of Gottingen I have been enabled to
examine and figure the original specimen of this type, which was described by Bohlsche
from Rio Janeiro (PL XXXVII. fig. 2). There is a very considerable difference between
it and that obtained by the Challenger at Bahia (PI. XXXVII. fig. 1), but their general
resemblance is so close that there can be no question of their belonging to the same
specific type. The cirri are very uniform in appearance, but the radial axillary has a
much greater length in Bohlsche's specimen than in the Challenger one; while the anambu-
lacral plating on the perisorne between the rays is reduced in the former to a very definite
nodule which intervenes between every two second radials, very much as was figured by
Miller1 in his Comatula Jimbriata ( = Antedon rosacea). In fact it seems to rest directly
upon the upper angles of the first radials (PI. XXXVII. fig. 2), and it may possibly
represent a true calyx-interradial rather than anambulacral plates, which was shown by
Dr. Carpenter2 to be the case with Miller's type. The lower and middle arm-joints of
Bohlsche's example overlap but little, and the basal ones after the eighth are distinctly
triangular in outline, but in the Challenger specimen they are quadrate from the first and
overlap considerably, so that the dorsal line of the arm is markedly serrate (PI. XXXVII.
fig. 3). In this form too the syzygial interval is often five or six joints, while it is
1 Op. cit, Frontispiece, tig. 2. - Phil. Trans., 1866, p. 716, pi. xxxiii. fig. 7, a, b.
REPORT ON THE CRINOIDEA. 183
usually two and nowhere more than four in the type specimen. The lower pinnules are
also different in the two cases. Those of the type have spiny projections at the distal
ends of the overlapping joints which are almost entirely absent in the Challenger specimen,
and the length of the first pinnule is both relatively and absolutely much greater in the
former than in the latter.
In many respects this species comes very near to some forms of Antedon rosacea ;
and it may be that a larger acquaintance with the variations of the two types will lead
to their union. I do not think this possible at present, however, as they seem to be
pretty clearly distinguished by the characters of the arm-joints. In the type specimen
of Antedon diibeni there are a few triangular joints immediately above the second syzygy
(PI. XXXVII. fio-. 2), but in that dredged by the Challenger these joints are shortly
quadrate, though the length gradually becomes equal to the width, as is the case in the
type, and the sutures are so little inclined that the outline of the joints is tolerably
square, becoming more elongated, however, in the outer part of the arms. This character
distinguishes Antedon diibeni, from the British form of Antedon rosacea, in which the
joints are subtriangular, or, at any rate, have much-inclined sutures till some way out on
the arms ; though they are more nearly square in the Mediterranean form. But in all
the numerous varieties of Antedon rosacea the syzygial interval consists very regularly
of two joints ; and this seems to be also the case in the type of Antedon diibeni (PI.
XXXVII. fig. 2), so far as can be judged from the first forty brachials, the remainder
having been lost, though Bohlsche mentions one to four joints as the length of the syzygial
interval ; while in the Challenger specimen the number varies from two to six joints.
For the present, therefore, I would regard Antedon diibeni as distinct from Antedon
rosacea. The latter species has not yet been identified on the coast of the United States,
and one must hesitate therefore before giving the South American coast as a locality for
a type which is found as far north as the Shetlands, though it does seem to extend to
Madeira and the Canaries, and possibly to the Gulf of Guinea ; but I am not yet clear as
to the latter point ; and it is not improbable that Greeff s example from this locality may
turn out to be identical with Antedon diibeni.
8. Antedon lineata, n. sp. (PI. XIII. figs. 4, 5).
Specific formula — A.y.
Description of an Individual— Centvo-dovsal almost completely covered by some
twenty-five cirri with about thirty short joints. The lower joints are rather stout and
the fifth slightly the longest, while all of them overlap slightly on the dorsal side, and
gradually acquire a faint dorsal keel with a forward projecting spine.
First radials partially visible ; the second and third both rather high in the middle
184 THE VOYAGE OF H.M.S. CHALLENGER.
and falling away at the sides. The former are oblong, slightly incised, and not united
laterally ; the axillaries pentagonal and somewhat wider than long. Ten arms ; the first
brachials rather incised by the second, which are relatively short and wide. The
following joints smooth and obliquely quadrate, becoming rather elongated towards the
end. Syzygies in the third, eighth, and twelfth brachials, and then at intervals of one
to four, usually two or three joints.
The second brachial bears a styliform pinnule of about a dozen longish joints ; and
the next two or three pinnules on each side are of the same character, but of
diminishing size. The following pinnules increase in length and stoutness, the third and
fourth joints being expanded and broadly V-shaped ; the later pinnules are slender and
filiform. Disk invisible • the pinnule-ambulacra have abundant sacculi at their sides
and also numerous small pieces of calcareous network, which do not, however, form
definite plates.
Colour in spirit, — the arms dirty white, and the pinnules grey with white bands at
the joints.
Spread probably about 18 cm.
Locality.— Station 320, February 14, 1876; lat, 37° 17' S., long. 53° 52' W.; GOO
fathoms; green sand ; bottom temperature, 37°'2 F.
Remarks. — This species may be at once distinguished from those previously described
in the same group by the spiny cirrus-joints and by the expansion of the third and
following joints in the genital pinnules, as shown in PI. XIII. figs. 5a, 5b. A similar
character presents itself in Antedon gracilis and in Antedon accela, both from Station
214, off the Meangis Islands (PI. XV. fig. 4 ; PI. XVI. fig. 2), and also in Hyocrinus
bethellianus from the Southern Ocean (Part I. pi. vc. fig. 18 ; pi. vi. fig. 1). But in all
these species the ventral side of the genital glands is more or less protected by calcareous
plating, which is not the case in Antedon lineata.
9. Antedon remota, n. sp. (PL XXIX. figs. 5-9).
Specific formula — A. — .
Centro-dorsal sharply hemispherical, bearing twenty to thirty cirri, with nearly
twenty joints ; the lower ones are longer than wide and dice-box-shaped, with expanded
distal ends, overlapping their successors both dorsally and ventrally. From the tenth
joint onwards they are short and laterally compressed with a faint dorsal keel.
First radials just visible ; the second nearly oblong, rather convex and considerably
incised by the rhombic axillaries, which are wider than long, with sharp distal angles, so
that the first brachials are not united laterally. Both the axillaries and the two lower
brachials have traces of lateral projections.
EEPORT ON THE CRINOIDEA.. 185
Ten arms, of smooth, obliquely quadrate joints. Syzygies in the third and eighth or
ninth brachials, and then at intervals of two to four joints.
The first two pairs of pinnules long, slender and delicate, composed of several
elongated joints and tolerably equal in length. The two lower joints of the distal
pinnules are expanded and flattened.
Disk and ambulacra naked ; sacculi abundant on the pinnules.
Colour in spirit, — dirty white.
Disk about 4 mm.; spread probably about 8 cm.
Locality.— Station 147, December 30, 1873 ; lat. 46° 16' S., long. 48° 27' E.;
1600 fathoms ; Diatom ooze; bottom temperature, 340-2 F. Four mutilated specimens
and fragments of a fifth.
Remarks. — This species is readily distinguished from Antedon tenella and Antedon
rosacea and their allies by the characters of the lower pinnules, which are all long and
slender, but of tolerably equal size. They are much broken in all the specimens, and it
is difficult to determine either their length or the number of their component joints at
all exactly (PL XXIX. figs. 5, 7, 8) ; but the remains of them are quite sufficient to
show their difference from the lower pinnules of Antedon angustipinna and Antedon
abyssorum (PL XXIX. figs. 2, 11). There is no such enlargement of the joints of the
genital pinnules as occurs in the former species (PL XXIX. fig. 3), to which Antedon
remota has some resemblance in the characters of the cirrus-joints, though their number
is less ; while they are much shorter than those of Antedon abyssorum (PI. XXIX.
fig. 10).
One individual is peculiar in having only two radials on one ray ; but the axillary
or second radial is altogether different from the other rhombic axillaries, for it is
broadly pentagonal in form, with a perfectly even proximal margin, and no indication
whatever of a backward projecting proximal angle (PL XXIX. fig. 6).
10. Antedon longipinna, n. sp. (PL XXX. figs. 1-3).
Specific formula — A.-y.
Centro-dorsal rather sharply hemispherical, and bearing about thirty cirri, which have
twenty to twenty-five slightly overlapping joints, mostly longer than wide, and especially
so near the base.
First radials just visible ; the second short, free laterally, and deeply incised by the
rhombic axillaries, which are about as wide as long and have a sharp distal angle
separating the first brachials.
(ZOOL. CHALL. EXP. — PART LX.— 1887.) Ooo 24
186 THE VOYAGE OF H.M.S. CHALLENGER.
Ten arms of smooth, obliquely quadrate joints. Syzygies on the third, eighth, and
twelfth brachials, and then on every alternate joint.
The first pair of pinnules are about 7 mm. long and consist of some eighteen elongated
and rather dice-box-shaped joints. The second pair but little smaller, and the following
ones all long, decreasing but slowly in size, and retaining the elongated joints, which
bear short and fusiform genital glands. The later pinnules are slender again, but not as
long as the first pair.
Disk and ambulacra naked ; sacculi moderately abundant.
Colour in spirit,- — dirty white.
Disk 4 mm.; spread probably about 4 cm.
Locality.— Station 320, February 14, 1876; lat. 37° 17' S., long. 53° 52' W.; 600
fathoms ; green sand ; bottom temperature, 37°'2 F. Three mutilated individuals.
Remarks. — The length of the cirrus-joints and the very regular alternation of the
syzygies beyond the twelfth brachial readily distinguish this species from Antedon remota.
It is strikingly characterised by the great length of the joints composing the lower
pinnules, as is well seen in the immature specimen shown in PI. XXX. fig. 1. Its genital
glands are but little developed ; but a somewhat older individual has tolerably large
testes, and in that represented on PL XXX. fig. 2, there are much swollen ovarian sacs,
extending over about three pinnule-joints, with thin walls through which large white ova
are visible.
11. Antedon tenuicirra, n. sp. (PI. XXX. figs. 4-8 ; PI. XXXIII. figs. 4, 5).
Specific formula — A.y.
Centro-dorsal hemispherical, bearing thirty or more long and slender cirri of fifteen
to twenty-five joints or more, the first three of which are very short and the rest much
elongated.
First radials just visible ; the second short, rounded and oblong, free laterally and
scarcely incised by the axillaries, which are widely rhombic, with open proximal angles.
Ten arms, of smooth obliquely quadrate joints. Syzygies in the third and eighth
brachials and then at intervals of two to five joints.
The first pair of pinnules are long, slender, and delicate, composed of numerous joints
which are but little longer than wide. The second pair is of about the same length, but
consists like its successors of stouter and more elongated joints, which become slender
again in the distal pinnules.
Disk and ambulacra naked ; sacculi moderately abundant.
Colour in spirit, — light brown.
Spread, 11 cm.
REPORT ON THE CRINOIDEA. 187
Locality.-— Station 219, March 10, 1875; lat. 1° 54' 0" EL, long. 146° 39' 40" E.;
150 fathoms ; coral mud. One specimen, and one varietal form.
Remarks. — Besides this one mutilated individual (PI. XXX. fig. 4) Station 219 also
yielded another, which I at first regarded as belonging to a different specific type; and
it was accordingly represented on PI. XXXIII. figs. 4, 5, as Antedon notata. Probably,
however, it would be better considered as a varietal form of Antedon tenuicirra ; though
it presents some not unimportant differences from the type specimen described above.
Although of larger size, it shows more of the first radials, while the axillaries have
sharper proximal angles, and the second radials are therefore more incised. There are
slight indications of lateral flattening upon the four lower brachials, and the joints of
the first pinnule are relatively longer than in the type. The cirri are both more
numerous and have a larger number of joints than in the type (PL XXX. fig. 4 ;
PI. XXXIII. fig. 4) ; but the joints have the same smooth elongated character in both
forms ; and until better-preserved material is available, it will probably be safer to regard
them as specifically identical. The smooth and delicate long-jointed cirri and the difference
in shape of the joints composing the first and second pinnules respectively separate
the type very clearly from the species previously described. All the pinnules are much
broken, but the basal portions of the first two are very different, as seen in PL XXX.
figs. 5, 6. The lowest joints of the first pinnule recall those of the corresponding
pinnules in Antedon quadrata and its allies (PL XXVII. figs. 8, 14), but the later
joints are distinctly longer than wide, though not greatly so. In the second
pinnule, however (PL XXX. fig. 6), the component joints are stouter, and all except the
first two are distinctly longer than wide, as is the case in its successors (PL XXX. fig. 7).
12. Antedon kevis, n. sp. (PL XXXI. fig. 6).
Specific formula — A.y.
Description of an Individual. — Centro-dorsal hemispherical, covered with a cluster
of about thirty cirri with some twenty -five to thirty joints, but few of which are longer
than wide. The distal joints have a faint dorsal keel which passes into a small opposing
spine on the penultimate.
Three radials visible ; the first short, and the second rather sharply convex ; axillaries
rhombic, about as wide as long. Ten arms ; first brachials nearly oblong, and but little
incised by the second, which are irregularly quadrate. The next joints are nearly oblong
till the second syzygy, and the following ones smooth and obliquely quadrate, gradually
becoming longer than wide. Syzygies in the third, eighth, and twelfth brachials, and
then at intervals of two joints.
188 THE VOYAGE OF H.M.S. CHALLENGER.
The lowest pinnules seem to be tolerably equal in length, consisting of cylindrical
joints which are relatively longer in the second than in the first pair.
Disk invisible ; pinnule-ambulacra naked, with abundant sacculi.
Colour in spirit, — the skeleton straw-coloured, with the perisome brownish.
Spread probably about 5 cm.
Locality. — Station 214, February 10, 1875 ; off the Meangis Islands; lat. 4° 33' N.,
long. 127° 6' E.; 500 fathoms; blue mud; bottom temperature, 4l°*8 F.
Remarks. — This little species differs from the two last described in having much
shorter cirrus-joints (PI. XXX. figs. 3, 4, 8 ; PI. XXXI. fig. 6). The lower pinnules are
much broken, but they appear to have been tolerably equal in length, and the joints of
the first pinnule are relatively shorter and stouter than those of the second ; while in
Antedon tenuicirra the reverse is the case. There is a certain amount of resemblance
between Antedon Isevis and Antedon remota ; but the former species has relatively longer
axillaries than occur in Antedon remota, and also more numerous cirrus-joints, which do
not overlap as is the case in that species (PI. XXIX. figs. 5, 6).
13. Antedon hirsuta, n. sp. (PI. XXXI. fig. 5).
be
Specific formula — A.-^-.
Description of an Individual. — Centro-dorsal conical, bearing about thirty-five cirri
in irregular vertical rows. The cirri have twenty-five to thirty joints, the lower ones
somewhat elongated and the later joints smaller, with slight dorsal keels.
Three radials visible ; the first short, the second oblong, rather convex and but little
incised for the widely rhombic axillaries. Ten arms ; the first brachials nearly oblong, and
the second relatively short and wide, with a very open proximal angle. The next few joints
oblong, and the following ones elongately triangular, gradually becoming more quadrate
and finally cylindrical, with slight lateral projections for the pinnule facets. The distal
edge of each joint bears a small fringe of spines which projects forwards over the base of
its successor so as to give the arms a somewhat serrate appearance. Syzygies in the
third, and then in the sixth or eighth brachials, after which they are rather irregular, but
generally at intervals of two or three joints.
The arms are mostly regenerated at the first syzygy ; but in the uninjured ones the
first two pinnules, which are stiff and tapering, consist of about twelve longish joints, and
appear to be tolerably equal, the first being perhaps a little the longer, and with stouter
joints, the lowest of which may be slightly flattened. The second pinnule has a genital
gland, and the following ones are at first shorter and more slender than those below them,
but have relatively longer joints, after which the length gradually increases.
REPORT ON THE CRINOIDEA. 189
Disk invisible ; pinnule-ambulacra naked, with abundant sacculi at their sides.
Colour in spirit, — white.
Spread about 7 cm.
Locality. — Station 145, December 27, 1873 ; near Marion Island; lat. 46° 43' 0" S.,
long. 38° 4' 30" E.; 140 fathoms ; volcanic sand. One specimen.
Remarks. — This species differs from Antedon Isevis in the serrate appearance of the
outer parts of the arms along their dorsal line, in the relatively greater width of the two
outer radials and of the second brachials, and in the flattened appearance of the basal
joints of the first pinnule. The syzygies too seem to be less regularly arranged than in
Antedon Isevis; but this may be due to the fact that most of the arms have been
regenerated at the third brachial in the single individual obtained, entangled in the cirri
of which was a specimen of Ophiolebes scorteus.
14. Antedon angustipinna, n. sp. (PI. XXIX. figs. 1-4).
Specific formula — A.-^ .
Description of an Individual. — Centro-dorsal nearly hemispherical, and almost
covered by about thirty cirri. These have some twenty -five joints with sharpened dorsal
edges, the lower ones slightly the longer and the remainder tolerably equal, but short.
Three radials visible ; the second oblong, almost free laterally, and strongly incised
to receive the rhombic auxiliaries, which are wider than long. Ten smooth and rounded
arms, the joints after the second syzygy being elongately triangular or quadrate.
Syzygies in the third, eighth, and twelfth brachials, and then on every alternate joint.
The second brachial bears a short stiff pinnule of five to eight joints, and there is
an even smaller one on the third brachial; but that on the fourth is longer, often
much so.
The next five or six pinnules on each side have longer and much stouter joints, and
support the genital glands, the remaining pinnules being slender and delicate.
Disk and ambulacra naked ; sacculi fairly abundant.
Colour in spirit, — yellowish-white.
Spread probably 5 cm.
Locality.— Station 320, February 14, 1876; lat. 37° 17' S., long. 53° 52' W.; 600
fathoms; green sand; bottom temperature, 37°'2 F. One specimen.
Remarks.— This curious little species from the South Atlantic is readily distinguished
by its short cirrus-joints and by the peculiarities of its lower pinnules. That on the
second brachial is unusually small, sometimes having no more than five or six joints,
all of them, however, longer than wide (PI. XXIX. fig. 2). The fourth brachial bears
190 THE VOYAGE OF H.M.S. CHALLENGER.
a much longer pinnule, the lower joints of which are usually, though not always,
enlarged for the support of the genital glands, as in its successors (PL XXIX. fig. 3).
After about the twelfth brachial, however, the glands become reduced in size and the
pinnule-joints smaller again (PI. XXIX. fig. 4). As in Antedon alternata every alternate
brachial above the twelfth is usually a syzygial joint (PI. XXIX. fig. 1 ; PI. XXXII.
figs. 5, 7).
15. Antedon abyssorum, n. sp. (PL XXIX. figs. 10-13).
Specific formula — A.—.
Centro-dorsal conical, bearing about thirty cirri of fifteen to eighteen joints, nearly
all of which are longer than wide.
Three radials visible ; the second partially free, rather convex and very deeply incised
for the axillaries. These are shield-shaped or rhombic, as long or longer than wide, with
much incurved distal edges. First brachials deeply incised by the irregularly quadrate
second brachials. Ten arms of nearly smooth joints, which are almost oblong up to the
second syzygy and then become obliquely quadrate. Syzygies in the third and eighth
brachials, then about the fourteenth, and afterwards at intervals of one to five joints.
The first pair of pinnules are slender and delicate, composed of ten or twelve
elongated joints, that on the second brachial being rather the longer. Those on the
fourth and the ten or twelve following brachials are longer with stouter joints, and
support the short and thick genital glands. The later pinnules are much elongated.
Disk and ambulacra naked ; sacculi tolerably abundant on the outer pinnules.
Colour in spirit, — dirty white.
Spread perhaps 6 cm.
Locality.— Station 147, December 30, 1873 ; kit, 4G° 16' S., long. 48° 27' K; 1600
fathoms ; Diatom ooze ; bottom temperature, 34°-2 P. Eleven specimens.
Remarks.- — This abyssal form agrees with Antedon angustipinna in the relatively
small size of the first pair of pinnules (PL XXIX. figs. 2, 11), but differs from that type
in several points.
There is a smaller number of cirrus-joints, but they are long and slender instead of
short and wide ; while there may be from one to five joints between the brachial
syzygies, and not one only as in Antedon angustipinna (PL XXIX. figs. 1, 10). In the
latter species those lower joints of the genital pinnules which support the glands are
considerably thickened ; but this is not the case in Antedon abyssorum (PL XXIX.
figs. 3, 12) ; while the later pinnules of this species are much longer than in Antedon
angustipinna (PL XXIX. figs. 4, 13).
EEPORT ON THE CRINOIDEA. 191
1G. Antedon abyssicola, n. sp. (PI. XXXIII. figs. 1, 2).
Specific formula — A.—.
Centro-dorsal subconical, bearing about fifteen cirri. These have eight to ten joints,
of which the second is longer than wide and the following joints much elongated, the
fourth being about the longest ; the remainder diminish to the penultimate, which is not
much longer than wide and has a very faint opposing spine.
Three radials visible ; the first and second about equal in length, the angles of the
former extending inwards and upwards for some distance, so that the second radials are
not united laterally. They are somewhat incised by the hinder angles of the rhombic
axillaries, the distal edges of which are much curved. Ten arms ; the first brachials
quite separated and more or less incised by the proximal angles of the irregularly
quadrate second brachials. The following joints are square till the second syzygy or
even slightly longer than wide ; the later joints obliquely quadrate and much longer
than wide, gradually becoming almost dice-box-shaped. Syzygies in the third, eighth,
and twelfth or thirteenth brachials, and then at intervals of two to four, usually three,
joints.1
All the pinnules are much broken, but they seem to have been slender and delicate
on the arm-bases.
Disk and ambulacra naked ; sacculi small and rare.
Colour in spirit, — white.
Disk about 3 mm.; spread perhaps 5 cm.
Localities.— Station 160, March 13, 1874; lat. 42° 42' S., long. 134° 10' K; 2600
fathoms ; red clay ; bottom temperature, 33°'9 F. One specimen.
Station 244, June 28, 1875 ; lat. 35° 22' N, long. 169° 53' E.; 2900 fathoms; red
clay ; bottom temperature, 3 5° "3 F. Two specimens.
Remarks. — This little species is one of very considerable interest, apart altogether
from the peculiarities of its calyx, for it is the only Comatula yet found at a greater
depth than 2000 fathoms. Bathycrinus, and perhaps also Hyocrinus, extend down to
2400 fathoms; Promachocrinus and Thaumatocrinus occur at 1800 fathoms, but with
the exception of Antedon abyssicola no other Comatulse have been found below 1600
fathoms, at which depth (Station 147) Antedon abyssorum, Antedon bispinosa and
Antedon remota were obtained. Antedon abyssicola has been dredged, however, at two
stations, one (Station 160) shortly before the Challenger reached Melbourne, where the
depth was 2600 fathoms, and the other in the deepest part of the North Pacific at
2900 fathoms (Station 244). Antedon abyssicola thus resembles Antedon alternata
in occurring at widely separated localities in the abyssal region, and it has some points
1 In fie. 1 the tenth and fourteenth, and in fig. 2 the sixth brachials are wrongly drawn as syzygial joints.
192 THE VOYAGE OF H.M.S. CHALLENGER.
of resemblance with the younger individuals of this type, as has been pointed out already.
The three specimens obtained were unfortunately all much mutilated, especially as
regards the cirri and pinnules ; but the peculiarities of the calyx are very characteristic
and serve to distinguish the type, though its precise relations to the other abyssal species
must remain somewhat uncertain in the absence of properly preserved individuals.
A considerable portion of the first radials appears externally, and they are somewhat
wider than the second radials, while their angles are considerably produced both upwards
and inwards, so that the second radials are altogether prevented from coming into lateral
contact. This is especially well shown in the southern form (PI. XXXIII. fig. 2), but it
is not so apparent in the larger individuals from the North Pacific, except in an inter-
radial view of the calyx. Both this character and also the great length of the arm-joints,
the later ones of which are almost dice-box-shaped, indicate that the type is an embryonic
one, as is well seen on a comparison of figs. 1 and 2 on PI. XXXIII. with the larvae
represented in figs. 3 to 6 on PL XIV. The extension of the first radials upwards and
inwards, so as to keep the second radials from coming into lateral contact, is a larval
character which is better developed in the fossil Eugeniacrinus, and reaches its maximum
in the allied genus Phyllocrinus.
A few poorly developed sacculi are present in the individual from 2600 fathoms
(Station 160), but I have not been able to find any indications of them in the two
specimens from the greater depth in the North Pacific (Station 244, 2900 fathoms).
5. The Milberti-gvou-p.
The first pair of pinnules comparatively small, and their component joints but little
longer than wide ; one or more of the second, third, and fourth pairs are longer and
more massive, with stouter joints than their successors.
Remarks. — This is a somewhat heterogeneous group, and I have had considerable
trouble in working out an arrangement of it which I can regard as even approximately
satisfactory. The definition given above would almost include such forms as Antedon
angustipinna (PL XXIX. figs. 2-4) and Antedon tenuicirra (PL XXX. figs. 5-7), which
have been described in the JeneWa-group ; while Antedon parvicirra, which I have
placed in the Milberti-gxovi\), though with some doubt, has many points of resemblance
with Antedon rosacea and Antedon dtibeni. Indeed Antedon milberti itself exhibits
traces of the wall-sidedness of the radials and lower brachials which is so marked in the
Basicurva-grovi]). Then again, the ubiquitous Antedon carinata differs in many
respects from Antedon serripinna, Antedon milberti, and the typical members of the
group, so that another group may have to be established for it at some future time.
The special character which distinguishes the Milberti-groi\]p is the large size of one
or more of the second, third, and fourth pairs of pinnules, which are borne respectively
REPORT ON THE CRINOIDEA. 193
by the fourth to ninth brachials. This feature is well shown in Antedon anceps, Antedon
milberti, and Antedon variipinna (PI. XXXV. figs. 2, 4 ; PI. XXXVI. figs. 1, 4-6), and
also in some species described by Bell in the " Alert " Report ; while it reappears in a more
marked degree in some of the multibrachiate species of Antedon (PI. XLVIII. figs, 2, 3).
In some cases, as in Antedon milberti, the second, third and fourth pairs are all of
greater size than the pinnules above and below them, sometimes the second and some-
times the third being slightly the largest. In Antedon anceps and in Antedon variipinna
the pinnules of the fifth or sixth brachials (or both) are considerably longer and stouter
than their fellows (PL XXXV. fig. 2 ; PL XXXVI. figs. 1, 4-G) ; and in Bell's species
Antedon carpenteri and Antedon pumila, the large pinnule is on the fourth brachial.
But in Antedon carinata and Antedon parvieirra the third and the following pairs of
pinnules are much more ecpual in size. At the end of the group I have placed two
abnormal species in which the pinnule on the third brachial is absent, though in other
respects they conform pretty well to the general type.
All the members of the Milberti-growp are limited to the Pacific and the Eastern
Archipelago, with the exception of Antedon carinata, which also extends into the Indian
Ocean, Red Sea, and the Western Atlantic. It was dredged off St. Lucia, in 278 fathoms,
by Captain Cole of the telegraph steamer "Investigator"; but all the remaining
members of the group are confined to the littoral zone. Most of them have been
obtained at depths of 20 fathoms or less ; but Antedon variipinna occurs at 36 fathoms
in the Arafura Sea.
The general relations of the various members of the Milberti-growp are shown in
the following table : —
-*&
A. A pinnule on the third brachial.
I. Second pair of pinnules the largest.
a. Cirrus-joints short.
1. Twenty-five cirrus-joints ; the first brachials much incised, . pinniformis, Carpenter.
2. Barely twenty cirrus-joints ; the first brachials not incised.
Second pinnule serrate, ..... serripinna, Carpenter.
Second pinnule with large processes on the lower joints, . carpenteri, Bell.
b. Twelve long cirrus-joints, ..... pumila, Bell.
II. Second and third, and sometimes the fourth, pairs of pinnules about
equal.
a. Twenty-five to forty cirrus-joints.
Radials and lower brachials tubercular; the lower pinnules
rounded . . . . . • • 1- milberti, Mull., sp.
Radials and lower brachials smooth ; the lower pinnules
carinate, ....... Ixvissima, Grube, sp.
b. Forty-five cirrus-joints ; syzygial interval seven to ten joints, . tessellata,1 Mull., sp.
c. Sixty cirrus-joints; syzygial interval three to seven joints, . perspiTiosa, Carpenter.
1 1 only know this type from the description of it which is given by Miiller ; but it is the only species recorded
in the list given on pp. 53-55 which I have not personally examined.
(ZOOL. CHALL. EXP. — PART LX. 1887.) 0°° 25
194 THE VOYAGE OF H.M.S. CHALLENGER.
III. Third outer pinnule distinctly larger than the second.
a. Over thirty cirrus-joints.
Third pinnule has smooth joints ; syzygial interval four to
seven joints, . . . . . .2. anceps,1 n. sp.
Joints of third pinnule with lateral processes ; syzygial interval
nine to twelve joints, . . . . .3. variipinna,1 Carpenter.
b. Twenty to thirty cirrus-joints. The third and next following
pinnules tolerably equal in length; arms more or less carinate, 4. carinata, Lamk., sp.
c. Not more than twelve cirrus-joints. The third pinnule rather the
longest, . . . . . . .5. parvicirra, n. sp.
B. Usually no pinnule on the third brachial.
I. Joints of second and third pinnules not specially elongated. Fifteen
cirrus-joints, . . . . . . . 6. informis, n. sp.
H. Second and third pinnules have much elongated joints with enlarged
and denticulate ends, ...... loveni, Bell.
1. Anteclon milberti, Miiller, sp. (PI. XXXV. figs. 4-6).
Specific formula — A. y.
1846. Comatula (Alecto) Milberti, Miiller, Monatsber. d. k. preuss. Akad. d. Wiss. Berlin, 1846,
p. 178.
1846. Comatula Jacquinoti, Miiller, Ibid., p. 178.
1849. Comatula {Aledo) Milberti, Miiller, Abhandl. d. k. Akad. d. Wiss. Berlin, 1847 [1849],
p. 255.
1849. Comatula Jacquinoti, Miiller, Ibid., p. 255.
1862. Comatida Milberti, Dujardin and Hupe, Hist. Nat. des Zoophytes, Echinodermes, Paris,
1862, p. 202.
1862. Comatula Jacquinoti, Dujardin and Hupe, Ibid., p. 202.
1867. Antedon Milbertii, Verrill, Trans. Connect. Acad. Arts and ScL, 1867, vol. i. p. 341.
1875. Comatula Ixvissima, Grube (pars), 53e Jahresber. der Schlesisch. Gesellsch. f. Vaterl.
Cult., 1875, p. 74.
1879. Antedon Jacquinoti, P. H. Carpenter, Trans. Linn. Soc. Lond. (Zool.), ser. 2, 1879, vol. ii.
p. 29.
1879. Antedon Milberti, P. H. Carpenter, Ibid,, p. 29.
1882. Antedon Ixvissima, Bell (pars), Proc. Zool. Soc. Lond., 1882, p. 533.
1882. Antedon jacquinoti, Bell, Ibid., p. 534.
1882. Antedon milberti, Bell, Ibid., p. 534.
1882. Antedon jacquinoti, P. H. Carpenter, Ibid., p. 746.
1882. Antedon Isevissima, P. H. Carpenter, (pars), Ibid., p. 746.
1882. Antedon milberti, P. H. Carpenter, Ibid., p. 746.
1884. Antedon Milberti, Bell, Bep. Zool. Coll. H.M.S. "Alert," London, 1884, p. 156.
1884. Antedon milberti, Bell, Proc. Linn. Soc. N.S.W., 1884, vol. ix. p. 497.
Centro-dorsal hemispherical, bearing some twenty to thirty cirri. These have twenty-
five to thirty -five or even nearly forty joints, most of which, and especially the lower
ones, are wider than long. The middle and outer joints have a more or less distinct
dorsal spine.
1 These are both tridistichate species ; see pp. 254, 256.
REPORT ON THE CRINOIDEA. 195
First radials almost or entirely invisible ; the second rather sharply convex, and
rising to a median tubercle at their junction with the wide axillaries. A similar but
smaller tubercle at the junction of the first two brachials. In large specimens these four
joints are sometimes slightly wall-sided, with straight edges and the margins of the dorsal
surface flattened.
Ten arms, reaching nearly three hundred joints ; the third and next following
brachials smooth, rounded and nearly oblong, with a tendency to alternating tubercular
elevations at their junctions. After the second syzygy the joints are shortly triangular
and slightly overlapping, gradually becoming nearly oblong, but always much wider than
long. Syzygies in the third and eighth or ninth brachials, and often also in the twelfth
or thirteenth ; others at intervals of three to nineteen joints, usually eight or ten, the
intervals being somewhat longer in the outer parts of the arms than in their first third.
The first pair of pinnules are about 8 mm. long and consist of some eighteen
moderately stout joints, of which some of the middle ones are longer than wide. The
pinnules of the next five or six brachials (fourth to ninth) are somewhat longer and stiffer,
with much stouter joints, sometimes the second and sometimes the third pair being the
largest. The fourth pair are occasionally much smaller than the third, and the fifth pair
are always much so, after which the length of the joints increases and the later pinnules
become long and slender.
Disk naked ; sacculi abundant.
Colour in spirit, — dark reddish-brown, bleaching to white.
Disk 10 or 12 mm.; spread 25 to 30 cm.
Localities.— Station 203, October 31, 1S74 ; lat. 11° 6' N., long. 123° 9' E.; 20
fathoms ; mud. One specimen.
Station 212, January 30, 1875 ; lat, 6° 54' N., long. 122° 18' K; 10 fathoms ; sand.
Two specimens.
Other Localities. — Ceram (Valenciennes) ; North Borneo (Grube) ; H.M.S. " Alert,"
1881, Port Molle (12 to 20 fathoms), Port Denison (3 to 4 fathoms), Prince of Wales
Channel (7 to 9 fathoms), Torres Strait (10 fathoms); Padan Bay in the Mergui
Archipelago (Dr. J. Anderson).
History. — Under this name I have united the two species that were found by Miiller in
the Paris Museum with the MS. names " Comatula Milberti" and " Comatula Jacquinoti "
respectively, which had been given to them by Valenciennes. They are each based upon
single specimens which I was able to examine carefully in 1876, and again in 1880 ; and
the subsequent study of a considerable amount of material obtained by H.M.SS.
Challenger and " Alert," and also by Dr. J. Anderson, F.K.S., of the Calcutta Museum,
has convinced me that the two types are really identical. Muller hardly ever made any
comparison of his species with one another, but simply contented himself with descriptions,
196
THE VOYAGE OF H.M.S. CHALLENGER.
leaving his readers to determine the real points of difference between his various species.
For this purpose I have analysed his descriptions of Comatula milberti and of Comatula
jacquinoti respectively, with the following result : —
Comatula milberti.
Twenty-five to thirty cirri of thirty-five spiny joints.
The spine "quer absteht."
First radials " ausserst niedrig."
Arm-joints "niedrig."
Syzygial interval eight or nine joints.
The second, third, and fourth pinnules are "die
grossten."
Colour, — brownish-black.
Spread approaching 2 feet.
Comatula jacquinoti.
Twenty-two cirri of thirty-five spiny joints.
The spine is " vorwarts gerichtet."
First radials "sehr niedrig."
Arm-joints " niedrig."
Syzygial interval three to six joints.
The three to four first pinnules are " starker."
Colour. — brownish-black.
Spread approaching 2 feet.
The preceding table shows that the differences between Comatula milberti and
Comatula jacquinoti, as described by Midler, are in reality exceedingly slight. The
number of cirrus-joints, the characters of the radials and of the arm-joints, the colour, and
even the size are respectively identical in the two types. Comatula milberti has twenty-
five to thirty cirri with the spines transverse, whde in Comatula jacquinoti there
are twenty-two cirri with the spines directed forwards. In Comatula milberti the
syzygial interval is eight or nine joints, and the second, third and fourth pinnules are
the largest, whde in Comatula jacquinoti the syzygial interval is three to six joints and
the first three or four pinnules are " starker." Neither of these characters, however, nor
even the combination of them, can be regarded as of specific value, especially when we
remember that each of Midler's species was based upon a single specimen. That of
Comatula jacquinoti had been obtained at Ceram by the expedition of d'Urville in the
"Zelee" (1841), whde the form which Midler described under the specific name milberti
had previously received it from Valenciennes in honour of M. Milbert of New York, who
had given it to the Paris Museum ; and it was possibly for this reason that the type was
labelled as coming from North America. Under these circumstances Valenciennes, and
after him Midler, were perhaps a httle predisposed to regard it as distinct from the
Comatula jacquinoti of Ceram, which Midler described along with it and in such nearly
identical terms. I feel quite confident, however, that Milbert's specimen was not obtained
anywhere on the Atlantic coast of North America. I have seen nothing like it among
the West Indian Comatulge dredged by the "Blake"; while the only species of Antedon
which have been found on the Atlantic coast of North America are Antedon tenella and
perhaps Antedon eschrichti (Stimpson). All its characters are those of the species of
Antedon which inhabit the Eastern Seas, where the type has been obtained at various
localities from the Mergui Archipelago to Eastern Australia ; and I have little doubt that
Mdbert's specimen had been brought to America from somewhere within this region.
REPORT ON THE CRINOIDEA. 197
Verrill 1 has referred it to the Caribbean fauna, but with a ? ; while Dujardin and Hupe\2
who must have seen it for themselves in the Paris Museum, refer to it as having come
"de l'Amerique septentrionale." We know nothing respecting any Comatulae on the
Pacific coast of Central and North America, and I strongly suspect that Milbert's specimen
must have been wrongly labelled.
Under the name of Comatula lasvissima, Grube 3 described in 1875 two ten-armed
examples of Antedon which he had obtained from Borneo ; and Professor Schneider,
Grube's successor at Breslau, has been good enough to send them to me for examination.
They agree pretty closely in the characters of their cirri and short arm-joints ; but, as is
indicated in Grube's diagnosis, their colour is altogether different, while one of them has
a tubercular junction between the two outer radials and also between the first two
brachials, a character which is altogether absent in the other individual. In the latter
too the joints of the lower pinnules are sharply carinate. This is not the case in the
form with tubercular radials, which I find to be a small individual of Antedon milberti ;
and Grube's specific name leevissima will therefore only apply to the other specimen,
which I propose to describe more fully at a future time.
Remarks. — The tubercular radials and the stout but rounded joints of the large lower
pinnules, together with the spiny cirri and the short arm -joints, thus combine to make
Antedon milberti an easily recognisable type. Although the second and third pairs of
pinnules are distinctly larger than the first, neither of them is especially characterised by
its greater size, as is the case in Antedon anceps and Antedon dubia (PI. XXXV. fig. 2 ;
PI. XXXVI. figs. 1, 4-6). Sometimes the one and sometimes the other is a little the
larger, while the third pair is occasionally nearly equal to the second, and in other
individuals considerably smaller, though always distinctly larger than its successor.
The grouping of the syzygies in the arms is somewhat irregular. The second one is
very often on the eighth or ninth brachial, and is followed by another four joints after-
wards ; but in some arms the second syzygy does not come till the twelfth or thirteenth
joint. The examination of a large number of arms shows the syzygial interval to vary
from three to nineteen joints. It is usually from eight to ten in the middle and outer
parts of the arms, though somewhat less in their lower portions.
In some individuals the axillaries and the lowest brachials have indications of
straight lateral edges and of the peculiar wall-sided character which has been described
above as distinctive of the Basicurva-groTxp. This is most marked in the specimen
obtained by the Challenger at Station 203, which differs frum all the other examples of
the type that I have seen in showing a considerable portion of the first radials externally.
Their length is more than half that of the second radials, and the tubercles which the
latter form with the pentagonal axillaries are less prominent than usual. Both joints
1 Echinodernis. Comparison of the Tropical Faunae of the East and West Coasts of America, Trans. Connect. Acad.
Arts and Sci., 1867, vol. i. p. 341.
2 Op. cit., p. 202. 3 53° Jahresber. der Schlesisch. Gesellsch.f. Vaterl. Cult., 1875, p. 74.
198 THE VOYAGE OF H.M.S. CHALLENGER.
and also the first two brachials have the margins of the dorsal surface flattened, with
straight lateral edges, and in some arms this character also extends on to the hypozygal
of the third brachial, but the wall-sidedness is always much less distinct than in the
Basicurva-grou-p, and in some examples is scarcely visible at all ; while there are no
indications of the flattening of the sides of the first pinnule which is so characteristic of
Antedon basicurva, Antedon valida, and their allies (see woodcut fig. 3, on p. 122).
Furthermore Antedon milberti has unplated ambulacra, and in all other respects it is
closely allied to Antedon anceps, Antedon serripinna, and the other species which I have
placed with it in the same group. At the same time the indications in this distinctly
littoral type of a peculiarity which is especially characteristic of Comatulae from the
continental and abyssal regions is a point of considerable interest.
Some of the examples of this species which were dredged at Mergui were infested
by a species of Myzostoma which Professor von Graff has been unable to determine
satisfactorily, owing to its state of preservation.
2. Antedon anceps, n. sp. (PI. XXXV. figs. 1-3).
Specific formula — A.(3).y.
Locality.— -Station 212, January 30, 1875; lat. 6° 54' N., long. 122° 18' E.; 10
fathoms ; sand.
Remarks. — Of the three individuals of this species which were dredged by the
Challenger, one has ten arms, while the other two have three and four distichal series
respectively. The type will therefore be described together with the remaining members
of the tridistichate group. But a few words may be said here about its ten-armed
variety. It has a considerable superficial resemblance to the less tubercular forms of
Antedon milberti, with which it agrees in the characters of its arm-joints ; but the third
outer pinnule (on 6 br.) is larger than the second (on 4 br.), as seen in PI. XXXV.
fig. 2, which represents the pinnules on the inner side of the arm, i.e., on the third and
following brachials. The cirri, too, have smooth joints, and so are very different from
the spiny cirri of Antedon milberti (PL XXXV. figs. 3, 4).
3. Antedon variipinna^ Carpenter (PL XXXVI. figs. 1-6).
Specific formxda — A.[3.(2)].y.
Locality. — Arrou Islands.
Remarks. — Most individuals of this species are distinctly tridistichate, but the two
from the Arrou Islands seem to owe this character to a regeneration after fracture at
1 A revised diagnosis of this species, together with its synonymy, will be fonnd on p. 256.
REPORT ON THE CRINOIDEA. 199
the syzygy in the third brachial of the primary arm, and they may therefore be
considered as members of the ten-armed series.
The terminal cirrus-joints of Antedon variipinna are more sharply carinate than
those of Antedon anceps (PI. XXXV. fig. 3; PI. XXXVI. fig. 1), and are sometimes
spinous, while the interval between the later syzygies of the arms is considerably longer
(PL XXXV. fig. 2; PL XXXVI. fig. 3). Furthermore the joints of the third pinnule
have more or less prominent lateral processes, while those of Antedon anceps are smooth
(PL XXXV. fig. 2 ; PL XXXVI. figs. 1, 4).
4. Antedon carinata, Lamk., sp. (PL III. figs. 1—3 ; PL XXXIV.).
b
Sp)ecif.c formida — A.y.
1811. Antedon gorgonia (?), de Freniinville, Bull. Soc. Philom. Paris, 1811, t. ii. p. 349.
1815. Aleeto carinata (?), Leach, Zool. Miscellany, London, 1815, vol. ii. p. G3.
1816. Comatula carinata, Lamarck, Hist. Nat. Anim. sans Vert. Paris, 1816, t. ii. p. 534.
1834. Comatula carinata, Griffith, Animal Kingdom (Cuvier), vol. xii., Mollusca and Radiata,
London, 1834, pi. viii. fig. 2.
1834. Comatula carinata, de Blainville, Manuel dActinologie, Paris, 1834, p. 249.
1843. Aleeto carinata, Mtiller, Archiv f. Naturgesch., 1843, Jahrg. ix. Bd. i. p. 135.
1849. Comatula (Aleeto) carinata, Miiller, Abhandl. d. k. Akad. d. Wiss. Berlin, 1847 [1849],
p. 252.
1862. Comatula carinata, Dujardin and Hupe, Hist. Nat. des Zoophytes, Echinoderrnes, Paris,
1862, p. 200.
1865? Antedon Braziliensis, Liitken, MS.
1867. Antedon Braziliensis, Verrill, Trans. Connect. Acad. Arts and Sci., 1867, vol. i. p. 341.
1868. Antedon Dubenii, Verrill, Ibid., p. 365 (with?).
1878. Antedon carinata, Pourtales, Bull. Mus. Comp. Zool., 1878, vol. v. p. 214.
1879. Antedon Brasiliensis, P. H. Carpenter, Proc. Roy. Soc, 1879, vol. xxviii. p. 386.
1879. Antedon carinata, P. H. Carpenter, Trans. Linn. Soc. Lond. (Zool), ser. 2, 1879,
vol. ii. p. 29.
1879. Antedon bicolor, P. H. Carpenter, Ibid., p. 29.
1879. Antedon carinatus (?), Rathlmn, Trans. Connect. Acad. Arts and Sci., 1879, vol. v.
p. 156.
1880. Antedon brasiliensis, P. H. Carpenter, Quart. Journ. Geol. Soc, 1880, vol. xxxvi. p. 41.
1881. Antedon carinata, P. H. Carpenter, Notes from the Leyden Museum, 1881, vol. iii.
p. 179.
1881. Antedon carinata, P. H. Carpenter, Bull. Mus. Comp. Zool., 1881, vol. ix. No. 4, p. 7.
1882. Antedon carinata, Ludwig, Mem. Acad. Sci. Bruxelles, 1882, t. xliv. p. 5.
1882. Antedon carinata, P. H. Carpenter, Journ. Linn. Soc. Lond. (Zool.), 1882, vol. xvi. p. 502.
1882. Antedon carinata, Bell, Proc. Zool. Soc. Lond., 1882, p. 534.
1882. Antedon carinata, P. H. Carpenter, Ildd., p. 746.
Centro-dorsal a thick, roughly circular disk, with the dorsal surface bare, and
bearing from twenty to thirty marginal cirri. These have from twenty to thirty
stout joints, all wider than long, the basal ones especially so. The penultimate joint
sometimes, but rarely, has a small opposing spine.
200 THE VOYAGE OF H.M.S. CHALLENGER.
The first radials partially visible ; the second short and oblong. Axillaries
triangular, twice the length of the second radials, and forming with them a more or
less distinct median tubercle. The first two brachials wedge-shaped (the first least so),
with sharp outer edges and a similar median elevation in their line of union. Ten
arms of about one hundred and sixty short joints, which are wide and nearly oblong
till the second syzygy, after which they are narrower and more triangular, gradually
becoming more oblong again and finally square at the ends of the arms. The middle
of the distal edge of each joint in the lower part of the arm is slightly raised, and
gradually develops into a keel or crest curving slightly forwards. This may follow
immediately after the median tubercle between the first and second brachials, or not
begin till after the twentieth joint, and varies very much in its development, gradually
becoming less marked towards the ends of the arms. Syzygies in the third and about
the eighth and twelfth brachials, and then at intervals of two to eight joints, usually
about four or six.
The lower pinnules are all of tolerably equal length. That on the second brachial
is about 12 mm. long, rather slender, and consists of some twenty trihedral joints,
most of which are longer than wide. The following pinnules have rather wider and
more flattened joints, with a sharp dorsal edge, and they gradually increase in stoutness
till that on the sixth brachial, which is the largest pinnule on the arm. The next few
pinnules decrease slowly in stoutness but increase in length, their outer joints becoming
relatively longer, but the basal ones remain wide for some distance ; the terminal
pinnules long and filiform.
Disk and ambulacra naked ; sacculi very abundant on the disk, arms, and
pinnules.
Colour in spirit, — very variable ; light brown, purple, or various combinations of
the two, either mottled or in broad or narrow bands ; other specimens are mottled
purple and white.
Disk 11 mm.; spread about 25 cm.
Locality. — Bahia, 7 to 20 fathoms.
Other Localities. — Off St. Lucia (278 fathoms); Venezuela; Pernambuco ; the
Abrolhos Islands; Rio Janeiro; Chile; Java (?); Ceylon; the Seychelles; Muscat;
Aden ; Red Sea ; Zanzibar ; Mauritius ; Madagascar ; St. Helena.
History. — This species has a wide distribution in the littoral zone of the tropical
and the southern subtropical seas ; and it is not improbably therefore identical with
the Alecto carinata of Leach,1 who defined his type very briefly from a specimen sine
patriot, in the British Museum. But the originals of Leach's species are not now to be
found in the national collection, although Professor F. J. Bell has made a careful search
1 Zool. Miscellany, 1815, vol. ii. p. 63.
REPORT ON THE CRINOIDEA. 201
for them ; and the identity of his Alecto carinata with the Comatula carinata from
Mauritius, which was described by Lamarck1 in the following year must therefore remain
uncertain. Lamarck referred to Antcdon gorgonia, de Freminville, as a possible
synonym of his species, and from this one may perhaps conclude that he had been
unable to get access to de Freminville's type. We have seen that he had ignored de
Freminville's generic name Antcdon, which had five years' precedence over Comatula,
and that his definition of this latter type differed but little from that of Antedon
which had been previously given by de Freminville. But the latter author gave no
figure nor formal description of Antedon gorgonia as distinguished from his definition
of the genus ; and if Lamarck was unable to see de Freminville's original specimen we
can understand his uncertainty respecting the possible identity of Comatula carinata
and Antedon gorgonia.
Lamarck's species was redescribed by Midler, and his diagnosis of it was copied by
Dujardin and Hupe in 1862. A few years afterwards Dr. Liitken gave the MS. name
Antedon braziliensis to a type which had been obtained at Rio Janeiro and has since
proved to be very abundant on the Brazilian coast at the Abrolhos Islands and also at
Bahia. Examples of it with Liitken's name attached were distributed to various
museums, and in 1867 the name Antedon braziliensis appeared in a comparative list of
the Echinoderms from the east and west coasts of Tropical America which was drawn up
by Verrill.2 The same author 3 in the following year doubtfully referred to Antedon
diibenii, Bohlsche, another example of this type from the Brazilian coast, which he
regarded, however, as different both from Antedon braziliensis, Liitken, MS., and from
the Antedon carinata of Mauritius and Zanzibar. The difference, however, seems to be
chiefly one of coloration, and it is now practically certain that Yerrill's and Liitken's
types alike are identical with the species from the Indian Ocean. Pourtales 4 wrote
as follows in 1878 : — " A species common on the coast of Brazil answers to the description
of the Comatula carinata Lamk. (Leach sp.). It is quoted as from Mauritius, and the
museum has specimens from Zanzibar differing only in some minor details from the
Brazilian ones." Rathbun,5 writing a few months later, referred to the Brazilian form as
Antedon carinatus (?), and made some comparisons between it and some examples from
Zanzibar, concluding with the remark that " the study of a large series of specimens
would probably serve to unite the BrazUian with the East African species beyond all doubt."
I was fortunately able to carry out this study in the autumn of 1880, when a careful
examination of the material which I found in several continental museums, from a con-
1 Op. cit., p. 534.
2 Trans. Connect. Acad. Arts and ScL, 1867, vol. i. p. 341.
3 Ibid., p. 365.
4 Reports on the dredging operations of the U.S. Coast Survey Steamer " Blake." Corals and Crinoids, Bull. Mus.
Comp. Zool., 1879, vol. v. No. 9, p. 214.
5 A list of Brazilian Echinoderms with Notes on their Distrihution, &c, Trans. Connect. Acad., 1879, vol. v. p. 156.
(zool. chall. bxp. — part lx. — 1887.) Ooo 26
202 THE VOYAGE OF H.M.S. CHALLENGER.
siderable variety of localities, led me to acquiesce in the conclusions of Pourtales and
Rathbun, which were also adopted by Ludwig 1 two years later.
Remarks. — Antedon carinata is thus very extensively distributed in tropical and sub-
tropical seas. Originally described from Mauritius, it has since been found at Ceylon, the
Seychelles, Madagascar, Zanzibar, Muscat, Aden, and in the Red Sea, The British Museum
contains specimens from St. Helena ; it is common all along the South American coast
from Rio Janeiro to Pernambuco, reappears at Venezuela, and was dredged abundantly in
278 fathoms off St. Lucia. As yet it is only known from Chile on the Pacific coast of
America ; and this is further south than any locality on the Atlantic coast at which the
type has yet been obtained. I have a strong suspicion too that an individual from
Norfolk Island, which I saw at Vienna in 1880 with the museum name Antedon
marmorata, is very closely allied to, if not identical with, Antedon carinata; but I
should prefer to leave the point undecided for the present, until I can make a more
detailed examination of the Vienna specimen. In the same year I found some very
typical examples of Antedon carinata in the museum at Hamburg, which were labelled
as having been obtained at Java. This of course is only separated by a part of the
Indian Ocean from Mauritius and the Seychelles ; but if the locality of these four
specimens is rightly given, it is curious that no other examples of Antedon carinata from
the eastern shores of the Indian Ocean should have occurred in any one of the numerous
collections of Coniatuke which I have examined. Thus, for example, it is not represented
in Dr. Anderson's collection from the Mergui Archipelago. It has been recently obtained
at Ceylon, however, but Mr. G. C. Bourne was unable to find any Comatulae at all on
the Coral reefs of the Chagos Islands, which occupy an intermediate position between
Java and the Seychelles, although he was good enough to make a special search for
them on my behalf. Under these circumstances, therefore, I must confess to a certain
amount of doubt respecting the presence of Antedon carinata at Java, as the Hamburg-
label records, and can only wait with interest for further information on the subject.
The characters of the centro-dorsal, arms, and lower pinnules distinguish Antedon
carinata very clearly from the other members of the M ilberti-growp. In fact, as hinted
above, it may become desirable at some future time to remove the type from this group
altogether. The lower pinnules are all of tolerably equal length, and only differ in the
proportions of their component joints. The stoutness of the joints increases up to the
third outer pinnule (on sixth brachial), and the next two or three pinnules are most
frequently almost equally stout, but in a few cases the size of the pinnule-joints decreases
from this point onwards. In full-grown individuals the width of the arm remains
uniform until the second syzygy (eighth brachial), after which the joints become more
triangular, and the width begins to decrease, while the median keel or crest becomes more
1 Verzeiclmiss der von Prut'. Dr. Ed. van Beneden an der Kiiste von Brasilien gesammlten Ecbinodermen, Mem.
Acad. Sci. Bruxelles, 1882, t. xliv. p. 5.
REPORT ON THE CRINOIDEA. 203
distinct. This varies greatly in the extent of its development, and is so slight in some
individuals which I have seen from Mauritius, the locality of Lamarck's original specimen,
that he would most assuredly never have given them the specific name carinata. There
is always more or less of a tubercular elevation on the junction lines of the two outer
radials and the two lower brachials respectively, and from the second of these onwards the
median dorsal line of the arm is more or less sharply indicated, owing to the way in which
the dorsal surface of each joint falls away from it, so that the arm has somewhat the
appearance of having been compressed laterally. The bases of many arms show little more
than this ; but in others the middle of the distal edge of each joint is distinctly raised, and
a sharp forward projecting crest or keel is gradually developed upon it (PI. XXXIV. figs.
4-7), and continues for some way out on the arms, till it becomes less and less distinct in
their terminal portions. In the few specimens which I have seen from Muscat and from
the Red Sea, this character, and also the tubercular elevations on the radials and lowest
brachials are considerably less distinct than in those from the Indian Ocean, Brazil, or the
Caribbean Sea ; while both in the African and in the Red Sea variety the terminal portions
of the arms have stiffer pinnules and a less feathery appearance than in the Brazilian
examples. The sacculi are extremely abundant in this species, and occur in considerable
quantity at the sides of the ambulacra both in disk and arms, which is by no means
always the case in other forms of Antedon. The ambulacra are often supported by delicate
rods and spicules of limestone, but there is never anything like a definite skeleton. The
colour is extremely varied. Some specimens are dark reddish-purple or light yellowish-
brown all over ; others have alternating bands of these two colours, each band covering
two or three arm-joints; in others again the bands are quite narrow, while some individuals
have a more or less mottled appearance, with the brown occasionally replaced by white.
The cirri of Antedon carinata are peculiar from the very general absence of an
opposing spine on their penultimate joint (PI. XXXIV. figs. 1-3). In two individuals,
one from Mauritius and one from Bahia, I have found a cirrus which shows signs of
having been broken and subsequently repaired, the distal portion of it being much smaller
than the base. This is worth recording, because I have generally found that regenera-
tion after fracture, though common enough in the arms, occurs but rarely in the cirri.
The centro-dorsal of this type is very characteristic (PI. III. figs, la, 3b ; PI. XXXIV.
figs. 1-3). It is a thick disk with a single or partially double row of marginal cirri, but
its dorsal surface is smooth and free from cirri, just as in Actinometra (PI. V. figs, lb, 2b,
2e, 46, 5b, 5c), though in young individuals it is more convex, with only a small cirrus-
free space at the dorsal pole. The ventral surface (PI. III. figs, lb, 3a) is marked by an
indistinct pentagonal impression which corresponds to' a similar marking on the under
surface of the radial pentagon (PI. III. fig. lc), and is of interest from its foreshadowing
to a certain extent the deeper bilobed impressions in the corresponding positions on the
centro-dorsal and radials of Antedon quinduplicava (PL IV. figs, lc, Id).
204 THE VOYAGE OF H.M.S. CHALLENGER.
Antedon carinata has a well-developed basal star, as shown in PI. III. figs. \c, 2a,
2b, 3a, 3b; and the articular surfaces of the radials, though relatively wider than is
generally the case in Antedon (compare PI. III. figs. 46, 5a, 6d), are considerably inclined
to the vertical axis of the calyx (PI. III. figs. Id, 3a, 3b); while there is a wide central
funnel, the opening of which is often not filled up by any calcareous network, so that the
ventral surface of the rosette is more or less visible through it (PI. III. figs. Id, 3a).
A few comparatively young specimens were obtained by the Challenger at Bahia.
They differ from the more mature individuals in the greater length of the arm- and
cirrus-joints, and in the more convex shape of the centro-dorsal, but a small portion of
which is free from cirri (PL XXXIV. fig. 3).
One or two examples of Antedon carinata which were taken at Bahia were infested
by Myzostoma gigas, which is also a common parasite of Antedon eschrichti in the
Circumpolar Seas.
5. Antedon parvicirra, n. sp. (PI. XXXVI. figs. 7, 8).
be
Specific formida — A.— .
Description of an Individual. — Centro-dorsal small and hemispherical, bearing
about forty cirri of ten or twelve joints, all but the lowest of which are longer than
wide, the terminal ones being somewhat compressed laterally ; the penultimate with a
small opposing spine.
First radials scarcely visible ; the second very short and quite free laterally.
Axillaries four times their length, and pentagonal, with a faint median elevation on
the proximal edge. Ten arms of about eighty joints, which are triangular at first, and
about as long as wide, gradually becoming obliquely quadrate. Syzygies in the third,
eighth, and about the twelfth brachials, and then at intervals of two or three joints.
The second brachial bears a slender pinnule about 4 mm. long, which consists of
some fifteen joints. The following pinnules increase gradually both in length and in
stoutness to those of the third pair (on sixth and seventh brachials), which bear long and
fusiform genital glands. The succeeding pinnules are of nearly the same length, and
then gradually diminish in stoutness.
Disk and ambulacra naked ; sacculi very abundant on the disk, arms, and pinnules.
Colour in spirit, — purplish-red, with frequent intervals of white on the arms.
Disk 6 mm.; spread about 10 cm.
Locality.— Station 208, January 17, 1875; lat. 11° 37' N., long. 123° 31' E.;
1 8 fathoms ; blue mud. One specimen.
Remarks. — The short cirri of this little species, with their somewhat compressed
terminal joints, together with the freedom of the rays and the relatively long joints of
REPORT ON THE CRINOIDEA. 205
the lower pinnules, give it a certain amount of similarity to Antedon rosacea and
Antedon diibeni (PI. XXXVII. figs. 1, 2). But it resembles Antedon carinata in the
large size and the tolerable equality of the pinnules on the sixth and following brachials
(PI. XXXVI. fig. 8), a point which distinguishes it altogether from Antedon rosacea and
Antedon diibeni, in which the first pinnule is the longest (PL XXXVII. fig. 3). The
small number of cirrus-joints separates it from Antedon anceps and Antedon variipenna,
which somewhat resemble it in the characters of the pinnules (PL XXXV. figs. 1-3 ;
PL XXXVI. figs. 1, 4-6).
6. Antedon informis, n. sp. (PL XXXIII. fig. 3).
Spec ific formula — A. -r .
Description of an Individual. — Centro-dorsal discoidal, with a smooth dorsal surface
and about a dozen marginal cirri. These have fifteen to eighteen joints which are as
wide or wider than long, most of them with a slight elevation in the middle of the dorsal
edge ; the penultimate with a faint spine.
First radials partially visible ; the second oblong, with a rounded dorsal surface, and
but slightly united laterally. Axillaries also rounded, short, and widely rhombic. Ten
arms ; the first few brachials nearly oblong ; the following ones rather wider than long,
somewhat overlapping, and almost triangular, gradually becoming oblicpiely quadrate. A
syzygy in the third brachial, and the next usually about the eleventh or twelfth, with
others at intervals of three to five joints.
The second brachial bears a comparatively small pinnule of about a dozen squarish
joints. There may be a similar but smaller one on the third brachial, or more generally
none at all. That on the fourth is considerably longer and stouter, but the following-
pinnules are smaller again.
Disk lost ; sacculi very abundant on both arms and pinnules.
Colour in spirit, — white.
Spread perhaps 8 cm.
Locality.— Station 208, January 17, 1875; lat. 11° 37' K, long. 123° 31' E.;
1 S fathoms ; blue mud. One imperfect specimen.
Remarks. — Of the five mature arms which remain in this much mutilated individual,
only one, the central one in the figure (PL XXXIII. fig. 3), has a pinnule on the third
brachial, and that but a small one. In the other four arms it is entirely absent, and in
one ray which has been completely regenerated there are no pinnules on the third, fifth,
and seventh brachials, although those on the other (outer) side of the arm are all present
as usual.
There is a similar absence of a pinnule on the third brachial in the unique specimen
206 THE VOYAGE OF H.M.S. CHALLENGER.
of Antedon loveni,1 which was described by Bell, though the fact seems to have escaped
his notice, for he makes no mention of it.2 The cirri and one arm are lost; but only one
of the remaining nine arms has a pinnule on the third brachial. The great size and
the elongated joints of the second and third pinnules in Antedon loveni are sufficient,
however, to prevent any confusion between it and Antedon informis.
Another species in which the third brachial bears no pinnule is Antedon manca
(PI. XLIV. figs. 2, 3), which will be described further on in the Bidistichate group.
Apart from the absence of the pinnule on the third brachial, Antedon informis is also
distinguished by the long interval between the first and second syzygies,3 and by the
peculiar minute spinelets on the cirrus-joints. I know of no described species with which
it is likely to be confounded.
All the ten-armed species of Antedon which were obtained by the Challenger and
" Porcupine " have now been considered, with one exception. This is the singular form
which I have called Antedon balanoides (PI. XXXIII. figs. 6, 7); and there are four
other species besides it which do not seem to fit into any of the groups established above.
All but one inhabit the Eastern Archipelago, and for the present they may be classified
as follows : —
A. The second and third brachials have pinnules.
I. The first pinnule is the largest. Cirrus-joints have two dorsal
spinelets, ....... bidens, Bell.
II. The lower pinnules tolerably equal.
Twenty cirrus-joints without spines ; syzygial interval three or
four joints, ...... adeems, Lamk., sp.
Twenty-five to thirty-five spiny cirrus-joints ; syzygial interval
nine or ten joints, ...... Ixvijnnna, Carpenter.
B. The second and third brachials have no pinnules.
Sixty cirri of thirty-five to forty joints on a conical centro-dorsal, . 1. balanoides, n. sp.
Twenty cirri of about twenty joints, ..... defecta, Carpenter, MS.
C. The third, fourth and fifth brachials have no pinnules. Eight or ten
cirri of twelve joints, ...... impinnata, Carpenter, MS.
The last of these is a little species which was obtained at Mauritius by Professor
Mobius, who was kind enough to show it to me when I visited Kiel ; and I found it to
be serving as host to Myzostoma excisum and Myzostoma carinatum, von Graff.
1 " Alert " Report, p. 158, pL x. fig. C (non A).
2 Bell's figure is also incorrect ; for the pinnules of the second, fourth, and sixth brachials are represented as being
placed on the inner instead of on the outer side of the arm. A similar error occurs in the figure of Antedon ■pumila on
the same plate, in which the first, second, fourth, and sixth, &c, brachials are all represented as bearing pinnules on
the inner side of the arm, an arrangement which never occurs in any Crinoid.
3 The central arm represented in the figure has a syzygy in the fourth brachial as well as in the third.
REPORT ON THE CRINOIDEA. 207
1. Antedon balanoides, n. sp. (PL XXXIII. figs. 6, 7).
Specific formula — A.y-
Description of an Individual. — Centro-dorsal a much elongated cone (5 '5 mm.),
bearing five converging double rows of cirrus-sockets, six or seven in each row. Its
lower portion has no functional sockets, but shows faint scars continuing the rows
downwards ; a deep interradial furrow between every two double rows.
Thirty-five to forty cirrus-joints, of which the basal ones are short. The next till
the tenth or twelfth are much longer than wide, and develop a faint dorsal keel sloping
backwards from the distal edge, where it projects beyond the base of the next joint ; the
following joints are shorter and become compressed laterally.
First radials concealed ; the second just in contact at their proximal angles, and
oblong, with a deeply incised distal edge, the centre being very short and raised to meet
the elevated blunt angle of the rhombic axillary.
Ten arms ; the first brachials have long outer sides and a short raised centre ; the
second irregularly quadrate and similarly raised in the centre. Arm-joints triangular,
wider than long. Syzygies about the third, eighth, and twelfth brachials, and then at
intervals of two to six, usually three, joints.
The second and third brachials have no pinnules ; the fourth bears a slender one of
fifteen joints and 7 mm. long. The next pair are generally stouter, and the following
ones less so, but of increasing length.
Disk naked ; sacculi tolerably abundant on the pinnule-ambulacra.
Colour in spirit, — the calyx a deep rose red ; the cirri and the ends of the arms white.
Disk about 6 mm. ; spread probably about 1 2 cm.
Locality.— Station 201, October 26, 1874; lat. 7° 3' N., long. 121° 48' E. ; 82
fathoms ; stones, gravel. A calyx and some arm-fragments.
Remarks. — Only a single and much mutilated example of this remarkable type was
obtained by the Challenger. Its centro-dorsal is not unlike that of Atelecrinus in shape
(PI. VI. figs. 5, 7 ; PI. XXXIII. fig. 6), but the cirri are more numerous and the double
rows more regularly arranged than in that genus. These characters also distinguish the
type from Antedon defecta of the above list, which was dredged by the " Blake " in the
Caribbean Sea.
The list given on p. 54 also includes six other ten-armed species of the " Blake " collec-
tion, viz. : —
Antedon armata, Pourtales.
Antedon brevipinna, Pourtales.
Antedon columnaris, Carpenter.
Antedon cubensis, Pourtales.
Antedon duplex, Carpenter, MS.
Antedon hageni, Pourtales.
208 THE VOYAGE OF H.M.S. CHALLENGER.
I have not inserted these in any of the above schemes, since my work on the " Blake "
Comatula? is not yet complete ; but they will be fully described and illustrated in the
Report on the Collection, which will be published as one of the Memoirs of the Museum
of Comparative Zoology at Harvard College.
Antedon, Series III.
Two articulated distichals.
Remarks. — This series includes all those multibrachiate species in which there are but
two distichal joints united by a bifascial articulation as the two outer radials are (PL XL.
fig. 1 ; PL XLIIL). The palmar and post-palmar series, when present, are normally of
the same character, though, like the distichals, they may be occasionally replaced by a
three-jointed series with a syzygy in the axillary. This, however, is far less common
than in Actinometra. The individuals of Antedon similis and Antedon occulta
represented in PL XL VII. fig. 3, and PL XLVIII. fig. 1, have their distichal and palmar
series very regularly constructed ; while in those of Actinometra elongata and of
Actinometra valida (PL LVII. fig. 2 ; PL LIX. fig. 3) two and four of the bidistichate
series respectively are replaced by sets of three joints, the axillary with a syzygy.
The bidistichate forms of Antedon make up the largest series, next to that of the
ten-armed type, of all those into which the species of this genus naturally arrange them-
selves. It may be divided for the purposes of formulation into three groups, according
as there are one, two, or three axillaries respectively above the radials, as is shown on
pp. 54, 55. Several species of course appear in two groups, owing to the occasional
total absence of palmar or post-palmar axillaries in some individuals ; while others may
sometimes have only ten arms. Thus, for example, there are now and then no distichal
series at all in certain individuals of Antedon brevipinna, Antedon duplex, Antedon
Jlexilis, and Antedon lusitanica (PL XXXIX. figs. 1, 3). In like manner Antedon
pourtalesi and Antedon quinquecostata always have one and sometimes two post-radial
axillaries ; while there are always two and sometimes three in Antedon palmata and
Antedon spinifera.
While, therefore, it is useful to have a system of formulation which gives some
information as to the extent of the ray-divisions, too much stress must not be laid upon
the presence of a palmar or post-palmar axillary as an aid to classification. In the list
given on pp. 54, 55, I have arranged the bidistichate species of Antedon in three groups,
according to the frequency of the ray-divisions. There are certain occasions in which
such a mode of grouping them is of considerable use ; but one must always bear in mind
that a species with (say) three post-radial axillaries which appears new at first sight, may
in reality be identical with one which is already known, but has never yet been found
with any axillary beyond the palmars, e.g., Antedon tuberculata (PL XLV. fig. 2). The
&
EEPORT ON THE CRISTOIDEA. 209
converse is also equally true. All the three Challenger specimens of Antedon occulta
have post-palmar axillaries (PI. XLVIII. fig. 1 ; PI. XLIX. fig. 3), but it is quite
possible that other examples of the type may be eventually discovered in which these
are absent. It would be premature, however, to describe them as new, simply because
they did not agree with any of the species which had no axillary beyond the palmars.
In all such cases as these, and we shall meet with them frequently, the general characters
of the type must be carefully taken into consideration, apart from the frequency of its
ray-divisions, which is often much greater in some individuals of a species than in others ;
thus, for instance, examples of Actinometra parvidrra have been described with thirteen
and with thirty-nine arms respectively, palmar axillaries being altogether absent in the
former and abundantly developed in the latter.
It is therefore desirable, so far as may be possible, to employ some other means of
classifying any particular series of multibrachiate Comatulaa than one which is based
solely on the number of post-radial axillaries ; and in the present case there is no difficulty
in effecting this object.
We have seen that the bidistichate species of Antedon fall into three sets according
as there are one, two or three axillaries above the radials. Each of these sets contains
species which belong to two very different types of structure, the one with and the other
without an ambulacral skeleton. Most of the forms which have a distichal but no palmar
axillary (A. 2) resemble the members of the Basicurva-gvou]) in the ten-armed series in
the presence of an ambulacral skeleton and in the straight-edged and wall-sided nature
of the radial axillaries and of the next following joints. In fact, two members of this
group (Antedon flexilis and Antedon lusitanica) have been already noticed in connec-
tion with the Basiciirva-growp, owing to the occasional absence of any distichal axillary
(PL XXXIX. figs. 1-3 ; PI. XLIL). In like manner Antedon quinquecostata and
Antedon pourtalesi always have a number of distichal series, but these are not always
followed by palmars ; whde I have seen some examples of Antedon spinifera with one or
more post-palmar series, and others in which there are no axillaries beyond the palmars.
All these species, together with others which will be described immediately, are the
bidistichate representatives of the Basicurva-group, and they form a very natural
assemblage which may be conveniently designated as the Spinifera -group. It thus includes
a variety of specific types which may have one, two, or three axillaries beyond the
radials, Antedon spinifera having sometimes two and sometimes three, and being the first
species in which an ambulacral skeleton was described. The remainder of the bidisti-
chate species of Antedon, which have neither an ambulacral skeleton on the pinnules nor
distinctly wall-sided rays, represent the 3£ilberti-gvou\) among the ten-armed species of
the genus in the characters and relations of their lower pinnules, and they may be
conveniently designated as the Pa hna to-group. Like the Sjnnifera-grou^ it includes
species with one, two, or three post-radial axillaries, but one or more of the pinnules
(ZOOL. CHALL. EXP. — PART LX. — 1887.) OoO 27
210 THE VOYAGE OF H.M.S. CHALLENGER.
oft the fourth and following brachials is considerably stouter and often much longer than
its predecessors, very much as in the MUfoerti-gromp of the ten-armed series.
This is very well shown in Antedon occulta and in Smith's figure of Antedon indica ; 1
aud the distinctions between the different species of the P«/?)i«.ta~group depend very largely
upon the number and position of these large pinnules, as shown in the key on p. 225.
Taking the Bidistichate Series as a whole, we find that its distribution, as at present
known, is of a somewhat limited character. Unless the bklistichate example of Antedon
lusitanica, which was dredged by the "Porcupine" from 740 fathoms in the East Atlantic, is
anything more than a mere individual variation, there is no certain evidence of the Series
being represented below 269 fathoms. The Challenger may have obtained Antedon similis
from 610 fathoms at Station 174, but I think it more probable that the depth for this
species and for the two associated with it was either 210 or 255 fathoms ; for the only
other Challenger station which yielded bidistichate species from below 100 fathoms was
No. 192 (140 fathoms) ; though they range down to 269 fathoms in the Caribbean Sea,
where they occur in considerable quantity. They are excessively abundant between the
meridians of 100° and 180° E. (Sumatra to Fiji), and one species occurs at Rodriguez.
But with the exception of the doubtful Antedon lusitanica, none have been found in the
open Atlantic, nor are they known anywhere outside the fortieth parallels of latitude.
The Caribbean Sea and the Eastern Archipelago, therefore, are their two principal locali-
ties; and it may be broadly stated that the species with plated ambulacra (5pm?yer«-group)
predominate in the former, while those with unprotected ambulacra and large stiff pinnules
on the fourth and following brachials (Palmat a- gvowp) are mostly confined to the Eastern
Seas. These are the only bidistichate species which belong to the strictly littoral fauna,
i.e., which have been found at depths of 20 fathoms or less. Three of them (Antedon
occulta, Antedon similis, and Antedon tuberculata) were dredged by the Challenger at
Station 174b,c, or r> (255, 610 and 210 fathoms). The depth was probably one of the two
lesser ones ; but either of them is considerably below the usual range of the Pahna ta-gronp.
With the exception of the aberrant Antedon macronema from South-eastern Australia,
no members of the Sjnnifera-grou]} have been met with above 80 fathoms either in
the Caribbean Sea or in the Eastern Archipelago. The " Blake " dredged them at
some twenty stations in the Caribbean Sea between 80 and 270 fathoms ; but all the
five typical species of the Challenger collection were obtained from 140 fathoms in
the Arafura Sea (Station 192), though one of them also occurred at Station 201 among
the Phdippine Islands (82 fathoms). The remaining species are Antedon lusitanica,
dredged by the "Porcupine" from 740 fathoms in the East Atlantic, and Antedon
macronema from 30 fathoms or less in Kingj George's Sound, Port Jackson, and Port
Stephens. The latter, however, is an abnormal species in many respects. The lateral
flattening of its rays is very variable in its extent, and never specially distinct ; while the
1 Zoology of Rodriguez ; Ecliinodermata, Phil. Trans., 1679, vol. clsviii. pi. li. fig. Zb.
REPORT ON" THE CRINOIDEA. 211
ambulacral skeleton of the pinnules, though well-developed as compared with that of
species which have no covering plates at all, is far less highly differentiated than in
Antedon jiexilis and in Antedon spinifera itself, which have more distinct side plates
than many species of Pentacrinidaa. It is worth notice that with the single exception of
Antedon denticulata from 49 fathoms, no member of the Basicurva-gcoup was obtained
by the Challenger from a less depth than 140 fathoms, while they range as far down-
wards as 1600 fathoms. The bidistichate species with the same characters of the rays
and ambulacra range from 80 to 740 fathoms, so far as is yet known ; and the tridistichate
species, which also have flattened rays and plated ambulacra, are likewise almost
exclusively limited to the continental and abyssal regions. These facts are of interest
because the Pentacrinida?, which also have an ambulacral skeleton on the pinnules, do
not occur at depths of less than 70 fathoms and range down into the abyssal fauna ;
and we may therefore not unreasonably infer that fossfl species like Millericrinus pratti,
which have the ambulacral skeleton still preserved on the pinnules, lived at depths of at
least 50 fathoms. The same conclusion may perhaps be drawn for those Comatulse such
as Solanocrinns costatus, Goldfuss, in which the axillary radials and the lower brachials
are very distinctly flattened and wall-sided.
The following key to the Spinifera-growp contains the names of four Caribbean species,
two of which, Antedon brevipinna and Antedon spinifera, have been described by
Pourtales and myself respectively.1 Antedon duplex is one of the hosts of the encysting
Myzostoma murrayi, von Graff, which also occurs on Antedon breviradia and Antedon
angustiradia of the Challenger collection, both species from the Eastern Archipelago.
Antedon pourtalesi is a fine species which I have dedicated to the memory of the late
Count Pourtales, and is the host of Myzostoma brevipes, von Graff.
6. The Spinifera-grovq)).
Bidistichate species with the radial axillaries and some of the following joints more
or less wall-sided, and a well-marked ambulacral skeleton on the pinnules : —
A. Over thirty cirrus-joints ; the later ones spiny.
I. The first pinnule much smaller than the second. Centro-dorsal a thick
disk or low rounded column, bearing two or three tiers of cirrus-
sockets usually without definite arrangement ; .eighty or ninety
cirrus-joints. First radials completely visible, . . . 1. macronema, Mull, sp.
II. The first pinnule as long as or longer than the second.
a. Centro-dorsal shortly columnar, with five double rows of cirrus-
sockets, separated by interradial ridges.
1. Twenty arms of sharply carinate joints; eighty ciirus-
joints or more, . . . . . .2. quinquecostata, n. sp.
2. Thirty arms, their joints bearing curved dorsal spines ;
forty to sixty cirrus-joints, .... spinifera, Carpenter.
1 Bull. Mus. Comp. Zool., 1867, vol. i. No. 6, p. Ill ; Ibid., 1881, vol. ix. No. 4, p. 8.
212 THE VOYAGE OF H.M.S. CHALLENGER.
b. Centro-dorsal more rounded ; the cirri without definite arrange-
ment.
1. Thirty to forty cirrus-joints; the radial axillaries long;
lower joints of the genital pinnules expanded, . . duplex, Carpenter, MS.1
2. About fifty cirrus-joints ; the radial axillaries short and
wide, . . . . . . .3. lusitanica, n. sp.1
B. Fifteen to twenty-five stout and usually smooth cirrus-joints.
I. Centro-dorsal shortly columnar or conical, with five double rows of
cirrus-sockets.
a. First radials visible; the arm-bases smooth and rounded, . 4. flexilis, n. sp.1
h. First radials entirely concealed ; the lower arm-joints have raised
distal edges, . . . . . . -5. patula, n. sp.
II. Cirri without definite arrangement.
a. The distichals and lower brachials have distinctly flattened sides.
The later cirrus-joints smooth.
1. Calyx and arm-bases almost smooth; lower joints of the
genital pinnules not specially distinguished, . . 6. robusta, n. sp.
2. Calyx and arm-bases irregularly tubercular.
a. The pinnules from the tenth to twentieth brachials
have the third to fifth joints flattened and expanded
laterally, ..... pourtalhi, Carpenter, MS.
/3. The genital pinnules comparatively slender, with
very slightly expanded joints, . . . brevipinna, Pourtales.1
b. The distichals and lower brachials are but slightly flattened
laterally ; blunt spines on the later cirrus-joints, . . 7. compressa, n. sp.
1. Antedon macronema, Mull., sp. (PI. IV. figs. 3, a-d; PI. XXXVIII. figs. 4, 5).
Specific formula — A. 2.—.
1846. Comatula macronema, Miiller, Monatsber. d. k. preuss. Akad. d. Wiss. Berlin, 1846,
p. 179.
1849. Comatula macronema, Miiller, Abhandl. d. k. Akad. d. Wiss. Berlin, Jahrg. 1847
[1849], p. 258.
1862. Comatula macronema, Dujardin and HupcS, Hist. Nat. des Zoophytes, Echinodermes,
Paris, 1862, p. 203.
1879. Antedon macrocnema, P. H. Carpenter, Trans. Linn. Soc. Lond. (Zool.), ser. 2, 1879,
vol. ii. p. 29.
1880. Antedon macrocnema, P. H. Carpenter, Journ. Linn. Soc. Lond. (Zool.), 1880, vol. xv.
p. 213, pi. xii. figs. 25, a-c.
1882. Antedon macronema, Bell, Proc. Zool. Soc. Lond., 1882, p. 534.
1882. Antedon macronema, P. H. Carpenter, Ibid., p. 746.
1885. Antedon macronema (misprinted mauonema), Bell, Proc. Linn. Soc. N.S.W., 1884
[1885], vol. ix. p. 497.
Centro-dorsal a thick disk or short column with a smooth dorsal pole and about
twenty-five to thirty-five cirri on its sides. These are very slender and reach 45 mm. in
length, the longest having nearly one hundred joints, most of which are wider than long.
1 Some individuals of these species have only ten arms ; see p. 54.
REPORT ON THE CRINOIDEA. 213
The middle and outer joints are laterally compressed so as to overlap dorsally, and
gradually develop a small spine.
The ends of the basal rays are often visible above the angles of the centro-dorsal.
The first radials are nearly oblong ; the second rather longer and more convex in the
centre. Their lower ends are united, but the outer ends are free with sharp lateral edges.
Axillaries pentagonal, also much rounded in the centre, with sharp straight edges and
small flattened sides. The rays may divide twice ; two distichals, the axillary not a
syzygy. The distichals and lower brachials are rounded like the radials, with similar
straight edges and small, flattened, outer sides. The inner side of the third brachial may
also show traces of flattening.
Eleven to fifteen short arms, consisting of about seventy shortly triangular joints, the
lower ones rounded ; the later joints gradually become compressed laterally and develop
a forward projecting spine which overlaps the base of the next joint and is much more
distinct in some individuals than in others. A syzygy in the third brachial, and another
between the tenth and twentieth. Others at intervals of three to eight joints, usually
four or five. The first pair of pinnules (on second and third brachials) are short and
stiff, consisting of eight or nine elongated joints, and not more than 5 mm. long. The
next pair are much stouter, with about a dozen rounded joints, and reach 10 mm. long.
The following pinnules are of about the same length, with broader and more flattened
basal joints, which are sometimes slightly carinate. There are six or eight pinnules of
this character on either side of the arm, but the greater width of the lowest joints is
distinct for some distance further, and the length of the pinnules does not increase.
Disk and brachial ambulacra but slightly plated ; the genital glands protected by an
imperfect plating which supports the ambulacral skeleton. This is not well differen-
tiated ; the covering plates rest on a continuous limestone band which is scarcely
segmented into side plates. Sacculi abundant on the arms and pinnules, with a few on
the disk.
Colour in spirit, — the calyx nearly white, and the arms a darkish purple.
Disk 7 mm.; spread about 13 cm.
Locality. — Port Jackson ; 30 to 35 fathoms.
Other Localities. — King George's Sound ; Port Stephens.
Remarks. — This is one of the most isolated, and therefore among the most easily
identified species of the genus.
Although the lower joints of the rays have sharp and straight edges as in the
Basicurva-gvou]), their dorsal and ventral surfaces approach one another so rapidly that
they have but very small lateral faces. Such as they are, however, these faces have the
usual flattened character, and as there is an ambulacral skeleton on the pinnules, the type
clearly belongs to the Spinifera-growp, rather than to that of Antedon palmata, Antedon
214 THE VOYAGE OF H.M.S. CHALLENGER.
flagdlata, &c, which it somewhat resembles in the very small size of the first pair of
pinnules as compared with their successors (PI. XXXVIII. fig. 4). The characters of these
pinnules, the shortness of the arms, the large number of cirrus-joints, and the presence
of the first radials externally, together with the very slightly wall-sided character of the
lower joints of the rays, render it very easy to recognise the type, which has really no
close allies among the other bidistichate species.
The disk is scarcely plated at all, and the brachial ambulacra but slightly so. The
genital glands contained in the expanded portions of the large lower pinnules are covered
by an imperfect pavement of ill-defined plates, above which the ambulacra are situated.
The covering plates are tolerably distinct, but the limestone band supporting them is
scarcely differentiated into side plates, except in some of the later pinnules.
The position of the side plates, however, is indicated by the sacculi, which are
also abundant on the brachial ambulacra and extend down on to the outer part of the
disk.
The centro-dorsal of Antedon macroncma varies considerably in its shape. Most
commonly it is a thick disk with a smooth dorsal surface and the cirrus-sockets arranged
irregularly on its sides, as shown in PL IV. fig. 3a ; but it is sometimes more nearly
hemispherical, and sometimes almost columnar, with the sockets disposed in alternating
vertical rows of three or four each.
A similar series of variations in the form of the centro-dorsal is characteristic of
Antedon scrobieulata from the Oxfordian of the Jura ; and the whole aspect of the calyx
of Antedon macronema is more similar to that of the Jurassic Antedon costata and
Antedon gresslyi than that of any other recent Antedon which I have seen. The great
difference between the fossil and the recent types is that the basals of the latter undergo
metamorphosis into a rosette ; while in the former they persist as prismatic rods between
the radials and the centro-dorsal. The positions of these are occupied in the recent form
by the rays of the basal star (PL IV. fig. 3c), the ends of which sometimes appear on
the exterior of the calyx (PL IV. fig. 3a). The general characters of the radials of
Antedon macronema have been already described on pp. 23 to 26, and it is not necessary
therefore to refer to them again.
The type was first discovered by Quoy and Gainiard in King George's Sound, and the
Challenger dredged it in Port Jackson, while there are examples in the Sydney Museum
from Port Stephens. I have never heard of its occurrence at Port Philip, however, though
I have seen various other Comatulse from that locality, where its presence might naturally
be expected.
Apart altogether from its resemblance to certain Jurassic Comatulse, Antedon
macronema is remarkable as being a link between the species with the rays flattened
laterally and an ambulacral skeleton on the pinnules, and those in which these characters
do not present themselves, even as slightly as they do in Antedon macronema ; while its
REPORT ON THE CRINOIDEA. 215
extension into the purely littoral fauna is quite exceptional, for the members both of
the Basieurva-growp and of the Spinifera-groixp are almost entirely confined to the
continental and the abyssal regions.
2. Antedon quinquecostata, n. sp. (PL III. figs. 6, a-d; PL XXXVIII. figs. 1-3).
Specific formula — A. 2. (2).—.
Centro-dorsal a shortly pentagonal column with five interradial ridges veiy
prominent at their ventral ends, and separated by more or less alternating double rows
of cirrus-sockets, three or four in each row. About thirty-five cirri which may reach
45 mm. long, with eighty to ninety joints. Some of the lower joints are much longer
than wide, but the following ones become shorter and laterally compressed, with a sharp
dorsal keel, which passes into a prominent spine in the short middle and later joints.
The angles of the first radials are just visible, being turned slightly outwards above
the interradial processes of the centro-dorsal ; the second are short and sharply convex,
rising to meet a strongly carinate backward process of the widely rhombic axillaries.
The rays may divide three times. Distichal and palmar series of two joints each, the
axillary not a syzygy. The two joints of each series, like the two outer radials, have
sharp median crests, which are continued out on to the arms. All these joints, and
especially those at the outer side of the ray, have straight lateral edges and flattened
sides. On some arms this character ceases at the second brachial, but in others it is
very visible on the third and even on the fourth.
There are usually twenty arms (but one example has twenty-one), with a sharp
median keel, and composed of one hundred and twenty or more joints, which become
much compressed laterally so that the later ones overlap rather sharply. A syzygy in
the third, and then not till the twentieth or twenty-fifth brachial ; others at intervals of
four to eleven (usually five to seven) joints.
The second brachial bears a moderately stout pinnule about 10 mm. long, and
consisting of some twelve or fifteen joints, most of which are longer than wide. The
first four or five are flattened on the outer side, where they meet the corresponding
pinnules of adjacent arms, and their inner edges are also slightly cut away. The
following pinnules are rather shorter, with more rounded joints, the two joints at the base
being wider than then- fellows on the lower part of the arm. Further out, however, this
is less marked, and the pinnules are somewhat carinate, but never specially long.
Disk moderately plated, and the arms rather more so ; the pinnule ambulacra have
covering plates and partly differentiated side plates. Sacculi rare or absent altogether.
Colour in spirit, — the skeleton yellowish-brown or whitish-brown, but the perisome
darker.
216 THE VOYAGE OF H.M.S. CHALLENGER.
Disk 7 mm.; spread about 20 cm.
Locality. — Station 192, September 26, 1874; near the Ki Islands; lat. 5° 49' 15" S.,
long. 132° 14' 15" E.; 140 fathoms; blue mud. Eight specimens and two fragments.
Remarks. — This remarkable species is very readily distinguished from every other
bidistichate Antedon, with the exception of Antedon spinifera from the Caribbean Sea,
Both types alike have long and spiny cirri ; but those of Antedon quinquecostata are
both more numerous and reach a greater length than in the Caribbean species. The
latter also has a shortly columnar centro-dorsal, with double rows of cirrus-sockets,
though there are frequently only two sockets in each row and not four, as is so often the
case in Antedon quinquecostata (PI. III. fig. 6cZ).
In Antedon spinifera too, the radials, distichals and lower brachials are by no means
so sharply carinate as in Antedon quinquecostata, and there is an alternating double
row of strong curved spines on the base of each arm. Generally also there are thirty
arms, owing to the presence of two palmar axillaries on each ray, while the number in
Antedon quinquecostata is typically twenty, though I have found a single palmar
series to be present in two separate individuals (PI. XXXVIII. fig. 1). Another point
of difference between the two species is that Antedon spinifera has very large and
abundant sacculi, while, if present at all, they are most scantily developed in
Antedon quinquecostata, as is also the case in some other species from the same
station.
The lateral flattening of the radial axillaries and of the next following joints is less
marked in this species than in the Basicurva-gvoup. The first distichal and the first
brachial, especially the two on the outer arms of each ray, are the joints which show it
most distinctly ; but it is sometimes to be traced as far as the fourth brachials of the
outer arms. The pinnules of adjacent second brachials, however, have their lower joints
flattened against one another, very much as in Antedon valida (PI. XV. figs. 5, 6),
though not cmite to the same extent. The two lower joints of the next few pinnules
are somewhat wider and more expanded than their fellows, but this feature disappears
in those further out on the arm (PI. XXXVIII. figs. 2, 3).
The radial pentagon of this type differs somewhat in character from that of most
other species of Antedon (PI. III. figs. 6, a-d). Its angles are produced outwards to
correspond with the interradial ridges of the centro-dorsal, each of which fits into a
notch between the everted lateral angles of two adjacent radials ; and under ordinary
circumstances these angles are the only parts of the first radials which are visible
externally. The dorsal surface of the radial pentagon is remarkable for showing no
signs of any basal star, as there is a very well developed one in Antedon spinifera. The
central opening is relatively large, and the rosette within it rather ill-defined, an
unusual condition in a tropical species of Antedon.
REPORT ON THE CRINOTDEA. 217
3. Antedon lusitanica, n. sp. (PI. XXXIX. figs. 1-3).
Specific formula — A. (2).—.
Locality.— H.M.S. "Porcupine," 1870, Station 17a; lat. 39r 39' N., long. 9° 39' W.;
740 fathoms ; bottom temperature, 49°"3 F.
Remarks. — The ten-armed form of this species has been already described on p. 109 ;
but its bidistichate variety finds a place here (PI. XXXIX. fig. 1). It resembles
Antedon quinquecostata and Antedon spinifera in having spiny and many-jointed cirri,
but they show no traces of any definite arrangement as is the case in those two species ;
while it differs from Antedon duplex, another Caribbean species, in the shape of the
axillaries and in the unmodified character of the genital pinnules, so far as can be
determined from the condition of their fragmentary remains.
4. Antedon jiexilis, n. sp. (PI. XLIL).
Specific formula — A. ( 2 ). ^.
Centro-dorsal columnar or slightly tapering, with its ventral angles produced into
marked interradial processes, and ten vertical rows of cirrus-sockets, three or four in each
row. Thirty to thirty-five cirri of twenty to twenty-five stout, but very smooth joints,
most of which are longer than wide ; the penultimate with an opposing spine.
Three radials visible ; each of them, and especially the axillary, rather sharply
convex, with a more or less distinct median tubercle. Axillaries subhexagonal and
considerably wider than the second radials, which are short and band-like and in close
contact laterally.
Ten to thirteen arms ; two distichals, the axillary not a syzygy. The first two
brachials or the two distichals, if present, have median tubercles like those of the radials,
but less prominent. The radial axillaries and the next two joints have sharp, straight
edffes and wall-like sides. The inner sides of the second and of the third brachial
(hypozygal) are also flattened, especially if the distichals are absent. The lower arm-
joints are somewhat discoidal, with very rounded surfaces ; and the following ones are
shortly triangular, with the edges slightly raised, but they gradually become more
smooth and relatively longer. A syzygy in the third, and another between the twelfth
and eighteenth brachials; others at intervals of three to seven, usually four or five
joints.
The first pinnules, which reach 13 mm. in length, have their outer sides somewhat
flattened against one another below, and consist of some forty short and wide
(ZOOL. CHALL. EXP. PAUT LX. — 1887.) 0°° 28
218 THE VOYAGE OF H.M.S. CHALLENGER.
joints. The following pinnules are of about the same length, but the joints gradually
increase in size and diminish in number, still remaining much wider than long. Tn the
pinnules of the fifteenth and several of the following brachials the fourth and fifth joints
are considerably wider than their fellows, but in the later pinnules the joints are longer
than wide.
Disk not much plated, except along the ambulacra. There is a strong covering
of plates over the genital glands, with numerous sacculi imbedded in it ; and the later
pinnules have a well-defined ambulacra! skeleton, the sacculi alternating with the side
plates.
Colour in spirit, — a light whitish-brown, with a brownish-gre}?- ventral perisome.
Disk 10 mm.; spread about 55 cm.
Locality. — Station 192, September 26, 1874 ; near the Ki Islands ; lat. 5° 49' 15" S.,
long. 132° 14' 15" E.; 140 fathoms ; blue mud. Four specimens.
Remarks. — Two of the four representatives of this fine species have but ten arms
each, and they thus find a place in the Basicurva -group, as has been already noticed on
p. 128. A third, that figured on PI. XLIL, has three distichal axillaries on separate
rays ; while the fourth has one normal distichal series and two others on a ray which
has been regenerated from the second radial.
The tendency of the centro-dorsal to assume a columnar shape, and the arrangement
of the cirri upon it in five double rows beneath the rays, are points of resemblance
between this species and Antedon quinquecostata from the same locality (PI. XXXVIII.
fig. 1); but they are very different in other respects. Antedon quinquecostata has slender
and spiny cirri composed of many joints ; while those of Antedon fiexilis consist of but
twenty-five smooth and relatively stout joints. The tubercular nature of its rays and
arm-bases is also a good distinctive character ; while it has extremely abundant sacculi,
although these structures are most scantily developed or even altogether absent in Antedon
quinquecostata. Not only do they alternate with the side plates of the pinnule-ambulacra,
but they are very abundantly distributed over the plated coverings of the genital pinnules
which are unprovided with ambulacra, as in Antedon incisa (PI. XXI. fig. 2a) and
Antedon angusticalyx (Part I. pi. liv. fig. 5).
Attached to the under side of the centro-dorsal of one specimen is a Brittle- Star which
seems to belong to the genus Opliiomusium, so far as I have been able to make out its
characters from a view of the dorsal surface only. But I cannot refer it to any species
of this genus, or to any other Ophiuran which was obtained by the Challenger. It has a
relatively large dorso-central, five small basals, and five large radials, the other ends of
which are tubercular, and fit in between the two large radial shields which are also more
or less tubercular on their line of junction. The arms of the Brittle-Star extend outwards
between the cirri of the Comatula and coincide in direction with its rays, while their
REPORT ON THE CRINOIDEA. 219
ends are more or less twisted round the bases of the cirri. The relations of the two
forms are thus somewhat closer than those of Ophiolebes scorteus with Antedon hirsuta,
which were noticed on p. 189; and the symmetrical arrangement of the large primary
plates on the Ophiurid disk, together with the position of its arms, gives a very singular
appearance to the centro-dorsal of the Comatula.
5. Antedon patula, n. sp. (PI. XLIIL).
Specific formula — A. 2.^.
Centro-dorsal subcorneal and flattened at the apex, with short and broad interradial
processes at its ventral angles. Five double rows of cirrus-sockets, three or four in each
row. About thirty cirri, of some twenty stout and smooth joints, most of which are
longer than wide, and somewhat compressed laterally, so as to have a sharp dorsal edge.
The first radials invisible ; the second short, closely united laterally and almost V-shaped
in side view. Axillaries widely rhombic, with a large and rounded backward projection
which is more or less tubercular in character. The rays divide twice, forming twenty
arms of about one hundred and sixty joints. Two distichals, the axillary without a
syzygy. The radial axillaries, the distichals, and the first three or four brachials have
sharp lateral edges and flattened sides. The surfaces of the distichals, and of the lowest
brachials rise to a more or less distinct tubercular projection. The fourth and following
brachials are short and nearly oblong, their surfaces rising considerably from the proximal
to the distal margins, which stand up rather prominently. Beyond the fifteenth brachial
the joints are more triangular, with a median ridge, and overlap slightly. A syzygy in
the third and then not till the eighteenth or twentieth brachial ; others at intervals of
four to eight, usually six or seven, joints.
The first pinnule is some 8 mm. long, and consists of about twenty-five short joints,
the lowest of which are broad and rather sharply flattened. The next pinnule is a
trifle longer and stouter, with a smaller number of larger joints, and in the next
following pinnules the joints gradually increase in size and become more carinate, the
third to the fifth being the widest, but they do not become longer than wide till some
way out on the arm.
Disk thickly plated, and also the arms, both along the ambulacra and in the inter-
muscular spaces. The genital pinnules have a covering of small plates, with the sacculi
scattered upon them, and the ambulacra of the later pinnules have well differentiated
side plates with intervening sacculi.
Colour in spirit, — the skeleton whitish-brown, and the disk darker.
Disk 10 mm.; spread about 35 cm.
Locality.— Station 192, September 26, 1874 ; near the Ki Islands ; lat. 5° 49' 15" S.,
long. 132° 14' 15" E.; 140 fathoms; blue mud. Two specimens.
220 THE VOYAGE OF H.M.S. CHALLENGER.
Remarks. — This is another robust species with a considerable resemblance to Antedon
Jlexilis (PI. XLII.) in its triangular arm-joints, expanded genital pinnules, and the
smooth, stout cirri arranged in ten vertical rows. But the centro-dorsal is somewhat
more conical than in that type and conceals the first radials entirely. The radial portions
of its margin are very deeply incised, so that its interradial angles are extremely
prominent, and they separate the lower lateral angles of the second radials, not of the
first, as in Antedon Jlexilis. In a side view of the calyx the second radials have an
almost V-shaped appearance, and are sometimes entirely invisible, owing to the manner
in which the axillaries project backwards into them. This gives a somewhat pear-shaped
appearance to the axillaries when seen " full," almost the whole of their length being
behind the line which joins their lateral angles.
Antedon patula also differs from Antedon jlexilis in the characters of the lower arm-
joints. In the latter type they are smooth and rounded and in no way specially
prominent (PI. XLII.). But in Antedon patula the distal edges of the fourth and each
of the following brachials, till the twenty-fifth or thirtieth, are raised into a sort of collar,
which stands up above the base of the next joint ; and as soon as the joints assume a
triangular shape they are marked by a distinct medio-dorsal ridge, which gives the arm
a carinate appearance, a character which is altogether absent in Antedon Jlexilis. The
joints of the genital pinnules are also somewhat carinate, and less enlarged than in
Antedon Jlexilis ; but there is the same plating over the genital glands as in that species,
though the sacculi are not quite so abundant.
6. Antedon robusta, n. sp. (PI. XLIV. fig. 1).
c
Specific formula — A. 2.^.
Description of an Individual. — Centro-dorsal a thick disk, 9 mm. wide, and bearing
about fifty cirri round its margin. These have from eighteen to twenty-three smooth,
stout joints, several of which are longer than wide. The penultimate forms a small
opposing spine.
First radials just visible at the angles of the calyx above the low interradial
processes of the centro-dorsal, which partly conceal the short second radials in their
median line. Axillaries widely rhombic, with an open distal and sharper proximal
angle, the latter rising to form a prominence with the second radial. Twelve arms,
there .being one bidistichate series on each of two rays. The first two brachials (or the
distichals when present) form a slight prominence as the outer radials do, and have
much flattened outer sides like the axillaries. The inner sides of the second, and both
sides of the third brachials, are also sometimes flattened. Arms long, of more than two
hundred smooth joints, the first few oblong, and the later joints more triangular. A
REPORT ON THE CRINOIDEA. 221
syzygy in the third brachial ; the next not till after the twentieth, and others at long
intervals (nine to twenty-five joints).
The second brachial bears a stout pinnule about 12 mm. long and composed of some
twenty joints, the lowest of which are much larger than their successors and of almost
prismatic shape, being flattened against the corresponding joints of the adjacent
pinnules. The third brachial has a similar but slightly smaller pinnule, and its
successors are of about the same length, but have broader and flatter joints. The later
pinnules gradually become elongated, but none of their lower joints are conspicuously
wider than the rest. In the styliform middle and outer pinnules the first joint is
flattened and expanded, with a curved distal edge.
Disk thickly plated, and also the arms, both along the ambulacra and in the inter-
articular regions. The genital pinnules are protected by irregular plates, and the
ambulacra of the later pinnules have well-defined side plates, with alternating sacculi,
which are also fairly abundant on the genital pinnules.
Colour in spirit, — the perisome dark blackish-brown, but the skeleton whiter.
Disk 12 mm.; spread nearly 50 cm.
Locality.— Station 192, September 26, 1874; near the Ki Islands; lat. 5° 49' 15" S.,
long. 132° 14' 15" E.; 140 fathoms; blue mud. One specimen.
Remarks. — This type is a larger and more massive species than Antedon flexilis or
Antedon patula, which resemble it in the characters of the stout cirri, the flattened
lower pinnules, and the thickly-plated ventral perisome. But the centro-dorsal is
altogether different and the calyx less tubercular than in these two species. The centro-
dorsal is a thick disk, with the cirri closely set round its margin in two or three tiers.
(PI. XLIV. fig. 1), while the wide dorsal surface, which is slightly concave, is entirely
free from them. The radial axillaries are relatively wider and much less prominent than
in Antedon patida (PI. XLIIL), and the first radials are not so entirely concealed as in
that type (PI. XLIV. fig. 1). In this respect, as also in the smoothness of the arms,
Antedon robusta approaches Antedon flexilis (PI. XLII.) ; but the lower joints of the
genital pinnules are not so markedly expanded as in that species.
The single specimen of Antedon robusta which was obtained by the Challenger has
two distichal axillaries, but it is quite possible that other examples may some time be
discovered which have only ten arms, as is the case in Antedon flexilis, two specimens
of which have three axillaries each, while the other two are only ten-armed. In that
case Antedon robusta would find a place in the Basicurva-gvoup by the side of Antedon
flexilis.
222 THE VOYAGE OF H.M.S. CHALLENGER.
7. Antedon compressa, n. sp. (PI. XLL).
Specific formula — A. 2.^.
Centro-dorsal a thick convex plate, with the dorsal pole free, and fifteen to twenty
marginal cirri. These have about twenty joints, a few of which are longer than wide,
and the later joints are somewhat compressed laterally, with blunt dorsal spines.
The first radials are concealed, and sometimes also parts of the second. These are short
and sharply convex, with a slight median ridge, and they meet one another laterally
beyond the angles of the centro-dorsal. Axillaries short and widely rhombic, also with a
median ridge, and forming a small tubercle with the preceding joints. The rays divide
twice ; distichals two, with a faint median ridge, the axillaries without a syzygy, short
and widely rhombic. The outer sides of the radial axillaries, distichals, and of the two
lower brachials are slightly flattened ; and the inner sides of the second and third
brachials are also somewhat flattened.
Fifteen to twenty arms, of one hundred and fifty or more joints ; the lowest are
nearly oblong, with indications of a dorsal keel and raised distal edges. The following
joints more triangular and distinctly carinate, gradually becoming quadrate and somewhat
compressed laterally, with a tendency to overlap. A syzygy in the third, and then in
the eleventh or twelfth brachial ; others at intervals of two to seven, usually four or five
joints.
The second brachial bears a tolerably stout pinnule some 10 mm. long, and consisting
of about eighteen short joints, the first few of which are much wider than their successors
and slightly prismatic. That of the third brachial is smaller, with the basal joints more
rounded, and not so wide ; and the following pinnules diminish in size till about the
tenth brachial, having fewer but relatively longer joints. Beyond this point the length
gradually increases again and the later pinnules are slender and delicate, with the two
basal joints flattened and somewhat expanded.
The disk is well plated along the ambulacra ; but the interpalmar areas only have a
few scattered granules ; the brachial ambulacra and interarticular spaces also well plated.
The side plates of the pinnule-ambulacra are fairly distinct, with intervening sacculi.
Colour in spirit, — very light brown ; the disk darker.
Disk 8 mm.; spread 28 cm.
Localities. — Station 192, September 26, 1874 ; near the Ki Islands ; lat. 5° 49' 15" S.;
long. 132° 14' 15" E.; 140 fathoms; blue mud. Two specimens.
Station 201, October 26, 1874; Philippine Islands; lat. 7° 3' N., long. 121° 48' E.;
82 fathoms ; stones, gravel. One specimen.
Remarks. — This species is in some respects a transitional form between Antedon
patula. and Antedon flexilis on the one hand, and the Palmata-gvou]) on the other. The
REPORT ON THE CR1NOIDEA. 220
lateral flattening of the lower brachials is scarcely more distinct than in Antedon
jiagellata or Antedon brevicuneata ; but the first pinnule has prismatic lower joints, and
the ambulacra of the disk, arms, and pinnules are well plated, though the interpalmar
areas of the disk are comparatively bare. The genital pinnules are not specially
distinguished, however, except by their shortness, and their glands are not protected by
any special pavement of plates as in Antedon jlexilis and Antedon patula, though there
is an ambulacral skeleton above them which is less completely differentiated than in the
slender later pinnules.
The presence of blunt spines on the later cirrus-joints also distinguishes this species
from the three just described, all of which have very smooth cirri (Pis. XLII., XLIIL;
PI. XLIV. fig. 1), and are altogether of a more robust nature.
The two individuals which the Challenger collected at Station 192 are essentially
similar in all their characters ; but a younger specimen from Station 201 has much
smoother joints at the bases of the arms, their distal edges being but little raised ; while
in some fragments of a larger form obtained at the same locality there is a tendency to
expansion in the third and fourth joints of some of the genital pinnules, which recalls
their condition in Antedon Jlexilis (PI. XLII.). The interpalmar areas of the disk are
also more plated than in the examples from Station 192.
7. The Palmata-gxovq).
Bidistichate species with an unplated disk and no definite ambulacral skeleton. The
sides of the lower brachials are scarcely, if at all, flattened. The first pinnule smaller
than its successors.
Remarks. — This group is not only extremety well-defined as regards its general
characters, but it is also distinctly limited in its distribution, both bathymetrical and
geographical.
The disk is either naked or bears but a few isolated plates, and there is no definite
ambulacral skeleton. The ambulacra may be supported by isolated rods and networks
of limestone, but they never form distinct covering plates like those of the Basicurva-
and $pmi/era-groups. In a few species like Antedon jiagellata and Antedon similis
(PL XLVII. fig. 1), which have the rays closely approximated, the lower brachials of
their outer arms are somewhat flattened laterally. But this condition is not a constant
one, and it does not affect the lower pinnules ; so that it is altogether different from the
flattening of the arm-bases in Antedon basicurva, Antedon valida, or Antedon robusta
(fig. 3 on p. 122 ; PI. XV. fig. 6 ; PI. XLIV. fig. 1).
We have seen that the Spinifeixt-growp among the bidistichate species corresponds
to the Basicurva-gvo'a'p of the ten-armed type ; and in like manner the ten-armed
Milberti-grouip is represented in the bidistichate series by the Palmata-gvouip, all the
224 THE VOYAGE OF H.M.S. CHALLENGER.
members of which have either the second or the third pinnule (or both) distinctly larger
than the first one. In a few species like Antedon elongata and Ante don jiagellata the
third pinnule is the largest, as in the ten-armed Antedon variipinna (PI. XXXVI.
figs. 1, 4-6). In others, again, like Antedon articulator and Antedon regalis, the second
and thud pinnules are of about equal size, as in the ten-armed Antedon parvicirra
(PI. XXXVI. fig. 8). But in Antedon palmata, and in the majority of the species com-
posing the group, the second pinnule is considerably larger than both the first and the third
(PI. XLVIII. fig. 2 ; PI. XLIX. fig. 4), just as in Antedon pinniformis of the Milberti-
group. The parallel between the two groups may be continued yet further; for the singular
Antedon informis, which is without a pinnule on the third brachial (PI. XXXIII. fig. 3)
has two representatives in the Palmata-grou]), viz., Antedon disciformis (PI. XXXIX.
fig. 4), and Antedon manca (PI. XLIV. figs. 2, 3). The first of these has no axillary
beyond the distichals, but palmars are present in Antedon manca as in most species of
the group ; and there are usually six arms to each ray, of which only the two outermost
usually have pinnules on the second brachials.
Besides Antedon disciformis, Antedon clemens and Antedon marginata (PI. XXXIX.
fig. 5 ; PL XL. fig. 1) are the only members of the group which have but one post-
radial axillary. Some forms, like Antedon articulata, Antedon palmata, and Antedon
conjungens (PI. XLV. fig. 1), always have two and occasionally three; while we are not
yet acquainted with examples of Antedon Jiagellata, Antedon gyges, and Antedon
occulta (PL XLVIII. fig. 1 ; PL XLIX. fig. 3) in which a post-palmar axillary does not
occur on one or more of the rays. I have no doubt, however, that simpler forms of
these species will eventually be found, and I have, therefore, made no use of the presence
or absence of a post-palmar axillary for the purpose of classification.
With the exception of the three species (Antedon occidta, Antedon similis, and
Antedon tubercidata), which were dredged by the Challenger at one of the three
Stations 174b, c, or D (255, 610 or 210 fathoms), all the members of the Palmata-
group belong to the littoral fauna ; and they are exclusively limited to the Western
Pacific and the Indian Ocean. They are extremely abuudant between the Friendly
Islands on the east, and the Mergui Archipelago on the west, ranging northwards as far
as Southern Japan, but not extending to the south beyond the tropic of Capricorn.
Isolated species occur at the Sandwich Islands on the east and also at Ceylon and
Rodriguez on the west ; while Antedon palmata, the type of the group, is common at
Aden and in the Red Sea. This is the furthest western limit of the group, which is
altogether unrepresented in the Atlantic ; for all the bidistichate species of Antedon
from the Caribbean Sea have plated ambulacra, and therefore belong to the Spinifera-
group.
The mutual relations of the various species composing the Palmata-grou-p are shown
in the following key: —
REPORT ON THE CRINOIDEA.
225
A. No pinnule on the third brachial.
I. Two post-radial axillaries ; the inner arms of each ray usually without
a pinnule on the second brachial, ....
II. One post-radial axillary; the second brachial always has a pinnule,
R The third brachial has a pinnule.
I. One post-radial axillary ; the rays quite free laterally.
a. Thirty cirrus-joints ; brachials very short ; sides of rays smooth, .
p. Twenty cirrus-joints ; brachials not specially short ; irregular pro-
jections at the sides of the rays, .....
II. Two or more post-radial axillaries.
a. Second pinnule larger than third.
1. The rays free laterally.
a. The second pinnule stiff and styliform, of twelve to
eighteen much elongated joints.
(i) Rays have marginal projections ; third pinnule not
greatly shorter than the second.
a. Forty cirri ; axillaries more than twice as long
as second radials, ....
/?. Twenty-five cirri; axillaries less than half as
long again as second radials,
(ii) Margins of rays smooth ; third pinnule considerably
shorter than the second,
b. The second pinnule has twenty-five or more joints,
which are not specially elongated,
(i) The lower pinnules are larger on the outer arms of
each distich ium than on the inner ones.
a. Third pinnule quite short,
ft. Third pinnule not specially short, nearly as long
as the second on inner arms,
(ii) The lower pinnules fairly uuiform in size on all the
arms.
a. The fourth and fifth brachials bear large and
tolerably equal pinnules,
/3. The pinnule on the fourth brachial larger than
that on the fifth.
Third pinnule smaller than the first ; second
syzygy about the twentieth brachial,
Third pinnule equal to the first ; second syzygy
about the thirteenth brachial,
2. The rays in close contact laterally.
a. Spiny cirri.
(i) The second pinnule not greatly larger than the
third ; no post-palmars ; the fifth brachial has
the first syzygy in arms which spring from a
distichal axillary, ....
(ii) The second pinnule considerably longer than the
third ; post-palmars ; the first syzygy always in
the third brachial.
a. Over thirty cirrus-joints ; the first pinnule not
much larger than the second. The lower
brachials have flattened sides, * j j
(ZOOL. CHALL. EXP. — PART hX. — 1887.)
1. manca, n. sp.
2. diseiformis, n. sp.
3. clemens, n. sp.
4. marginata, n. sp.
5. tubereulata, n. sp.
spicata, Carpenter.
indica, Smith, sp.
protecta, Liitken, MS.
6. conjungens, n. sp.
xquipinna, Carpenter.
Iwvicirra, Carpenter.
imparipinna, Carpenter.
regime, Bell.
gyges, Bell.
Ooo 2a
226
THE VOYAGE OF H.M.S. CHALLENGER.
j3. Not over twenty-five cirrus-joints ; the first
pinnule much smaller than the second,
b. Cirri carinate, but not spiny.
(i) No post-palmars ; lower brachials flattened ; two
radials visible.
a. Fourth pinnule altogether smaller than the
third, . .■ .
ft Fourth pinnule nearly similar to the third,
(ii) Post-palmars ; axillaries almost concealed, .
b. Second and third pinnules about equal in size.
1. Thirty-five to forty cirrus-joints, the later ones distinctly spiny,
2. Twenty-five to thirty cirrus-joints, the later ones with pointed
keels ; lower brachials flattened, ....
c. Third pinnule larger than the second.
1. Spiny cirri.
a. Rays well separated ; no post-palmars ; second syzygy
about the fourteenth brachial,
b. Rays in close contact, and slightly flattened laterally ;
postpalniars ; second syzygy about the twentieth
brachial, ......
2. Cirri not spiny ; second syzygy about the twentieth brachial,
palmata, Mull., sp.
brevicuneata, Carpenter.
7. similis, n. sp.
8. occulta, n. sp.
articulata, MulL, sp.
9. regalis, n. sp.
elongata, Mull., sp.
flagellata, Mull., sp.
bimaeidata, Carpenter.
1. Anteclon manca, n. sp. (PL XLIV. figs. 2, 3).
Specific formula — A. 2. 2. y.
Description of an Individual. — Centro-dorsal a thick disk, with a flattened dorsal
surface and about twenty marginal cirri. These have twenty-five to thirty joints, a few
of which are longer than wide, and develop a dorsal spine from the eighth onwards.
First radials concealed ; the second oblong, and quite free laterally ; auxiliaries
pentagonal. Two distichals and two palmars, the axillaries not syzygies ; but the
pal mars are only developed on the outer pair of every four secondary arms, so that there
are normally six arms to each ray, viz., 2,1,1,2. But one palmar axillary is undeveloped,
giving twenty-nine arms only. They have about one hundred smooth and rounded
joints, the first few discoidal, and the following ones triangular, about as long as wide,
but becoming quadrate further out. A syzygy in the third brachial, and the next about
the eighteenth or twentieth ; others at intervals of three to nine, usually five or six
joints.
The third brachial has no pinnule at all, while that of the second brachial is always
absent on the innermost of every two arms springing from a palmar axillary, and
sometimes also on the arms which are borne directly on the distichal axillaries. But it
is sometimes present on these latter arms, and always on the two outer arms of each ray,
though varying in size, consisting of twelve or fifteen joints, most of which are longer
than wide.
REPORT ON THE CRINOIDEA. 227
The pinnules of the fourth and fifth brachials are sometimes twice its length, reaching
12 mm., and consisting of about eighteen elongated joints, with spines at their distal
ends. The next pinnule is not half the size of this pair, and is smaller than that on the
second brachial, while the next pair are the smallest on the arm. after which the length
of the pinnules increases slowly.
Disk very much incised, and quite naked ; sacculi very abundant on the arms and
pinnules.
Colour in spirit, — the skeleton reddish-brown, and the perisome rather darker.
Disk 8 mm.; spread 12 cm.
Locality.— Station 192, September 26, 1874 ; near the Ki Islands ; lat, 5° 49' 15" S.,
long. 132° 14' 15" E.; 140 fathoms; blue mud. One specimen.
Remarks. — This is a very singular species, the general relations of which are with
Antedon marginata and Antedon sequipinna; but it is distinguished from them both,
and from all the other bidistichate forms of Antedon, by the peculiar distribution of the
pinnules. The arrangement of the arm-divisions seems to be like that of Pentacrinus
maclearanus, palmar axillaries being only developed on the two outermost of the four
secondary arms on each ray, so that the grouping of the arms is 2,1,1,2. In one ray,
however, a palmar axillary is missing, so that the total number of arms is twenty-nine,
and not thirty as it would otherwise be.
There is never any pinnule on the third brachial, as is also the case in Antedon
disciformis (PI. XXXIX. fig. 4) and in the ten-armed Antedon informis of the
Milberti-groi\]) (PI. XXXIII. fig. 3) ; and that of the second brachial is also absent in
some arms. It is undeveloped on the inner arm of every pair which springs from a
palmar axillary, and it is occasionally also absent on the two inner arms of the ray which
are borne directly on the distichal axUlaries ; though it is sometimes present, as in the
two central arms of the lowest ray represented in PI. XLIV. fig. 2, while it is always to be
found on the two outer arms of the ray. Of the two single arms which are borne on the
inner faces of the two distichal axillaries, one may have a pinnule on the second brachial
and the other not ; and there appears to be no constancy as to its occurrence in this
position. When present, it is somewhat smaller than its fellow on the outside of the ray.
The large pinnules of the fourth and fifth brachials are tolerably equal on all the arms,
that on the fourth being perhaps a little the longer ; but the pinnule of the sixth brachial
is much smaller again, and the next pah- still more so, barely reaching 4 mm.
The disk has a large pentagonal perisome (PI. XLIV. fig. 2), but is much incised,
and the anal tube appears to be quite at its margin, so far as one can judge from the
mutilated condition of the specimen. The sacculi do not appear to occur on the disk,
but they are well developed on the arms, and especially so on the pinnule-ambulacra.
228 THE VOYAGE OF H.M.S. CHALLENGER.
2. Antedon disciformis, n. sp. (PI. IV. figs. 2, a-d ; PI. XXXIX. fig. 4).
Specific formula — A.2.-J-.
Centro-dorsal a thick pentagonal disk, with an irregular row of marginal cirri and the
dorsal surface free. Fifteen to twenty cirri, of twenty-five to thirty joints, several of
which are longer than wide. The fourth or fifth joints project beyond their successors
on the dorsal side, and the following joints gradually develop a sharp forward-projecting
spine at their distal edge ; as the joints shorten this comes to be placed further and
further back, and is both shorter and more upright.
First radials mostly concealed ; the second oblong and quite free laterally ; axillaries
pentagonal, nearly twice their length. The rays are well separated and may divide
twice. Two distiehals ; the first nearly oblong, and the axillary without a syzygy.
Fifteen to twenty arms of about one hundred and twenty smooth and rounded joints,
the first few discoidal and their successors triangular, about as wide as long, gradually
becoming more quadrate. A syzygy in the third, and then between the eighth and
fourteenth brachials ; others at intervals of one to five joints.
The second brachial has a slender pinnule of about eighteen short joints, but little
longer than wide ; but the third brachial has no pinnule. The next pair are rather
stouter and much longer than the first pinnule, reaching 12 mm., and consisting of
twenty elongated joints, the apposed edges of which are somewhat produced towards the
ventral side. The pinnule on the sixth brachial may be of nearly equal size or distinctly
smaller, and its successors diminish in length to about the tenth brachial, and then
increase, becoming exceedingly slender in the outer parts of the arms.
Disk naked and rather incised, with a few sacculi, which are very abundant along
the ambulacra of the arms and pinnules.
Colour in spirit, — the skeleton almost white, and the perisome grey or brownish.
Disk about 8 mm.; spread probably 15 cm.
Locality. — Zebu Reefs. Six specimens and one fragment.
Remarks. — This species, while resembling Antedon manca in its spiny cirri and in
the absence of a pinnule on the third brachial, differs from it altogether in having no
palmar axillary, and in the constant presence of a pinnule on the second brachial. The
rays being quite free laterally, it stands rather near to Antedon marginata (PI. XL.),
resembling it also in the elongated joints, in the great size of the pinnules on the fourth
and fifth brachials, and in the absence of a palmar axillary, though sharply distinguished
from it by the want of a pinnule on the third brachial and by the very spiny cirri.
The extreme flatness of the centro-dorsal and the limitation of the cirri to its margin,
so as to leave the dorsal surface free (PI. IV. fig. 2a), recall the characters of Actinometra ;
REPORT ON THE CRINOIDEA. 229
but the high articular faces of the radials, which are much wider below thau above
(PI. IV. figs. 2a, 2b), are those of a typical Antedon. The lower parts of the fossaj
lodging the great ventral muscles are cut off from their upper portions as seen in fig. 2a ;
and the same peculiarity appears both on the proximal faces of the second radials and on
the distal faces of the axillaries. The ventral surface of tha centro-dorsal (PL IV.
fig. 2d) is marked by five minute radial pits, corresponding to the ventral ends of the
radial axial canals, which are seen on the under surface of the radial pentagon (PI. IV.
fig. 2c), just as I have described in some forms of Antedon plxalangium and of Antedon
rosacea}
3. Antedon clemens, n. sp. (PI. XXXIX. fig. 5).
Specific formula — A.2.-T-.
Description of an Individual. — Centro-dorsal hemispherical, and bearing some
twenty-five cirri, which have about thirty tolerably uniform, smooth joints, the
penultimate with a small spine.
First radials not visible ; the second slightly united laterally and the axillaries
pentagonal. One ray does . not divide at all ; three divide once, and one twice. The
latter has two distichals, the second axillary without a syzygy. Eleven arms of smooth
triangular joints, which are much wider than long and gradually become quadrate.
Syzygies in the third and then in the eighth to twelfth brachials, with others at intervals
of one to nine, usually four or six joints.
The first pinnule is 5 mm. long, with about twenty joints which diminish greatly in
size after the first five or six. The next pair are much longer, with a smaller number of
stouter joints, several of which are considerably longer than wide, and the third pair are
smaller again ; the more distal pinnule of each pair is smaller than the proximal one.
Those of the eighth and following brachials have slight dorsal keels on their lower joints.
Disk naked and rather incised ; sacculi abundant on the pinnules.
Colour in spirit, — the perisome purplish-grey, and the skeleton brownish -white.
Disk 7 mm.; spread about 15 cm.
Locality.— Station 212, January 30, 1875; Celebes Sea; lat. 6° 54' N., long. 122°
18' E.; 10 fathoms ; sand. One specimen.
Remarks. — This is a unique specimen in every way. I have never before met with
any individual which combined in such a singular manner the characters of two other
genera besides that to which it naturally belongs. One of the rays does not divide at all.
as in Eudiocrinus.
1 Trans. Linn. Soc. Lond. (Zool.), ser. 2, 1879, vol. ii. p. 78, pi. iv. figs. 7, 16.
230 THE VOYAGE OF H.M.S. CHALLENGER.
It has a pinnule on the second brachial, just as in Eudiocrinus varians (PI. VII.
figs. 3,4); but the next joint has none, thus affording a parallel to the condition of
Antedon informis and Antedon disciformis. On the other hand, one ray does not
divide at all till the fourth joint beyond the primary radial, as is often the case in
Metacrinus moseleyi and in Metacrinus rotundus, though the axillary is the syzygial
joint, and not the second of the series as in those types ; but the second and third bear
pinnules just as in Metacrinus.
The only other bidistichate Antedon, besides Antedon clemens, which has a pinnule
on the third brachial and no palmars developed is Antedon marginata (PL XL.). It is
altogether a larger species than Antedon clemens, however, with relatively longer and
more quadrate brachials, and more numerous cirrus-joints ; while the second pair of
pinnules of Antedon clemens are relatively longer than the first and third pairs than is
the case in Antedon marginata, and the sides of the rays are smooth, without the lateral
processes which are characteristic of that type.
4. Antedon marginata, n. sp. (PI. XL.).
Specific formxda — A. 2. -r-.
Description of an Individual. — Centro-dorsal saucer-shaped, and bearing some
twenty-five cirri on its sides, each of about twenty joints, a few of which are somewhat
longer than wide. The terminal joints are rather compressed laterally and have a faint
keel, passing into the dorsal spine of the penultimate.
First radials just visible; the second oblong, and quite free laterally; axillaries
pentagonal, about one and a half times their length. The rays are well separated and
may divide twice. Two distichals, the axillary without a syzygy. Both radials and
distichals are rather convex, rising sharply to the middle of their apposed edges. The
outer edges of all the pieces at the sides of the rays, from the second radial to the second
brachial inclusive, are marked by irregular projections towards the ventral side.
Fourteen arms, of about one hundred and fifty joints, the lower ones thick disks, and
their successors more triangular, but wider than long, gradually becoming quadrate
and more discoidal again in the middle of the arm. A syzygy in the third, and
then between the eighth and thirteenth brachials ; others at intervals of three to six
joints.
The second brachial bears a comparatively slender pinnule of about twenty joints,
most of which are longer than wide, and the third has a similar but smaller pinnule.
The next pair of pinnules are not much longer than the first one, reaching 10 mm., but
they have only ten or twelve very stout and rather elongated joints, which terminate
somewhat abruptly. That of the fourth brachial is the larger of the two, and the next
REPORT ON THE CRINOIDEA. 231
pair are of the same character, but less stout, though not much shorter. The following
pinnules become more slender and gradually increase in length.
Disk naked and moderately incised ; the ambulacra have lines of sacculi at their sides,
which become very prominent towards the margin of the disk, and give off branches to
the first pair of pinnules. The sacculi are large and abundant along the ambulacra of
both arms and pinnules.
Colour in spirit, — the skeleton reddish-white, with dark red lines at the articulations ;
perisome grey or purplish-grey.
Disk 15 mm.; spread 20 cm.
Locality.— Station 208, January 17, 1875; off Manila ; lat. 11° 37' N., long. 123°
31' E.; 18 fathoms; blue mud. One specimen.
Remarks. — This type is readily distinguished from the only two other bidistichate
species of Antedon in which the rays divide but twice. It differs from Antedon disci-
formis (PI. XXXIX. fig. 4) in the presence of a pinnule on the third brachial, the
shorter arm-joints, and the smaller number of cirrus-joints (PL XL. fig. l); while the
second pair of pinnules are relatively much stouter than in Antedon clemens, and the
number of cirrus-joints is smaller than in that type (PI. XXXIX. fig. 5). In the freedom
of its rays and in the irregular processes at their sides it resembles Antedon tuberculata
(PL XLV. fig. 2), but differs from it in having a smaller number of cirri and no palmars
developed, so that there are only fourteen arms instead of thirty, while the lengths
of the first pinnules are much more nearly equal than is the case in that type (PL XLV.
fig. 3).
The second pair of pinnules of this unique specimen of Antedon marginata terminate
so abruptly that they seem to have been broken by some accident and not completely
repaired. The diameter of the joints suddenly decreases and there are from one to four
quite small joints at the end of a large and stout one which is considerably longer than
wide.
The disk of this specimen is remarkable for the abundance of sacculi upon it.
There is a line of them along each side of the ten secondary ambulacra ; and branches
proceed direct from these lines to the primary pinnules as seen in PL XL. fig. 2,
thus marking the course of their water-vessels, which, however, have no tentacular
extensions. This character recalls the arrangement of the pinnule-ambulacra on the disk
of Metacrinus as figured on pi. xxxix. fig. 2 and pi. xliii. fig. 3 of Part I. But in
Antedon marginata the lines of sacculi and the water-vessels are unaccompanied by the
other ambulacral structures, and the lower pinnules are non-tentaculiferous, as in Antedon
rosacea and other types.
232 THE VOYAGE OF H.M.S. CHALLENGER.
5. Antedon tuberculata, n. sp. (PI. XLV. figs. 2, 3).
Specific formula — A.2.2.-^-.
Description of an Individual. — Centro-dorsal saucer-shaped, bearing about forty
rather stout cirri, of twenty to twenty -five tolerably uniform joints, few of which are
longer than wide ; a small spine on the penultimate.
Only two radials visible ; the second short and rather convex, not united laterally ;
axillaries more than twice their length and widely pentagonal, the junction of the two
joints rather tubercular. The rays are well separated and may divide three times ; the
distichal and palmar series each of two joints, with a tubercular junction, and the axillary
not a syzygy. The palmar axillary is usually only developed on the outer pair of every
four secondary arms, giving six arms to the ray, viz., 2,1,1,2. The outer edges of all
the pieces at the sides of the rays from the radial to the palmar axillaries are marked by
small tubercles, which project somewhat towards the ventral side. Thirty-one arms of
one hundred and twenty or more joints, the first eight or ten of which are thick disks,
and the following ones shortly triangular, gradually becoming more discoidal again. A
syzygy in the third brachial, and the next between the seventh and twenty-first, generally
about the twelfth or fourteenth ; others at intervals of five to ten, usually seven joints.
The first pair of pinnules are about 9 mm. long, and consist of some twenty-five
longish joints ; the next pair much stouter and very stiff and tapering, reaching 15 mm.,
and composed of about a dozen joints, all of which, except those at the two ends, are
much longer than wide. The succeeding pair are of the same character, but rather
shorter than the second pair ; and the fourth pair are much smaller and less stiff.
Disk lost ; sacculi abundant on the ambulacra of both arms and pinnules.
Colour in spirit, — the skeleton purplish-white, with occasional dark purple bands ;
the perisome greenish-grey.
Disk probably about 12 mm.; spread about 180 mm.
Locality. — Station 174 1 (b, c, or d), August 3, 1874; near Kandavu, Fiji; lat.
(about) 19° 6' S., long, (about) 178° 18' K; 255, 610, or 210 fathoms; Coral mud;
bottom temperature (at 610 fathoms), 39° F. One specimen.
Remarks. — This species, as well as Antedon spicata and Antedon indica, are
distinguished by the characters of the second pair of pinnules, which are well shown in
Smith's figure 2 of the last mentioned type. They are considerably longer than the first
pair, though composed of a smaller number of joints. But these joints are of very large
size, some of them reaching 1*5 mm. in length. They decrease gradually in diameter
1 The exact locality, and consequently the exact depth, is not recorded.
2 Zoology of Rodriguez, Echinodermata, Phil. Trans., 1879, vol. clxviii. pi. li. fig. 36.
REPORT ON THE CRINOIDEA.. 233
from the base to the tip of the pinnule, so as to give it a remarkably stiff and tapering
appearance (PI. XLV. fig. 3). There is some indication of this in Antedon marginata,
but its large pinnules are less stiff, with relatively shorter joints, which are more uniform
in diameter, so that the pinnule lacks the tapering and styliform appearance which is so
marked in Antedon spicata and Antedon tuberculoid. Its cirri too are both smaller
and have fewer joints than those of Antedon tuberculata, while the second radials
and the axillaries are more equal in length, and portions of the first radials are visible
(PL XL. fig. 1 ; PL XLV. fig. 2).
Antedon tuberculata has many points of resemblance with Antedon spicata from the
Banda Sea, and it may be that a larger knowledge of both types wdl eventually lead to
their union. The cirri of Antedon tuberculata are both considerably more numerous and
reach a larger size than in Antedon spicata, though the actual number of joints composing
them is the same in both forms. The second radials of Antedon tvherculata are short as
compared with the axillaries, not reaching half their length ; while in Antedon spicata
the axillaries are short as compared with the second radials. The arms of the latter type
are also longer than in Antedon tuberculata, and the muscle-plates more prominent at
the sides of the ambulacra.
Antedon indica differs from both these types in the slighter development of marginal
projections at the bases of the rays, and in the marked difference in the characters
of the second and third pairs of pinnules. The latter are not so stiff as in Antedon tuber-
cidata, but are considerably smaller than the second pair, consisting of a number of small
joints, like the first pair.
6. Antedon conjungens, n. sp. (PL XLV. fig. 1).
Specific formida — A. 2.2. 2. y.
Centro-dorsal a thick slightly convex disk, bearing about twenty-eight cirri round its
margin. They have twenty to thirty uniform joints, the later ones somewhat compressed
laterally, with a sharp dorsal edge which passes into the spine of the penultimate.
First radials not visible; the second widely hexagonal, partly united laterally;
axillaries pentagonal. The rays, which are free from the second radials, divide thrice
and occasionally four times ; each series of two joints, the axillary without a syzygy.
Rather over forty arms of about one hundred and fifty joints, the first few discoidal, and
their successors shortly triangular, gradually becoming cpiadrate, but always much wider
than long. A syzygy in the third brachial, and the next between the fourteenth
and twentieth, generally about the fourteenth or fifteenth ; others at intervals of five
to eleven, usually seven or eight, joints.
Of the four or more arms borne on each distichal axillary the two outer ones have
(ZOOL. CHALL. EXP. PART LX. — 1887.) OoO 30
234 THE VOYAGE OF H.M.S. CHALLENGER.
inuch larger lower pinnules than the inner arms. That of the fourth brachial may reach
15 mm., with nearly thirty joints, the lowest of which are very stout, but not specially
long. The corresponding pinnule of the inner arms is about two-thirds of its length, with
fewer and smaller joints, and the pinnule of the next joint is of nearly equal size, while
on the outer arms it is considerably smaller than its predecessor. The nest pinnule is
about equal to it, reaching 10 mm., with twenty joints, but on the inner arm it is
markedly smaller than that on the fifth brachial. In like manner the first pinnule
reaches 12 mm. on the outer arms, with nearly thirty joints, less stout than those of the
second pinnule, but still of considerable size at the base, while on the inner arms it is
small and slender. That of the third brachial is always quite small. The disk is very
deeply incised, almost to the level of the radial axillaries. The outer sides of the
distichal and palmar joints are much produced towards the ventral surface, so that each
of the five divisions of the disk, as seen from the ventral side, has more or less distinct
bony margins. Sacculi abundant along the pinnule-ambulacra.
Colour in spirit, — the disk is grey, and the skeleton white, more or less mottled with
purplish- or reddish-grey in bands and patches.
Disk 17 mm.; spread 20 cm.
Locality. — Zebu Eeefs. Two specimens.
Remarks. — This species may be readily distinguished by the characters of its lower
pinnules, which have more numerous and much shorter joints than those of Antedon
tuberculata and its allies, while they are not of equal size on all the arms. Excepting
in one ray of each specimen there is no axillary beyond the palmars, and so there are
normally eight arms, four on each distichal axillary. In the two outermost of these four
arms, the first three pinnules are much larger than their fellows on the inner pair. This
is especially the case with the second pinnule (on fourth brachial), so that while on the
inner arms it is about equal to that on the next joint, it is one-half longer and con-
siderably stouter on the outer arms.
A somewhat similar variation is presented by one of the types which have been
distributed by the Godeffroy Museum under the MS. name Antedon protecta,
Liitken. Thus in one individual, which I owe to the kindness of Professor Loven, the
first two pinnules on the outer pair of every four tertiary arms are greatly larger than
the corresponding pinnules on the inner arms. The second one has twenty-five joints
and reaches 12 mm., nearly three times the length of its fellow on the inner arm.
In this type, however, the third pinnule on both inner and outer arms alike has
little more than a dozen joints, and is only some 4 mm. long. The small size of this
third pinnule is remarkable, not only as distinguishing the type from Antedon
conjungens, in which it is at least half the length of the second pinnule, if not more, but
also in the whole group of species with large second pinnules.
EEPORT ON THE CRINOIDEA. 235
Each of the two examples of Antedon conjungens which are described above has
normally eight arms to the ray, but a palmar axillary is occasionally absent ; while in the
D ray of each individual post-palmars are developed, one axillary in the one, and two in
the other specimen.
For the present therefore the formula of the type must be A. 2. 2. 2.-^-, though it is
quite likely that examples of it will be eventually discovered in which post-palmars are
absent, so that the last figure must then be put within brackets.
7. Antedon sirnilis, n. sp. (PI. XLVII. figs. 1-3).
be
S}dccific formula — A.2.2.y.
Description of an Individual. — Centro-dorsal a thick disk, bearing about forty
marginal cirri. These have some twenty-five tolerably uniform joints, the later ones
compressed, with a sharpened dorsal edge which passes into the penultimate spine.
The first radials are entirely concealed, and also part of the second, which are
closely united laterally. Distichals are present all round the calyx, and palmars on
four of the rays, the two outer secondary arms bearing palmar axillaries, so as to give
six arms to the ray, viz., 2.1.1.2. Each series is of two joints, the axillary without
a syzygy. The distichals and palmars of adjacent rays are closely appressed, with sharp
lateral edges and flattened sides. Twenty-eight long and slender arms of about one
hundred and fifty joints, the lower ones discoidal and the rest shortly triangular.
gradually becoming cruadrate, but always much wider than long. A syzygy in the third
brachial, and the next between the sixteenth and twenty-first, with others at intervals
of eight to sixteen joints.
The first pinnule is some 7 mm. long, with about twenty joints, which are but little
longer than wide. The second is much stouter, consisting of nearly twenty -five rather
longer joints, and reaching 14 mm. The fifth brachial has a similar but rather smaller
pinnule, and that on the sixth is 6 mm. long with nearly twenty joints. The fourth
pinnule is but little shorter, with a dozen joints.
Disk naked and rather incised ; sacculi abundant on the pinnule-ambulacra.
Colour in spirit, — a mixture of brownish-white and greenish -grey.
Disk 12 mm.; spread 16 cm.
Locality. — Station 174 l (b, c, or d), August 3, 1874; near Kandavu, Fiji; lat.
(about) 19° 6' S., long, (about) 178° 18' E.; 255, 610, or 210 fathoms; coral mud;
bottom temperature (at 610 fathoms), 39° F. One specimen.
Remarks. — This species stands very close to Antedon brevicuneata, which was
brought from Amboina to the Leyden Museum, and it is with some hesitation that I
1 The exact locality, and consequently the exact depth, is not recorded.
236 THE VOYAGE OF H.M.S. CHALLENGER.
have separated them. The large size of the centro -dorsal and the lateral flattening of
the rays appear in both types. But in Antedon similis the greater part of the second
radials is concealed (PI. XLVII. fig. 1), which is not the case in Antedon brevicuneata ;
while the lower pinnules are smaller in the latter type, though it is individually of
larger size and has palmar axillaries developed on all the secondary arms instead of on
the outer arms of each ray only. The fourth pinnule of Antedon similis is similar to
and of almost the same length as the third ; whereas in Antedon brevicuneata it is a
good deal shorter and has a smaller number of joints. It is in the proportions of these
pinnules and the characters of the second radials that the chief difference between the
two types presents itself.
One of the arms of this specimen bore a Myzostoma -cyst of a somewhat peculiar
type. It was entirely independent of the arm- and pinnule-joints, but consisted of a
number of relatively large granules of limestone, irregularly aggregated together on the
ventral surface of the arm.
8. Antedon occulta, n. sp. (PI. XL VIII. figs. 1, 2; PI. XLIX. figs. 3, 4).
be
Specific formula • — A. 2. 2. 2.-^-.
Centro-dorsal a thick disk, reaching 6 mm. in diameter, and bearing thirty -five to
forty-five marginal cirri. These have twenty-five to thirty tolerably uniform joints,
the later ones compressed laterally with a slight dorsal keel which passes into a faint
spine on the penultimate.
The first radials are entirely concealed, together with the greater part of the second
and also part of the axillaries. The rays may divide four times, and the lower joints
of adjacent rays are in close lateral contact and somewhat flattened, but are not specially
straight-edged. Each division is of two joints, the axillary without a syzygy and often
somewhat unsymmetrical. Thirty-six to forty-eight arms, of about one hundred and
seventy smooth joints, the first few quadrate and the following ones shortly triangular,
gradually becoming quadrate again, but remaining much wider than long till near the
end of the arm. A syzygy in the third brachial and another between the thirteenth
and thirtieth ; others at intervals of seven to seventeen joints.
The lower pinnules of the inner arms are generally rather smaller than those on the
outer arms of each distichal group, and more especially than those on the outer arms of
the rays. The first one may be 7 to 9 mm. long, with twenty to twenty -five joints, the
lowest of which are rather wide. The second pinnule may have thirty joints, the first
half of which are very stout, and reaches 10 or 15 mm. The third is sometimes nearly
equal to it, but is more usually considerably smaller both in length and in stoutness,
while its successor on the seventh brachial is always much smaller than the pinnule on
the fifth.
REPORT ON THE CRINOIDEA. 237
Disk naked and more or less incised ; saceuli abundant at the sides of the pinnule-
ambulacra.
Colour in spirit, — the skeleton brownish-white, and the perisome the same or
greenish -grey.
Disk 12 mm.; spread reaching 22 cm.
Locality. — Station 174 l (b, c, or d), August, 3, 1874; near Kandavu, Fiji; lat.
(about) 19° 6' S., long, (about) 178° 18' E.; 255, 610, or 210 fathoms; coral mud;
bottom temperature (at 610 fathoms), 39° F. Three specimens.
Remarks. — These three individuals, which are somewhat variable in their characters,
but apparently belong to the same specific type, were obtained at Station 174 together
with the single example of Antedon similis. They all agree in the presence of one or
more post-palmar series, in the great development of the centro-dorsal, so as to partly
cover the axillaries, and in the absence of the sharp straight edges to the distichal and
palmar joints on the outer sides of the rays, which are so marked in Antedon similis
(PI. XLVII. fig. 1). They are therefore pretty clearly distinguished both from this
type and from its close ally Antedon brevicuneata. But they vary considerably in the
characters of their lower pinnules. Those at the outer side of each distichal group, and
more especially the outer pinnules of the rays, are generally rather longer and stouter
than the corresponding pinnules on the inner arms ; but I have been unable to make out
any great constancy in this arrangement, and it is much more marked in one of the two
specimens with the lower pinnules exposed than it is in the other. The third pinnule is
generally much smaller than the second (PI. XLVIII. fig. 2 ; PI. XL1X. fig. 4), but is
sometimes nearly or quite equal to it in size, a character which may occur on the inner as
well as on the outer arms. In the individual which shows the greater inequality of the
pinnules on the inner and outer arms (PL XLIX. figs. 3, 4), they are generally stiffer
and more styliform than in the more regular example (PI. XLVIII. figs. 1, 2). The latter
thus presents an approach toward Antedon conjungens, while the former rather resembles
Antedon protecta. These two species, however, have much less closely approximated
rays and a smaller centro-dorsal, which leaves the second radials visible as well as the
axillaries (PL XLV. fig. 1).
9. Antedon regalis, n. sp. (PL XLVL).
be
Specific formula — A.2.2. y.
Description of an Individual. — Centro-dorsal a thick disk, bearing about forty cirri
of twenty-five to thirty joints. The middle and outer joints are somewhat compressed
1 The exact locality, and therefore the exact depth, is not recorded.
238 THE VOYAGE OF H.M.S. CHALLENGER.
laterally, developing a bluntly pointed keel, which passes into the dorsal spine of the
penultimate.
The angles of the first radials just visible ; the second short and partly united
laterally ; axillaries wide, more than twice their length, and almost triangular. The
rays may divide three times, each division of two joints, the axillary without a syzygy.
The first few joints above the radial axillary on the outer side of the ray have their
outer edges curved and folded ; while the lower brachials, both of the inner and of the
outer arms, have their apposed sides flattened against one another. Twenty-seven long and
tapering arms, of about one hundred and eighty joints, the lower ones discoidal and their
successors shortly triangular, becoming more quadrate in the middle, and in the terminal
third more nearly square, elongating slightly towards the end. A syzygy in the third
brachial ; the next between the fifteenth and eighteenth, with others at intervals of
eight to eighteen joints.
The first pinnule on the outer side of the ray may reach 8 mm., with twenty-seven
joints, but on the inner arms it is generally somewhat smaller. That of the third brachial
is about equal to it. The second pinnule is also rather larger on the outer than on the
inner arms, reaching 1 5 mm., with about thirty joints, of which the first third are moderately
stout, and the remainder more slender and somewhat elongated. The pinnules of the
next three brachials (fifth to seventh) are of nearly equal size, but the fourth pair are only
about half their length, with fifteen joints, and the next pair are still smaller.
Disk naked and much incised ; sacculi abundant at the sides of the pinnule-ambulacra.
Colour in spirit, — dark purple, with greenish-white spots on the disk.
Disk 20 mm.; spread about 30 cm.
Locality. — Tongatabu reefs. One specimen.
Remarks. — This fine specimen is not unlike Antedon articulata, Muller, but has a
smaller number of cirrus-joints, with less well defined spines than occur in that species.
In fact the spines are hardly anything more than a small pointed process in the middle of
the sharp dorsal keel. The fourth pinnule is relatively smaller and the second syzygy
nearer the disk than in the type of Antedon articulata ; and there are less than thirty
arms, instead of nearly forty, or even more, as palmar axillaries are not always developed,
and there are no post-palm ars at all.
Antedon, Series IV.
Three distichals, the first two articulated, and the third axillary with a syzygy.
Remarks. — The tridistichate species of Antedon are less numerous than those in
which only two distichals are present, but the two series have many points of
resemblance, both in their distribution and in their modifications of structure.
REPORT ON THE CRINOIDEA. 239
We have seen that, alike in the ten-armed and in the bidistichate series, two very
distinct morphological types are to be found, the one with the rays flattened laterally
and a complete ambulacral skeleton, and the other without these characters. The first
of these, represented by the Basicurva- and Spinifera-gromps, is restricted almost
entirely to the continental and abyssal regions, and is especially characteristic of the
Pacific Ocean and Eastern Archipelago. One abnormal form occurs in the littoral
fauna of the Southern Australian Seas ; a few typical but isolated species have been
found at considerable depths in the Atlantic and Southern Sea ; while three more range
from 80 to 270 fathoms in the Caribbean Sea. Among the tridistichate species of
Antedon there are seven which have wall-sided rays and an ambulacral skeleton. Two
of them occur in the Philippine Archipelago ; a third was obtained at two (or perhaps
three) stations in the South Pacific ; the Challenger dredged two more at a station near
Ascension (Station 344), one of which was also found off Tristan da Cunha (Station 135g) ;
and yet two more appear in several of the " Blake " dredgings in the Caribbean Sea at
depths of 80 to 270 fathoms. These seven forms may be spoken of collectively as the
Granulifera -group, after the name of a Caribbean species which was described by
Pourtales in 1878 ; and they have essentially the same distribution, both bathymetrical
and geographical, as the Basicurva- and ^:»« (/era-groups, which are also distinguished
by wall-sided rays and an ambulacral skeleton.
On the other hand, the tridistichate species of Antedon, which have unprotected
ambulacra and no flattening of the lower ray-joints (Savignyi-group), belong for the
most part to the littoral fauna of Northern Australia and the great Eastern Archipelago,
ranging northwards to Hong Kong and Japan. Individual species occur here and there
on the western shores of the Indian Ocean ; but I am not aware that there is any
tridistichate Antedon in the Atlantic or in the Caribbean Sea which has simple rays and
unprotected ambulacra.
8. The Gra?iulifera-grou]).
Tridistichate species with plated ambulacra and the lower parts of the rays flattened
laterally.
Remarks. — Four of the five members of the Granidifera-grou-p which are
considered in this Report are constructed upon the same type as Antedon granulifera
itself.
Neither the lateral flattening of the rays, nor the plated disk and ambulacral
skeleton of this species, seem to have attracted the attention of Count Pourtales when he
examined it, though they have since turned out to be characters of primary systematic
value. He described the type ' as having " three brachials between primary and
1 Bull. Mus. Comp. Zool., 1878, vol. v. No. 9, p. 215.
240 THE VOYAGE OF H.M.S. CHALLENGER.
secondary axials, one between secondary and tertiary." This would now be stated as
" three distichals, the axillary a syzygy, and two palmars united by syzygy " ; but in
neither case is the syzygy at all easy to recognise, and bis omission to notice it is
therefore not surprising. He found, however, that " sometimes there are syzygia in the
first and second joints of the arms." This, interpreted by the light of later morpho-
logical work, would mean that the first two brachials are united by syzygy,1 and that
the third may also be a syzygial joint. Antedon granulifera thus presents a type of
structure which we have not yet studied. There is no syzygy between the two outer
radials or between the first two distichals, and yet the first two joints above the
distichal axillary are united by syzygy, instead of by tbe usual bifascial articulation ;
while the normal syzygial union in the third brachial may or may not persist. If a
palmar series is present it consists of two joints united by syzygy, just as the first two
(p )5?'
brachials are ; and the formula of the type thus becomes A. 3. — is—. This also holds good
for Antedon distincta of the Challenger collection, as seen in PL LI. fig. 1. The
unique specimen of this fine species has its full complement of ten distichal series,
which are all normal in character, and it has already been noticed on p. 55 as illustrating
an exceptional type of arm-structure. But in Antedon granulifera some of the distichal
series are usually absent, so that the arms spring directly from the radial axillaries.
This is frequently also the case in Antedon angusticalyx and Antedon inaequalis, which
are constructed on the same type as Antedon distincta and Antedon granulifera, except
that there is normally no axillary above the distichal (PI. L. fig. 1 ; PL LI. fig. 2).
But in all three species alike the first syzygy of an arm which starts directly from the
radial axillary is in the third brachial, the two preceding joints being united bifascially.
We thus meet with a reversion from the abnormal grouping of the syzygies, which is
most fully developed in Antedon distincta (PL LI. fig. l), to that of the simple ten-
armed type with the first syzygy in the third brachial.
Antedon multispina affords another excellent instance of the same kind. Four
individuals of this type were obtained by the Challenger, three of them having only ten
arms. But in the fourth (PL LXIX. figs. 1, 2) there are two tridistichate series, and in
each of the four arms which are thus produced the first two brachials are united by
syzygy. This is not the case, however, in the tridistichate varieties of Antedon
rosacea, Antedon variipinna, and Antedon anceps (PL XXXV. fig. 1), which retain the
third brachial as a syzygial joint above the intercalated distichal axillary, and thus
remain normal in character.
The syzygial union of the two lowest brachials above the distichal axillaries of
Antedon multispina thus gives an important clue to its affinities, which is of the greater
1 My reasons for considering this union as a syzygy between the first two brachials, and not as a syzygy in the
first brachial, will be found in Part I. pp. 51, 52.
BEFOPT ON THE CBINOFDEA. 241
value, as there is no tridistichate Antedon of the normal type with a syzygy in the
third brachial which has flattened rays and protected ambulacra.
A still more striking instance of reversion to the more generalised condition is some-
times met with both in Antedon angusticalyx and in Antedon insequalis. The tridistichate
series may be replaced by a bidistichate one ; but this apparently unimportant variation
is always accompanied by a change in the grouping of the syzygies above the distichal
axillary. Except in one or two abnormal species like Actinometra pulchelld, the first
two brachials are never united by syzygy when they follow two articulated distichals ;
and so when a bidistichate series occurs as a variation in Antedon angusticalyx or
Antedon insequalis the first two brachials are articulated, and there is a syzygy in the
third. This is well seen in the two lateral rays of the figured specimen of Antedon
angusticalyx1 (PL L. fig. 1).
There is one species of the Granulifera-gcowp which presents a different type of arm-
structure from the rest. Antedon porrecta is also tridistichate, but the second joint
above the distichal axillary is a syzygial one (PI. LII. fig. 3). There are often no
palmars ; but when they do occur the series consists of two joints, the axillary a syzygy,
so that the formula of the type becomes A.3.2{(p..)br} . This is the only species of
Antedon with a syzygy in the second brachial, though the character is a common one in
Actinometra, as seen in PI. LX.; PI. LXII. fig. 3 ; and PI. LXVI. figs. 1, 4.
The species of the Granulifera-gTowp may be classified as follows : —
A. A syzygy between the first two brachials.
I. Calyx and arm-bases not spinous. The first two pinnules about equal,
with compressed and carinate joints ; the genital pinnules have un-
equally expanded joints.
a. Primary arms of adjacent rays in close lateral contact. Palmars
usually absent ; the second syzygy generally not beyond the
fifteenth brachial.
1. Cirri smooth and without an opposing spine. First radials
invisible; the second very short and deeply incised. The
lower joints of the distichal pinnule not specially wide, 1. angusticalyx, n. sp.
2. Cirri somewhat carinate, with an opposing spine. First
radials partly visible. The lowest joint of the distichal
pinnule much wider than the rest, . . .2. inxqualis, n. sp.
b. The distichal axillaries of adjacent rays partially separated by the
pinnule of the preceding joint. Palmars usually present ; the
second syzygy from the twentieth to the twenty-fifth brachial.
1. The lower pinnules rather stout, .... .7>'aM"^/era,Pourtales.
2. The lower pinnules comparatively slender, . . 3. didincta, n. sp.
II. Calyx and arm-bases spinous. The first pinnule much longer than the
second, with stout joints, the lowest of which have their inner edges
cut away. The genital pinnules have uniformly expanded joints, . 4. multispina,2 n. sp.
B. A syzygy in the second brachial, . . . • • .5. porrecta, n. sp.
1 The fourth brachial is the syzygial joint in one arm of the right-hand ray.
2 Some forms of this species have only ten arms ; see p. 117.
(ZOOL. CHALL. EXP. PAKT LX. 1887.) 000 31
242 THE VOYAGE OF H.M.S. CHALLENGER.
1. Antedon angusticolyx, n. sp. (PI. II. figs. 4, a-d ; PL L. figs. 1, 2 woodcut on
p. 246, fig. 5, B ; also Part I. pi. liv. fig. 5 ; pi. lv. fig. 6).
Specific formula — A. 3. "o.y.
Centro-dorsal a truncated hemisphere, marked by indistinct interradial ridges which
are produced upwards into rather prominent processes between the radials. Twenty to
twenty-five cirri, of eighteen to twenty-three smooth joints, most of which are slightly
longer than wide ; the penultimate without an opposing spine.
First radials entirely concealed ; the second quite short, especially in the middle line,
barely in contact above the angles of the centro-dorsal. They are deeply incised by the
tubercular backward processes of the axillaries, which sometimes almost reach the centro-
dorsal. Three distichals, the junction of the first two somewhat tubercular, and the
axillary a syzygy, in close contact with its fellow on the next ray. The outer radials
and the three distichal joints have sharp straight edges, and both sides flattened, and the
first two or three brachials may show the same characters, but to a less extent.
Fourteen to twenty arms, of over one hundred joints, of which the lowest are nearly
oblong, their successors triangular, and wider than long, gradually becoming longer and
more quadrate.
The first syzygy is in the third brachial when the primary arms do not divide, and
the next between the eleventh and fifteenth. When distichals are present the first two
brachials are usually united by syzygy, and the next syzygial joint is from the
seventh to the twenty-fifth brachial, usually about the twelfth or fourteenth. After
this an interval of four to twelve, generally six or seven joints, between successive
syzygia.
The distichal pinnule is about 9 mm. long, and consists of some thirty short, carinate
joints, the lowest of which, though thick, are not specially wide. The next two or three
pinnules are of about the same length, with relatively longer terminal joints, and the
lower ones somewhat Matter. The following pinnules are a little shorter, with the first
two joints smaller than in the proximal pinnules ; but the third joint and from two to
four of its successors are broad and flattened, with the outer edges much produced towards
the ventral side. Traces of this expansion may be visible as far as the twenty-fifth
brachial, after which the joints become elongated and the pinnules more slender.
The disk is much incised and completely plated, as are also the brachial ambulacra
and the interarticular spaces. The genital glands are covered by closely set plates in
which sacculi are embedded. These are small and inconspicuous on the pinnule-
ambulacra, which have well-defined side plates.
Colour in spirit, — light whitish-brown;
Disk 7 mm.; spread 15 cm.
REPORT ON THE CR1NOIDEA. 243
Locality. — Station 214, February 10, 1875 ; off the Meangis Islands; lat. 4° 33' N.,
long. 127° 6' E.; 500 fathoms; blue mud; bottom temperature, 41°-8 F. Several
specimens, some with cysts of Myzostoma tenuispinum.
Remarks. — This species is readily distinguished from all the other tridistichatc forms
of Antedon, with the exception of Antedon inasqualis (PI. LI. fig. 2), with which it has
many characters in common. It does not reach the size of that species, however, and
differs in various respects from its younger stages, as will be explained further on.
The cirri have rather elongated joints, which are unusually smooth, hardly any trace
of an opposing spine appearing on the penultimate. The centro-dorsal is much flattened
at the dorsal pole and has more or less distinct indications of interradial ridges on its
sides, which are produced upwards into rather prominent processes at the angles (PI. II.
fig. 4a). It is considerably wider than the radial pentagon (PL II. fig. Ad), so that the
first radials are entirely concealed by it, with portions of the second as well. Both
edges of these latter joints are thus strongly curved in the adult calyx, the proximal
edges occupying the hollows between the interradial processes of the centro-dorsal,
while the distal edges are incised to receive the strong backward processes of the
axillaries (PL L. fig. 1). The first two distichals, or in their absence the first two
brachials, have a similarly tubercular junction.
Antedon angusticahjx is a species of considerable interest from its presenting several
of the characters which are distinctive of three species of Antedon that were found
associated in the South Pacific, near the Kermadec and the Fiji Islands respectively
(Stations 170a and 174). Two of these, with only ten arms {Antedon basicurva and
Antedon incisa), are characterised by having less than thirty smooth cirrus-joints, and
some of the lower joints of the genital pinnules expanded on the outer side so as to
form a protection for the genital glands, which are also covered by a strong anambulacral
plating (PL XXI. figs. 2a, 2b). Antedon insequalis (PL LI. fig. 2), which also occurred
at both Stations (Nos. 170a, 174), is a tridistichate species possessing these same
peculiarities ; while Antedon angusticalyx, which closely resembles it in the characters
of the arm-divisions and genital pinnules (PL L. figs. 1, 2), represents the tridistichate
type in the North Pacific. But the sides of its rays are less distinctly flattened than in
the three species from the South Pacific ; while those of the ten-armed species (Antedon
acosla), which is associated with it, have no flattening at all (PL XVI. fig. 1), though
the cirri and genital pinnules have much resemblance to the corresponding parts of
Antedon basicurva and Antedon incisa.
Some of the characters of Antedon angusticahjx and Antedon iniequalis appear in
Antedon granulifera of the Caribbean Sea. But this type usually has two post-radial
axillaries, i.e., distichals and palmars, and the rays are less closely in contact than is the
case in the Pacific species.
244 THE VOYAGE OF H.M.S. CHALLENGES.
2. Antedon inasqualis, n. sp. (PI. II. figs. 5, a-d ; PL LI. fig. 2 ■ woodcut, p. 246,
fig. 5, A ; also Part I. pi. liv. fig. 8).
Specific formula — A. 3. — 9~~-"y-
Centro-dorsal hemispherical, rather flattened at the dorsal pole, and bearing twenty
to twenty-five cirri. These have about twenty joints, a few of which are longer than
wide ; the later ones are somewhat compressed laterally and more or less distinctly
carinate ; the penultimate with an opposing spine.
First radials partially visible above the angles of the centro-dorsal; the second
short, sharply convex, and closely united laterally. Auxiliaries short, broadly pentagonal,
and very convex iu the centre, forming a median tubercle with the second radials.
Three distichals with a syzygy in the axillary, which is in close contact with its
fellow on the next ray, and another syzygy between the first two brachials. These five
joints, and in a less degree also the two outer radials and the third brachials, are in
close lateral contact and very distinctly wall-sided, with sharp edges and the margins
of the dorsal surface a little depressed. The second, and occasionally also the third,
brachial may likewise be slightly flattened on both outer and inner sides. One
specimen has two palmars united by syzygy, and another two with the axillary a
syzygy.
Eleven to twenty arms of some one hundred and twenty joints, the lowest nearly
oblong, and the following ones triangular, as long as wide, and gradually becoming more
cpaadrate. The pieces of the calyx and the lower parts of the arms often have some-
what prominent edges. On the arms, which start directly from the radial axillary, the
third brachial is a syzygial joint, and the next syzygy is between the fourth and
the thirteenth brachials ; but when distichals are present the first two brachials are
generally united by syzygy, and the next syzygial joint is from the seventh to the
tenth brachial. After this there is an interval of two to fifteen, usually four to seven,
joints between successive syzygia.
The second distichal bears a small pinnule, 7 mm. long, which consists of some
twenty to twenty-five short joints, the lowest of which, and especially the first, are
wide, trihedral, and flattened against the arm, while the remainder are slightly carinate.
The next pinnule (on second brachial) is a trifle longer, with relatively long terminal
joints, and the basal ones less wide and more carinate. The third and following
brachials have stdl longer and stouter pinnules (12 mm.), with the outer edges of the
third and the two to four following joints much produced towards the ventral side, so as
to give them a broad and flattened appearance. The length of the pinnules decreases
somewhat after the sixth brachial, but the expansion of their lower joints is traceable
till the fifteenth or twentieth, after which they become more slender, with only the two
lower joints wider than long. Disk much incised and completely plated, as are also the
REPORT ON THE CRINOIDEA. 245
arms, both along the ambulacra and at their sides. A pavement of anambulacral plates
eovers the genital glands. The pinnule-ambulacra have well-defined side plates, alter-
nating with and partly concealing the sacculi, which are mostly small.
Colour in spirit, — light whitish-brown.
Disk 10 mm.; spread 20 cm.
Localities. — Station 170a, July 14, 1874, near the Kermadec Islands; lat. 29° 45' $.,
long. 178° 11' W., 630 fathoms; volcanic mud; bottom temperature, 39°-5 F. Twelve
specimens ; two of them with cysts of Myzostoma tenuispinum and Myzostoma
willemoesii.
Station 174 (b, c, or d), August 3, 1874, near Kandavu, Fiji ; lat. (about) 19° G' S.,
long, (about) 178° 18' E.; 255, 610, or 210 fathoms;1 coral mud ; bottom temperature
(at 610 fathoms), 39° F. Five specimens, one with Myzostoma- cysts, and some free
individuals.
Doubtful— Station 175, August 12, 1874, near Kandavu, Fiji; lat, 19° 2' S., long.
177° 10' E.; 1350 fathoms; Globigerina ooze; bottom temperature, 36° F. One
broken specimen.
Remarks. — This species is rather closely allied to Antedon angusticalyx, but reaches
a considerably larger size. The cirri are often slightly carinate, with a tolerably distinct
opposing spine on the penultimate.2 The radial pentagon is relatively larger than in
Antedon angusticalyx, so that it completely covers the centro-dorsal (PL II. figs. 4a,
id, 5a, 5a7), and the whole of the second radials, together with more or less continuous
portions of the first, are thus visible on the exterior of the calyx. The axdlaries are
relatively short, and have no such large tubercular projections into the second radials as
are visible in Antedon angusticalyx (PI. L. fig. 1 ; PL LI. fig. 2).
The difference between the calyces of the two types, which are so closely similar in
other respects, comes out very clearly if a young individual of the larger Antedon
insequalis be compared with a mature Antedon angusticalyx of equal size. The first
radials of the former are completely visible, forming a narrow but continuous band
between the centro-dorsal and the second radials, which plates are not incised by the
short axillaries ; whereas in Antedon angusticalyx the first radials are entirely concealed,
and the second are rather deeply incised by the tubercular backward projections of the
axillaries.
The characters of the distichal pinnules afford another good distinction between the
two types. Those of Antedon angusticalyx have somewhat carinate joints, the lowest
of which are rather wider than the rest, though not markedly so (fig. 5, b), but in Antedon
insequalis the lower joints are generally more rounded and less carinate, while the first,
1 The exact station, and consequently the exact depth, are not recorded.
2 The two cirri which remain on the figured specimen are rather more smooth than is usually the case.
246
THE VOYAGE OF H.M.S. CHALLENGER.
A B
Fig. 5. — The lowest pinnules of Ante-
don incequalis (A) and of Antedon
angusticalyx (B). x 3.
or sometimes the first and second, is considerably wider than its successor (fig. 5, a). It is
produced towards the ventral side, so that it has a large flattened lateral surface corre-
sponding to those of the first and second distichals, which are both relatively and
absolutely larger than the same parts in Antedon angusti-
calyx (fig. 5, b), and are also divided by a groove into two
portions at different levels, which is not the case in the
latter species.
The number of arms which may be present in Antedon
in&qualis varies very considerably, just as in Antedon
angusticalyx. Several individuals have twenty, but fourteen
to sixteen is a not uncommon number, and two specimens
have only eleven ; so that it is quite possible that this may
really be a dimorphic species, and that it should find a place
with the ten-armed series near to Antedon basicurva, as
well as in the tridistichate group. Palmar series occur iu
two specimens. In one there are two palmars united by syzygy, just as is naturally
the case in Antedon distincta (PL LI. fig. l), while the other presents the type of
Antedon porrecta, viz., two palmars, the axillary a syzygy (PL LI. fig. 2; PL LII. fig. 3).
This, however, is an abnormal variation owing to the intercalation of a joint above the
distichal axillary ; because the first two brachials are united by syzygy in the ordinary
way, instead of being articulated with a syzygy in the second one as in Actinometra
sentosa and Actinometra midtiradiata (PL LXVI. figs. 1, 4).
One example of Antedon insequalis and some fragments of Pentacrinns naresianus
were sent me with the label of Station 175; but there is no record in the Station Book of
their having been dredged at this Station (1350 fathoms), though there are two Comatulse
mentioned which reached me with the corresponding label. These are the ten-armed
Antedon breviradia and Antedon acutiradia (PL XL figs. 3, 5), which have the general
facies of abyssal forms ; and as no other Antedon with more than ten arms has been
obtained from a greater depth than 750 fathoms, I much doubt whether Antedon
inasqualis really was obtained from 1350 fathoms at Station 175.
Both Stations 170a and 174 yielded examples of Antedon insequalis with the cysts
of Myzostoma tenuispinum, which also occurred on the allied species Antedon angusticalyx
at Station 214. One individual from Station 170a, with four cysts of this Myzostoma,
had another of Myzostoma willemoesii ; while at Station 174 an individual was found
with cysts of Myzostoma tenuispinum, and also a combined cyst of this species and
Myzostoma willemoesii, which type likewise occurred at Station 170a on Antedon
basicurva.
REPORT ON THE CRINOIDEA. 247
3. Antedon distincta, n. sp. (PI. LI. fig. 1).
Specific formula — A. 3.— . — .
Description of an Individual. — Centro-dorsal a thick disk, with the angles somewhat
produced and some twenty -five cirri on its sides. These have about twenty rather stout
joints, of which the sixth and seventh are the longest. The following joints are shorter
and gradually develop a dorsal keel.
The first radials not visible ; the second are short and much curved, and the axillaries
subtriangular, both joints being very convex, with a rather sharp dorsal edge. Three
distichals, the first two very convex at the junction and the axillary a syzygy. The
first two brachials, or the two palmars when present, united by syzygy. The outer
radials and the first two distichals are in close lateral contact, with sharp edges and
flattened sides ; and in the middle line of the ray this character is continued on to the
distichal axillaries and the next four or five joints. But the outer sides of the distichal
axillaries are prevented from meeting those of adjacent rays by the pinnules of the second
distichal joints, which are placed very near the dorsal surface.
Twenty-seven arms, the first twelve joints nearly oblong and the following ones more
triangular, gradually becoming longer than wide. A syzygy between the first two
brachials, and the next about the twenty-fifth ; others at intervals of five to fourteen,
usually seven to ten, joints.
The distichal pinnule is about 8 mm. long and rather slender, composed of some thirty
small, compressed, and slightly carinate joints. The following pinnules are similar,
slightly decreasing in size to about the sixth brachial, after which the next eight or ten
pinnules on each side have the lower joints carinate and expanded on the outer side,
with the upper ones more styliform. This character gradually dies out, and the pinnules
become more elongated and slender.
Disk and brachial ambulacra well plated. The pinnule-ambulacra have well-defined
side plates which alternate with the sacculi.
Colour in spirit, — light whitish-brown, with patches of brownish-grey.
Spread 20 cm.
Locality. — Station 210, January 25, 1875; off the Panglao and Siquijor Islands;
lat, 9° 26' N., long. 123° 45' E.; 375 fathoms; blue mud; bottom temperature, 54°-l F.
One specimen.
Remarks. — Only a single example of this elegant species was obtained by the
Challenger ; and it seemed so different from all the other species of Antedon then known
that I proposed to call it Antedon distincta. Several months afterwards I received
Bourtales' description * of some new Comatula3 dredged by the first " Blake " expedition,
1 Bull. Mus. Comp. ZobL, 1878, vol. v. No. 9, p. 215.
HT^
248 THE VOYAGE OF H.M.S. CHALLENGER.
amongst which was Antedon granulifera with " three brachials between primary and
secondary axials, two between secondary and tertiary." The first part of this statement
clearly indicates that the type has three distichals, of which the axillary is a syzygial or
double joint. But it was impossible to tell from Pourtales' description whether the two
palmare are articulated or united by syzygy, though the latter condition seemed probable
from his further note that " sometimes there are syzygia in the first and second joints of
the arms." When the "Blake" collection came into my hands I found not only that
Antedon granulifera has the same grouping of the arms as Antedon distincta, but also
that it has an ambulacral skeleton and the rays flattened laterally, two characters of
which no hint was given in Pourtales' description. In fact, these two species, though so
widely separated geographically, are in reality very closely allied, the chief point of
difference between them being the greater size of the lower pinnules in Antedon
granulifera.
Antedon distincta differs from Antedon angusticalyx and Antedon miequalis in the
long interval between the first and second syzygies of the arms, and also in the
separation of the distichal axillaries of adjacent rays by the pinnules on the preceding-
joints, which are attached nearer to the dorsal surface than usual. This is less marked
in Antedon granulifera, though it agrees with Antedon distincta in the long syzygial
interval. On the other hand, the joints of the genital pinnules of Antedon distincta are
more uniformly expanded than in Antedon granulifera, which rather resembles Antedon
angusticalyx and Antedon inwqualis in this respect. But in all four species alike the
outer side of each pinnule-joint is more expanded than the inner one, just as in Antedon
hasicurva and Antedon incisa (PI. XXI. fig. 2), while in the tridistichate variety of
Antedon multispina the large joints of the pinnules are broadly V-shaped and similarly
expanded on both sides.
Antedon granulifera seems to be fairly abundant in the Caribbean Sea; but it
exhibits a good deal of variation in its characters, which will be fully discussed in the
report on the " Blake " Comatulse.
4. Antedon midtispina, n. sp. (PL XIII. figs. 1-3 ; PI. XIV. figs. 5-7 ; PL L.
figs. 3-6 ; PL LXIX. figs. 1-4).
Specif c formula — A.f — J. y.
Localities.— Station 135g, October 18, 1873; off Tristan da Cunha; lat. 37° 10'50"S.,
long. 12° 18' 30" W.; 550 fathoms ; hard ground. One mutdated specimen.
Station 344, April 3, 1876 ; near Ascension; lat. 7° 54' 20" S., long. 14° 28' 20" W.;
420 fathoms ; volcanic sand. Four broken individuals and three Pentacrinoids.
REPORT ON THE CRINOIDEA. 249
Remarks. — Three individuals of this species, and also three larvse, all with ten arms,
were obtained at Station 344, near Ascension, and have been already described.1 But
another individual from the same station must be noticed here from its having two
tridistichate series. It resembles Antedon ancjustieahjx and Antedon iniequcdis in the
syzygial union of the two lowest brachials, but it differs both from them and from the
other tridistichate species in its spiny calyx and in the characters of the pinnules. The
first pinnule (PI. LXIX. figs. 2, 3) consists of rather massive joints with their inner edges
cut away a little and the outer sides slightly flattened, presenting, in fact, the same
characters, though in a less prominent form, as the first pinnules of Antedon valida,
Antedon incerta, and their allies among the ten-armed species of the Basicurva-gvoi\-p
(PI. XV. figs. 5, 6 ; PI. XVIII. fig. 5). The first pinnule of Antedon multispina is
much larger than its successor, a character which distinguishes the type both from
the species just mentioned and from the other members of the Grramdifera-growp,
from which it also differs in the uniformly expanded shape of the large joints of the
genital pinnules.
Station 135g, off Tristan da Cunha, yielded a single mutilated Antedon (PI. L.
figs. 3-6), which after some consideration I have decided to refer to this species, though
I was at first inclined to place it elsewhere. The cirri are generally similar to those of
the more northern form (PI. XIII. fig. 1 ; PL L. fig. G), but may have as many as
thirty-five joints, while the number does not exceed thirty in the smaller and premature
individuals from near Ascension. The latter do not show the first radials externally
(PI. LXIX. figs. 1, 2), but they are visible in the larger calyx of the southern variety
(PI. L. fig. 3), which is also less distinctly spinous than that of the northern individual,
and the same is true of the arm- and pinnule-joints.
The first pinnules of the southern form have somewhat the same flattened appearance
on their outer sides as is traceable in that from Ascension (PI. LXIX. figs. 2, 3), and
is more marked in the typical members of the Basicurva-gvowp (PI. XV figs. 5, G ;
PI. XVIII. fig. 5). But it is so slight as to be hardly recognisable except by a trained
eye, and the same may be said of the lateral flattening of the lower brachials. In fact this
variety of Antedon midtispina is a good connecting link between the Basicurva- and
Gramdifera- groups on the one hand, and the ordinary Comatuke with normal rays and
unplated ambulacra on the other, for the plating of the disk is very incomplete (PI. L.
fig. 4) and the ambulacral skeleton of the pinnules by no means well differentiated.
There are thirteen arms in this individual, owing to the presence of three distichal
series. One of these is only two-jointed, and the first syzygy above it is in the third
brachial (PI. L. fig. 3), just as in the case of Antedon anyusticalyx already referred to
on p. 241 (PI. L. fig. 1). But of the two arms which follow each tridistichate series one
has the normal syzygy (for this type) between the first two brachials, while in the other
■See pp. 117-119.
(ZOOL. CHALL. EXP.— PART LX. — 1887.) OoO 32
250 THE VOYAGE OF H.M.S. CHALLENGER.
these two joints are articulated and the first syzygy is in the third brachial ; so that we
here get an approach to the characters of the Savignyi-gvowp, next to be described,
while the bidistichate series indicates a similar variation towards the Pcdmata-groii]).
In this single individual, therefore, we meet with the characters of one ten-armed and
three multibrachiate types of Comatuhe, and its true affinities would have been a matter
of some doubt, but for the presence of more normal individuals of the same type at
another station.
Three Pentacrinoid larvae were also obtained off Tristan da Cunha, but at a consider-
ably greater depth (1000 fathoms) than the mature Antedon (Station 135e, October 18,
1873 ; Lit. 37° 21' 0" S., long. 12° 22' 30" W.; 1000 fathoms ; hard ground, shells, gravel).
The best preserved of them is represented on PL XIV. fig. 8. It appears to belong to a
ten-armed species, as is naturally to be expected at such a great depth ; and it has many
points of resemblance with the " cold area " larva which I have referred conjecturally to
Antedon hystrix (PL XIV. fig. 2). The basals are high and the first radials very wide,
while the two following joints are relatively long and narrow ; though a considerable
number of brachials are developed above them. These show no traces of an ambulacral
skeleton, however, as is the case in the youngest larva of Antedon midtispina (PL XIV.
fig. 5), which has only about the same number of arm-joints, though the calyx is
relatively much more developed than that of the abyssal larva.
5. Antedon porrecta, n. sp. (PL LII. figs. 3-5).
Specific formida — A.3.2 {( p.)br} .—.
Centro-dorsal a thick disk with the interradial angles slightly produced, and from
twenty to thirty long and stout cirri on its sides. They have from forty to fifty joints,
nearly all of which are wider than long, and produced on the dorsal side into a strong
pointed process. The first radials are invisible except at the angles of the calyx ; the
second and third both rather convex and slightly tubercular at the junction, the second
short, united laterally, and the axillaries broadly pentagonal, about two and a half times
their length. Three distichals and sometimes two palmars, each axillary with a syzygy.
These joints are very convex and have their inner sides flattened against one another ;
but this is less marked at the outside of the rays where the hypozygals of the distichal
axillaries and of the second palmars (or brachials) are kept apart by the large pinnules
on the preceding joints.
Over twenty arms, of compressed triangular joints, which become elongated and
quadrate towards the end. From the third brachial onwards the middle of the distal
edge of each joint is raised into a strong plate, the front face of which is hollowed.
Beyond about the tenth or fifteenth joint this gives place to an overlap of the usual
character, which extends far out on the arm.
REPORT ON THE CRINOIDEA. 251
A syzygy in the second brachial, and the next between the sixth and fourteenth ;
others at intervals of three to twelve joints, the intervals often becoming shorter towards
the end of the arm.
The second distichal and the first palmar (or brachial) bear tolerably equal pinnules
of about fifteen stout joints, the five lowest of which are rather broad and trihedral, with
flattened outer faces and the inner sides slightly bevelled away. The second and third
brachials have smaller pinnules with fewer joints, the basal ones being more compressed ;
and the following pinnules are larger again, with broader lower joints, the outer edges of
which are expanded towards the ventral side. This arrangement gradually dies away
in the outer parts of the arms, and the joints become more elongated.
Disk slightly incised and well plated, like the brachial ambulacra ; the pinnule-
ambulacra have well-defined side plates and small sacculi.
Colour in spirit, — dark grey-brown.
Disk 15 mm.; spread probably 30 cm.
Locality. — Station 344, April 3, 1876; near Ascension; lat. 7° 54' 20" S., long.
14° 28' 20" W.; 420 fathoms ; volcanic sand. Four broken specimens.
Remarks. — This species cannot well be mistaken for any other, as it is the only
tridistichate Antedon with a syzygy in the second brachial (PI. LII. fig. 3); though the
type is common enough in the genus Actinometra (PI. LX.; PI. LXII. fig. 3). The
four specimens obtained were all much mutilated, the arms having broken away at
the syzygy in the second joint above the distichal axillary. In some cases this was the
hypozygal of the palmar axillary, but as only a few arms are preserved it is impossible
to determine their number, or whether palmar axillaries occurred in all the specimens.
Under these circumstances therefore I have thought it best to enclose the palmar sign
within brackets in the specific formula.
The length of the cirri and the strong dorsal processes on their numerous joints are
also good distinctive characters of the type. One cirrus, as shown in the figure (PI. LII.
fig. 3), has been fractured and subsequently regenerated, a somewhat rare occurrence, as
I have already remarked on p. 203. The characters of the pinnules of Antedon porrecta
are the same as those of Antedon basicurva and its allies, though in a less marked
degree. The lower joints of the genital pinnules are expanded towards the ventral side
so as to protect the genital glands, which have but a slight covering of anamljulacral plates,
while the first two pinnules have massive lower joints with the outer sides flattened just
as in Antedon valida and Antedon incerta (PI. XV. figs. 5, 6 ; PI. XVIII. fig. 5).
Apart from its general specific characters Antedon porrecta is remarkable as being
one of the few Atlantic species of the genus which have an amljulacral skeleton. It
was obtained in the neighbourhood of Ascension, together with the dimorphic Antedon
multispina, which also occurs near Tristan da Cunha, and they thus serve as a con-
252 THE VOYAGE OF H.M.S. CHALLENGER.
necting link between Antedon lusitanica and the Caribbean species (Antedon. granulifera,
Antedon spinifera, &c.) of the North Atlantic, and Antedon bispinosa of the Southern
Ocean.
9. The Savignyi-gvou-p.
Bidistichate species with an unplated ■ disk and no definite ambulacra! skeleton ; the
bases of the rays are not flattened laterally.
Remarks. — I have associated this group with the name of Savigny, as its earliest
described representative was brought by him from the Red Sea and named after him by
Midler ; while it is one of those which appear both with and without palmar series, and it
therefore has a wide range of alliances. It also occurs at Muscat and at Kurrachee, but
is not known to extend further eastwards.
Antedon reynaudl has been described from Ceylon, and I have seen some undescribed
species from Zanzibar in the continental museums. But all the remaining types of the
group belong to the littoral fauna of the eastern seas from Japan to Sydney, with the
exception of Antedon angustiradia, which was found by the Challenger at 140 fathoms
in the Arafura Sea (Station 192).
Some forms of this group have no palmars above the distichals ; while in others there
are palmar series, consisting sometimes of two and sometimes of three joints. I have not
thought it necessary, however, to separate these latter species as a distinct group. They
all belong to the same general tridistichate type, and may be classified as follows : —
A. Three distichals, not succeeded by palmars.
I. The centro-dorsal bears ten vertical rows of cirri with sixty or seventy
joints. The distichal pinnule longer than its successors, . .1. angustiradia, n. sp.
II. Not more than forty-five joints in the cirri, which are without definite
arrangement. The distal pinnule generally smaller than its suc-
cessors.
a. The joints of the lower pinnules without lateral processes.
1. Forty to forty-five cirrus-joints, which are mostly spiny ;
usual syzygial interval seven to ten joints, . . reynaudi, Mull., sp.
2. Twenty-five to thirty-five cirrus-joints ; usual syzygial
interval three to seven joints.
a. Twenty-five to thirty cirri with strong spines on
the later joints ; second syzygy about the
eighteenth brachial. Distichals always present
and sometimes palmars, . . . savignyi, Mull., sp.
(Z. Twenty cirri, the later joints not spinous ; second
syzygy not beyond the fourteenth brachial.
Distichals sometimes absent, . . .2. amoqps} n. sp.
b. The joints of the lower pinnules have lateral processes at their
ends, . . . . . . .3. variij)iniia,1 Carpenter.
1 These species may have only ten arms ; see p. 194.
REPORT ON THE CRINOIDEA.
253
R Palmar series developed above the distichals.
I. Two palmars, the axillary not a syzygy.
a. The joints of the lower pinnules have lateral processes at their
ends, . . .
b. The joints of the lower pinnules have no lateral processes.
1. Twenty-five to thirty-five cirrus-joints.
a. Twenty cirri, the later joints not spinous ; second
syzygy not beyond the fourteenth brachial.
Tolerably equal pinnules on second distichal
and second brachial, ....
(3. Twenty-five to thirty cirri with strong spines on
the later joints ; second syzygy about the
eighteenth brachial. Distichal pinnule smaller
than that of second brachial, .
2. Forty-five to fifty-five cirrus-joints.
a. Cirrus-joints smooth and longer than wide ; no
spine on penultimate, ....
[i. Cirrus-joints wider than long ; the later ones
with faint tubercles, and the penultimate with
a spine, .....
II. Three palmars, the axillary a syzygy.
a. Forty-five cirrus-joints; the later ones short and spiny,
b. Nearly sixty cirrus-joints, the later ones longer than wide and
quite smooth. The terminal joints of the lower pinnules
much smaller than the basal ones, ....
3. variipinna,'1 Carpenter.
4. quinduplicava, n. sp.
savignyi, Mull., sp.
acuticirra, Carpenter.
ludovici, Carpenter.
jili'dlberti, Mull., sp.
bipartipinna, Carpenter.
1. Antedon angustiradia, n. sp. (PI. XLV. fig. 4).
Specific formula — A. 3.— .
Description of cm Individual. — Centro-clorsal columnar, its sides bearing ten vertical
rows of cirri, usually four in each row, which alternate more or less with those of
adjoining rows. They are long and slender, reaching 25 mm. in length, with sixty or
seventy joints, a few of which are longer than wide. The distal edges of the outer half
have a forward projecting spine which becomes more marked in the shorter terminal
joints. Three radials visible ; the first short, and depressed at their lateral edges, the
second oblong, twice their length, quite free laterally, rather convex, and rising to the
middle of their junction with the pentagonal axillaries. The rays are quite free and may
divide a second time ; three distichals, the axillary with a syzygy. The first distichals
(or brachials) nearly oblong and quite free laterally ; the second quadrate, with a slightly
angular base.
Fourteen arms, of about one hundred joints, at first triangular and then quadrate, the
later ones becoming narrow and elongated. A syzygy in the third, and then between
the twelfth and fifteenth brachials, with others at intervals of one to six, usually four or
five, joints.
1 This species may have only ten arms ; see p. 194.
254 THE VOYAGE OF H.M.S. CHALLENGER.
The second joint above the radial axillary, whether brachial or distichal, has a pinnule
about 10 nam. long, composed of some thirty elongated joints, the basal ones being rather
stout. The next few pinnules are much shorter and less stout at the base, with fewer
joints, and then' successors increase again, becoming long and filiform in the outer parts
of the arms.
Disk naked and much incised ; the rays and the lowest pinnules somewhat webbed
by perisome ; sacculi abundant on the disk, arms, and pinnules.
Colour in spirit, — the skeleton a very light brown, and the perisome darker.
Disk 9 mm.; spread about 9 cm.
Locality. — Station 192, September 26, 1874, near the Ki Islands; lat. 5° 49' 15" S.,
long. 132° 14' 15" E.; 140 fathoms; blue mud. One specimen.
Remarks. — This type is unfortunately represented only by a single and much-
mutilated individual, the disk of which bears the cysts of Myzostoma inflator, von Graff.
It may be readily distinguished, however, from the other members of the group by the
abundance of long and spiny cirri, and by the complete freedom of the rays, the second
radials not coming into lateral contact at all.
One of the distichal series is only two-jointed, as shown on the right of the figure
(PL XL V. fig. 4); while in that on the left side the two elements of the axillary have
almost the appearance of being articulated and not united by syzygy. If this be really
the case, it is a somewhat anomalous condition, especially as the lower joint, the normal
" hypozygal," bears no pinnule.
2. Antedon anceps, n. sp. (PL XXXV. figs. 1-3).
Specific formula — A. ( 3 ). -r-.
Centro-dorsal a low convex disk with about twenty cirri on its sides. These have
twenty-five to thirty-five tolerably uniform joints, few of which are longer than wide,
the later ones being slightly carinate.
First radials partially visible ; the second short and partly united, forming a more
or less distinct tubercle at the middle of their junction with the widely pentagonal
axillaries. Ten to fourteen arms, distichal series being sometimes absent altogether ;
when present, they consist of three joints, the axillary with a syzygy. The arms have
about one hundred and fifty joints, the earlier ones triangular and much wider than
long, their successors becoming more quadrate and finally almost oblong, with a slight
tendency to overlap. A syzygy in the third, and then between the eighth and twelfth
brachials, with others at intervals of two to nine, usually four to seven, joints. The
first pinnule, whether distichal or brachial, is considerably smaller than its successor on
REPORT ON THE CRINOIDEA. 255
the same side. In arms which spring direct from the radial axillary the largest pinnules
are those of the fifth, sixth, and seventh brachials, which may reach 12 mm. long, and
consist of twenty smooth joints, most of them longer than wide, and the later ones
carinate. On the inner arm borne by a distichal axillary, the largest pinnules are those
of the fourth and fifth brachials; while on the outer arms these are little, if at all,
larger than that of the second brachial. But the third brachial always bears a small
pinnule.
Disk naked and much incised ; sacculi very abundant on the disk, arms, and
pinnules.
Colour in spirit, — the skeleton white, with patches or bands of a faded purple, and
the perisome darker.
Disk 8 mm.; spread 17 mm.
Locality. — Station 212, January 30, 1875 ; off Samboangan, Philippine Islands;
lat. 6° 54' N., long. 122° 18' E.; 10 fathoms ; sand. Three specimens.
Remarks. — This is another of those dimorphic species which may or may not have
distichal series ; and it has therefore been assigned a place among the ten-armed forms of
Antedon, as noticed on pp. 194, 198. It is readily distinguished from the other members
of the Savignyi-gvowp, which have no palmar series nor lateral processes on the pinnules
like Antedon variipinna (PI. XL VIII. fig. 3).
Antedon reynaudi has more numerous and spiny cirrus-joints and a longer syzygial
interval ; while there is a larger number of cirri in Antedon savignyi, also with spiny
joints, and the second syzygy is further from the calyx instead of being within the first
fourteen brachials as in Antedon anceps (PL XXXV. figs. 1, 2). The arrangement of
the lower pinnules of this type is rather peculiar. On the outer side of the ray the
distichal pinnule, when present, is much smaller than that of the second brachial above
it. That of the third brachial is again small ; but the next pair are little if at all larger
than that on the second. On the other hand, if there is no distichal pinnule, owing to
the arms springing directly from the radial axillary, the pinnule on the second brachial
is smaller than that on the fourth, and this again is smaller than those of the next three
joints (PI. XXXV. fig. 2). An arrangement intermediate between these two is found on
the inner arm of each pair borne on a distichal axillary, in which the fourth and fifth
brachials have the largest pinnules, that on the second, as the first pinnule on the arm,
being distinctly smaller.
256 THE VOYAGE OF H.M.S. CHALLENGER.
3. Antedon variipinna, Carpenter (PI. XXXVI. figs. 1-6; PI. XLVIII. figs. 3-5;
PI. XLIX. figs. 1, 2).
Specific formula — A.[3.(2)].-r-.
1882. Antedon variipinna, P '. H. Carpenter, Journ. Linn. Soc. Lond. (Zool), 1882, vol. svi.
p. 506.
1882. Antedon erenulata, P. H. Carpenter, Ibid., p. 507.
1882. Antedon decipiens, Bell, Proc. Zool. Soc. Lond., 1882, p. 534.
1882. Antedon irregidaris, Bell, Ibid., p. 534.
1882. Antedon decipiens, P. H. Carpenter, Ibid., p. 746.
1882. Antedon erenulata, P. H. Carpenter, Ibid., p. 746.
1882. Antedon irregularis, P. H. Carpenter, Ibid., p. 746.
1882. Antedon variipinna, P. H. Carpenter, Ibid., p. 746.
1884. Antedon decipiens, Bell, Bep. Zool. Coll. H.M.S. "Alert," Lond., 1884, p. 159, pi. xi.
figs. B, a.
1884. Antedon irregidaris, Bell, Ibid., p. 161, pi. xiii. figs. A, a-c.
Centro-dorsal a low and slightly convex disk, bearing from fifteen to thirty cirri on
its sides. They have twenty to thirty-five joints, some of the lower ones being longer
than wide. The later joints are usually somewhat compressed laterally and rather
sharply carinate in consequence, but they sometimes bear well-marked spines.
The first radials are never altogether invisible in a side view, and are often com-
paratively large and granulated externally. The second are short, wide, and laterally
united, forming more or less of a prominence at the middle of their junction with
the broadly pentagonal axillaries. The rays generally divide twice and sometimes three
times, the distichal series consisting of three joints, the axillary with a syzygy, and the
palmars (when present) of two joints, the axillary without a syzygy. The dorsal surface
of these radial, distichal. and palmar joints is often considerably arched.
The arms vary in number from eleven (probably even ten) to twenty-five or more,
and may have one hundred and eighty joints. The first six or eight brachials are
relatively short and wide, nearly oblong in outline, and often much rounded dorsally.
The following joints are more triangular, with a variable tendency to overlap dorsally,
and their broader ends project alternately on opposite sides of the arm to a greater or
less extent. Further out they become more quadrate again, but remain relatively short
and wide and more or less overlapping till almost the very end of the arm. A syzygy in
the third brachial, and the next between the tenth and fifteenth, wTith others at intervals
of six to twelve joints, usually nine or ten.
The first pinnules are comparatively small, and consist of about twenty short joints,
the lowest of which are broad and slightly carinate. The distichal pinnule (if present) is
smaller than that on the second brachial, and so is that on the third brachial. The
following pinnules may reach nearly 15 mm. in length, with as many as twenty-five
joints, which are both longer and stouter than in the lower pinnules. The relative sizes
REPORT ON THE CRINOIDEA. 257
of the pinnules vary greatly. In the outer arm of each distichal pair the largest pinnules
are generally those of the fourth and fifth brachials ; but in the inner arm they are on
the fifth and sixth, while in arms which are borne directly on the radial axillaries the
third pair of pinnules (on sixth and seventh brachials) are usually the longest. All but
the lowest of these large pinnules have strong and blunt lateral processes at their distal
ends. The next pair of pinnules are generally considerably smaller, with relatively
shorter joints, which gradually become more elongated in the slender distal pinnules, but
lose the lateral processes at their ends.
Disk naked and much incised. Sacculi small, but abundant.
Colour in spirit, — ashy-grey, white, or pale flesh colour, with frequent bands or
patches of purple or yellowish-brown ; sometimes purple with whitish bands.
Disk 10 mm.; spread 20 cm.
Localities. — Station 186, September 8, 1874, Prince of Wales Channel ; lat. 10° 30' N.,
long. 142° 18' E.; 8 fathoms; coral mud. Two specimens.
Arrou Islands. Two specimens.
Other Localities. — Canton; Borneo. H.M.S. "Alert" 1881, — Torres Strait; Prince
of Wales Channel; Dundas Strait (17 fathoms) ; Arafura Sea (Station 160, 32 to 36
fathoms).
Remarks. — This is in some respects the most remarkable species of Antedon that I
have yet seen. For it has a very considerable range of variation and has been described
under four different names. The first of these, by which it must henceforth be known,
was given by myself in 18821 to a tridistichate and bipalmar Antedon from Canton in
the Hamburg Museum, with sharply spinous cirrus-joints, serrate arms, and a tolerably
regular inequality in the relative sizes of the pinnules at the bases of the inner and outer
arms of each ray, these lower pinnules having projections at the distal ends of their
component joints. At the same time I described another new species, Antedon erenulata,2
from Borneo, as having some of these peculiarities, but remarked that " it is alto-
gether a larger species than Antedon variipinna, from which it is readily distinguished
by its crenulated first radials, tubercular arm-bases, and smoother arms, while the
inequality in the sizes of the lower pinnules is not of the same character in the two
species." The Challenger collection contains two individuals from the entrance to Prince
of "Wales Channel in Torres Strait which agree with the two just mentioned in several
points, but have no palmar series at all, while one of them has spines on the cirri, though
those of the other are only carinate. At first sight, however, it did not seem advisable
to unite these two forms in one specific type, the one having palmar series and the other
not, though I now know that I was wrong. The same course was taken two years later
1 Journ. Linn. Soc. Lond. (Zool.), 1882, vol. xvi. p. 506. ' Ibid., p. 507.
(ZOOL. CHALL. EXP. — PART LX. — 1887.) 0°° 33
258 THE VOYAGE OF H.M.S. CHALLENGER.
by Bell,1 who described the two allied species Antedon decipiens and Antedon irregu-
laris, the former with spiny cirri and no palmars, and the latter with palmars but
unarmed cirri. But their other characters, especially the short arm-joints and the
lateral projections on the lower pinnules, agree very closely with those of Antedon
crenulata. Bell appears to have regarded the absence of palmars in Antedon decipiens
and of cirrus-spines in Antedon irregularis, which has palmars, however, as sufficient to
separate both these types from Antedon crenidata. They had been dredged by H.M.S.
" Alert " on the north-east coast of Australia ; and when in August last I began to revise
the tridistichate species of Antedon in the Challenger collection, the descriptions of which
had been written five or six years before, I found that a form closely allied to Antedon
decipiens and an example of Bell's Antedon irregularis, but without palmars, had been
figured on PI. XLVIII. figs. 3-5 and PI. XLIX. figs. 1, 2 respectively. Both alike had
been obtained in Prince of Wales Channel, and had formerly seemed to me, as the
" Alert" specimens from the same locality did to Bell, to represent two different specific
types which could not be referred either to Antedon variipinna or to Antedon crenidata.
A third form from the Arrou Islands also appeared to be new, and 1 figured it under
the name of Antedon dubia (PL XXXVI. figs. 1-6), not being quite clear in my
own mind as to whether its tridistichate condition is the normal one or merely due to
regeneration of a ten-armed form, as is so often the case in Antedon rosacea and other
species.
Lately, however, I have made a critical study of all the " Alert " material, and have
also reconsidered my descriptions of Antedon variipinna and Antedon crenulata. The
result is that I find myself unable to discover any characters which are sufficiently
constant to be of specific value as distinguishing Antedon irregularis and Antedon dubia
from Antedon decipiens, or any of these three from Antedon variipinna and Antedon
crenidata. Bell2 had himself remarked after describing Antedon irregularis — " This
species has some resemblance to Antedon decipiens ; but it may be distinguished from it
by (a) the absence of spines from the joints of the cirri, (/3) the broader lower pinnules,
and (y) the greater length of the more distal pinnules." He gave no details, however,
respecting the relative sizes of the lower and distal pinnules respectively in the two
types, and after examining his material I find a difficulty in attributing the difference to
anything more than the size of the individual specimens, those of Antedon decipiens
being generally smaller than those of Antedon irregularis. The presence of spines on
the cirrus-joints of Antedon decipiens, and their absence on the more numerous joints of
the cirri in Antedon irregularis, seemed, however, to be good specific characters. But
when I came to examine the grey specimens from Prince of Wales Channel, which Bell
had provisionally regarded as a variety of the white individuals obtained at the same
locality, on account of their cirri being " rather more numerous and more jointed," I found
1 "Alert " Report, pp. 159-162. . ! " Alert " Report, p. 1C2.
REPORT ON THE CRINOIDEA.
259
the cirrus-joints to be also unprovided with definite spines, though they have the same
sharply carinate appearance as those of Antedon irregularis.
The first radials of these individuals are also mostly concealed, as is the case in
Antedon irregularis, though in the type of Antedon decipiens from the Arafura Sea
they are "quite distinct" as described and figured by Bell;1 but they are much less
distinct in the white individuals from Prince of Wales Channel. In all the specimens
from this latter locality, therefore, the first radials resemble those of Antedon irregularis
rather than the radials of Antedon decipiens; but some of them had spiny cirri as in
the type of Antedon decipiens, while in the others the joints are only sharply carinate
as in Antedon irregidaris. The arms and pinnules of all these specimens, however, are
most like those of Antedon decipiens.
It would seem impossible, therefore, to make any distinction between the two species
in the characters of either the arms, the radials, or the cirri ; and this conclusion is
confirmed by the following considerations. The two individuals from the Arrou Islands,
which I formerly referred to a new species, Antedon dubia, have about thirty-five cirrus-
joints, with the later ones carinate as in Antedon irregularis (PL XXXVI. fig. 1 ;
PL XLIX. fig. 1). But they have relatively large and conspicuous first radials with a
sculptured surface (PL XXXVI. fig. 1), exactly as in Bell's figured specimen of Antedon
decipiens, which, like these, has no palmars. The arm-bases of the smaller individual
from the Arrou Islands resemble those of Antedon decipiens, while those of the larger
one show more of the characters of Antedon irregidaris. On the other hand, Antedon
variipinna and Antedon crenulata both have palmar series and thirty or more spiny
cirrus-joints, while the first radials are fairly distinct, those of Antedon crenulata being
more or less sculptured. Neither species has specially rounded arm-bases, like those of
Antedon irregularis, though the general outline of the joints is the same in all the
types.
The variations in the characters of all these different forms may be conveniently
expressed by letters as follows : —
Number of cirrus-joints,
. 30 to 35, A.
25,
a
Characters of
cirrus-joints,
Distinctly spiny, B.
Sharply carinate,
b
First radials,
Distinct, C.
Mostly concealed,
c
Palmar series,
Present, D.
Absent,
d.
Arm-bases,
Much rounded, E.
Less rounded,
e
Arms, .
Serrate, F.
Fairly smooth,
f.
1 " Alert " Report, pi. xi. fig. b.
260
THE VOYAGE OF H.M.S. CHALLENGER.
We then get the following expressions to denote the eight forms of this specific type,
five of which have been regarded as representing different species : —
Name.
Characters.
Locality.
1. Antedon variipinna,
ABCDeF.
Canton.
2.
, crenulata, .
ABCDef.
Borneo.
3.
, decipiens, type ("Alert"), .
aBCdef.
Arafura Sea.
4- ,
, decipiens, var. (" Alert "), .
Abcdef.
Prince of Wales Channel.
5. ,
, decipiens, var. (Challenger),
ABcdeF.
Prince of Wales Channel.
6.
, irregularis (Challenger),
AbcdEF.
Prince of Wales Channel.
7-
, irregularis ("Alert "),
AbeDEF.
Torres Strait and Prince of Wales Channel
8.
, dubia (Challenger),
AbCdEF.
Arrou Islands.
With these facts before us it is difficult to avoid the conclusion that we are dealing
with but one specific type ; and this conclusion is confirmed by the fact that in all these
different forms the general shape of the arm -joints and the characters of the pinnules
are respectively identical, though the latter vary considerably in the degree of their
development. The distal arm -joints have the same shape throughout the whole series,
as shown in the Challenger examples from the Arrou Islands and from Torres Strait
(PL XXXVI. fig. 3 ; PL XLVIII. fig. 5). On the other hand, the alternating lateral
projections of the joints in the lower parts of the arms is very marked in the form from
Torres Strait, which Bell called Antedon irregularis (PL XLIX. fig. 1), and it is fairly
distinct in those from the Arafura Sea and from the adjacent Arrou Islands (PL XXXVI.
fig. 1). But it is comparatively insignificant in the other form from Torres Strait
(PL XLVIII. fig. 5), which has much less convex radial and distichal series than the
irregularis-fonn from the same locality (PL XLIX. fig. 1).
Another universal character of all the different varieties which I have referred to this
species is the large size of the pinnules on the fourth and the two or three following
brachials, and the lateral projections at the distal ends of their component joints
(PL XXXVI. figs. 1, 4, 5, 6; PL XLVIII. fig. 3; PL XLIX. fig. 2). The distichal
pinnule, when present, is comparatively small ; but its successor on the second brachial
is somewhat larger, though that on the next joint is smaller again. Beyond this point,
however, there is much variation. The pinnules of the next three or four brachials are
considerably longer and stouter than that of the second, being the largest pinnules on
the arm (PL XXXVI. figs. 4-6 ; PL XLVIII. fig. 3). In those arms which spring
directly from the radial axillary, so that there is no distichal pinnule, the largest pinnules
are generally those of the sixth and seventh brachials. When, however, a distichal
axillary is present, the arm borne on its inner face usually has its largest pinnules on the
fifth and sixth brachials ; while on the outer arm they are on the fourth and fifth. But
this arrangement is very far from being a constant one. The next two pinnules after
the large pair may also be of considerable size and composed of somewhat elongated joints
EEPOE.T ON THE CRINOIDEA. 261
(PI. XLVIII. fig. 3). But in other cases they show a considerable alteration both in the
size and in the character of their component joints, as seen on the left side of fig. 1 on
PI. XXXVI.
The double row of lateral projections on the joints of these proximal pinnules is
developed in rather a singular manner. Their basal joints are somewhat flattened
against the arm, and the upper edge of their broad dorsal surface is sharpened, and more
or less carinate, while its distal end is marked by a median process of variable prominence,
as is well seen in PL XXXVI. figs. 4-6. As the following joints lose their flattened
appearance, and become more rounded, the carination of the upper edge develops into a
strong blunt process at the distal end of the joint on its inner side ; while the medio-
dorsal prominence passes into a corresponding process on the outer side (PI. XLIX.
fig. 2). There is much variation, however, in the exact nature and mode of development
of these processes.
The frecpiency of the ray-divisions of this species, and therefore the number of arms,
is subject to great fluctuations. A second post-radial axillary only occurs in the single
specimens which I named Antedon variipinna and Antedon crenidata respectively, and
sometimes also in the form which was described by Bell as Antedon irregularis. A
large number of individuals were obtained by the " Alert," and the majority of them
have two or more palmar series, though in others, as in the Challenger specimen (PI.
XLIX. fig. 1), palmars are entirely absent. Bell gives the number of arms as ranging
from eleven to twenty-two, but seems to have overlooked one example in which there
are twenty-five. The occurrence of an individual with only eleven arms makes it cpuite
possible that a ten-armed variety of this protean type may be eventually discovered.
In fact, the two individuals which I formerly called Antedon dubia are not improbably
of this nature. The one has two distichal series, and the other only one. But in each
case they result from regeneration of the arm at the syzygy in the third joint above the
radial axillary. This may perhaps have originally supported a distichal axillary ; or it
may have given rise to one arm only, which was replaced by two after fracture, as is
so often the case, an excellent instance of it having been described by Dr. Carpenter
in Antedon rosacea.1 Under these circumstances I have therefore thought it safer
to assign Antedon variipinna a place among the ten-armed species, to which it can
definitely be referred if ever an individual is found in which distichal series are entirely
absent. No harm will be done if it never turns up, and should it do so, it will run less
risk of being baptised as a new species, having undergone that process too frequently
already.
There is one point relating to the extremely variable characters of this species, which
seems to me to be of special importance. The variations which I have noticed above
are not altogether due to difference of locality. Varieties Nos. 5 and 6 were found
1 Phil. Trans., 1866, p. 725, pi xxxviii. fig. 8, B.
262 THE VOYAGE OF H.M.S. CHALLENGER.
associated by the Challenger in Prince of Wales Channel. No. 7, which is only No. 0
with palmar series, was found associated with it by the " Alert " both in Torres Strait
and in Prince of Wales Channel ; and in the latter locality No. 4 was obtained as well.
This repeated occurrence of two or more varietal forms of Antedon variipinna in one
and the same locality recalls the fact, that of the five varieties of the protean Actinometra
parvicirra 1 which were dredged by Professor Semper among the Philippine Islands,
two occurred at Ubay and two at Bohol ; while examples of Pourtales' two species,
Actinometra pulchella and Actinometra alata, which I have been compelled to unite
under the former name,2 were frequently found by the " Blake " to be living together at
the same locality in the Caribbean Sea. It is evident therefore that the cause of these
remarkable variations in one and the same specific type must be attributed to something
more than a mere change of local conditions.
The single example of Antedon variipinna, var. 5, which was obtained by the
Challenger in Prince of Wales Channel, was serving as host to fourteen individuals of
Myzostoma, which Professor von Graff has referred to the following species — Myzostoma
dentatum, Myzostoma JUiferum, and Myzostoma quadriferum. The name of the host is
given in his Report3 as Antedo n bidentata, P. H. Carpenter, this being the MS. name
which I had applied to the species before I became convinced of its identity with Antedon
variipinna, or had the opportunity of identifying it with Antedon decipiens, Bell.
4. Antedon quinduplicava, n. sp. (PI. IV. figs. 1, a-d ; PL XLVII. figs. 4, 5).
Specifa formxda — A.3.2.y.
Description of an Individual. — Centro-dorsal a thin disk, bearing about eighteen
cirri on its sloping sides. They have thirty tolerably equal joints, the last few rather
compressed, and faintly carinate ; the penultimate with a slight spine.
First radials just visible ; the second rather closely united, forming a median
prominence with the pentagonal axillaries ; and there is a similar but less marked pro-
minence on the first two joints above the axillary. The rays may divide three times ;
three distichals with a syzygy, and two palmars without one. Sixteen arms of one
hundred and fifty or more smooth joints, all but the terminal ones being wider than
long ; the lower ones subtriangular and the later ones quadrate or almost oblong. A
syzygy in the third, and then between the eighth and fourteenth brachials ; others at
intervals of four to ten, usually seven or eight, joints.
The distichal pinnule is about equal to that on the second brachial. That on the
third brachial is smaller again, while those on the fourth and fifth are both longer and
1 Trans. Linn. Soc. Lond. (Zool.), ser. 2, 1879, vol. ii. pp. 52, 53.
2 Bull. Mus. Comp. Zcbl., 1882, vol. ix. No. 4, p. 10.
3 Zool. Chall. Exp., 1884, part xxvii. p. 17; ibid., 1887, part lxi. p. 7.
REPORT ON THE CR1NOIDEA. 2(53
stouter, reaching 11 mm., with about eighteen smooth joints, most of them longer than
wide, and the lower ones carinate. The next pair are generally smaller again. But in
arms borne on the radial axdlary the sixth, and occasionally the seventh, brachials may
have large pinnules like those of the two preceding joints.
Disk naked and much incised ; sacculi abundant.
Colour in spirit, — the skeleton brownish-white and the perisome darker.
Disk 7 mm.; spread 16 cm.
Locality. — Station 212, January 30, 1875; lat. 6° 54' N., long. 122° 18' E.; 10
fathoms ; sand. One specimen and one fragment.
Remarks. — I have had some doubts as fro the propriety of separating this species from
Antedon anceps (PI. XXXV. figs. 1 -3), which occurred at the same station. The general
characters of the cirri, calyx, and the large lower pinnules are the same in both types.
One individual of Antedon anceps has only ten arms ; but another has three, and a third
four distichal series. The outer parts of the arms are rather serrate and the distichal
pinnule is distinctly smaller than that on the second brachial above it ; on the other hand
the two forms which I refer to Antedon quinduplicava each have palmar series, nearly
smooth arms, containing longer syzygial intervals, and a distichal pinnule of about the
same size as that on the second brachial.
Considering the remarkable series of variations in the characters of Antedon
variipinna, I think it quite possible that we are here dealing with another case of the
same kind ; but in the absence of the necessary intermediate links I prefer to keep
Antedon quinduplicava separate from Antedon anceps for the present. The only species
that approaches them at all closely is Antedon savignyi, in which, curiously enough,
palmars may or may not be present. But its more numerous and spiny cirri readily
distinguish it from them both.
One of the two individuals of Antedon quinduplicava which was dredged by the
Challenger was a mere fragment which had lost its cirri, disk, and most of its arms. As
it was practically useless in this condition, I made a preparation of its calyx, with a
somewhat surprising result. Each of the radial areas on the ventral surface of the centro-
dorsal is marked at its proximal end by a large bilobate pit (PI. IV. fig. Id), so that every
two pits are separated by an interradial ridge as seen in fig. la. These pits seem to be
nothing but an unusual development of the radial pits which occur round the lip of the
centro-dorsal in so many Comatulse, as seen in PL IV. fig. 2d, and receive the lower ends
of the axial radial canals ; and so in fact they are. But their capacity is increased by the
presence of corresponding pits on the under surface of the radial pentagon (PI. IV. fig. lc)
into which the axial canals, contained between the inner faces of the radials and the
spouts of the rosette, open directly. A possible explanation of this arrangement has
already been discussed on pp. 8, 9. The only Comatula in which I have found any
264 THE VOYAGE OF H.M.S. CHALLENGER.
large cavity of a similar kind within the calyx is the type hitherto known as Actinometra
robusta, of which I wrote as follows :T — "Just above the dorsal surface of the radial the
axial furrow occupying the median line of its internal face gives off a large horizontal
diverticulum into the substance of its calcareous tissue, which extends outwards for some
distance between the central canal and the dorsal surface of the radial ; and, like the axial
furrow, or canal as it is in the natural condition when the rosette is in situ, encloses a
dorsal extension of the body-cavity or coelom. "
In Actinometra robusta, therefore, the radial axial canal, though it terminates
blindly at the top of the centro-dorsal, communicates with a large cavity in the lowest
part of the radial ; while in Antedon quinduplicava this cavity is outside and below the
first radial, between it and the centro-dorsal.
The presence or absence of this cavity may prove to be a point of some importance
in assisting a future decision, as to whether Antedon quinduplicava is or is not identical
with Antedon anc&ps, a question which must wait till a further supply of material
is obtained.
Note on Antedon Jluctuans.
Since the printing off of pp. 94-96, which contain the description of Antedon
jluctuans, I have had occasion to revise the tridistichate species of Antedon that have
been classified as having articulated radials. Among these is Antedon elegans, which was
described by Bell in 1884 from three specimens obtained by the " Alert " at Port Molle
in Queensland.2
I had made a cursory examination of the greater part of the Comatulse dredged by
the " Alert " some time previously, but had not been able to identify any representative
of the type which appeared in my working list of new Challenger species as Antedon
Jluctuans. Subsequently, however, as pointed out on p. 95, I recognised this type in
an imperfect specimen from Torres Strait, which was in too mutilated a condition for
description with the rest of the " Alert " collection.
Bell noticed expressly3 that among the species of Antedon which he did describe in
the report, " in no case is the radial axillary a syzygy."
I have already pointed out, however, on p. 98, that there is a syzygy between the
second and axillary radials of Antedon microdiscus, which is one of Bell's new species ;
and I now find that the same is the case in all three examples of his Antedon elegans,
which I examined in August last for the purpose of definitely making out its relation
to the other tridistichate species of Antedon with articulated radials, before drawing up
a classification of the group.
1 Trans. Linn. Soc. Lond. (Zool.), ser. 2, 1879, vol. ii. pp. 86, 87.
1 " Alert" Report, p. 162. pi. xiii. figs, b, b<i. 3 Ibid., p. 155.
KEPOET ON THE CEINOIDEA. 265
Bell's first formula for Antedon elegans 1 indicated that it had three distichals and
sometimes three palmars, with syzygies in the axillaries ; and I classified it accordingly.2
His subsequent description3 of the palmars says, however, that "if the arms divide
ao-ain there are generally two joints, when the axillary is not a syzygy ; but there may
be three joints, and then the axillary is a syzygy." His figared specimen has four
palmar series of two joints and one of three joints, and he gives the specific formula as
including both varieties |A.3.pr.~). This is all very well in cases where two palmars
occur on the outer, and three on the inner arms of the ray, as in Actinometra nobilis
(PI. LXV. fig. 1), but if it is done in every case where the arm-divisions are not quite
regular, the formulae would become so complex that we should do better without them.
It is extremely rare for any tridistichate Comatula to have its secondary and subsequent
arm-divisions all exactly uniform ; and sometimes, as in Actinometra parvicirra (PL
LXI. figs. 1, 5), there is the same variation in the distichal series. Hence all that we can
do is to go by the majority of the distichal or palmar series respectively ; and as Bell
recognised this fact by omitting any mention of the two-jointed palmar series in
Antedon microdiscus, I wonder that he thought it necessary to refer to the abnormal
three-jointed series in Antedon elegans. His formula also omits any reference to the
post-palmar series which occur on one of his specimens.
The corrected formula for Antedon elegans thus becomes A.R.3.2.(2).— , which is
exactly the same as that given above for Antedon Jluctuans ; and the two species are
in fact identical. Under these circumstances the type must be known for the future as
Antedon elegans, Bell, although its most important distinctive character was omitted in
his diagnosis. It is noteworthy that of the three examples obtained by the " Alert " at
Port Molle, one is very considerably different from the other two, both in colour and in
the amount of serration of the arms ; while the Challenger's dredgings at Station 190
yielded four examples of the same type, three alike and one different.
The "Alert" found an intermediate form in Torres Strait; Semper's Philippine
collection contains representatives of the type ; and I have lately found a most valuable
series of varying forms of this species among the Comatula? dredged by Dr. Anderson
in the Mergui Archipelago. In these last, as in the examples obtained by the " Alert "
and Challenger, the ambulacra of the disk are very strongly plated, and also the inter-
palmar areas at their sides, though this is less marked in the Philippine variety. I find
the same extensive plating on the disk of another species from Mergui which has a
syzygy between the two outer radials and a formula A.R.2.2.2.y. It thus differs
altogether from Antedon elegans, Antedon multiradiata, and Antedon microdiscus in
having but two articulated distichals, instead of three, with a syzygy in the axillary.
i Proc. Zool. Soc. Lond., 1882, p. 534. 2 Ibid, pp. 746, 747. 3 " Alert" Report, p. 162, pL xiii. fig. B.
(ZOOL. CHALL. EXP. — PART LX. — 1887.) OoO 34
266 THE VOYAGE OF H.M.S. CHALLENGER.
In each of these four species the disk is strongly plated, just as it is in the Basicurva-,
Spinifera-, and Granu /{/era-groups (PI. IX. fig. 2). But in all these three latter types
the pinnule -ambulacra have a well-defined skeleton and the lower parts of the rays are
flattened laterally ; whereas in Antedon elegans and its three allies, which we may
conveniently call the Elegans-gxowp, these characters are absent, the plating of the
perisome being confined to the disk. This seems to be a constant peculiarity of the
multibrachiate species of Antedon which have a syzygy between the two outer radials ;
and I have not seen any species possessing a plated disk, but no ambulacral skeleton on
the pinnules, which has articulated radials.
The Comatula tessellata, Miiller,1 seems, however, to be of this character. It has
ten arms with forty-five cirrus-joints, the later ones spiny, and was described as having
the " Haut der Scheibe mit kleinen Knochenplattchen bedeckt." Midler's type is (or
was) in the museum at Bamberg, but I have never been able to get a sight of it, and it
is the only one of all the described species of Antedon (if indeed it be an Antedon)
which I have not personally examined. It may of course be an Actinometra, though the
great number of its cirrus-joints rather precludes this supposition ; and there is also a
possibility that its two outer radials may be united by syzygy, though Miiller said
nothing to that effect. But its true position must remain uncertain for the present ;
and I would simply draw attention to the fact that the mode of union of the two
outer radials must be carefully examined when an attempt is made to determine the
systematic position of a multibrachiate Antedon with a plated disk ; for the probabibty
is that it belongs to Series I.
Genus 5. Actinometra, Miiller, 1841 ; emend. P. H. Carpenter, 1887.
1758. Asterias, Linnseus (jiars), Systema Naturre, 10th ed., Holmire, 1758, t. ii. p. 663.
1783. Asterias, Retzius (pars), K. Svensk. Vetensk. Akad. Handl., Ar 1783, t. iv. p. 241.
1805. Asterias, Retzius (pars), Dissertatio, sistens Species Cognitas Asteriaruru, Lunda3, 1805, p. 34.
1816. Comatula, Lamarck (pars), Histoire Naturelle des Aniinaux sans Vertebres, Paris, 1816, t. ii. p. 530.
1830. Comatula (Astrocoma), de Blainville (pars), Diet. d. Sci. Nat., 1830, t. lx. p. 229.
1830. Alecto, Cuvier, Regne Animal, Paris, 1830, t. iii. p. 228.
1834. Comatula (Astrocoma), de Blainville (pars), Manuel dActinologie, Paris, 1834, p. 248.
1835. Comatula, Agassiz (jiars), Mem. de la Soc. d. Sci. Nat. de Neucbatel, 1835, t. i. p. 193.
1835. Comaster, Agassiz, Ibid., p. 193.
1840. Comatula, Miiller (pars), Monatsber. d. k. preuss. Akad. d. Wiss. Berlin, 1840, p. 91.
1841. Actinometra, Miiller, Ibid., 1841, p. 180.
1841. Alecto, Miiller (pars), Ibid., 1841, p. 182.
1843. Alerto, Miiller (pars), Abhandl. d. k. Akad. d. Wiss. Berlin, Jahrg. 1841 [1843], p. 203.
1843. Actinometra, Miiller, Ibid., p. 226.
1843. Alecto, Miiller (liars), Archiv f. Naturgesch., 1843, Jahrg. ix. Bd. i. p. 131.
1843. Actinometra, Miiller, Ibid., p. 132.
1843. Asterias, Miiller, Ibid., p. 133.
1 Abhandl. d. h. Alcad. d. IFiss. Berlin, Jahrg. 1847 [1849], p. 251.
REPORT ON THE CRINOIDEA. 267
1846. Aetinometra, Muller, Monatsber. d. k. preuss. Akad. d. Wiss. Berlin, 1846, p. 178.
1846. Comatula, Muller (pars), Ibid., p. 179.
1849. Comatula (Alecto), Muller (pars), Abhandl. d. k. Akad. d. Wiss. Berlin, Jahrg. 1847 [1849], p. 246.
1849. Comatula (Aetinometra), Muller, Ibid., p. 246.
I860. Comatula, Bronn (pars), Klassen und Ordnungen des Tkierreichs, 1860, Bd. ii. p. 233.
1862. Comatula, Dujardiu and Hupe (pars), Hist. Nat. des Zoophytes, Echinoderrues, Paris, 1862, p. 192.
1862. Actiiwmetra, Dujardin and Hupe, Ibid., p. 208.
1862. Comaster, Dujardin and Hupe (pars), Ibid., p. 211.
1864. Aetinometra, Liitken, Vid. Meddel. nat. Foren. Kjtfbenhavn, 1864, p. 218.
1865. Alecto, E. C. and A. Agassiz (pars), Seaside Studies, Boston, 1865, p. 121.
1866. Aetinometra, Bohlsche, Archiv f. Naturgesch., 1866, Jabrg. xxxii. Bd. i. p. 90.
1866. Phanogenia, Loven, Ofversigt k. Vetensk.-Akad. Forbandl., 1866, No. 9, p. 231.
1869. Antedon, Pourtales (pars), Bull. Mus. Comp. Zool, 1869, vol. i. No. 11, p. 355.
1869. Comatula (Aetinometra), J. A. Herklots, Bijdragen tot de Dierkunde, 1869, ix. p. 10.
1875. Comatula, Grube (pars), 53e Jahresber. der Schlesiscb. Gesellsch. f. Vaterl. Cult, 1875, p. 74.
1875. Aetinometra, Grube, Ibid., p. 75.
1877. Aetinometra, P. H. Carpenter, Journ. Linn. Soc. Lond. (Zool.), 1877, vol. xiii. p. 442.
1878. Antedon, Pourtales (pars), Bull. Mus. Comp. ZobL, 1878, vol. v. No. 9, p. 214.
1879. Aetinometra, P. H. Carpenter, Trans. Linn. Soc. Lond., ser. 2, 1879, vol. ii. p. 27.
1879. Aetinometra, P. H. Carpenter, Proc. Roy. Soc, 1879, vol. xxviii. p. 385.
1879. Phanogenia, P. H. Carpenter, Ibid., p. 394.
1879. Antedon, Rathbun (pars), Trans. Connect. Acad., 1879, vol. v. p. 157.
1880. Aetinometra, P. H. Carpenter, Quart. Journ. Geol. Soc, 1880, vol. xxxvL p. 41.
1880. Aetinometra, Claus, Grundziige der Zoologie, 4th ed., 1880, Bd. i. p. 335.
1880. Phanogenia, Claus, Ibid., p. 335.
1880. Actiiwmetra, P. H. Carpenter, Journ. Linn. Soc. Lond. (Zool.), 1880, vol. xv. p. 198.
1881. Aetinometra, P. H. Carpenter, Notes from the Leyden Museum, 1881, voL iii. p. 176.
1882. Aetinometra, Ludwig, Mem. Acad. Sci. Bruxelles, 1881 [1882], t. xliv. p. 6.
1882. Aetinometra, P. H. Carpenter, Journ. Linn. Soc. Lond. (Zool.), 1882, vol. xvi. p. 514.
1882. Aetinometra, P. H. Carpenter, Bull. Mus. Comp. Zool., 1882, vol. ix. No. 4, p. 13.
1882. Aetinometra, Bell, Proc. Zool. Soc. Lond., 1882, p. 533.
1882. Aetinometra, P. H. Carpenter, Ibid., p. 747.
1884. Aetinometra, BeU, Rep. Zool. Coll. H.M.S. "Alert," Lond., 1884, p. 155.
1884. Aetinometra, P. H. Carpenter, Proc. Roy. Soc. Edin., 1884, vol. xii. p. 369.
1885. Aetinometra, P. H. Carpenter, Zool. Chall. Exp., part xxxii. vol. ix. 1884 [1885], p. 137.
1885. Aetinometra, Quenstedt, Handbuch der Petrefactenkunde, Aufl. 3, Tubingen, 1885, p. 913.
1885. Aetinometra, Bell, Proc. Linn. Soc. N. S. Wales, 1884 [1885], vol. ix. p. 498.
1885. Aetinometra, Ludwig, Leuuis, Synopsis der Thierkunde, Dritte Auflage, Hannover, 1885, Bd. ii.
p. 948.
Definition. — Centro-dorsal usually discoidal, and bearing fifteen or twenty marginal
cirri, rarely more; sometimes pentagonal or stellate with no trace of cirri, but occasionally
hemispherical and almost covered by them. Outer faces of the radials relatively wide,
with small muscle-plates, and nearly or cpute parallel to the vertical axis of the
calyx.
Disk with an excentric mouth and a variable number of unequal ambulacra, at least
two of which enclose the anal area in a horseshoe-shaped curve. Some of the arms,
generally only the hinder ones, may be much shorter than the rest, ungrooved, and non-
268 THE VOYAGE OF H.M.S. CHALLENGER.
tentaculiferous. Neither arms nor pinnules have any distinct ambulacra! skeleton, and
sacculi are altogether absent. Some of the lower pinnules have terminal combs.
History. — This generic name was proposed by Midler1 in 1841 for a fine specimen of
Comattda Solaris, Lamarck, which he had examined at Vienna in the previous year. He
had not then seen the type of Lamarck's species, and seems to have assumed that it was
an eudocyclic form like Pentacrinus and the three European Comatulae, i.e., that the disk
bore five ambulacral grooves converging upon a central mouth.
This is not the case, however, in reality, for Muller discovered on a later visit to Paris2
that the disk has the same peculiarity in Lamarck's types of Comatula Solaris, as in the
large Vienna specimen " welche generisch von andern durch die Bildung ihres Scheitels
verschieden zu sein schien. Auf dem Scheitel der mit blumenartigen Kalkblattchen
bedeckt ist, ist keine Spur von den Furchen zu sehen die bei den Comatulen von den
Armen zum Munde fiihren. Auch ist dort nichts vom Munde zu sehen. Die mitte der
Bauchseite nimmt eine Bohre ein. Die Arme haben die ventrale Furche der Comatnlen,
die Furchen der 10 Arme mtinden aber in gleichen Abstanden in eine die Seheibe am
Bande umziehende Cirkelfurche. Diese eigenthiimliche Bildung liesse sich durch eine
unsymmetrische Vergrosserung desjenigenlntertentacularfeldes worin die Afterrorhe steht
liber den ganzen Scheitel, und auf Kosten der anderen Intertentacularfelder erklaren, so
dass der Mund aus der Mitte des Scheitels ganz an die Seite zwischen je 2 Armen
gerath." Owing to the dry state of the Vienna specimen the exact position of the mouth
could not be determined ; and the same difficulty presented itself with the types of
Asterias multiradiata and Asterias pectinata, Betzius, which Muller examined in the
Betzian collection at Lund, and found to present "ganz dieselbe Bildung des Scheitels" as
the Vienna specimen.3
In the absence of better-preserved material Muller hesitated to make a definite generic
separation of these three Comatulae from the ordinary endocyclic species. But in 1844
he visited the Paris Museum and there found several Comatulae in spirit with the same
arrangement of ambulacra on the disk as he had described in Actinometra imperialis and
in the two Betzian species, i.e., a circular furrow extending round the greater part of the
margin of the disk, with the ambulacra of the primary arms opening into it at tolerably
regular intervals, very much as in PI. LVII. fig. 3. The number of ambulacra converging
on the excentric mouth would thus be less than five, and in fact was reduced to three in
two of the three individuals first seen by Muller. This character, and not the position
of the mouth, was regarded by him as the most distinctive peculiarity of Actinometra.
For he found that some species may have a central anal tube and excentric mouth " ohne
dass die Ambulacra ihre symmetrische Vertheilung auf die 5 Armstamme einbiissen."4
1 Monatsber. d. k preuss Akad. d. Wiss. Berlin, 1841, p. 180. 2 Ibid., 1846, p. 178.
3 Archivf. Naturgesch., 1843, Jahrg. ix. Bd. i. pp. 132, 133.
4 Monatsber. d. k preuss. Ahul. d. Wiss. Berlin, 1846, p. 177.
REPORT ON THE CRINOIDEA. 269
The number of ambulacra reaching the peristome was thus the sole character by which
Miiller proposed to separate Actinometra from the majority of the Comatulse then known
to him ; and the discovery that five symmetrically distributed ambulacra might converge
on an excentric mouth led him to regard the grouping of the ambulacra as a character of
less systematic value than he had previously attributed to it, so that the name Actinometra
was reduced from generic to subgeneric rank.
It is curious that Miiller should have attached so much importance to the number of
ambulacra converging on the peristome, and so little to the excentric position of the
mouth and the accompanying enlargement of the anal interradius which he had described
so clearly. For whether the number of primary ambulacra be three, four, or five, as he
figured in Comatula Solaris, Comatula waHbercji, and Comatula multiradiata respectively,
the mouth is always excentric, and the anal tube in the middle of the horseshoe-shaped
curve formed by the two posterior ambulacra. The Comatula multiradiata which he
figured1 was not the dry Eetzian type bearing this specific name which he had already
referred to Actinometra, but a spirit specimen in the Paris Museum which had been
identified with the Comatula multiradiata of Lamarck. It has an excentric mouth,
but five primary ambulacra which Miiller described as distributed symmetrically to the
different groups of arms,2 and it was therefore referred by him to the subgenus Alecto.
Except as regards " die Bildung des Scheitels," however, his specific description of Alecto
midtiradiata was simply a reproduction of that which he had given of the dry Asterias
multiradiata, Retzius. He had stated expressly that this showed the same horseshoe-
like distribution of the ambulacra as his type species of Actinometra ; and his subsequent
reference of it to Alecto is therefore difficult to understand. The number of Actinometra
species thus became reduced to three, viz. — (l) the type, Comatula {Actinometra) Solaris,
Lam., sp., with which Miiller was inclined to unite Asterias pectinata, Retzius; (2)
Comatula {Actinometra) ivahlbergi, Midi.; and (3) Comatula {Actinometra) rotalaria,
Lam., sp. All three of these had been previously referred by Midler to Alecto, which
name he used in place of Comatula, Lamarck, as being one of older date ; but when,
later on, he referred them to a subgenus Actinometra in which the number of ambulacra!
grooves joining the excentric mouth is less than five, he used Alecto as a subgeneric
name for the species with five grooves, irrespective of the position of the mouth. Fifteen
species were definitely referred to this latter type in Muller's final memoir, and three
to Actinometra, the remaining seventeen being simply mentioned as Comatula, without
any further detail.
More than a dozen years elapsed after the publication of Muller's systematic work
1 Abhandl. d. k. Ahad. d. Wiss. Berlin, 1847 [1849], p. 245.
2 Muller's diagram of the disk of this specimen is somewhat idealised, for it only represents forty arms disposed
in five groups of eight each ; whereas their number is really forty-nine, and the arrangement of the five primary
ambulacra at the peristome is by no means so symmetrical as shown in his diagram.
270 THE VOYAGE OF H.M.S. CHALLENGER.
on the Comatulse before the characters of Actinometra again came under discussion.
Messrs. Dujardin and Hupe followed the general lines of Midler's classification, but
made some important alterations in it. Leach's name Alecto was abandoned altogether
in favour of the later name Comatula, Lamarck ; and Actinometra was restored to the
generic rank which Midler had first proposed for it. But the French authors1 found
some difficulty in defining it properly, remarking that " ce genre ne differe guere des
vraies Comatules que par la position de l'anus au centre et de la bouche au bord du
disque. II en resulte que les gouttieres ambulacraires, au lieu de se rendre a la bouche
en suivant la direction des bras comme chez les Comatules, s'inflechissent et suivent le
contour du disque." Dujardin and Hupe stated, however, that the mouth of Comatula
was only " ordinairement au centre,"2 so that its excentric position could not be regarded
as especially distinctive of Actinometra, though this has since proved to be the case.
The restoration of the latter type to a distinct generic position was nevertheless a
considerable step in advance ; but the mode in which the French authors disposed of
some of Miiller's species was very singular.
The type species of the genus, Actinometra im<perialis of Midler, was subsequently
discovered by him to be identical with Comatula Solaris, Lamarck, or Alecto Solaris
as he called it at first. But in his concluding memoir 3 it appeared as Comatula
(Actinometra) Solaris, and Miiller further expressed the opinion that Asterias pectinata,
Betzius, which he had also found to be an Actinometra, is merely a varietal form of the
same type. Dujardin and Hupe, however, regarded these three forms at Vienna, Baris,
and Lund as respectively representing three different species. They referred the
Lamarckian type at Baris to Comatula, but the other two forms to Actinometra;
although Miiller had expressly pointed out both in the Monatsbericht4 (1846) and in the
Abhandlungen5 (1849) that Lamarck's originals were specifically identical with the type
of his Actinometra.
Dujardin and Hupe gave no reason for their restoration of a specific name which
Miiller had withdrawn in favour of that established at an earlier date by Lamarck ; and
one can only conclude, therefore, that they had overlooked Miiller's final references to
the type, confining themselves to quoting those of 1841 and 1843, which were made
before his visit to Baris.
On the other hand the French authors left Asterias multiradiata, Betzius, in the
genus Actinometra, to which it had been originally assigned by Miiller, though he
subsequently withdrew it. But it would almost seem as if this were due to their not
having consulted Miiller's later writings, to which they made no reference. For no place
was assigned in their classification to the individuals in the collections of Beron and of
*&•
1 Op. cit., p. 208. • Ibid., p. 194.
3 Abhandl. d. k. Akad. d. Wiss. Berlin, 1847 [1849], p. 248. 4 Loc. cit., p. 178.
5 Loc. cit, p. 248.
REPORT ON THE CRINOIDEA. 271
Quoy and Gaimard, which Miiller had identified with Asterias (Actinometra) multi-
radiata owing to their having not three but two palmars with the axillary a syzygy ;
though he called them Alecto on account of the symmetrical distribution of the
ambulacra on the disk of one specimen.1
In fact, the French authors seem as a rule to have only quoted MuUer's complete
Memoir " Ueber die Gattung Comatula, Lam., und ihre Arten " when dealing with
species which were not described by him in either of his two preliminary communications
of 1841 and 1843 respectively. They recognised that the Alecto ivahlbergi of 1843
was in 1849 transferred by Miiller to Actinometra; but they quite ignored the fact that
he also transferred Comatula rotatoria, Lam., to the same generic type, although it was
described as an Actinometra on the very same page (256) of the final memoir as Comatula
(Actinometra) ivahlbergi, and they simply refer to it as Alecto rotalaria, Miiller, 1843.
The French authors then recognised four species of Actinometra; but only one of these
(Actinometra ivahlbergi) was understood by them in the same sense as it had been by
Miiller. One of his species was restored by them to Comatula,; while, on the other
hand, they retained in Actinometra a type which he had erroneously transferred back again
to Alecto. They also regarded the Asterias pectinata as a distinct species, instead of classi-
fying it with Comatula Solaris, Lamarck. This course is likewise adopted in the following
pages, though the two forms are not placed in different genera, as was done by Dujardin
and Hupe, but in one only, viz., Actinometra, just as they appear in Muller's memoir.
Nearly twenty years elapsed after Dujardin and Hupe wrote before the genus
received much further notice. Isolated species were described by Bohlsche and Grube
respectively, but no formal definition of its characters was ever published. Dr. Liitken,
however, had the opportunity of examining a large number of Comatulse which were
collected in the Eastern Archipelago for the Godeffroy Museum ; and he was led to the
conclusion that the essential character of Actinometra, as distinguished from Antedon, is
the excentric position of the mouth,2 and that the number of ambulacra reaching the
peristome is a character of no importance whatever, instead of being one of generic
value, as Miiller had supposed. Liitken further discovered that the proximal pinnules
of all the exocyclic Comatulse are provided with a terminal comb (PI. LIII. figs. 3-6 ;
PI. LVI. figs. 2, 4 ; PI. LXIII. figs. 5,7; PI. LXV. fig. 7 ; PI. LXVII. figs. 2, 4 ;
PI. LXVIII. fig. 3), but that this is absent in the endocyclic species. The constant
association of these two characters enabled him to recognise Actinometra as a good
generic type ; and various species of the genus were distributed from the Godeffroy
Museum bearing Liitken's MS. names. Unfortunately, however, he was prevented by
other engagements from ever publishing his descriptions of these species, or even a
precise diagnosis of the genus.
i Abhandl. d. k. Akad, d. Wits. Berlin, 1847 [1849], p. 261.
2 See his Note in Tram. Linn. Soc. Lond. (Zool.), ser. 2, 1879, voL ii. p. 18.
272 THE VOYAGE OF H.M.S. CHALLENGER.
During the winter of 1875-76 circumstances enabled me to examine the numerous
Comatulse which had been collected by Professor Semper among the Philippine Islands
some years before. Among them were a dozen examples of one common species, together
with several other well-defined types ; and I was soon led to conclude, as Liitken had
previously done, though unknown to me, that the essential difference between Antedon
and Actinometra depends upon the position of the mouth and not upon the number of
ambulacra reaching the peristome. In reply to the inquiries which I addressed to him
upon the subject, Dr. Liitken was good enough to communicate to me his own observa-
tions, and also his discovery of the constant presence of a terminal comb on the lower
pinnules of exocyclic Comatulse. A few months later, by the kindness of Professor
Perrier, I was enabled to verify Liitken's conclusion for myself on the fine collection of
Comatulse in the Paris Museum ; while at the same time I succeeded in making out
various other points of difference between Antedon and Actinometra, and referred seven-
teen of the Comatu Za-species then known to the latter genus.1
A subsequent study of the large collections obtained by the Challenger, the " Alert,"
and the " Blake," and also of the Comatulse in the principal museums of the continent,
has afforded ample verification of the earlier conclusions which had been reached by Dr.
Liitken and myself; and it has likewise enabled me to make out some other distinctive
characters both of Antedon and of Actinometra.
The genus Phanogenia was established by Loven2 in 1866, for a remarkable
Comatula from Singapore which has a stellate centro-dorsal bearing but slight traces
of cirri, a nearly central mouth, and a terminal comb on the lower pinnules. Several
examples of Actinometra of different species, with a centro-dorsal like that of Phanogenia,
were dredged by the Challenger (PL LVII. fig. 1 ; PI. LXIII. fig. 6 ; PL LXV. figs. 1-6 ;
PL LXVII. fig. 1 ) ; and I have seen others in different museums, many of them with the
mouth unusually near the centre of the disk, and the ambulacra almost as uniformly
distributed as in Antedon ; but the interpalmar area containing the anal tube is always
considerably larger than its fellows. This is also the case in Loven's original specimens
of Phanogenia, which I therefore referred to Actinometra in 1882,s for it became no
longer possible to distinguish Phanogenia as a separate genus by the characters of its
centro-dorsal only, as I have pointed out on pp. 13-16.
Remarks. — In by far the greater number of individuals which belong to the genus
Actinometra the mouth is situated at some distance from the centre of the disk, which
is occupied by the anal tube, and it is occasionally very close to the margin (PL LVII.
fig. 3; PL LXII. fig. 4; PL LXIV. fig. 2; PL LXVIII. fig. 1 ; see also Part I., pi. lv.
fig. 2 ; pi. lvi. figs. 7, 8). The peristome is usually fairly open and somewhat elongated
1 Trans. Linn. Soc. Lond. (Zool.), ser. 2, 1879, vol. ii. p. 27.
- Ofversigt h. Vetensk. Ahad. Forhandl., 1866, No. 9, p. 231.
3 Notes from the Leyden Museum, 1882, vol. iii. p. 195.
REPORT ON THE CRINOIDEA. 273
from side to side, as is well seen in PI. LXVIII. fig. 1 ; but it is sometimes very much
restricted and not readily distinguishable from one of the larger ambulacra (PI. LXIV.
fig. 2). In a few exceptional cases the mouth is practically central, just as it is in
Antedon, though in other individuals of the same species it is nearly marginal (PI. LXII.
figs. 2, 4).
In the endocyclic Crinoids the position of the mouth on the ventral side corresponds
very closely with that of the centre of radiation on the dorsal side ; though it is some-
times a little in front of the centre of the disk, as is well seen in Atelecrinus (PI. VI.
figs. 4, 6). But in any case the interambulacral area of the disk in which the anal tube
is situated corresponds precisely with an interradius of the skeleton, and the ambulacrum
opposite to it passes directly on to the joints of the corresponding ray, whether it is
undivided as in Eudiocrinus (PI. VI. fig. 2), or forks as in Atelecrinus (PI. VI. figs. 4, 6)
and Antedon (PL IX. fig. 2 ; PL XL. fig. 2 ; PL XLVII. fig. 2). In the latter genus the
displacement of the mouth, if it is not quite central, is always in the direction of this
anterior ambulacrum, and its position may therefore be described as radial.
This is also true of a great many forms of Actinometra (Fig. 6, a). Thus for example
in the disk of an abnormal individual of Actinometra jimbriata, represented on PL LXII.
fig. 2, a median vertical plane would be radial in front of the nearly central mouth, and
interradial behind, where it would cut the anal tube. The same is the case in the more
typical specimen which is shown in fig. 4 of the same plate. In each alike there is a
short but wide anterior ambulacrum, which forks twice and so sends a branch to each of
the four arms on the anterior ray. Four more grooves leave the peristome in each
individual. But whereas in the one (fig. 2) each primary groove supplies all the arms
of one ray, just as in Antedon (PL IX. fig. 2), this is not the case in the other (fig. 4).
For each of the two antero -lateral grooves supplies but two arms of the corresponding-
ray ; and the two remaining arms receive their ambulacra as offshoots of the two primary
grooves, which supply the two postero-lateral rays, and together form a sort of horse-
shoe enclosing the anal interradius. This is much larger than in the endocyclic forms
(PL LXII. fig. 4), as the two hinder ambulacra curve outwards from one another towards
the margin of the disk, and so greatly reduce the size of the remaining interambulacral
areas. The anal tube is at the centre of the disk, and a vertical plane, cutting mouth
and anus, would pass along the short ambulacrum in front of the mouth, which is there-
fore radial in position, just as in Eudiocrinus and Antedon (PL VI. fig. 1 ; PL XL.
fig. 2). This condition may be traced, though less clearly, in the disk of Actinometra
elongata (PL LVII. fig. 3).
On the other hand, there are a great many forms of Actinometra, as shown in the
diagram (Fig. 6, b), which have a distinctly interradial mouth. Two grooves start from the
sides of the peristome, instead of one from its anterior border (Fig. 6, a), and supply the
oral arms of the two corresponding rays (a2, Bj) ; while the ambulacra of their aboral arms
(ZOOL. CHALL. EXP. — PART LX. 1887.) OoO 35
274
THE VOYAGE OF H.M.S. CHALLENGER.
may start directly from the peristome (b2 on Fig. 6, b), or come off from one of the large
furrows which supply the posterior and the two lateral rays (Aj on Fig. 6, b). The furrow
on the right or western side of the disk supplies the radi E and D, as in the species with
a radial mouth (Fig. 6, a) ; but it is longer and has a larger curve than in these forms, as
the d ray is exactly behind the mouth (Fig. 6, b), and the corresponding posterior furrow
thus extends round three-fifths of the disk instead of round but one half of it. The
shape of the peristome in a species with interradial mouth is well seen in Actinometra
regalis (PI. LXVIII. fig. 1); while the origin of the primary ambulacra is also seen in
Actinometra belli, though the peristome of this specimen is so much contracted that the
position of the mouth on the disk is only indicated by the point of convergence of the
ambulacra (PI. LXIV. fig. 2).
I was led to think at one time that the situation of the mouth, whether radial or
interradial, might serve as a character of specific value. But wider experience has
Fig. 6. — Diagrams showing the different positions of the mouth in Actinometra ; A, with a radial mouth ; B, with an inter-
radial mouth. The dotted lines mark the interambulacral regions of the disk. Alt Aj, Ej, E2, the five pairs
of secondary ambulacra.
shown me that too much reliance must not be placed upon it. For although the mouth
generally has a constant position in any given species, it sometimes happens that some
individuals of the species have an interradial mouth and others a radial one. Thus, for
example, the mouth is radial in nearly all the specimens of Actinometra pectinata which
I have seen, as it almost invariably is in the Solaris-gvou]) ; but it is interradial in the
three individuals dredged by the Challenger, one in Torres Strait, and two at Samboangan.
On the other hand the mouth is interradial in the examples of Actinometra lineata which
were dredged at Bahia, but in another individual which I have seen from Barbados its
position is radial.
There is another character which very commonly presents itself in Actinometra, and
seems to be correlated with the excentric position of the mouth. I refer to the very
KEPOET ON THE CRINOIDEA. 275
frequent absence of the ambulacral groove and its associated tentacular apparatus on
more or fewer of the arms. This is well shown both in Actinometra belli and in
Actinometra regalis (PL LXIV. fig. 2; PL LXVIII. fig. 1), and also in Actinometra
magnified (Part I., pi. lvi. fig. 7). The number of arms is very great in the last-
named species, and there are some without grooves on every ray, a condition which
also occurs in Actinometra nobilis. But as a general rule the ungrooved arms are
those which come off from the posterior part of the disk. Thus, for example, some or
all of the four posterior arms are very frequently ungrooved in the ten-armed types,
Actinometra Solaris and Actinometra pectinata ; while in other individuals of the
same species all the arms are provided with grooves, just as in Antedon. The same is
the case in the multibrachiate forms. I have seen one individual of Actinometra
parvicirra in which nineteen out of thirty-one arms were entirely devoid of an
ambulacral groove and tentacular apparatus, while in other specimens there is a groove
on every arm.
It is then the potential, rather than the constant presence of ungrooved arms which
must be regarded as one of the distinguishing characters of Actinometra ; and the same
may be said of another peculiarity which is frequently associated with it, viz., the
difference in length of the anterior and posterior arms. This is less apparent in the
ten-armed than in the multibrachiate species, in which, however, it is sometimes very
distinct, e.g., Actinometra belli, Actinometra nobilis, and Actinometra regalis. The
anterior arms are much longer, taper more slowly, and contain far more joints than the
posterior arms, though these often have their genital glands better developed than the
anterior arms. In Actinometra simplex the tentaculiferous anterior arms have one
hundred joints, while there are only forty-five in the hinder arms, which have no
ambulacral groove nor tentacles. The two characters are not always associated, however,
for in the single specimen of Actinometra elongata all the arms are grooved and tenta-
culiferous, but the posterior ones have only fifty-five joints and reach but 4-5 cm. long,
while the anterior arms with one hundred and twenty joints reach 11 cm.
This species is also remarkable for the presence in the later pinnules of the posterior
arms of those curious brown cellular bodies that I have supposed to be sense-organs
(PL LVII. fig. 4). I found them first in some specimens of Actinometra parvicirra
from the Philippines,1 and have since detected them in an example of this species from
Banda, in Actinometra elongata from the same locality, and in Actinometra simplex
from the Admiralty Islands ; while they also occur in examples of Actinometra meri-
dionalis from two localities on the American coast. They are not always present in
either species and are generally confined to the pinnules of the hinder arms, sometimes
to one or two arms only ; but in one case I found them on all the arms except the
two immediately adjoining the mouth. I know not what these brown " ovoid bodies "
1 Trans. Linn. Hoc. Lond. (Zool.), ser. 2, 1879, vol. ii. p. 40, pi. ii. fig. 6, o, b.
276 THE VOYAGE OF H.M.S. CHALLENGER.
may be ; but as they occur in four species of Actinometra, one American and
three from the Eastern Archipelago, and are unknown in Antedon, they provide us
with another potential character of the former genus which has a certain systematic
value.
The presence of a terminal comb on the lower pinnules is, however, an absolutely
constant character of Actinometra. It varies much in its development (PI. LIII.
figs. 3-6 ; PI. LVI. figs. 2, 4 ; PL LXI. figs. 8-10 ; PL LXIII. figs. 5, 7 ; PL LXVI.
figs. 3, 5 ; PL LXVII. figs. 2, 4 ; PL LXVIII. fig. 3), but it is always present ; and this
peculiarity, together with the invariable absence of sacculi on the ventral perisome,
enables single arms of Actinometra to be recognised with the utmost certainty.
The arms and pinnules of this genus are never provided with the ambulacra! skeleton
which is so well developed in many species of Antedon; and the character which is so
often associated with this, viz., the lateral flattening of the lower parts of the rays, is also
entirely absent in Actinometra. This indeed is only to be expected, for the three groups
of A ntedo ?i-species which present these combined characters are almost entirely hmited
to the abyssal and continental regions, while Actinometra is essentially a shallow-water
genus, having only been obtained nine times at depths exceeding 200 fathoms.
In certain localities, however, e.g., Cape York and Port Curtis in Queensland, species
of Actinometra occur with the disk very completely plated, although it may be entirely
membranous in the same species elsewhere. This is especially noteworthy in the
cases of Actinometra Solaris, Actinometra pectinata, and Actinometra paucicirra ; but
however well plated the disk may be, there is no ambulacral skeleton on the arms and
pinnules, any more than there is in those species of Antedon like Antedon elegans and
Antedon multiradiata, which have the two outer radials united by syzygy and a thickly
plated disk (PL IX. fig. 2 ; Part I., pi. lv. figs. 3, 4). The essential characters of the
radials of Actinometra have been fully explained on pp. 24-26, and need not therefore
be further discussed.
The centro-dorsal is very often only a thin flattened disk, with an imperfect double
row of cirrus-sockets round its margin (PL IV. fig. 4a; PL V. figs, lb, id, 2b, 2d, 2e, 2d;
PL LII. figs. 1, 2 ; PL LIII. fig. 1, 2, 15 ; PL LXII. figs. 1, 2 ; PL LXIV. figs. 1, 3)
There are not often more than about twenty functional cirri on the centro-dorsal at the
same time ; but this number is sometimes exceeded (PL LX. figs. 1-3 ; PL LXVI. fig. 4).
On the other hand, the centro-dorsal is occasionally reduced to the condition of a mere flat
plate without any trace of cirrus-sockets (PL LIV. figs. 1-8), and it is often separated
from the radial pentagon by more or less definite slits (PL LVII. fig. 1 ; PL LXI. fig. 1 ;
PL LXIII. fig. 6; PL LXV. figs. 1, 5, 6; PL LXVII. fig. 1). It has been pointed out above
that the new genus Phanogenia was established by Lov^n for a species of Actinometra
possessing these characters ; and the nature of the change which produces them has
already been noticed on pp. 13-16. It need not therefore be further considered here,
REPORT ON THE CRLNOIDEA. 277
but as it never occurs in Antedon, it affords another good potential character of the genus
Aetinometra.
The distribution of the genus, both in space and in time, has been discussed already
on pp. 35-40.
Classification. — The species of Aetinometra, like those of Antedon, fall into certain
very well defined series, as shown above on pp. 57-59. But the various series do not
altogether correspond in the two genera. Each has a ten-armed, a bidistichate, and a
tridistichate series. But the first two of these are epiite small in Aetinometra, although
they together contain three-fourths of the described species of Antedon. On the other
hand, the tridistichate series of the former genus is remarkable for the number and
variety of the forms which it includes, though it is poorly developed in Antedon. There
are, however, three tridistichate species of Antedon with a considerable number of arms ;
but in all three alike the two outer radials are united by syzygy, and each arm-division
above the distichal axillary consists of two or three joints, in the latter case with a
syzygy in the axillary (Pis. VIII., IX.; PI. XXXVII. fig. 4). This combination of
characters does not present itself in any Aetinometra yet known. In all the four
tridistichate species of this genus which have the two outer radials united by syzygy,
each arm-division above the distichal axillary consists of two joints, which are themselves
united by syzygy, as the radials are (PI. LV. fig. 1 ; PI. LVI. fig. 3 ; PI. LVII. fig. 1).
In another group the two outer radials and the first two distichals, palmare, and
brachials are respectively united by syzygy (PI. LIV. figs. 1,2); while in a third the
radials and brachials have the same characters, but distichals are undeveloped, so that
there are only ten arms (PI. LIII. figs. 2, 15). Neither of these three types is represented
in the genus Antedon at all, and they are all strictly limited to the Eastern Archipelago,
ranging from Mergui on the west to Fiji on the east, but scarcely passing the limits of
the tropics. Neither of them is at all rich in species, though the range of variation
within the limits of a single specific type is in some cases very considerable. They may
be classified as follows : —
Series I.
The two outer radials, and the first two brachials respectively united by syzygy.
Ten arms, . . . . . • • ■ • • • 1- Solaris.
Two distichals united by syzygy, . . . • • • • -2. Paucicirra.
Three distichals, the axillary a syzygy, . . • • • ■ ■ 3- Typica.
278 THE VOYAGE OF H.M.S. CHALLENGER.
1. The Solaris-growp.
Ten arms. The two outer radials and the first two brachials respectively united by
sj^ygy.
A. Less than twenty-five cirrus-joints. The basal joints of the lower pinnules
usually have more or less prominent keels.
Nine to fifteen cirrus-joints, . . . . . . .1. pectinata, Retz., sp.
Seventeen to twenty-three cirrus-joints, . . . . .2. Solaris, Lam., sp.
B. Thirty to thirty-five cirrus-joints. The basal joints of the lower pinnules not
specially distinguished, ....... brachiolata, Lam., sp.
Seven other species besides the three contained in the above list have been referred
to this group at different times, viz., Actinometra hamata, Herklots, Actinometra
imperialis, Midler, Actinometra intermedia, Bell, Actinometra purpurea, Miiller, sp.,
Actinometra robusta, Liitken, MS., Actinometra rosea, Miiller, sp., and Actinometra
strota, P. H. Carpenter, MS. The first four are synonyms of either Actinometra
pectinata or Actinometra Solaris, while the only character on which Miiller relied
as separating Comatula rosea from Comatula brachiolata was the absence in the
latter of a terminal comb on the oral pinnules ; and I have since found this comb to
be present in the type specimens, both at Berlin and at Vienna. Alecto purpurea
is a small form which was described by Miiller in 1843, before he had himself examined
Lamarck's types at Paris, but it afterwards appeared to him to differ from Actinometra
Solaris only in showing two radials externally instead of three ; and he thought that the
difference might possibly be due to the immaturity of the specimen, which he regarded
as probably a young form of Actinometra Solaris. Both the Paris and Vienna
specimens of this type have twenty cirrus-joints ; and Miiller described Alecto
purpurea as having only twelve, so that it should probably be referred to Actinometra
pectinata.
Comatula (Actinometra ?) hamata was the name given by Herklots ' to a specimen
in the Leyden Museum from Cape Bantano, which was figured by Kuhl and van Hasselt,
but is not, however, sufficiently distinct from the general type of Actinometra Solaris to
justify the establishment of another species. I believe uow that the same may be said
of Actinometra imperialis, Miiller, of Liitken's MS. species Actinometra robusta, of the
form which I have hitherto called Actinometra strota, and also of Actinometra inter-
media, Bell. This may seem to be a somewhat comprehensive statement ; but it is the
result of a careful and often-repeated examination of a very large amount of material,
including the type specimens at Lund, Paris, Berlin, Vienna, Leyden, Copenhagen, and
in the British Museum, besides numerous isolated specimens in other collections. My
1 Echinodermes peintes d'apres nature par les soins de Kuhl, van Hasselt, et Sal. Miiller, Bijdragen tot de
Dierkunde, 1869, Bd. ix. p. 10, pL is.
REPORT ON THE CRINOIDEA. 279
conclusions, however, are chiefly based on the very large series of specimens belonging to
the Solaris-group which were collected by H.M.S. "Alert" at various localities in Torres
Strait and on the Queensland coast. They are now in the British Museum, and have
been placed unreservedly at my disposal by Professor F. J. Bell, to whom my best
thanks are due for the ready way in which he has always done bis utmost to facilitate
my work.
Two years after Midler first described Actinometra imperialis from his own examina-
tion of the Vienna specimen with an excentric mouth, he gave a description of Comatula,
or as he called it, Alecto Solaris, which had been drawn up by Dr. Troschel as the result
of his own examination on Midler's behalf of the Lamarckian types of Comatula Solaris
at Paris. Midler knew nothing about the structure of the disk in these last, nor even
that the two outer radials are united by syzygy ; and it was not till he subsequently
visited Paris himself that he found Lamarck's specimens to be both specifically and
generically identical with that which he had previously called Actinometra impenalis.
He therefore withdrew the specific name imperialis, replacing it by Solaris, Lamarck,
which he redescribed.1 All subsequent writers have accepted this identification, with
the exception of Dujardin and Hupe, who described the two forms, not only as distinct
species, but also as distinct genera.2
The " Neue Beitrage " in which Troschel's diagnosis of Alecto Solaris was published,
also contained a careful description, by Midler himself, of a specimen in the Retzian
collection at Lund which had been referred by Retzius to the Asterias pectinata of
Linnaeus, an identification which was subsequently adopted in Gmelin's edition of the
Systema Naturae. Milder 3 described it under the name of Asterias pectinata, Retzius,
recognising, however, its generic identity with his own Actinometra imperialis, though
he still regarded the latter as distinct from Comatula Solaris, Lamarck. But after
examining Lamarck's type, he united the two species and also came to the conclusion that
Asterias pectinata, Retzius, " scheint eine Farbenvarietat dieser Art zu sein .... Die
Farbenzeichnung ist aber sehr eigenthiimlich. Auf der Riickseite der Arme sehr
regelmassig zwei schwarze Langslinien. die in der Mitte durch eine helle Linie getrennt
sind."4
I certainly cannot attach any specific importance to this peculiar marking, which I
have seen in several individuals from very different localities, amongst others in that
from Hong Kong which served as the basis of my own redescription of Actinometra
Solaris in 1882.6 In this form, however, there are over twenty cirrus-joints. Midler
gave the number as twenty in the Vienna specimen,0 while Troschel's description
&
1 Abhandl. d. k. Akad. d. Wiss. Berlin, 1847 [1849], p. 248. 2 Op. cit., pp. 200, 209.
3 Archivf. Naturgesch., 1843, Jahrg. is. Bd. i. p. 133.
4 Abhandl. d. k. Akad. d. Wiss. Berlin, 1847 [1849], p. 249.
6 Journ. Linn. Soc. Lond. (Zool.), 1882, vol. xvi. p. 514.
' Monatsber. d. k. preuss. Akad. d. Wiss. Berlin, 1841, p. 181.
280 THE VOYAGE OF H.M.S. CHALLENGER.
mentioned thirty (which is clearly a misprint for twenty) in Lamarck's type at Paris.1
But the number of cirrus-joints in the original of Asterias pectinata, Retzius, was stated
by Miiller as only thirteen, and that of the cirri as sixteen.2 I have lately examined a
large number of specimens from eighteen different localities, all of which agree in having
no more than sixteen cirri with a small number of joints, and in a generally less robust
appearance than that of the type specimens of Actinometra Solaris. In the majority of
cases the number of cirrus-joints is eleven or twelve (PL LIII. fig. 15), but it may fall as
low as nine, though it occasionally rises to fifteen. This reduction in number is not due
to immaturity, as in the case of the small specimen represented in PI. LIII. fig. 1, the
very youthful condition of which is indicated by the shape of its arm-joints, as compared
with those of the larger individual shown on the same plate (fig. 2). But it is a character
of much constancy, accompanied by others which will be noticed immediately ; and as
such it will serve, I think, for the separation of the two species Actinometra pectinata and
Actinometra Solaris.
To those who know the extent of the variation in the number of cirrus-joints within
the limits of individual species of Antedon, the difference between an average of twelve
and another of eighteen or twenty may seem altogether insufficient to serve as a basis of
specific distinction. But it must be remembered that as the number of cirrus-joints in
Actinometra rarely exceeds twenty-five, it has but a very slight range of possible varia-
tions. The number does vary in each of these two types — from nine to fifteen in the
one and from seventeen to twenty-three in the other, — while there are other characters
which also help to separate them. There may be as many as twenty-five cirri in
Actinometra Solaris, which is a much more robust species than Actinometra pectinata,
i.e., it has stouter joints, both in the cirri and in the arms and pinnules. Thus, for
example, taking an immature individual of Actinometra Solaris with about the same
" spread " as an Actinometra pectinata from the same locality, I found the corre-
sponding pinnules with nearly the same number of joints to be one-third longer in the
former than in the latter, while the arm-joints were also wider.
The two species present a parallel series of colour-variations. The)7 are sometimes a
deep purple, and sometimes brown in various shades, greyish, reddish, and blackish, or
occasionally more or less mottled purple and brown. Miiller described the medio-dorsal
line of the arms in Actinometra pectinata as marked by a white band with a dark one on
either side. I have seen this also in examples of Actinometra Solaris from Hong Kong
and Billiton. In the dry state the arms of Actinometra pectinata have a slightly raised
ridge in the medio-dorsal line which is hidden by the white band just mentioned, when
this is present. But the ridge always exists and is very frequently visible in spirit
specimens, though sometimes, as in that dredged by the Challenger, it is scarcely visible
till the arm is dried. It is never so marked in Actinometra Solaris, however, as it may be
1 Archivf. Naturgesck, 1843, Jahrg. ix. Bd. i. p. 135. 2 Ibid., p. 133.
REPORT ON THE CRINOTDEA. 281
in Actinornetra pectinata, and is sometimes almost indistinguishable, even on the dry
arm.
In Midler's own description of Actinornetra Solaris he noticed that the lower joints of
the second pinnule "zeichnen sich durch ihre Erweiterung aus."1 He had made nearly the
same statement in his previous description of Asterias pectinata;- and when re-describing
the former species, from a specimen in the Hamburg Museum,3 I pointed out that in
the pinnules of the fourth to seventh brachials the second and third joints are wide, with
strong and expanded dorsal keels, mentioning at the same time the variations of this
character which I had found in the original types of the species at Paris and Vienna.
Neither of these forms had any indication of a keel on the lower joints of the first pair of
pinnules; and this character, together with the larger number of cirrus-joints (over twenty),
then appeared to me to constitute the special marks of Actinornetra Solaris as distinguished
from Actinornetra pectinata, with its thirteen cirrus-joints and traces of keels on the
basal joints of both the first pinnules, in addition to those on the second pair.
Since examining all the "Alert " collection I find that this view will still hold good,
except for one point, the occasional presence of a distinct keel on the pinnule of the third
brachial in Actinornetra Solaris. I have only found it in three individuals from Billiton
and in one from Port Molle. As a rule, however, there is no more indication of it than
is shown in PI. LIII. fig. 10, and the base of this pinnule, like that of its predecessor on
the second brachial, is not specially marked (PL LIII. figs. 3, 4). On the fourth and
fifth brachials, however, the case is different. The second and third, with sometimes
the fourth and even the fifth joints of their pinnules, have large and prominent keels
(PL LIII. figs. 11, 12), traces of which may generally be found on the pinnule of the
sixth and sometimes on that of the seventh brachial. In Actinornetra pectinata, on the
other hand, there are never more than two joints, and sometimes only one, which has a
definite keel, and this keel may appear on the pinnule of the second brachial, as in some
individuals from Bohol. It is usually present on those of the third to fifth brachials, and
sometimes on that of the sixth as well (PL LIII. figs. 17-20). But as a rule it is absent
in the latter case, and I have never seen any individual with the other characters of
Actinornetra pectinata, which has any sign of a keel on the pinnule of the seventh
brachial. Broadly speaking, then, we may say that there are not more than two, and
sometimes only one carinate joint on the lower pinnules of Actinornetra pectinata ; that
keels are generally present on the pinnules of the third, fourth, and fifth brachials, and
sometimes on those of the second and sixth, but never on that of the seventh brachial.
On the other hand, Actinornetra Solaris generally has two carinate joints, and occasion-
ally sometimes three or even four, on the pinnules of the fourth, fifth, and sixth brachials,
sometimes on those of the third or seventh, but never on that of the second.
i AbJmndl. d. k. Ahad. d. Wiss. Berlin, 1847 [1849], p. 248. 2 Archivf. Naturgesch., 1843, Jahrg. ix. Bd. i. p. 134.
3 Journ. Linn. Soc. Lond. (Zool.), 1882, vol. xvi. p. 515.
(ZOOL. CHALL. EXP. — PART LX. — 1887.) 0°0 36
282 THE VOYAGE OF H.M.S. CHALLENGER.
When I re-described Actinometra Solaris in 1882, I added a diagnosis1 of the form
which had been long known in the catalogues of the Godeffroy Museum as Actinometra
robusta, Liitken, MS. The chief character distinguishing it from Actinometra Solaris,
apart from its generally more robust nature, seemed to me to be the entire absence of
any expanded keels on the lower joints of its second and third pairs of pinnules. The
examination of the " Alert " collection has shown, however, that this distinction will no
longer hold good. The " Alert " dredged large specimens at Prince of Wales Channel,
Port Molle, and Port Curtis, which are indistinguishable from Actinometra robusta in
almost every other character but those of the lower pinnules. All of them have three,
and that from Port Molle as many as five pinnules with keeled basal joints ; and for
reasons which will appear immediately, we have, I think, no other course open to us but to
refer them all, together with Actinometra robusta, to one and the same type, Actinometra
Solaris. When describing the Comatulse obtained by the "Alert," Bell proposed, in the
following terms,2 to establish a new species, Actinometra intermedia: — "As Mr.
Carpenter has pointed out, it appears to be possible, in part at any rate, to distinguish
A. Solaris from A. robusta by the character of the keels, which, in the former, are so
strikingly developed on the basal joints of the second pinnule. Basing myself on the
theory that the keel is constantly present on the basal joints of the second pinnule of
A. Solaris, and that it is never found on those of A. robusta, I venture to think that, in
the case of A. intermedia, we have to do with a form in which constantly the keels are
never as well developed as in A. Solaris, and never so slightly as in A. robusta, while at
the same time there are considerable differences in the extent of the development of the
keel, not only within the limits of the species, but even of the individual."
I have made a careful examination of the half dozen specimens which Bell referred
to Actinometra intermedia, and I find it impossible to differentiate them from Actino-
metra Solaris. They present a great amount of variation in the carination of the basal
pinnules, but not more so than I have found in a number of specimens collected by the
Challenger in Torres Strait, which I now refer to Actinometra Solaris, though, like
Bell, I formerly considered them as representing a new species (which I called Actino-
metra strota), intermediate between Actinometra Solaris and Actinometra robusta.
The Challenger specimens from Booby Island and Albany Island, and Bell's Actinometra
intermedia from the latter locality, agree in every respect except colour. The lower
pinnules are sometimes almost as slightly keeled as in the ?'obusta-form (PI. LIU.
figs. 3-6); while, on the other hand, they may have all the characters of the pinnules in
the typical Actinometra Solaris (PI. LIU. figs. 9-12), and the development of the keel
is not constant in any individual specimen. They all agree, however, in having from
eighteen to twenty cirrus-joints, and in the indistinct nature of the medio-dorsal ridge ;
1 Journ. Linn. Soc. Lond. (Zool.), 1882, vol. xvi. p. 517.
! "Alert" Report, p. 166.
REPORT ON THE CRINOIDEA. 283
while when the lower pinnules have keels, they are developed like those of Actinometra
Solaris, i.e., on two or three joints (PI. LIII. figs. 11, 12), and not on one only as is so
usual in Actinometra pectinata (PL LIII. figs. 17-20).
We have already seen that there are similar variations, though of a somewhat more
extensive character, among the largest specimens of all which would naturally be
referred to Actinometra robusta ; and I do not think therefore that any other course is
possible than to consider Actinometra intermedia and Actinometra robusta as identical
with Actinometra Solaris. The variation in the extent of development of the keel —
from almost nothing but a mere sharpened dorsal edge to large projections on two or
three joints of from two to five pinnules — then becomes itself a character of specific
value, just as Bell pointed out for Actinometra intermedia. For so far as my experience
goes the basal pinnules of Actinometra pectinata are much more uniformly carinate than
in Actinometra Solaris. With the possible exception of the immature specimen which is
the type of Alecto purpurea, Muller, I have never seen any individual which would be
referred to Actinometra pectinata on account of its cirrus-characters, with so slightly
developed keels on the lower pinnules, as are shown in PI. LIII. figs. 5, 6 ; and I know of
none which are absolutely keelless, like the form which I described as Actinometra
robusta, but now refer to Actinometra Solaris.
The habit of Actinometra brachiolata is only known vaguely as "Australia"; but
the geographical range of Actinometra Solaris and Actinometra pectinata is very much
better defined. They are limited to craite shallow water, 12 fathoms or less, in the
Eastern Archipelago, scarcely extending, however, beyond the limits of the tropics.
Actinometra pectinata ranges as far west as Java and Singapore, and has also been
found among the Philippines and Moluccas. It likewise occurs along the north-west
coast of Australia, in the Arafura Sea, and on the Queensland coast from Cape York to
Port Curtis. I have seen examples of Actinometra Solaris from the China Sea and from
Hong Kong, Singapore, and Billiton ; but I know of no other localities for it between
Java and Torres Strait. Like Actinometra pectinata, it is abundant at Cape York and
down the Queensland coast to Port Curtis, in lat. 24° S. But with this exception, I
have never heard of either of these two species occurring to the south of the Tropic of
Capricorn, abundant as they are on the northern shores of the continent. They re-
present perhaps the most characteristic type of the Crinoid fauna of the Eastern
Archipelago, not extending eastwards to Fiji, nor even to Mergui on the west ; though
these localities have representatives respectively of the other two groups of Actinometra-
species which have the radials united by syzygy. A form like Actinometra paucicirra
occurs at Mergui, while Actinometra typica extends from Malacca to Fiji.
But so far as my present knowledge goes not one of these three types of structure
is represented among the Comatulas of Southern Australia. Actinometra paucicirra
abounds at Cape York, was found by Jukes on the reef of Atagor, and is recorded from
284 THE VOYAGE OF H.M.S. CHALLENGED.
Port Molle ; but it is not known from any locality further south. Port Molle is likewise
the southern limit of two other widely distributed species, (l) Antedon milberti, which
extends as far west as Mergui. (2) Actinometra parvicirra, which is still more widely
distributed in the Eastern Archipelago, and also occurs on the coast both of South
Africa and of Peru. Then again two members of the large Pa/mcrfa-group occur at
Port Molle, but the group is not represented further south.
Thus then, not one of these common tropical species enters into the fauna of
Southern Australia ; and, on the other hand, the two Coruatulse which are especially
characteristic of this region do not extend into the tropics. Antedon macronema occurs
at King George's Sound, Port Jackson, and Port Stephens, while Actinometra
trichoptera has been found at King George's Sound, Port Philip, and Port Jackson ; but
neither of them reaches Port Curtis.
These facts confirm in a very striking manner the views of Gimther and Bell
respecting the independence of the marine fauna of Southern Australia, as compared
with that of the north-eastern and northern shores of that continent. The following
statement which was made by Bell1 as the result of his studies of the Asterids, Ophiurids,
and Urchins, is equally true of the Crinoids : — " The species found on the northern
and north-eastern shores of Australia have a wide range eastward and westward, but
gradually disappear as we pass southwards. In fine, an Australian Echinoderm-fauna,
as conterminous with the Australian shores, does not exist."
The Crinoid fauna of Western Australia is still almost completely unknown ; but
from what little I have seen of it, I believe it to be essentially identical with that of the
Eastern Archipelago.
1. Actinometra pectinate/,, Retzius, sp. (PI. LIU. figs. 15-22).
Specific formula — a. R.-g ■ — .
1758. Asterias pectinata, Linnaeus (pars), Systema Naturae, 10th ed., Holmiae, 1758, t. ii. p. 663.
1783. Asterias pectinata, Eetzius (pars), K Svensk. Vetensk. Akad. Handl., Ar. 1783, t. iv. p. 241.
1788. Asterias pectinata, Linnaeus (pars), Systema Naturae, ed. 13, cura J. F. Gmelin, Lipsias,
1788, t. i. pars vi. p. 3166.
1805. Asterias pectinata, Eetzius (pars), Dissertatio sistens Species Cognitas Asteriarum, Lundaa,
1805, p. 43.
1843. Alecto purpurea, Miiller, Archiv f. Naturgesch., 1843, Jahrg. ix. Bd. i. p. 132.
1843. Asterias pectinata, Miiller, Ibid., p. 133.
1849. Comatula (Actinometra) Solaris, var. ? Miiller, Abhandl. d. k. Akad. d. Wiss. Berlin,
Jahrg. 1847 [1849], p. 249.
1862. Comatula purpurea, Dujardin and Hupe, Hist Nat. des Zoophytes, Echinodermes, Paris,
1862, p. 202.
1862. Actinometra pectinata, Dujardin and Hup6, Ibid., p. 210.
i "Alert" Report, p. 175.
REPORT ON THE CRINOIDEA. 285
1879. Adinometra pectinata, P. H. Carpenter, Trans. Linn. Soc. Lond. (Zool.), ser. 2, 1879,
vol. ii. p. 27, pi. v. figs. 5-9 ; pi. viii. figs. 5-8.
1879. Comatida purpurea, P. H. Carpenter, Ibid., p. 27.
1879. Adinometra purpurea, P. H. Carpenter, Proc. Roy. Soc, 1879, vol. xxviii. p. 386.
1882. Adinometra affinis, Liitken, MS., Journ. Linn. Soc. Lond. (Zool.), 1882, voL xvi. p. 517.
1882. Adinometra affinis, P. H. Carpenter, Proc. Zool. Soc. Lond., 1882, p. 747.
1882. Adinometra pedinata, P. H. Carpenter, Ibid., p. 747.
1882. Adinometra purpurea, P. H. Carpenter, Ibid., p. 747.
1884. Adinometra Solaris, BeO. {pars), Rep. Zool. Coll. H.M.S. "Alert" Lond., 1884, p. 164.
1884. Adinometra sp. juv., Bell, Ibid., p. 170.
Centro-dorsal a thin disk, bearing from ten to eighteen marginal cirri. These have
nine to fifteen joints, usually eleven or twelve, the later ones smooth and generally but
little longer than wide.
First radials more or less visible ; the second very short, closely joined laterally, and
united by syzygy to the widely pentagonal axillaries. Ten arms, which are widest about
the tenth or twelfth brachial. They consist of some hundred and fifty subtriangular
joints, nearly twice as wide as long, and provided with a tolerably distinct median ridge,
the later joints becoming more quadrate. The posterior arms are sometimes non-
tentaculiferous and taper more rapidly, with only about sixty joints.
A syzygy between the first two brachials, and another in the third ; the next syzygy
is anywhere between the eighth and twelfth brachials, with others at intervals of two
to ten joints, generally four or five, but sometimes only two in the lower parts of the
arms.
The first pair of pinnules (on second and third brachials) may reach 14 mm. long, with
forty-five joints, mostly shorter than wide. The terminal comb is sometimes very large,
more than half the joints taking part in its formation. The second pair of pinnules are
shorter, with fewer and smaller joints, but are also combed ; the third pair still shorter
with stouter and wider joints, the dorsal edge sharply serrate, but not forming a true
comb. The following pinnules are long again with stouter joints. The second, or some-
times the second and third, joints of the pinnules on the fourth and fifth brachials have
the dorsal edge raised into a prominent keel. This is generally also present on the
pinnule of the third brachial, sometimes on that of the sixth, and rarely on that of the
second.
The mouth is generally radial ; the disk sometimes quite naked, and sometimes a
good deal plated, both along the ambulacra and between them.
Colour in spirit, — purple ; reddish, greyish, or blackish-brown, passing into brownish-
white. The arms of brown specimens may have a median band of white between two
darker ones.
Disk 1 3 mm. ; spread 1 8 cm.
Localities. — Cape York, September 7, 1874; channel between Albany Island and
Somerset; 8 to 12 fathoms. One specimen.
2S6 THE VOYAGE OF H.M.S. CHALLENGEE.
Samboangan; 10 fathoms. Two specimens.
Other Localities. — Indian seas (Eetzius) ; Australian seas (Peron and Lesueur) ;
Singapore ; Java ; the Moluccas ; North Celebes ; Banka ; Billiton ; Bohol ; North-west
Australia; the ArafuraSea; Dundas Strait ; Warrior Eeef ; Thursday Island ; Prince of
Wales Channel ; Fitzroy Island ; Port Molle ; Port Curtis.
Remarks. — Midler associated this species with the name of Eetzius,1 referring only to
the latter author's famous dissertation which was published in 1805. The name, how-
ever, had been used by Linnaeus in the tenth edition of the Systema Naturae (1758),
the first in which species were characterised ; 2 and it was also employed by Eetzius3 in
1783. Linnaeus referred to Asterias pectina ta the two ten-armed Comatulas figured by
Linck,4 which are the British and the Mediterranean varieties of Antedon rosacea, and
also the Stella chinensis of Petiver. But except for his mention of the type as belonging
to the Indian seas, there is no evidence of his having associated the specific name
p&otinata with any particular form from this locality. This, however, was done by Eetzius
in 1783 and again in 1805, when he separated Asterias tenella from Asterias pectinata,
and his type specimens of both species are still in existence. He added a description in
Swedish to the Linnaean diagnosis of Asterias pectinata, and his reference to the number
and characters of the cirri indicates that he was not speaking of a European Comatula,
but of the specimen from the Indian seas in the Eetzian collection ; while he eventually
only included under this name one of Linck's two species, Decacnemos barbata, from
the Mediterranean, remaining in doubt as to the position of the British Decacnemos
rosacea.5
Lamarck made no allusion whatever to Eetzius' two descriptions of Asterias pectinata,
although the first one was quoted in Gmelin's edition of the Systema Naturae on the
same page (3166) as that to which Lamarck referred in the case of Asterias tenella.
De Blainville also left it without notice as an eastern species, though he quoted Adams'
use of the name for the British Comatula ; and it remained in obscurity till Midler's visit
to Lund in 1841. After examining Eetzius' type specimen, he gave a careful and
perfectly recognisable description of it, one of the best, in fact, which he ever wrote.6
He eventually came to the conclusion, however, that it seemed to be a colour variation
of Comatula Solaris, Lamarck,7 and he put Asterias pectinata into the synonymy of this
type, but with a (?). I believe myself that the two species really are distinct ; but should
it ever become necessary to unite them under one name, that name must be pectinata
and not Solaris. Lamarck's description of Comatula Solaris is as insufficient as that of
1 Archivf. Naturgesch., 1843, Jahrg. ix. Bd. i. p. 133. 2 Op. cit, t. ii. p. 663.
3 K. Svensk. Vetensk. Akad. Handl., At. 1783, t. iv. p. 241. 4 Op. cit, Tab. xxxvii. figs. 64, 66.
6 Op. cit, p. 34. 6 Archivf. Naturgesch., 1843, Jahrg. ix. Bd. i. p. 133.
7 Abhandl. d. k. Akad. d. Wiss. Berlin, 1847 [1849], p. 249.
REPORT ON THE CR1NOIDEA. 287
Asterias pectinata by Linnaeus, and the essential characters of the latter species were
described by both Retzius and Miiller before the latter author visited Paris and saw the
Lamarckian types for himself, so that his diagnosis of them did not appear till six years
after he had properly described Asterias pectinata.1
Dujardin and Hupe deserve the credit of having definitely restored Asterias
pectinata, Retzius, to specific rank on the basis of Midler's description of it, though they
erroneously state that it corresponds to Comatula pectinata and Comatula barbata of
other authors.2 Both these names were given to varieties of the European Antedon
rosacea, which are altogether different from the type of Asterias pectinata from the
Indian seas in the Retzian collection at Lund.
Remarks. — Little need be said about this species, as its essential characters have been
fully discussed already. The centro-dorsal rarely conceals the first radials entirely
(PI. LIII. fig. 15), and it is sometimes relatively smaller than in any other Comatula;
while each of the radial areas on its surface has a deep marginal hollow which corresponds
to one on the surface of the radial above it.3 It thus shows an approximation towards
the characters of Actinometra typica and the other sj>ecies which have more or less
definite openings around the margin of the centro-dorsal (PI. LVII. fig. 1 ; PI. LXIII.
fig. 6 ; PI. LXV. figs. 1-6 ; PI. LXVII. fig. 1). The three Challenger specimens are
remarkable for having an interradial mouth, as it is radial in most examples of the
type that I have seen, just as in Actinometra Solaris.
The Copenhagen Museum contains a specimen from Java which bears the MS. name
Actinometra affinis, Liitken. I was at first inclined to regard it as distinct from
Actinometra pectinata ; but since examining the " Alert " collection I have no doubt that
the two forms are identical. The Java specimen is remarkable for the carination of the
lower joints of the first pinnule, as in some individuals from Bohol in Semper's collection;
while it has eleven arms, owing to one of the normal second brachials being replaced by an
axillary, i.e., there are two distichals united by syzygy, just as in Actinometra paucicirra
(PI. LIV. figs. 1, 2).
This, of course, is what might naturally be expected from the characters of the
type.
1 Troschel's description of Alecto Solaris in 1843 omits all mention of the syzygies in the radials and lower brachials,
and so is useless for the recognition of the species ; while it appears two pages later than Miiller's more detailed descrip-
tion of Asterias pectinata, which noticed this point and also the presence of the keel on the second pinnule, of which
Troschel said nothing.
2 Op. cil, p. 209.
3 See Trans. Linn. Soc. Lond. (Zool.), ser. 2, 1879, pp. 67, 89-91, pi. v. figs. 6-9 ; pi. viii. figs. 5-8.
288 THE VOYAGE OF H.M.S. CHALLENGER.
2. Actinometra Solaris, Lamarck, sp. (PI. V. figs. 4, a-c ; PI. LIII. figs. 1-14;
Part I. pi. liv. figs. 10, 11 ; pi. lv. fig. 2).
Specific formula. — a.R. "o-^-
1816. Comatula Solaris, Lamarck, Histoire Naturelle des Animaux sans Vertebres, Paris, 1816,
t. ii. p. 533.
1834. Comatula Solaris, de Blainville, Manuel dActinologie, Paris, 1834, p. 249.
1841. Actinometra imperialis, Muller, Monatsber. d. k. preuss. Akad. d. Wiss. Berlin, 1841,
p. 181.
1843. Actinometra imperialis, Muller, Archiv f. Naturgesch., 1843, Jahrg. ix. Bd. i. p. 132.
1843. Alecto Solaris, Muller, Ibid., p. 135.
1843. Actinometra imperialis, Muller, Abhandl. d. k. Akad. d. "Wiss. Berlin, 1841 [1843],
p. 226.
1849. Comatula (Actinometra) Solaris, Muller, Abhandl. d. k. Akad. d. Wiss. Berlin, 1847
[1849], p. 248.
1862. Comatula Solaris, Dujardinand Hupe, Hist. Nat. des Zoophytes, Echinodermes, Paris, 1862,
p. 200.
1862. Actinometra imperialis, Dujardin and Hupe, Ibid., p. 209.
1869. Comatula (Actinometra?) hamata, Herklots, Bijdragen tot de Dierkunde, 1869, Bd. ix.
p. 10, pi. ix.
1879. Actinometra Solaris, P. H. Carpenter, Trans. Linn. Soc. Lond. (Zool.), ser. 2, 1879, vol. ii.
p. 27, pi. i. fig. 2 ; pi. v. figs. 1-4.
1879. Actinometra robusta, P. H. Carpenter, Ibid., p. 27, pi. v. figs. 10-15.
1881. Actinometra Solaris, P. H. Carpenter, Notes from the Leyden Museum, 1881, vol. iii.
p. 192.
1882. Actinometra Solaris, P. H. Carpenter, Journ. Linn. Soc. Lond. (Zool.), 1882, vol. xvi.
p. 514.
1882. Actinometra robusta, P. H. Carpenter, Ibid., p. 517.
1882. Actinometra albonotata, Bell, Proc. Zool. Soc. Lond., 1882, p. 535.
1882. Actinometra Solaris, Bell, Ibid., p. 535.
1882. Actinometra albonotata, P. H. Carpenter, It>id., p. 747.
1882. Actinometra robusta, P. H. Carpenter, Ibid., p. 747.
1882. Actinometra Solaris, P. H. Carpenter, Ibid., p. 747.
1884. Actinometra Solaris, Bell (pars), Rep. Zool. Coll. H.M.S. " Alert " Lond., 1884, p. 164,
pi. xvi. fig. A, a.
1884. Actinometra Solaris, var. albonotata, Bell, Ibid., p. 165.
1884. Actinometra intermedia, Bell, Ibid., p. 166, pi. xvi. fig. A, b.
1884. Actinometra robusta, Bell, Ibid., p. 167, pi. xvi. fig. A, c.
1884. Actinometra strota, Bell, Ibid., p. 167.
1885. Actinometra Solaris, Bell, Proc. Linn. Soc. N. S. W., 1884 [1885], vol. ix. p. 498.
1885. Actinometra intermedia, Bell, H/id., p. 498.
Centro-dorsal a thin disk, bearing from ten to twenty-five marginal cirri. These have
from seventeen to twenty-four joints, usually about twenty, the later ones smooth and
.sometimes much longer than wide.
First radials mostly concealed ; the second very short, closely joined laterally, and
united by syzygy to the widely pentagonal axillaries. Ten arms, which have more or
REPORT ON THE CRINOIDEA. 289
less distinct alternate tubercular elevations at their bases, and are widest about the
twelfth joint, reaching 5 mm. They may have nearly two hundred subtriangular joints,
twice as wide as long, with but little or no indication of a median ridge, the later ones
becoming more quadrate. The posterior arms are sometimes non-tentaculiferous, with
fewer joints, and taper more rapidly. A syzygy between the first two brachials, and
another in the third ; the next anywhere between the eighth and twelfth, with others at
intervals of three to ten, generally four or five joints.
The first pair of pinnules (on second and third brachials) may reach 25 mm. long,
with about sixtjr joints, most of which are a good deal longer than wide, the latter half
forming a large terminal comb. The second pair are shorter with fewer and smaller
joints, but are also combed. The third pair are still shorter, with smaller basal joints,
but the succeeding ones are wider, and somewhat saucer-shaped. Their dorsal edges
often stand up prominently, but do not generally form a true terminal comb. The
following pinnules are long again, with wider and more massive joints, which project
laterally beyond their successors and are more or less sharpened along the medio-dorsal
line.
The second and the two or three following joints of the pinnules on the fourth and
fifth brachials have a sharpened dorsal edge, which is generally produced into a more or
less prominent keel. The pinnule of the sixth brachial usually has the same character as
the two previous ones, while keels may also be present on the second or more of the lower
joints of the pinnules of the third and seventh brachials. Occasionally, however, there is
no keel at all.
Mouth radial ; disk sometimes quite naked, and sometimes very extensively plated
both alonsj the ambulacra and between them.
Colour in spirit, — deep purple or rose-colour, more or less relieved by patches of white
and brown ; light reddish-brown, the arms with a median band of white between two
dark ones ; or greyish-white with dark spots.
Disk 15 to 25 mm.; spread 25 to 30 cm.
Localities. — Cape York, September 7, 1874; Channel between Albany Island and
Somerset; 8 to 12 fathoms. Several specimens.
Station 187, September 9, 1S74; off Booby Island; lat. 10° 36' S., long. 141° 55' E.;
6 fathoms ; coral mud. Abundant.
Other Localities. — " Indien " (Mus. Wien) ; Australian Seas (Peron and Lesueur) ;
Hong Kong ; China Sea ; Singapore ; Billiton ; Prince of Wales Channel ; Albany Island;
Port Molle ; Port Curtis.
Remarks. — This is a large and extremely handsome species when fully developed.
The centro-dorsal of the form which has been hitherto known as Actinometra rdbusta
may reach 7 mm. in diameter ; and with the exception of this type and of Antedon
(ZOOL. CHALL. EXP. — PART LX. — 18S7.) OoO 37
290 THE VOYAGE OF H.M.S. CHALLENGER.
eschrichti, there are few recent Comatulse with a calyx which at all approaches that of
many fossil species in size.
The centro-dorsal of the' large Vienna specimen has lost all trace of its cirrus-sockets
on one side, and is almost reduced to a level with the radials ; while iu an "Alert"
specimen from Port Molle the sockets are all obliterated, leaving nothing but a thin flat
plate, very much as in some forms of Actinometra paucicirra (PI. LIV. figs. 1-4). The
calyx of the form from Cape York, which I have hitherto called Actinometra strota, is
represented in figs. 4, a-c, on PI. V. Except for the almost entire absence of a basal
star (fig. 4c), it is not greatly different from that of the individual from Singapore which
I figured in 1879 ; x but it is very much smaller than the calyx of Actinometra robusta,
which reaches 7 mm. in diameter, while 5 mm. is the maximum size of the Challenger
specimens ; and none of them show any trace of the curious diverticulum of the axial
canal into the substance of the radial which occurs in that variety.2
The large " Alert " specimen from Port Molle is also remarkable for having the disk
perfectly soft and membranous ; while in others from Port Curtis and Torres Strait it
is covered with minute polygonal plates, and the ambulacra are also strongly plated
as shown in Part I., pi. liv. figs. 10, 11. The ambulacral plating ceases entirely,
however, where the arms come off at the margin of the disk, and they have nothing at
all like the ambulacral skeleton which is often so fully developed in Antcdon and in the
Pentacrinidse.
A large number of this species were obtained by the Challenger at Booby Island,
including several in a more or less immature condition. One of the smallest of these is
represented on PL LIII. fig. 1, and is noteworthy for the great relative length of its
arm-joints, as compared with those of the adult (fig. 2).
Bell has given the varietal name albonotata to a specimen from Albany Island with
twenty to twenty-five cirri and but slightly keeled basal joints on the lower pinnules, in
which the coloration consists of white spots on a dark ground.3 But the colour variations
of this species, even in one and the same locality, are almost as numerous as those of
Antedon carinata; and I cannot see much use in giving varietal names even to
forms which may seem to reach "the extreme limit of the species." Differences of colour
are in fact quite useless for the purpose of specific discrimination among the Comatulse.
2. The Paucicirra-gvowp.
Bidistichate species, with twenty arms or more ; the two outer radials and the first
two joints after each axillary respectively united by syzygy.
Remarks. — This group includes only two species, which are closely allied to the
1 Trans. Linn. Soc. Lond. (Zool.), ser. 2, 1879, vol. ii. pi. v. figs. 1-4.
2 Ibid., p. 86. 3 "Alert" Report, p. 1C5.
REPORT ON THE CRINOIDEA. 291
Solaris-group, but have twenty or more arms. The two outer radials, the two distichals,
the two palmars (if present), and the first two brachials are respectively united by syzygy.
One of these species was obtained at Mergui by Dr. J. Anderson, F.R.S., and is as yet
undescribed. The other is Actinometra paucicirra, which is abundant in Torres Strait,
and nearly always has its complete set of ten distichal axillaries (PI. LIV. figs. 1, 2).
On the rare occasions when one of these is absent there is still the syzygy between the
two lower brachials, just as in Actinometra Solaris and Actinometra pectinata (PI. LIII.
figs. 2, 15); while when distichals are abnormally developed in these forms there are
two joints united by syzygy, just as in Actinometra paucicirra.
Actinometra paucicirra, Bell (PL IV. figs. 6, a, b ; PI. V. figs. 3, a-c ; PI. LIV.;
also Part I., pi. lv. fig. 1).
d.(p).br.
Specific formula — a.R.^-^- — '.(—
1879. Actinometra Juhesii, P. H. Carpenter, Proc. Roy. Soc, 1879, vol. xsviii. p. 390.
1882. Actinometra ■paucicirra, Bell, Proc. Zool. Soc. Lond., 1882, p. 535.
1882. Actinometra jukesi, P. H. Carpenter, Ibid., p. 747.
1882. Actinometra, paucicirra, P. H. Carpenter, Ibid., p. 747.
1884. Actinometra jukesi, Bell, Rep. Zool. Coll. H.M.S. "Alert," Lond., 1884, p. 168.
1884. Actinometra paucicirra, Bell, Ibid., p. 169.
1885. Actinometra jukesi, Bell, Proc. Linn. Soc. N.S.W., 1884 [1885], vol. ix. p. 498.
Centro-dorsal small and discoidal in young individuals, and bearing five to ten
slender marginal cirri. These have twelve to eighteen smooth joints, most of which
are longer than wide, the penultimate with a slight spine. In mature individuals the
centro-dorsal is a fiat pentagonal plate, flush with the radials and devoid of all trace
of cirri.
Three radials visible, the two outer ones united by syzygy. The second are widely
oblong but free laterally in young individuals, trapezoidal and closely united in the adult.
Axillaries nearly pentagonal in the young, but almost triangular in the adult. Eighteen
to twenty-three arms, but generally twenty, each ray bearing two distichal series, which
consist of two joints, united by syzygy- The palmar series, if present, of the same
character.
The first two brachials are united by syzygy, and the two outside arms of each ray
also have a syzygy in the third brachial. The next syzygy is in the eighth or tenth
brachial, the latter being common in the outer arms, and others follow at intervals of one
to six, usually three or four joints. About one hundred and fifty brachials, the first five
or six nearly oblong, the following ones more triangular and wider than long, becoming
more quadrate towards the ends. The second brachial bears a long tapering pinnule with
a large terminal comb. It may reach 20 mm. long, and consists of about sixty short joints,
292 THE VOYAGE OF H.M.S. CHALLENGER.
the lowest rather wide and stout, with prominent dorsal edges. The next four or five
brachials bear similar pinnules with smaller combs and decrease rather rapidly in size,
two or sometimes three of the basal joints becoming rather strongly keeled. The
succeeding pinnules have stouter joints and increase a little in length.
The mouth is radial and almost marginal ; and the anal area is more or less thickly
covered with irregular plates, but the ambulacra are unprotected.
Colour in spirit, — reddish- or greyish-brown, bleaching to white, often with a dark
medio-dorsal line.
Disk 18 mm.; spread 20 cm.
Localities. — Cape York, September 7, 1874; Channel between Albany Island and
Somerset; 8 to 12 fathoms. Several specimens.
Station 187, September 9, 1874; off Booby Island ; lat. 10° 36' S., long. 141° 55' E.;
6 fathoms ; coral mud. Several specimens.
Arrou Islands. Tbree specimens.
Other Localities. — H.M.S. "Alert," Prince of Wales Channel; Albany Island, 3 to
4 fathoms; Port Molle, 12 fathoms.
Remarks. — This is one of the most easily recognisable species of Actinometra, owing
to the entire absence of cirri in its adult state (PI. LIV. figs. 1-7). After making a
preliminary examination of the Challenger material I found that the type was already in
the national collection, having been dredged by Jukes on the north-east coast of Australia ;
and I therefore proposed to call it " Actinometra Jukesii" under which name it was
noticed in my Preliminary Report.1 When the " Alert " collection reached the British
Museum, it proved to contain several examples of this type which Professor Jeffrey Bell
left undescribed, as he knew that I should publish its diagnosis in the present Report.
The "Alert" also dredged some smaller individuals which resembled Actinometra jukesii
in every respect except for the presence of half a dozen short cirri on the small centro-
dorsal. It did not strike me, however, during my first and somewhat cursory examina-
tion of the " Alert " material, that these might be young individuals of Actinometra
jukesii, like those which I had already noticed in my Preliminary Report ; and they were
eventually described and figured by Professor Bell under the name of Actinometra
paucicirra,2 by which the species must in future be known.
Two other immature specimens, differing somewhat from the type, were obtained by
the Challenger at the Arrou Islands ; and the centro-dorsal of one of them is figured on
PI. IV. fig. 6, a, under the specific name of aruensis, which should be changed to
paucicirra, as I am now convinced that they belong to this species, though I regarded
them as distinct when the plate was lettered, six years ago. The changes which take
place in the centro-dorsal of the young Actinometra paucicirra during its growth to
1Proc. Roy. Soc, 1879, vol. xxviii. p. 390. 2 "Alert" Report, p. 169, pi. xvii. fig. A, a.
REPORT ON THE CR1NOIDEA. 293
maturity have been already described on p. 14, and are illustrated on PI. LIV. The
young centro-dorsal is a rounded plate with a flattened ventral surface bearing relatively
large basal grooves (PI. IV. fig. 6a). These lodge the well-developed basal star, an
isolated ray of which is seen in PI. IV. fig. 6b. In the adult, however, the ventral
surface of the centro-dorsal is much more convex, as it fits into the inverted funnel
formed by the ring of radials (PI. V. fig. 3c), no part of it being visible in a side view of
the calyx (fig. 3b). Its dorsal surface is flush with that of the radials, which is often
marked by the origin of a dark medio-dorsal line extending outwards over the calyx and
the bases of the arms (PL LIV. fig. 2). These are most frequently twenty in number.
Among fifty individuals I have only found one which had not got its full complement of
distichal axillaries, one ray (which wants a second radial) being entirely without them, so
that the number of arms is reduced to eighteen. Thirty-two examples have twenty arms ;
nine have twenty-one, seven twenty-two, and one twenty-three. The palmar series, when
present, always resemble the distichals in consisting of two joints which are united by
syzygy (PI. LIV. fig. 2).
The arrangement of the syzygies at the bases of the arms is somewhat peculiar.
There is always one between the first two brachials, even in the case where distichals
are absent, so that the type then reverts to that of the Solar is -group, in which the
distichal series, when abnormally present, resemble those of Actinometra pancicirra.
In both the members of the Solaris- group there is also a syzygy in the third brachial
(PI. LIII. figs. 1, 2, 15); and this is sometimes the case in Actinometra maculata
(PI. LV. fig. 2). It appears in one of the two arms of the single abnormal ray of the one
individual of Actinometra 'paucicirra which has no distichal series. In normal individuals,
however, the third brachial is very regularly a syzygial joint in the two outer arms of
the ray, the normal sequence of the syzygies thus being 1-2, 3, 11, 15 ; whereas on the
inner arms it is 1-2, 9, 13 (PL LIV. figs. 1, 2). This is a very distinct peculiarity
of the species; but the syzygies are rather obscure in young individuals and it seems
therefore to have escaped the notice of Bell, who makes no reference to it either in his
diagnosis or in his figure of an immature specimen.
The two outside arms of each ray in young individuals are often much smaller than
the inner pair (PL LIV. fig. 10). This is especially distiuct in those from the Arrou
Islands, in one of which, with a spread of 20 cm., the outside arms on some of the rays
are so small as to look like unusually developed pinnules. But their true nature is shown
by the fact that they bear small pinnules themselves. In the youngest of these small
arms there is a relatively large pinnule on the second, and a very small one on the third
brachial; but there are none on the next four joints, though they reappear again on the
eighth. This is altogether in accordance with the mode of development of the pinnules
in other Cornatulse, which I have described elsewhere.1
lBull. Mus. Comp. Zool., 1881, vol. ix. No. 4, pp. 14, 15.
294 THE VOYAGE OF H.M.S. CHALLENGER.
The amount of carination of the lower pinnules varies considerably, just as it does in
Actinometra solan's (PI. LIII. figs. 3-22). As a general rule the first pair of pinnules
have their basal joints somewhat produced towards the dorsal side, and in the next two
pairs the second and third joints have rather prominent keels, traces of which are some-
times visible as far as the twelfth or fifteenth brachial. The terminal comb, which is
very well developed on the basal pinnules, becomes gradually smaller and disappears
about the sixth or seventh brachial.
The visceral mass of Actinometra paucicirra, like that of Actinometra Solaris, which
occurs at the same locality, is somewhat readily detached from the calyx, and it was
occasionally dredged in an isolated condition. It is not so completely plated as that of
Actinometra Solaris often is. For the ambulacra are unprotected, and the interradial
areas are covered by larger and more nodular plates than in the latter species (Part I.,
pi. liv. figs. 10, 11 ; pi. lv. fig. 1). Both species, however, may sometimes have the
calcareous deposits considerably reduced in extent, though they are rarely entirely absent
(Part I., pi. lv. fig. 2). The figured specimen of Actinometra paucicirra shows a small
Anilocra living in the anal tube.
One tetra-radiate individual of this species occurred among all those dredged by the
Challenger. An examination of the disk shows that the anterior ray A is missing, so
that the mouth comes to be interradial, between the radii E and B, while the anus as
usual lies between C and D. The only other species which presents the same arrange-
ment of the arm divisions as occurs in Actinometra paucicirra is a new form from Mergui,
which differs from it in having normally two, and sometimes three, postradial axillaries,
and also in the presence of some thirty cirri on the centro-dorsal.
3. The Typica-grouj).
Tridistichate species with the radial axillaries and all the post-distichal axillaries
united to the preceding joints by syzygy.
Remarks. — This group contains four of those abnormal species in which the two
outer radials and the first two joints above the distichal and every subsequent axillary
are respectively united by syzygy ; while the distichal series itself consists of the usual
three joints, with the axillary a syzygy. They are all confined to the Eastern Archipelago
and Western Pacific, three of the four being purely littoral species ; while Actinometra
typica was also obtained by the Challenger in the neighbourhood of Fiji, from a depth of
over 200 fathoms. The rays of this species, and also those of Actinometra multibrachiata
divide very frequently, the number of postradial axillaries being sometimes as many as
seven (PI. LVI. fig. 3 ; PI. L VII. fig. 1) ; whereas in Actinometra distincta there is no
REPORT ON THE CRINOIDEA. 295
axillary beyond the palmar (PI. LV. fig. 1). The mutual relations of the four species
may be expressed as follows : —
A. Two post-radial axillaries. Twelve cirrus-joints, . . . .1. distineta, n. sp.
I>. Three or more post-radial axillaries.
L Centro-dorsal stellate, and without functional cirri, . . .2. tijpica, Loven, sp.
II. Centro-dorsal bears functional cirri.
a. Three or four post-radial axillaries. Cirri few, . . . novx-guinese, Mull., sp.
b. Six or eight post-radial axillaries. Cirri well developed, . . 3. mullibrachiata, n sp.
1. Actinometra distineta, n. sp. (PL LV. fig. 1).
Specific form ula — a. R. 3 . -Lrr1. —.
Description of an Individual. — Centro-dorsal a thick rounded disk, bearing about
thirty marginal cirri with a dozen joints, nearly all of which are longer than wide. The
later joints have faint spines.
First radials just visible ; the two outer ones short, wide, and united by syzygy.
The second are also closely joined laterally. The rays may divide three times ; three
distichals, the third axillary with a syzygy ; the two palmars and the first two brachials
are respectively united by syzygy.
Thirty-six arms, of triangular joints which are wider than long and overlap slightly,
the terminal ones becoming more quadrate. The anterior arms are long, slender, and
slowly tapering, of one hundred and twenty to one hundred and fifty joints, while the
posterior are short and taper rapidly, with only sixty to eighty segments.
A syzygy between the first two brachials ; the next about the eighth or tenth, with
others at intervals of two or three joints.
The pinnules decrease in length from that on the second distichal to those of the
fourth and fifth brachials, and then become larger again. The first eight or nine pinnules
on each side have a terminal comb, which may occur at intervals as far as the thirtieth
brachial.
Mouth interradial and the anal tube almost marginal ; a few small calcareous nodules
on the disk.
Colour in spirit, — brownish-white, with dark spots on the medio-dorsal line of the rays
and their branches as far as the first brachials ; there are also lateral spots on the junction
lines of the outer radials and lower distichals.
Disk 11 mm.; spread about 12 cm.
Locality. — Samboangan ; 10 fathoms. One specimen.
Remarks. — Except for the characters of the rays and their subdivisions, this little
species presents no special peculiarities, the shape of the arm-joints being that which is
296 THE VOYAGE OF H.M.S. CHALLENGER.
most common in the genus, while the lower joints of the pinnules are not carinate or
otherwise distinguished. The absence of any axillary above the palmar separates it
altogether from the multibrachiate species next to be described (PI. LVI. fig. 3 ; PI. LVII.
fig. 1), while it is distinguished from Actinometra paucicirra by the greater number of
distichal joints. All the arms seem to be tentaculiferous ; but there may be nearly twice
as many joints in the anterior as in the posterior arms.
2. Actinometra typica, Loven, sp. (PI. LVII. fig. 1).
Specific formula — a.E. 3. ' "2 " — '.
1866. Phanogenia typica, Loven, Ofversigt k. Vetensk.Akad. Fbrhandl., 1866, No. 9, p. 231.
Actinometra stellata, Liitken, SIS., Museum Godefi'roy.
1S79. Phanogenia typica, P. H. Carpenter, Trans. Linn. Soc. Lond. (Zool.), ser. 2, 1879, vol. ii.
p. 20.
1879. Actinometra stellata, P. H. Carpenter, Proo. Roy. Soc, 1879, vol. xxviii. p. 386.
1881. Actinometra typica, P. H Carpenter, Notes from the Leyden Museum, 1881, vol. iii.
p. 195.
1882. Actinometra, typica, Rell, Proc. Zool. Soc. Lond., 1882, p. 535.
1882. Actinometra typica, P. H. Carpenter, Ibid., p. 747.
Centro-dorsal stellate, with little or no trace of cirrus-sockets, and nearly flush with
the radials, from which it is separated by distinct clefts, sometimes being even below
their level. A syzygy between the two outer radials, which are both short and wide,
the second being almost completely united laterally ; but beyond this point the rays are
quite free. Sometimes as many as seven postradial axillaries ; the distichal series
normally of three joints with a syzygy in the axillary, while the palmar and subsequent
series are each of two joints united by syzygy. The first two brachials are united by
syzygy and there may also be one in the third brachial. The next is about the eighth
or tenth brachial, and others follow at intervals of two joints.
Eighty or more relatively short and slender arms of slightly overlapping, triangular
joints. The first pinnule, on the second distichal, is long but rather slender, and composed
of numerous short joints. The next, which is normally on the second brachial, is of the
same character, but smaller, and the next few pinnules are of decreasing length, becoming
longer and stouter again about the sixth brachial. The lowest pinnules have a well-
defined terminal comb which extends to about the twelfth brachial, and occasionally
appears further out on the arms. The joints of the middle and later pinnules are fringed
with strong spines.
The mouth is usually subcentral and radial, with the primary ambulacra arranged
very much as in Antedon, but the anal interradius is considerably the largest, with the
anus near its margin. Both interradial and interpalmar areas are often much plated.
Colour in spirit, — brown.
REPORT ON THE CRINOIDEA. 297
Disk 20 mm.; spread reaching 25 cm.
Locality. — Station 174b, c, or d, August 3, 1874; near Kanclavu, Fiji; lat.
(about) 19° 6' S., long, (about) 178° 18' E.; 255, 610, or 210 fathoms;1 coral mud;
bottom temperature at 610 fathoms, 39° F. One mutilated specimen.
Other Localities.- — Malacca; Jobie ; Zebu; Fiji; Kingsm ill Islands.
Remarks. — This species is the one for which the genus Phanogenia was established
by Loven 2 on account of its stellate centro-dorsal and exposed first radials. The same
peculiarity was noted by Dr. Liitken in an Actinometra of the Godeffroy collection from
Fiji, to which he gave the MS. name Actinometra stellata; and duplicates of the type
have been distributed from the Godeffroy Museum under this name. Having examined
some of these duplicates, aud also by the kindness of Professor Loven his original
specimens of Phanogenia, I came to the conclusion, as Dr. Liitken had previously done,
that the two types are identical. Loven's generic name thus becomes a synonym of
Actinometra, while his specific name is that by which the type must be known for the
future. It is a sufficiently remarkable species, apart altogether from the peculiarities of
its radials and centro-dorsal. For the mouth is at no great distance from the centre of
the disk, and the arrangement of the ambulacra in five primary divisions is almost as
regular as in the Endocyclic Crinoids. The anal interradius is therefore by no means
so large and conspicuous as it usually is in Actinometra (PI. LVII. fig. 3 ; PL LXVIII.
%• 1)-
Loven described the two outer radials of this type as articulated bifascially ; 3 but
I believe them to be really united by a syzygy of much the same character as occurs
in Pentacrinas and Rhizocrinus, viz., with the apposed faces almost smooth and devoid
of the radial striatum which is so marked in the syzygies of Antedon. The result is
that the junction line of the two joints is simple, instead of being more or less inter-
rupted as in the syzygies of the later ray-divisions in this type and in most other
Comatulse. Loven gave a sketch of the distal face of a second radial in Actinometra
typica i which seems to have a median vertical ridge like that which he figures in the
corresponding part of Antedon eschrichti.5 In reality, however, there is not an
articular ridge with a fossa on either side of it for the reception of a muscular or liga-
mentous bundle, but merely a division between the two sides of the joint-face, which
has a slight general convexity ; and there is a corresponding concavity, which is divided
into two parts by a median line, on the proximal face of the axillary radial. If the
two joints were really articulated each face would have a median ridge and lateral
fossae instead of fitting into one another by a slight curvature. The median line
1 The exact station, and consequently the exact depth, is not recorded.
2 O/versigt. k. Vetensk. Akad. Forhandl, 1866, No. 9, p. 231.
3 Ibid., p. 228. * Ibid., p. 230, fig. c. » Ibid., p. 230, fig. k.
(ZOOL. CHALL. EXF. — PART LX. — 1887.) Ooo 38
298 THE VOYAGE OF H.M.S. CHALLENGER.
represented by Loven in his figure of the second radial also appears on an undoubted
syzygial face from further out on the ray ; and I have no doubt whatever that the
union of the two outer radials is really a syzygial one, though the usual radiating ridges
and furrows, which are so characteristic of syzygies in Comatulse, are not present on the
apposed faces. Traces of them are sometimes visible, however, as a series of little
elevations which radiate outwards from the central canal and produce the appear-
ance of a syzygial face with its ridges interrupted at intervals. But in other
cases the apposed faces are almost smooth, just as in the syzygies of Pentacrinus and
Rhizocrinus. The syzygies further out on the rays, however, are more normal in
character.
There is a considerable amount of variation in the general features of this species.
The form which comes nearest to it is Actinometra novse-guinese ; but this is not known
to have more than four post-radial axillaries, while Actinometra typica may have
as many as seven. Furthermore the centro-dorsal of Midler's unique specimen of
Actinometra novse-gninese was described by him as having "15 Ranken und mehr" ;
though it shows traces of clefts at the sides and approximates therefore towards the
condition reached in Actinometra typica. Even in this last it ma}7 bear a few rudi-
mentary cirri, as in the specimen figured by Loven ; 1 and there is a considerable amount
of variation in the extent to which it is sunk within the radial funnel.
As in other species of Actinometra the tridistichate series is not unfrequently
replaced by a bidistichate one. This occurs on both sides of two rays in the Challenger
specimen (PL LV1I. fig. 1) ; and the Copenhagen Museum contains one anomalous
individual from Fiji in which eight out of the ten distichal series consist of but two
joints. I believe them to be articulated, and not united by syzygy, as one would
rather expect them to be. But then, it sometimes happens that there are three joints
in a palmar or post-palmar series, instead of the normal two ; and the first two of these
are syzygially united, a condition which is altogether anomalous for a three-jointed
series (see Rules 2, 6). On the other hand, however, it seems only natural that the
terminal faces of the two joints borne on any axillary should have the same character,
so that the normal syzygy of the one is accompanied by the abnormal syzygy of the
other.
The range of this species, as at present known, extends from Malacca through the
Philippine Group, to Fiji, in all of which localities it belongs to the purely littoral
fauna. It was, however, obtained by the Challenger at a depth of over 200 fathoms,
viz. 210, 610, or 255 fathoms. I imagine for many reasons that it did not occur at the
greatest of these depths, no Actinometra having been yet obtained from below 600
fathoms.
1 Ofversigt k. Vetensk. Alcad. Forhandl, 1866, No. 9, p. 230, fig. a.
REPORT ON THE CRINOIDEA 299
3. Actinometra mult ib rack lata, n. sp. (PI. LVI. figs. 3, 4).
Specific formula- — a. R.3.- " ^ — ' • — •
Description oj an Individual. — Centro-dorsal a pentagonal disk with incurved sides
which project somewhat over the smooth radials. Its dorsal surface is deeply hollowed
in the centre, and bears about twenty-five cirri. These have fourteen to sixteen joints,
nearly all of which are longer than wide, the penultimate with a small spine. Three
radials visible. The first are raised at the angles, but deeply hollowed in the centre
where they fail to meet the upper surface of the centro-dorsal. The two outer radials are
short, wide, and united by syzygy. The second are only partly united laterally and the
whole of the rays above them are cpiite free. Three distichals, the axillary a syzygy, and
two palmars, united by syzygy. There may be six subsequent divisions which are normally
similar to the palmars.
Arms slender, but very numerous, thirty or more to each ray, composed of some
hundred and fifty triangular and overlapping joints with very spinous edges. The first
two brachials are united by syzygy, and there is sometimes another in the third brachial ;
the next is about the tenth or twelfth, and others follow at intervals of two or three
joints.
The first pinnule on the second distichal is very long and slender, reaching 20 mm,
with numerous short joints; the next one, normally on the second brachial, is of the same
character, but much smaller, and the next few are of decreasing length, after which there
is but little increase. The lowest pinnules have a well-defined comb which extends to
about the twelfth brachial, and sometimes appears quite far out on the arms ; the joints
of the middle and later pinnules are very spiny.
Colour in spirit, — dark brown.
Spread probably nearly 30 cm.
Locality. — Banda ; 17 fathoms. One mutilated specimen.
Remarks. — Only a single mutilated example of this remarkable species was obtained,
but its characters are sufficiently distinct to show that it cannot be referred either to
Actinometra novas-guineas or to Actinometra typica. It resembles the latter form in
the frequency of its ray-divisions, but differs from it altogether in having a relatively
large centro-dorsal bearing over twenty well-developed cirri. On the other hand there
are probably three times as many arms as in Actinometra novas-guineas, each of the rays
which are preserved bearing thirty or more; while the centro-dorsal is larger with more
numerous cirri than occur in that type. Its angles rest upon the raised marginal portions
of the radials, which are deeply hollowed in the centre and do not therefore come in
contact with the upper surface of the centro-dorsal, so that it overhangs them consider-
ably when the calyx is viewed from the dorsal side.
300 THE VOYAGE OF H.M.S. CHALLENGER.
Two of the rays only are preserved, the three others having broken away above the
radial axillaries. One of them seems to have been of much smaller size than the rest, to
judge from the two joints of it which remain (PL LVI. fig. 3). I am not sure, however,
that this is simply a case of regeneration from the primary radials. For the small first
radial in this case differs somewhat from its fellows ; and its flattened central portion,
which bears the small second radial, projects outwards a little beyond the general line of
the radial pentagon. There may be some meaning in this asymmetry ; but as the disk is
lost it is impossible to determine whether it is related in any way to the position of the
anus.
The character of the union between the first two distichals of this type has puzzled
me greatly. It has very much the appearance of a syzygy, but I have not liked to attempt
to settle the point by an actual examination of the joint-faces, as it would have involved so
much further mutilation of an already imperfect specimen. This has been possible, how-
ever, in Actinometra typica, which has the usual bifascial articulation between the first
two distichals ; and as there are so many points of resemblance between this species and
Actinometra rnultibrachiata, it is by no means improbable that the latter may be in the
same condition, though the external characters seem to indicate the presence of a syzygy.
This would be a new type of structure altogether ; though it occurs abnormally in the
distal parts of the rays of Actinometra typica (PI. LVII. fig. l), as I have already
pointed out.
For the present, therefore, it will be safer to leave Actinometra rnultibrachiata in the
same group with Actinometra typica, until the question can be definitely settled by the
examination of more material.
Actinometra, Series II.
The two outer radials articulated. Ten arms.
Remarks. — This is a small series and contains but one group, which may be called
the Echmoptera-growp, after the name of its first described species. The Mullerian type
of this species is in the University Museum at Berlin, where it was deposited by Captain
Wendt many years ago, though no locality was recorded for it. I was permitted to
examine it in 1880, and have since come to the conclusion that it is a Caribbean species,
and not improbably identical with one of the many variations of the type which was
described by Pourtales in 1869 under the name Antedon meridionalis, A. Agassiz, MS.,
from the coast of South Carolina. This conclusion is strengthened by the fact, for which
I am indebted to the kindness of Professor E. von Martens, that the Berlin Museum also
contains some other Caribbean Echinoderms which were deposited by Captain Wendt.
Three of the five remaining species of the Echinop>tera-gvou^ are also members of
the Caribbean fauna, viz., Actinometra pulchella, Pourtales, sp., Actinometra rubiginosa,
REPORT ON THE CRINOIDEA. 301
Pourtales, sp., and Actinometra blakei, an MS. species of my own, which is the host of
Myzostoma areolatum, von Graff. These will all be described and illustrated in my
Report on the " Blake " Comatulae, where I hope to work out the relations of Coma-
tula cchinoptera, Miiller, to the later described Actinometra meridionalis.
Actinometra pidchella is also a bidistichate species, while tridistichate series
occasionally occur in Actinometra rubiginosa. The unique specimen of Actinometra
coppingeri, Bell, has but twelve arms ; and it is therefore not improbable that this
species should be referred, like Actinometra rubiginosa, to the ten-armed series, as well
as to the tridistichate one. It was obtained by the "Alert" at Flinders, Clairmont.
Another specimen dredged by the "Alert" at Port Molle has been referred by Bell to
Actinometra cumingi, Mull., sp., which was brought to the Berlin Museum from Malacca.
But no member of this group was obtained by the Challenger in the Eastern Archipelago ;
and I have not seen more than half a dozen other individual representatives of it in
addition to the three already mentioned, although I have visited all the chief continental
collections. The type is evidently a rare one in the Eastern Archipelago, although so
abundant in the Caribbean Sea and on the Brazilian coast. Most of the species of
Actinometra which occur in the Eastern seas belong to the tridistichate type ; and the
ten-armed forms are almost exclusively represented by the Solaris-group, which is rich
in individuals, though poor in species. But it does not occur at all in the Caribbean
Sea where the Echinopt era-group is so extensively developed.
4. The EcJiinoptera-grouv).
Actinometra meridionalis (A. Agassiz), Pourtales, sp. (PL IV. figs. 4, a-c ; PL LVI.
figs. 1, 2).
Specific formula — a. — .
1865. Aleeto meridionalis, A. Agassiz, MS., Seaside Studies, Boston, 1865, p. 121,
figs. 153, 154.
1866. Antedon meridionalis, Verrill, Proe. Bost. Soc. Nat. Hist., 1866, vol. x. p. 339.
1869. Antedon meridionalis, Pourtales, Bull. Mus. Comp. Zobl., 1869, vol. i. No. 11, p. 355.
1878. Antedon meridionalis, Pourtales, Ibid., vol. v. No. 9, p. 214.
1879. Comatida meridionalis, P. H. Carpenter, Trans. Linn. Soc. Lond. (Zool.), ser. 2, 1879,
vol. ii. pp. 20, 27.
1879. Antedon meridionalis, Ratlibun, Traus. Connect. Acad., 1S79, vol. v. p. 157.
1881. Actinometra meridionalis, P. H. Carpenter, Bull. Mus. Comp. Zobl., 1881, vol. ix. No. 4,
p. 6.
1882. Actinometra meridionalis, Ludwig, Mem. Acad. Sci. Bruxelles, 1882, t. xliv. p. 6.
1882. Antedon meridionalis, Bell, Proc. ZooL Soc. Lond., 1882, p. 533.
1882. Actinometra meridionalis, P. H. Carpenter, Ibid., p. 747.
Locality. — Bahia ; 7 to 20 fathoms. Fourteen specimens.
Other Localities. — South Carolina ; West of Tortugas, and off French Reef in Florida
302 THE VOYAGE OF H.M.S. CHALLENGER.
Strait, 35 to 45 fathoms ; Caribbean Sea, abundant between 50 and 262 fathoms ;
off Cape Frio, Brazil, 35 to 45 fathoms (Hassler).
Remarks. — This species has been figured for the sake of comparison with the other
ten-armed species of Actinometra which belong to the Solaris-group (PI. LII.
fio-s. 1, 2, 15). The essential difference between them is the absence in the Echinoptera-
oroup of the syzygies between the two outer radials and the two lower brachials
respectively, these joints being articulated as in most species of Antedon.
The examples of this species obtained by the Challenger at Bahia were the first
representatives of the genus which I had seen from the Brazilian coast ; and as I could
not identify them with any form then described, I proposed to call the type Actinometra
brasiliensis. The calyx is figured under this name on PL IV., which was printed off
before I received the " Blake " collection containing the original examples of Actinometra
meridionalis, and also a very large series of variations on the same general type. The
exact number of specific forms belonging to this type which are represented in the
Caribbean Sea, is a point which I propose to work out in my "Blake" report; and for
the present therefore the Challenger species may be known under the name Actinometra
meridionalis, though it is quite possible that this may have to be discarded in favour of
Actinometra echinoptera, Mliller, sp.
The dimorphism of the arms which is so common in the eastern forms of the genus
is very well marked in some of the Challenger specimens of Actinometra meridionalis,
the hinder arms being ungrooved and consisting of but half the number of joints wbich
occur in the anterior arms. But none of them possess the problematical "sense-organs"
on the pinnules which occur both in some individuals from Cape Frio and in others from
French Eeef on the Florida coast.
The calyx is that of a very typical Actinometra. The centro-dorsal is small and
discoidal (PL IV. fig. 4a), and the articular faces of the radials are set vertically with
small muscle-plates, so that their ventral aspect shows a widely open central funnel
(fig. 4c) ; while there is a well-developed basal star, an isolated ray of which is shown in
PL IV. fig. 46.
Actinometra, Series III.
Two articulated distichals.
Remarks. — This corresponds to Series III. of the Antedon-species, and comprises the
multibrachiate forms in which there are but two distichal joints united by a bifascial
articulation like that between the two outer radials. There may be no further division
as in Actinometra maculata, Actinometra simplex, and in some forms of Actinometra
pulchella (PI. LII. fig. 2; PL LV. fig. 2; PL LIX. fig. 1). But in other examples of
Actinometra pidchetta and in Actinometra stelligera there are palmar series like the
REPORT ON THE CRINOIDEA. 303
distichals (PI. LII. fig. 1 ; PI. LVIII. fig. 1), and I have seen a Philippine specimen with
two other divisions of the same character. On the other Land Actinometra rotalaria
and Actinometra valida have the palmar and subsequent series of three joints, the
axillary with a syzygy (PI. LIX. figs. 2, 3). In these two species, and also in Actino-
metra simplex (PI. LIX. fig. 1), the first syzygy in the free arm is that in the third
brachial ; but in Actinometra pulchella, Actinometra maculata, and Actinometra
stelligera the first two brachials above the last axillary are normally united by syzygy
(PI. LII. figs. 1,2; PL LV. fig. 2 ; PI. LVIII. fig. 1). This is also the case in Actino-
metra Solaris, Actinometra paucicirra, and Actinometra typica (PI. LIILfig. 2 ; PI. LIV.
fig. 1 ; PL LVII. fig. 1) ; but all these forms have a syzygial union between the two
outer radials, which is not the case in those belonging to Series III.
This series thus falls into two very well defined groups according as there is a
syzygial union or a bifascial articulation between the first two brachials of the free arm.
The first of these is altogether unrepresented in Antedon, having a general formula —
a.2.(2.2.2.)."9-. — and may be called the Stelligera- group, after a comparatively large and
well-defined species from Fiji and Samoa (PL LVIII. fig. l). It also includes the widely
distributed Actinometra pulchella ; but as this is a dimorphic type which also occurs in
the ten-armed Echinoptera-gvoxrp, the use of its name to denote a multibrachiate group
might lead to confusion.
The more normal type of bidistichate species which have the first syzygy in the third
brachial of the free arm is but poorly represented in Actinometra, though it includes a
considerable number of A ntedon- species. It is confined entirely to the Eastern Archi-
pelago, not occurring at all in the Caribbean Sea, where every bidistichate Actinometra
belongs to the protean type of Actinometra pulchella. There may be no palmars at all,
as in Actinometra elongata and Actinometra simplex (PL LVII. fig. 2 ; PL LIX. fig. 1),
or there are three with the axillary a syzygy, as in Actinometra valida, which has a
further division of the same character (PL LIX. fig. 3). This being the best-developed
species of the four members of the group which were obtained by the Challenger, it may
be conveniently called the Valida-gxowp,
5. The Stelligera-grou-p.
Two articulated distichals. The palmars and subsequent series, when present, are of
the same character ; but the first two brachials are united by syzygy.
Remarks. — This is a very well defined group, although its type of structure is
extremely anomalous and does not occur at all in Antedon, all the bidistichate species
of which have the first two brachials articulated, whereas they are united by syzygy in
Actinometra stelligera and its allies (PL LVIII. fig. 1).
304 THE VOYAGE OF H.M.S. CHALLENGER.
This type has normally one and sometimes two post-distichal axillaries ; while there
are three in an undescribed species (Actinometra nigra) which was brought by Professor
Semper from the Philippines. On the other hand there are none in Actinometra
maculata (PI. LV. fig. 2), and this is usually also the case in Actinometra pulchella,
though palmars may be occasionally present (PI. LII. fig. 1). This remarkable type
occurs on both sides of the Atlantic, and possibly also in the Eastern Archipelago ; but
all the remaining members of the group are limited to the latter region and to the
Western Pacific. Two of them, Actinometra maculata and Actinometra stelligera, were
obtained by the Challenger, but the group is not represented in the " Alert " collection,
though Dr. Coppinger dredged in two localities, Bowen and Torres Strait, where
examples of it had previously been obtained.
The following scheme shows the mutual relations of the four species belonging to the
Stelligera-giovLY> : —
A Arm-joiuts triangular and of moderate length, . . . .1. pulchella, Pourtales, sp.
B. Arm-joints relatively short, becoming rather saucer-shaped.
I. The first, and parts of the second radials concealed.
a. No post-distichal axillaries. The later cirrus-joints bluntly
spinous, . . . . . . .2. maculata, n. sp.
6. One or two post-distichal axillaries. The cirri scarcely
spinous, . . . . . . .3. stelligera, n. sp.
II. The first radials partly visible, and the second entirely so. Three
post-distichal axillaries, ...... nigra, Semper, MS.
1. Actinometra pulchella, Pourtales, sp. (PI. IV. figs. 5, a-c; PI. LII. figs. 1, 2).
Specific formulas — a.y.; and — a.2.(2)."2.-,-.
1878. Antedon alata, Pourtales, Bull. Mus. Comp. Zool., 187S, vol. v. No. 9, p. 215.
1878. Antedon pulchella, Pourtales, Ibid., p. 216.
1881. Actinometra pulchella, P. H. Carpenter, Ibid., 18S1, vol. ix. No. 4, p. 10.
1882. Antedon alata, Bell, Proc. Zool. Soc. Lond., 1882, p. 532.
1882. Actinometra pulchella, Bell., Ibid., p. 535.
1882. Actinometra -pulchella, P. H. Carpenter, Ibid., p. 717.
1884. Actinometra pulchella, P. H. Carpenter, Proc. Roy. Soc. Edin., 1884, vol. xii. p. 369.
Localities. — H.M.S. " Porcupine " : Station 25, July 27, 1870 ; near Cape St. Vincent ;
lat. 37° 11' N., long. 9° 7' W.; 374 fathoms; rock; bottom temperature, 53°-5 F. One
mutilated specimen.
Station 31, August 1870; lat. 35° 56' N., 7° 6' \V.; 477 fathoms; clay; bottom
temperature, 50° "5 F. One mutilated specimen.
H.M.S. Challenger: August 1873, St. Paul's Rocks. One specimen.
Station 192, September 26, 1874; near the Ki Islands; lat. 5° 49' 15" S, long.
132° 14' 15" E.; 140 fathoms ; blue mud. One doubtful specimen.
REPORT ON THE CRINOIDEA. 305
Other Localities.— S.S. "Dacia," 1883; lat. 34° 57' N., Long. 11° 57' W.; 533
fathoms.
The "Talisman," off Rochefort ; 1500 metres.
The Caribbean Sea ; abundant from 73 to 278 fathoms.
Remarks. — This singular species will be fully described and it3 variations illustrated
in the Report on the Comatulse of the " Blake " dredgings. It was first obtained by the
"Porcupine" in 1870, though I never saw the type till 1883, nearly five years after
it had been described by Pourtales from the dredgings of the " Hassler " expedition
at Barbados in 1872; and the Challenger had taken it at St. Paul's Rocks in the
following year. The " Hassler" specimens were described by Pourtales under the specific
name alata ; but at the same time he described an apparently different form from an
unknown Caribbean locality as Antedon pulchella ; l and when I subsequently found
reason, after examining the rich material obtained by the "Blake" in 1878-79, to unite
the two forms under one specific name,2 pulchella seemed more appropriate than alata.
I therefore described the type as Actinometra pulchella. It has been found at over
thirty localities in the Caribbean Sea, ranging from 73 to 278, and possibly to 380,
fathoms ; while it presents a very singular instance of dimorphic specific characters.
Some individuals have ten arms, each with a syzygy in the third brachial ; but others
have twenty, with two articulated distichals and the first two brachials united by syzygy.
The " Blake " material contains numerous intermediate conditions between these two
extremes, e.g., individuals with twelve or fifteen arms, owing to the distichal series only
occurring on some of the rays. The Challenger specimen from St. Paul's Rocks has
twenty arms, with its full complement of ten distichal series. In the figured " Porcupine "
example, however, there are but nine distichal axillaries ; so that the number of arms
would only be nineteen, but for the presence of a single palmar axillary, which brings
the total up to twenty (PI. LII. fig. 1).
This species is often an extremely difficult one to make out, owing to the obscurity
of the syzygial union between the first two brachials, as long as the arms remain whole ;
but when they drop away and the syzygial faces are exposed there can be no mistake
about the characters of the type. In some cases they have broken at the syzygy in the
third brachial ; though this is not always a syzygial joint, except perhaps in the two
outer arms of the ray.
The Challenger's discovery of this species at St. Paul's Rocks extended its geographical
range very considerably, and lias probably also brought its bathynietrical range up to less
than 70 fathoms, the specimen having been obtained at some depth between 10 and 80
fathoms. In like manner the presence of this species among the " Porcupine " collection,
from 374 and 477 fathoms near the entrance to the Straits of Gibraltar, brings it into the
1Bull. Mus. Comp. Zo'61., 1878, vol. v. No. 9, pp. 215, 216. -Ibid., 1881, vol. ix. No. 4, p. 10.
(ZOOL. CHALL. EXP. — PART LX. 1888.) Ooo 39
306 THE VOYAGE OF H.M.S. CHALLENGER.
European fauna, and adds nearly 200 fathoms to its bathymetrical range, which is
still further increased to 533 fathoms by the dredgings of the telegraph ship " Dacia,"
some four degrees more to the west. Judging from a figure published in La
Nature1 I imagine that it was also obtained by the "Talisman" in 1500 metres off
flochefort.
Among the numerous Comatulas which were dredged at Station 192 in the Arafura Sea
was a single mutilated specimen which has given me very great trouble (PI. LII. fig. 2).
Three of the rays which are preserved have bidistichate series, and the first two brachials
above the axillaries are clearly united by syzygy, the radiating ridges being very distinct
on the exposed distal faces of two of the first brachials. But I have had much difficulty
in determining the nature of the union between the two outer radials and the two
distichals respectively ; and after repeated changes of opinion, I have come to the con-
clusion that there is a bifascial articulation in each case. The specific formula thus
becomes the same as that of Actinometra pulchella, and in the absence of better
preserved material it seems best to refer the individual in question to this protean species.
The eastern form has fewer cirrus-joints, with larger and blunter spines than may occur
in the Caribbean type ; and the characters of the lower pinnules do not seem to be quite
the same in the two cases. But I have been unable to make out any differences which
would serve to separate the two forms specifically, though it is quite possible that they
may reveal themselves when better preserved material is examined. On the other
hand there is no a priori reason why Actinometra pulcliella, which occurs on both sides
of the Atlantic, should not also inhabit the Eastern seas. Another common Caribbean
species, Antedon carinata, is widely distributed through the Indian Ocean and also
occurs in the Pacific ; while Antedon quinquecostata, which was dredged by the
Challenger at Station 192, together with the doubtful form under consideration, is very
closely allied to the Caribbean Antedon spinifera.
On the whole, then, it appears most probable that the specimen obtained by the
Challenger in the Arafura Sea really does belong to Actinometra pidchella, though one
would like to see a more perfect specimen before definitely making such a large addition
to the geographical range of the Caribbean type. It is also possible, on the other hand,
that we are here dealing with a varietal form of Actinometra macidata from Torres
Strait (PI. LV. fig. 2) ; but I rather doubt this being the case, as its arm-joints are
relatively longer than those of that type, and the terminal cirrus-joints are more com-
pressed laterally. The Copenhagen Museum contains a form from Bowen with very
much the same characters, which bears the MS. name Actinometra fusca, Liitken. This
may be either Actinometra pidchella or Actinometra macidata, but the question of its
specific identity must be left for a future decision.
1 See H. Filhol, Explorations sons-marines, La Nature, 1884, 12 Ann. p. 32!).
REPORT ON THE CRINOIDEA. 307
2. Actinometra maculata, n. sp. (PI. V. figs. 1, a-cl ; PL LV. fig. 2).
opecijic jormula — a. 2. -.7.77.
Centro-dorsal a wide disk bearing twenty-five to thirty-five cirri of about twenty
joints. A few of the lower joints are longer than wide, but the remainder are short,
laterally compressed, and bluntly spinous.
The first radials are concealed, and also part of the second, which are closely united
laterally. Two distichals, the axillary not a syzygy.
Twenty arms, of about one hundred and twenty shortly triangular joints, which
overlap slightly and are sometimes almost saucer-shaped. The first two brachials
are united by syzygy, and there may be another in the third brachial. The next
is about the tenth or twelfth joint, and others follow at intervals of two or three
joints.
The second brachial has a pinnule some 15 mm. long, with a well-defined terminal
comb ; and the length gradually decreases to the pinnules of the eighth or tenth brachials,
which are stouter, but not specially short, and have no comb.
The two basal joints of the first four pinnules on each side are more or less carinate.
Mouth radial ; disk naked.
Colour in spirit, — dark reddish-brown, somewhat mottled with patches of yellowish-
green.
Disk 12 mm.; spread 14 cm.
Locality. — Station 186, September 8, 1874 ; Prince of Wales Channel ; lat. 10° 30' S.,
long. 142° 18' E.;, 8 fathoms ; coral mud. Two specimens, one much mutilated.
Remarks. — This elegant species differs from Actinometra 'ptdcliella in the shortness
of its arm-joints, a character in which it resembles Actinometra stelligera (PL LV. fig. 2 ;
PL LVTII. fig. 1). It is possibly identical with Lutken's MS. species, Actinometra
fusca, from Bowen, in the Copenhagen Museum.
The calyx has many resemblances to that of the allied species Actinometra stelligera
(PL V. figs. 1, 5, a-cl). In both cases the centro-dorsal is wider than the radial pentagon,
so as to conceal part of the second radials (figs, la, 5a), while the basal star is very well
defined (figs. \c, 5cl). The under surface of the centro-dorsal in Actinometra maculata
is marked by five indistinct elevations which correspond in position with the low ridges
beneath the basal grooves in its internal cavity.
308 THE VOYAGE OF H.M.S. CHALLENGER.
3. Actinometra stelligera, n. sp. (PI. V. figs. 5, ci-cl; PI. LVIII. figs. 1,2; also Part
I. pi. lvi. fig. 8).
br he
Specific formula — a. 2. 2. (2 ). -77. — r .
Actinometra tenax, Lutken, MS., Museum Godeffroy.
1880. Actinometra stelligera, P. H. Carpenter, Journ. Linn. Soc. Lond. (Zool)., 1880, vol. xv.
pi. xii. fig. 26.
Centro-dorsal a wide and rather thick disk bearing some thirty marginal cirri, with
twenty joints, a few of the lower ones being longer than wide. From the twelfth
onwards they are wider than long, sometimes with slight indications of a blunt dorsal
spine, which is more marked on the penultimate.
First radials concealed, and also part of the second, which are closely united laterally.
Two distichals, two palmare, and sometimes two post-palmars ; each division of two
joints, the axillary not a syzygy.
Thirty to forty arms, consisting of about one hundred and twenty slightly over-
lapping triangular joints, which are much wider than long, especially in the middle and
outer parts of the arms.
The first two brachials are united by syzygy ; and in the two outer arms of each ray
the third brachial is generally a syzygial joint. The next syzygy is in the tenth or
twelfth brachial, and others follow at intervals of two or three joints.
The second brachial has a pinnule about 16 mm. long, with a well-defined terminal
comb ; and the length gradually decreases to those of the eighth and ninth brachials,
which are short and have no comb. The two basal joints on the pinnules of the third
and the six or seven following brachials are more or less distinctly keeled.
Mouth generally radial ; the anal area often rather thickly plated.
Colour in spirit, — reddish or blackish-brown.
Disk 15 mm.; spread 18 cm.
Locality. — Station 174b, c, or d, August 3, 1874; near Kandavu, Fiji; lat. (about)
19° 6' S., long, (about) 178° 18' E.; 255, 610, or 210 fathoms;1 coral mud; bottom
temperature at 610 fathoms, 39° F. Seven specimens.
Other Localities. — Tonga ; Fiji ; Samoa ; Reef of Atagor (Jukes).
Remarks. — I believe this fine species to be identical with the type which has been
distributed by the Godeffroy Museum under the name of Actinometra tenax, Liitken,
but I did not discover the fact till after some of the plates illustrating its structure had
been lettered and printed off. The name which it now bears relates to the appearance
of the basal star, which stands out in white from a brownish background when the
centro-dorsal is removed from the radials. The ends of the star sometimes appear
1 The exact station, and consequently the exact depth, are not recorded.
REPORT ON THE CRINOIDEA. 309
externally between the angles of the radials, as seen in PI. V. fig. 5b. The radials
somewhat resemble those of Actinometra maculata, in not completely covering the
centro-dorsal (PI. V. figs, la, 5a) ; but the ventral pair of muscle-fossee on their
articular faces is even more reduced than in that type (PI. V. figs, lb, 5b, 5c). The two
species are closely allied, however, and may eventually prove to be connected by inter-
mediate forms. Actinometra stelligera is the larger of the two, and has a greater
number of arms, palmars being always developed, and sometimes post-palmars also ;
while there are no post-distichal axillaries in the two examples of Actinometra maculata,
which also has rather more spinous cirri. The two species further present the same sort
of difference in the carination of the basal pinnules as occurs between Actinometra
Solaris and Actinometra pectinata. In Actinometra maculata there may be keels on
the pinnules of the second to ninth brachials, whereas in Actinometra stelligera there
is no sign of carination on the basal joints of the first pinnule, though there may be
on that of the fifth on the same side (10th br.).
Reversions to the more normal type of arm-structure sometimes occur. Thus, for
example, the outer arm of the right hand ray in the figured specimen of Actinometra
stelligera (PI. LVIII. fig. 1) has the first two brachials articulated like the radials and
distichals ; whereas in the other arm borne on the same distichal axillary, and in three
similar arms of the centre ray, these two joints are united by syzygy. Two curious
abnormalities of the disk have also come under my notice. In one case there are two
mouths and two anal tubes, as shown in Part I. pi. lvi. fig. 8 ; while in the other the
anal tube is close up to the peristome, a little to one side of the median line, and not
central as is usually the case.
The depth at which this species was dredged is not known with certainty ; but it
was probably either 210 or 255 fathoms, the third depth at this locality being an
improbable one for an Actinometra, especially as the type belongs to the littoral fauna
at Fiji, Samoa, and Tonga. There is a closely allied, if not identical, species from Zebu
in the Museums at Dresden and Vienna. Semper's Philippine collection also contains a
fine species belonging to this group, which differs from Actinometra stelligera in the
presence of a third axillary beyond the distichals, and in the relatively smaller size of
the centro-dorsal, so that not only the second radials, but also portions of the first, are
visible externally. It is the type to which I have occasionally referred as Actinometra
nigra, Semper, MS., and is remarkable for the great development of the branches of the
axial cords of the arms and of the par-ambulacral network which is connected with them
in the ventral perisome, and also for the large size of the radial blood-spaces beneath
the ambulacra, the existence of which in Antedon rosacea has recently been denied by
Messrs. Vogt and Yung.1 Figures illustrating these points were given in Part I. pp. 121,
122, and pi. lxi. fig. 6.
1 Traite d'Anatomie compare pratique, 1886, Livr. vii. p. 538.
310 THE VOYAGE OF H.M.S. CHALLENGER.
6. The Valida-grorxp.
Two articulated distichals ; the first arm-syzygy in the third brachial.
Remarks. — Two somewhat different types of structure are comprised in this group.
viz., forms with two palmars like the distichals, and forms with three palmare of which
the axdlary is a syzygy. In addition to these there are also the species, like Actinometra
elongata and Actinometra simplex, which have normally no palmar series at all (PI. L VII.
fig. 2 ; PI. LIX. fig. 1). With one exception, which is in the National Collection, these
are the only species of the genus which have such a simple ray-structure ; and I do not
know of any other form which has its subsequent arm-divisions of the same character as
the distichals. This is in remarkable contrast to the number of Antedon-species which
have the same general formula and belong to the Spinifera- and Palmata-gvoups.
On the other hand a few Actinometra-sipecies like Actinometra rotalaria and Actino-
metra valida have one or more arm-divisions beyond the distichal axillary, each consisting
of three joints with the axillary a syzygy, an arrangement which does not occur in
Antedon. Some of the individuals which have been distributed by the Godeffroy Museum,
under Liitken's MS. names Actinometra intricate!, and Actinometra trachygaster, are of
this character ; but other specimens bearing the same names are tridistichate, and therefore
resemble Actinometra parvicirra (PI. LXI. figs. 1, 5). The two types are so intimately
connected, however, that it is impossible to consider them as representing separate
groups. Thus, for example, I have described examples of Actinometra parvicirra in
which half the distichal series consisted of two, and the other half of three joints ; and a
specimen in the Vienna Museum presents a similar variation. Then again, two (or more)
three-jointed distichal series occur in the unique specimens of Actinometra elongata and
Actinometra valida, and in the figured one of Actinometra rotalaria (PL LVII. fig. 2 ;
PI. LIX. figs. 2, 3) ; while in the two last two-jointed palmar series may also present
themselves as a variation on the normal three-jointed type.
Under these circumstances it is clear that these variations in structure are not
morphologically equivalent to the changes in the position of the arm-syzygies which
characterise the Stelligera- and Fimbriata- groups (PI. LVIII. fig. 1 ; PI. LXII. fig. 3), the
former having a syzygy betiveen the first two brachials, while the latter has a syzygy in
the second brachial ; and until the discovery of hitherto unknown species renders the
number of forms comprised in the P~a^'<ia-group much more considerable than it is at
present, we shall do best to include in it all those bidistichate species which have the first
arm-syzygy in the third brachial, whether the palmar series consists of two or of three
joints.
All the members of the group, as at present constituted, are confined exclusively to
the Eastern Archipelago, including the Fiji and the Friendly Islands.
EEPOET ON THE CEINOIDEA. 311
The following key shows the mutual relations of the species described in this
Report : —
A. No post-distichal axillaries.
First radials visible ; arm-joints relatively long, . . . . 1. elongata, n. sp.
First radials concealed ; arm-joints short and wide, . . . .2. simplex, n. sp.
B. Three palmars, the axillary a syzygy.
No post-palmars ; ten cirri of ten or twelve joints, . . , .3. rotalaria, Lam., bd.
Post-palmars like palmars ; fifteen cirri of fifteen joints, . . . i. ralida, n. sp.
1. Actinometra elongata, n. sp. (PI. LVII. figs. 2-4).
Specific formula — a. 2
Description of an Individual. — Centro-dorsal a small thin disk, bearing about ten
cirri of twelve or fourteen joints, a few of which are longer than wide. Three radials
visible ; the second partly united laterally, the remainder of the rays being well separated.
Two distichals, the axillary without a syzygy.
Eighteen arms, which are all tentaculiferous, but dimorphic. The anterior arms taper
slowly, reaching 11 cm. in length, and consist of one hundred and twenty quadrate
segments, the middle and later ones of which are very long. The posterior arms reach
only 4'5 cm., and taper rapidly, with about fifty-five shorter but still quadrate joints.
A syzygy in the third brachial ; the next between the sixth and tenth, with others at
intervals of about three joints.
The pinnules diminish in length from the first one on the second brachial, which
reaches 8 mm., to those of the fifth and sixth, and then increase again, becoming very
long and slender at the ends of the arms. The first six or eight have a slight terminal
comb, which occurs at intervals to far out on the arm. The later pinnules of the
posterior arms have " ovoid bodies " on their dorsal edge.
Mouth nearly radial ; disk naked.
Colour in spirit, — -greenish grey.
Disk 11 mm.; spread nearly 20 cm.
Locality. — Banda ; October 1, 1874.
Remarks. — This is a singular type in many ways. It differs altogether from the
majority of species of Actinometra in the great length of its arm-joints, which is
especially evident in the longer anterior arms (PL LVII. fig. 2) ; though the joints of the
posterior arms are also relatively long. The only form which comes at all near it in this
respect is the tridistichate Actinometra quadrata (PI. LXII. fig. 1). The great difference
in length between the anterior and posterior arms is the more remarkable, as they are all
tentaculiferous, none of them being unprovided with an ambulacral groove, as is so often
312 THE VOYAGE OP H.M.S. CHALLENGER.
the case. But the distal pinnules in at least five of the posterior arms are provided with
the curious ovoid bodies on their dorsal aspect which I have noticed in some forms of
Actinometra parvicirra.1 In the latter type the pinnules which bear these bodies are
generally non-teutaculiferous ; but this is not the case in Actinometra elongata.
The centro-dorsal of this form is very thin, with much reduced cirrus-sockets, and is
evidently in process of transformation into the P/iano^enia-condition shown on the same
plate (PI. LVII. figs. 1, 2). One of the arms has been broken at the syzygy in the third
brachial, and the new epiz)'gal is an axillary, as is so frequently the case. This fact may
possibly indicate that when pah ars are developed in this type, there are normally three
with a syzygy in the axillary, so that it would then be allied to Actinometra rotalaria
and Actinometra valida (PL LIX. figs. 2, 3).
The terminal comb on the oral pinnules is rather a small one, but it may occur at
intervals to some way out on the arms. The disk is very large and prominent, without
any trace of calcareous deposits, and the radial position of the mouth is not very distinct
(PI. LVII. fig. 3).
2. Actinometra simplex, n. sp. (PI. LIX. fig. 1).
Specific formula — a. 2.—.
Description of an Individual. — Centro-dorsal a thin disk bearing about fifteen
marginal cirri with fourteen to seventeen segments, a few of which are longer than broad,
First radials concealed, and also portions of the second, which are partly united laterally.
Two distichals, the axillary without a syzygy.
Eighteen arms ; the anterior with one hundred joints, as compared with forty-five in
the posterior arms, some of which are non-tentaculiferous. The joints are short, sub-
triangular, and slightly overlapping, becoming more elongated at the ends of the anterior
arms. A syzygy in the third brachial, and the next about the tenth brachial, with
others at intervals of two to four joints.
The second brachial bears a pinnule about 7 mm. long, and the following pinnules
diminish to those on the fifth and sixth brachials, afterwards increasing again. The
terminal pinnules of the anterior arms are very long and slender, those of the posterior
arms being shorter and stouter. The first four pinnules on each side have a small
terminal comb, which is found at intervals till near the ends of the arms.
Mouth interrarlial; a few calcareous granules on the disk.
Colour in spirit, — the skeleton a dull green, and the ventral perisome deep brown.
Disk 8 mm.; spread 9 cm.
Locality. — The Admiralty Islands; 16 to 25 fathoms. One specimen.
1 Trans. Linn. Hoc. Lond. (Zool.), ser. 2, 1879, vol. ii. p. 40, pL ii. fig- 6, o.b.
REPORT ON THE ORINOIDEA. 313
Remarks. — This is a curious little species, which differs altogether from Actinometra
elongata in the shortness of the arm-joints and in the non-appearance of the first radials
externally. It has many resemblances to Actinometra parvicirra, but is separated from
that type by its smaller number of distichal joints. It presents, however, the same
difference in the lengths of the anterior and posterior arms as occurs both in Actinometra
parvicirra and in Actinometra elongata; but some of the hinder arms are non-tenta-
euliferous, which is not the case in Actinometra elongata. Their distal pinnules may
have dark spots in the centre of the dorsal surface which appear to be rudimentary forms
of the "ovoid bodies" that occur in Actinometra parvicirra and Actinometra elongata.
They are comparatively small and insignificant, and do not occur on the pinnules of the
anterior arms.
3. Actinometra rotalaria, Lamarck, sp. (PI. LIX. fig. 2).
Specific fo rmula — a. 2.3.—.
1816. Comatula rotalaria, Lamarck, Histoire Naturelle des Animaux sans Vertebres, Paris,
1816, t. ii. p. 534.
1834. Comatula rotularia, de Blainville, Manuel d'Actinologie, Paris, 1834, p. 249.
1841. Aledo rotalaria, Miiller, Monatsber. d. k. preuss. Akad. d. Wiss. Berlin, 1841, p. 184.
1843. Aledo rotalaria, Miiller, Archiv f. Naturgesch., 1843, Jalirg. ix. Bd. i. p. 136.
1849. Comatula (Actinometra) rotalaria, Miiller, Abhandl. d. k. Akad. d. Wiss. Berlin, Jalirg.
1847 [1849], p. 256.
1862. Comatula rotalaria, Dujardin and Hupe, Hist. Nat. des Zoophytes, lichinodermes, Paris,
1862, p. 204.
1879. Actinometra rotalaria, P. H. Carpenter, Trans. Linn. Soc. Lond. (Zool.), ser. 2, 1879,
vol. ii. p. 27.
1882. Actinometra rotalaria, Bell, Proc. Zool. Soc. Lond., 1882, p. 535.
1882. Actinometra rotalaria, P. H. Carpenter, Ibid., p. 747.
Centro-dorsal a small, thin disk, bearing about ten cirri, of ten or twelve joints, none
(if which are much longer than wide.
First radials just visible ; the second closely united laterally. Two distichals, the
second axillary without a syzygy, and three palmars, the third axillary with a syzygy,
Twenty to thirty arms, of about eighty subtriangular and overlapping joints ; some of
the hinder arms may be non-tentaculiferous.
Syzygies in the third, tenth, and fourteenth segments, with others at intervals of
three or four joints.
The second palmar, when present, has a moderately long pinnule with rather stout
lower joints. The next pinnule is nearly as long, but that of the third brachial is much
smaller ; and the next pair are also small, after which the pinnules increase considerably
in both length and stoutness. The terminal comb is rather small, and does not extend
beyond the sixth brachial.
(zool. CHALL. EXP. PART LX. — 1888.) Ooo 40
3U THE VOYAGE OF H.M.S. CHALLENGER.
Mouth apparently radial ; disk naked.
Colour in spirit, — light brownish- white.
Disk G'5 mm.; spread 8 cm.
Locality. — Samboangan ; 10 fathoms. Two specimens.
Other Localities. — Australia (Peron and Lesueur).
Remarks. — I believe these two specimens to be identical with the form to which
Lamarck gave the name Comatula rotalaria. According to Midler's diagnosis,1 "Die
Radien bestehen aus 2 durch Syzygie verbundenen Gliedern. Auf diese folgen
uumittelbar wieder d ia, die wieder mit Syzygie versehen sind. Dann folgt nur
noch selten weitere Veiasteluug, also 20 Arme die Grundzahl."
When I visited the Paris Museum in 1876 I found that it contained no specimen
bearing Lamarck's name, but that a form which had been brought from Australia by
Peron and Lesueur, and appeared to be the original type of Lamarck's species as redefined
by Muller, was labelled Comatula brevicirra, Troschel.
The first radials are not very distinct, but they are undoubtedly present, and there is
no syzygy either between the two outer radials, or between the two distichal joints, as
described by Muller according to Troschel's diagnosis ; while the two palmar series which
are present each consist of three joints, the axillary with a syzygy.
The two individuals obtained by the Challenger at Samboangan present the same
characters and also retain their cirri, which are lost in the Lamarckian type. As i3
sometimes the case in Actinometra pectinata, there are five pairs which are placed
interradially or nearly so (PI. LIX. fig. 2), and have only ten or twelve joints. Tri-
distichate se.ies occur abnormally in both examples, whde there are sometimes only
two palmars instead of three.
The only type which resembles Actinometra rotalaria in the characters of its arm-
divisions is Actinometra valida (PI. LIX. fig. 3), which is altogether a larger form with
more arms and more cirri. Actinometra simplex is also bidistichate but has no palmars,
while the cirri are longer and more numerous (PL LIX. fig. 1).
4. Actinometra valida, n. sp. (PI. LIX. fig. 3).
Specific formula — a. 2. 3. 3.—.
Description of an Individual. — Centro-dorsal a thin circular disk, bearing about
fifteen cirri, which have some fifteen tolerably uniform joints ; the terminal ones laterally
compressed with a faint dorsal spine.
First radials just visible ; the second closely united laterally. The rays are wide, and
1 Abhandl. d. k. Akad. d. Wus. Berlin, Jahrg. 1847 [1849], p. 256.
REPORT ON THE CRINOIDHA. 315
the adjacent distichal series are in close contact. The distichals, palmars, and lower
brachials have rather flattened sides. Two distichals without a syzygy, three palmars,
and sometimes three post-palmars, the axillary with a syzygy.
Forty-six arms of subtriangular and somewhat overlapping joints, which become more
discoidal towards the middle of the arms and squarer towards the ends ; one hundred and
twenty joints in the anterior, and eighty in the posterior arms.
Syzygies in the third, tenth, and fourteenth brachials, and afterwards at intervals of
three or four joints.
The second joints after the distichal and subsequent axillaries bear long and rather
stout pinnules, the first one reaching nearly 25 mm. The pinnule of the third brachial
is smaller than that on the second, but the following ones are stouter, with rather large
joints. The terminal comb is small and much obscured by perisome.
Mouth radial ; disk naked, with several non-tentaculiferous arms.
Colour in spirit,— dark greyish-green.
Disk 21 mm.; spread probably 22 cm.
Locality.— Station 186, September 8, 1874 ; Prince of Wales Channel ; lat. 10° 30' N.,
long. 142° 18' E.; 8 fathoms; coral mud. One specimen.
Remarks. — This is a fine individual which is allied to Liitken's MS. species Actino-
metra trachygaster, and Actinometra intricata from Fiji, Tonga, and Samoa. I propose
to describe these at some future time, when it will be necessary to fix their characters
more precisely ; for I have seen specimens bearing these names which do not altogether
correspond with Liitken's types in the Copenhagen Museum. Actinometra valida is
much larger than Actinometra rotalaria, having an additional axillary, and also larger
and more numerous cirri ; while the rays are wide and generally in close lateral contact,
the sides of their lower joints being somewhat flattened, though much less so than in
Antedon. At first sight there appears to be no terminal comb on the lower pinnules.
This is due to its being obscured by the thickness of the perisome, but it becomes more
apparent in the dry state, though it is nothing like so well developed as in many smaller
individuals of other species.
Actinometra, Series IV.
Three distichals, the first two articulated, and the third axillary with a syzygy.
Remarks. — More than half the described species of Actinorru tra belong to this
series, which, both in the abundance and in the variety of its specific forms, presents a
very strong contrast to the corresponding series in Antedon. The articulation of the
two outer radials, as compared with their syzygial union in the Typica-gvoxnp, which is
also tridistichate, is associated with the fact that the first two joints beyond the distichal
316 THE VOYAGE OF H.M.S. CHALLENGER.
and all subsequent axillaries are articulated, and not united by syzygy as in Actino-
metra typica and its allies. Furthermore, the first syzygy in the free arms is not
between the first two brachials, as in Actinometra typica and Antedon inwqualis, but
in the second brachial as in Actinometra Jimbriata and Actinometra multiradiata
(PI. LXII. fig. 3 ; PL LXVI. fig. 1), or in the third as in Actinometra parvicirra and
Actinometra divaricata (PI. LXI. figs. 1,5; PL LXIII. fig. 6).
The numerous species of this series thus fall into two very well defined groups, each
of which contains forms with no axillary beyond the distichal, and others with two or
sometimes with three.
The first arm-syzygy in the second brachial, . . . . . .0. Jimbriata.
The first arm-syzygy in the third brachial, . . . . . .7. parvicirra.
7. The Fimbriata-grou]}.
Tridistichate species with a pinnule on the first brachial and a syzygy in the second.
The palmar and post-palmar series, when present, consist of two joints, the first bearing
a pinnule, and the second axillary with a syzygy.
Remarks. — The position of the first brachial syzygy in this group is altogether an
anomalous one. In ordinary Comatulee the third and fourth joints of the primitive arm
become closely united by suture, eventually forming a syzygy, while the pinnule of the
former remains undeveloped, like that of the first brachial. But in the Fimbriata-
group the first joint above the distichal and every subsequent axillary, whether it be a
palmar or a free brachial, bears a pinnule ; and the syzygial union occurs between the
primitive second and third brachials, instead of between the third and fourth (PL LX.
fig. 1 ; PL LXII. fig. 3), i.e., there is a syzygy in the second instead of in the third
brachial of the mature arm. When, however, there are no distichals, so that the arms
spring directly from the radial axillary, we usually find a reversion to the more primitive
type, with a syzygy in the third brachial, i.e., the epizygal of this syzygy bears a single
arm, instead of being an axillary (PL LX. fig. 2 ; PL LXVI. fig. 1). I have very rarely
met with any instance of a pinnule on the first and a syzygy in the second brachial
above a primary radial axillary. As a general rule this arrangement only occurs after
a secondary or tertiary axillary (i.e., distichal or palmar).
The Fimbriata-gmu]) is only represented in the genus Antedon by a single species,
Antedon porrecta. It includes a considerable variety of specific forms, all of which,
with two exceptions, are limited to the Indian Ocean, the Eastern Archipelago, and
the North-west Pacific. Actinometra lineata and Actinometra discoidea were dredged
by the "Blake" in the Caribbean Sea, while the former was likewise found by the
Challenger at Bahia. It possibly ranges down to 88 fathoms, while Actinometra
REPOftT ON THE CRINOIDEA. 317
discoidea occurs between this depth and 118 fathoms. But all the remaining species of
the group belong to the purely littoral fauna of the Eastern seas. One of them is the
only described Actinometra which I have not personally examined. It is the Actino-
metra borneensis of Grube, whose type-specimen has disappeared since his death, and I
am, therefore, uncertain about its proper place in the following scheme.
A. Arm-joints short and discoidal, . . . . . . 1. fimbriata, Lamarck, sp.
B. Arm-joints shortly triangular, becoming more quadrate or discoidal.
I. No palmars.
a. Less than twenty-four cirrus-joints, . . .2. coppingeri, Bell.
b. More than twenty-four cirrus-joints, . . . borneensis, Grube.
II. Two palmars, the axillary a syzygy.
a. Less than thirty cirrus-joints ; no post-palmars, . . 3. mulliradiata, Linn., sp.
b. More than thirty cirrus-joints ; post-palmars, like the palmars, 4. sentosa, n. sp.
C. Arm-joints triangular, nearly as long as wide, . . . .5. lineata, n. sp.
D. Arm-joints almost quadrate, . . . . . .6. discoidea, Carpenter, MS.
1. Actinometra fimbriata , Lamarck, sp. (PI. LXII. figs. 2—4).
Specific form ula — a. 3. 2br. —r .
1816. Comatula fimbriata, Lamarck, Hist. Nat. des Animaux Bans Vertebres, Paris, 1816, t. ii.
p. 534.
1S34. Comatula fimbriata, de Blainville, Manuel d'Actinologie, Paris, 1834, p. 249.
1841. Atecto fimbriata, Midler, Monatsber. d. k. preuss. Akad. d. Wiss. Berlin, 1841, p. 185.
1843. Atecto fimbriata, Midler, Archiv f. Naturgesch., 1843, Jahrg. ix. Bd. i. p. 136.
1849. Comatula (Aleeto) fimbriata, Miiller, Abhandl. d. k. Akad. d. "Wiss. Berlin, Jahrg. 1817
[1849], p. 258.
1862. Comatula fimbriata, Dujardin and Hupe, Hist. Nat. des Zoophytes, tfchinodennes, Paris,
1862, p. 204.
1879. Actinometra fimbriata, P. H. Carpenter, Journ. Linn. Soc. Lond. (Zool.), 1877, vol. xiii.
p. 443.
1879. Actinometra fimbriata, P. H. Carpenter, Trans. Linn. Soc. Lond. (Zool.), ser. 2, 1879, p. 27.
18S2. Actinometra fimbriata, Bell, Proc. Zool. Soc. Lond., 1882, p. 535.
1882. Actinometra fimbriata, P. H. Carpenter, Ibid., p. 747.
Centro-dorsal a moderately thick disk bearing fifteen or eighteen marginal cirri of
twenty to twenty -four joints, the lower ones longer than wide, and the later joints more
or less spinous, sometimes considerably so.
First radials barely visible ; the second short, wide, and closely united laterally.
Three distichals, the axUlary a syzygy.
Eighteen to twenty arms, which are all tentaculiferous and sometimes dimorphic,
the anterior having one hundred and seventy joints as compared with one hundred and
fifty in the posterior arms. The joints are all short and wide, with nearly transverse
articulations, so as to become almost oblong after about the thirtieth, and generally
318 THE VOYAGE OF H.M.S. CHALLENGER.
overlapping a little. The later joints become more square, and finally somewhat
elongated. A syzygy in the second brachial, except after the radial axillary, when it is
in the third ; the next from the thirteenth to twenty-sixth, usually about the sixteenth
brachial ; others at intervals of three to twelve joints, generally six or seven.
The second distichal and the first brachial bear tolerably equal pinnules about 10
mm. long, the first one being a little stouter at the base. Their lowest joints may be
slightly carinate. The next pair are somewhat shorter and the following pair more so.
The lowest pinnules have a fairly large terminal comb, which occurs on all the pinnules
as far as the tenth brachial and sometimes even to the twentieth or thirtieth.
Mouth radial ; the disk may have a few calcareous nodules.
Colour in spirit, — blackish- or reddish-brown.
Disk 14 mm.; spread 25 cm.
Localities. — Banda, 17 fathoms. Three specimens.
Station 208, January 17, 1875; lat. 11° 37' N., long. 123° 31' E.; 18 fathoms; blue
mud. One specimen.
Other Localities. — Sunda Strait (Regnault) ; Australian Seas (Peron and Lesueur) ;
Angio, Java ; Nicobar Islands ; Madagascar (?).
Remarks. — The Lamarckian type of this species is a dry specimen with twenty arms
which was brought by Peron and Lesueur from the Australian Seas ; but the name
Comatula Jimbriata was also applied by J. S. Miller to the common ten-armed Antedon
of Milford Haven, which is usually called Antedon rosacea. Johannes Miiller examined
Lamarck's original in the Paris Museum, where he also found three spirit specimens pre-
senting the same characters which had been obtained by Regnault in 1S29. Miiller gave
Trincomalee as the locality for this form ; ! but when I visited the Paris Museum in 1876
I found it labelled as having come from Sunda Strait. It bore the MS. name Comatula
brevicirra, Troschel ; while Peron's example, the type of the species, still bore the same
designation, Comatula multiradiata, Lamarck, as it did when Miiller examined it in 1844.
The later cirrus-joints of this specimen bear several small spines on their dorsal border. But
they are much more distinct in some cirri than in others ; while in Regnault's specimen
they are of smaller size and appear on fewer joints. In the Challenger individual from
the Philippines there is a small spine at the distal edge of the fifth cirrus-joint ; and in the
following joints it gradually develops into a crest bearing a variable number of spinelets,
which sometimes give rise to a double opposing spine on the penultimate. Two of three
forms from Banda have a similar armature on the cirri ; but in the third there is little or
no trace of it (PI. LXII. fig. 3). This species appears to be one in which palmare are not
developed, so that the number of arms does not exceed twenty, and may be less. The
latter condition is unusual, however, distichal axillaries being generally developed all
1 Abhandl. d. k. Akad. d. Wiss. Berlin, Jahrg. 1847 [1849], p. 258.
REPORT ON THE CRINOIDEA. 319
round the calyx, though four of them are missing in one of Reguault's specimens. The
Philippine example presents a curious abnormality, the second distichal of one ray being
axillary, though not a syzygy ; while one of its two arms having been broken and
regenerated has developed a palmar series. But with this exception, I have never seen
any specimen which presents the general characters of Actinometra jimbriata and possesses
palmar series. Its most important distinctive character is the shape of the lower brachials.
The first half-dozen joints are nearly oblong in outline, as in almost all Comatulse ; but
their successors do not become triangular or quadrate as is generally the case. For they
remain short and wide, with nearly equal sides, so that their ends are much less oblique
than usual (PL LXII. fig. 3). It is this character more especially which distinguishes
Actinometra Jimbriata from Actinometra coppingeri and Actinometra midtiradiata (PI.
LX. figs. 1,2; PI. LXVI. fig. 1). By the twenty-fifth brachial, or sooner, the joints are
almost perfectly oblong, and they remain as thick disks till near the end of the arm, where
they become squarer and finally slightly elongated. The joints of the middle and lower
parts of the arms overlap one another to a greater or less extent, and their edges are
fringed with small spines ; but there is much variation in both characters.
This thickly discoidal shape of the arm-joints appears to be their highest form of
development. A study of regenerated arms of different sizes shows that the joints are
at first elongated as they are in the Pentacrinoid, and that their gradual increase in width
makes them at first quadrate, then triangular, and finally more or less distinctly oblong,
this being the shape which is characteristic of the Pentacrinidae and of many fossil
Crinoids. We may perhaps say then that Actinometra cop>pingeri and Actinometra
multiradiata, with their more triangular joints at the bases of the arms (PI. LX. figs.
1, 2; PL LXVI. fig. l), are permanently immature forms of Acti, w met ra Jimbriata
(PL LXII. fig. 3), standing to it in the same relation as Antedon quadrata to Antedon
eschrichti.
The mouth of Actinometra Jimbriata is radial, being usually distinctly excentric. and
sometimes quite close to the margin of the disk, the anal tube being central or nearly so
(PL LXII. fig. 4), while the hinder ambulacra embrace it in a horseshoe-like curve.
But in the Philippine specimen the mouth is almost central, the anal tube greatly
reduced, and the ambulacra grouped like those of Antedon. The two primary ambulacra
of the B ray are separately connected with the peristome, the outer one supporting but a
single arm, as distichals are undeveloped, while the posterior one is connected to the
peristome by a short trunk which is common to it and to the single groove that supplies
the whole of the postero-lateral ray C (PL LXII. fig. 2).
The lower pinnules of this individual have somewhat carinate basal joints, but the
extent of the carination varies greatly, and it seems to be almost entirely absent in one
of the Banda specimens, though present in the others. It occurs in a form from
Madagascar, which, so far as I can judge from my notes of its other characters, appears
320 THE VOYAGE OF H.M.S. CHALLENGER.
to be identical with Actinometra jimbnata, as I have redefined it above. This example
was brought to the Paris Museum by Rousseau in 1841, and I found it bearing the
museum name Comatula coccodistoma ; but it differs from the other examples of Actino-
metra Jimbriata that I have seen in the interradial position of the mouth. The Copen-
hagen Museum contains three specimens which also seem to belong to this type. Two
are from Angio in Java, and have overlapping arm -joints ; while the third was obtained
by the " Galathea " at the Nicobar Islands, and has a less marked overlap.
2. Actinometra coppingeri, Bell (PI. LX. figs. 1, 2).
1882. Actinometra coppingeri, Bell, Proc. Zool. Soc. Lond., 1882, p. 535.
1882. Actinometra coppingeri, P. H. Carpenter, Ibid., p. 747.
1884. Actinometra coppingeri, Bell, Bep. Zool. Coll. H.M.S. "Alert," London, 1884, p. 168,
pi. xvi. ng. B.
Sjjccific formula — a. 3. 2br.-^.
Centro-dorsal a wide flat plate, bearing some fifteen to twenty-five marginal cirri of
fifteen to twenty-two tolerably uniform joints, with traces of double dorsal spines.
Second radials closely united laterally and only partially visible. Three distichals, the
third axillary with a syzygy.
Twelve to twenty arms, the fifth and following joints almost triangular, much wider
than long, gradually becoming blunter and more cmadrate. The first syzygy is usually
in the second brachial, but is in the third if the arm springs directly from the radial
axillary. The next may be between the fifth and fourteenth brachials ; and others
follow at intervals of three to nine, usually four or five joints.
The second distichal bears a pinnule about 1 5 mm. long ; that on the first brachial
is slightly shorter, and its successors diminish gradually to about the third pair. The
terminal comb may stop at the eighth brachial or go on to the fifteenth.
Mouth radial ; all the arms tentaculiferous. A few calcareous granules on the disk.
Colour in spirit, — reddish- or blackish-brown.
Disk 16 mm.; spread 15 cm.
Localities. — Banda, 17 fathoms. One specimen.
Samboangan, 10 fathoms. One specimen.
Other Localities. — H.M.S. " Alert," Flinders, Clairmont ; Singapore; Amboina ; the
China Sea.
Remarks. — The essential difference between this species and Actinometra Jimbriata
lies in the more triangular shape of its arm-joints, which become quadrate and eventually
cuboid, but are never so nearly oblong as is the case in Actinometra jimbriata (PI. LX.
figs. 1,2; PI. LXII. fig. 3). The two Challenger specimens from Banda and Samboangan
REPORT ON THE C'RINOIDEA. 321
respectively differ somewhat in their characters, and I was at first inclined to regard
them as specifically distinct ; but they are linked together by another form from
Singapore, which was kindly given to me by my friend Professor Charles Stewart.
I cannot separate these three specimens from the type which was described by Bell
as Actinometra coppvngeri} It is represented by a single individual with twelve arms,
owing to the presence of two distichal axillaries ; and as one of these is clearly due to
regeneration at the syzygy in the third joint above the radial axillary, Bell was to a
certain extent justified in saying that the normal number of arms " is probably ten."
The epizygal of this syzygy may, however, have been an axillary originally, and the
second axillary, which is figured by Bell, is so well developed that I believe it to be a
normal one. Furthermore, in all the four arms borne upon these two distichal axillaries,
whether regenerated or not, the first brachial bears a pinnule, and the second is a
syzygial joint. These characters escaped the notice of Bell, whose figure is incorrect,
as it shows a pinnule on the second brachial and a syzygy in the third joint above the
distichal axillary. On the strength of this figure I assigned a place to Actinometra
coppingeri in the Parvicirra-group, and gave a different name to the Challenger species.
But when I came to examine Bell's type for the purpose of determining its relations to
Actinometra parvicirra, I was surprised to find it identical with the form which I had
been accustomed to call Actinometra steivarti ; so that it adds another to the list of
species which were dredged both by the " Alert" and by the Challenger.
The arms are largest in the Challenger individual from Samboangan, but its cirri
are considerably smaller than in the other two, especially in that from Singapore, which
approaches it most nearly in the characters of the arms. The latter also has the longest
lower pinnules, and the terminal comb may extend to nearly the twentieth brachial ; while
it is rarely found after the eighth brachial in the Samboangan form which has twenty
arms. That from Singapore has eighteen, and the Banda one only fourteen, as three of
the rays have no distichals at all, and the first syzygy is therefore in its normal position
in the third brachial (PI. LX. fig. 2).
The museums at Berlin and Copenhagen each contain a specimen which I believe
to belong to this type. There are not more than eighteen cirrus-joints, as in the examples
from Banda and Singapore ; though that from Samboangan may have twenty or twenty-
two. This limitation in the number of cirrus-joints in specimens from five different
localities seems to indicate that the type is not identical with Actinometra borneensi.s,
Grube, which has twenty-two to twenty-eight joints.2 Grube also says of the arm-joints
" Die Glieder sind etwas klirzer als breit, und laufen nur anfangs in leichten Zick-Zack
weiterhin parallele." His type specimen has unfortunately disappeared. Professor
Schneider has been unable to find it at Breslau, and it is equally unknown at Berlin,
1 "Alert" Report, p. 168, pi. xvi. fig. B.
2 53e Jahresber. der Schlesisch. Gesellsch. f. Voted. Cult, 1875, p. 75.
(ZOOL. CHALL. EXP. —PART LX. — 1888.) OoO 41
322 THE VOYAGE OF H.M.S. CHALLENGER.
where some of his other types are. I imagine, however, that it differs both from
Actinometra jimbriata and from Actinometra coppingeri, the middle arm-joints becoming
more oblong, and not remaining quadrate as in the latter type ; while the obliquity of
their surfaces in the lower joints seems to separate this form from Actinometra Jimbriata,
which it rather resembles in the number of its cirrus-joints. But there is a difficulty
in coming to a satisfactory conclusion about this point in the absence of Grube's type-
specimen.
3. Actinometra multiradiata, Linn., sp. (PL LXVI. figs. 1-3).
Specific Jormula — a.3.2.[p.(p').6r.]. — .
1758. Asterias multiradiata, Linnaeus, Systema Naturae, ed. 10, Holrnise, 1758, t. ii. p. 663.
1783. Asterias multiradiata, Retzius, K. Svensk. Vetensk. Akad. Handl., Ar 1783, t. iv. p. 241.
1788. Asterias multiradiata, Linnaeus, Systema Naturae, ed. 13, Lipsire, 1788, pars vi. p. 3166.
1805. Asterias multiradiata, Retzius, Dissertatio, sistens species cognitas Asteriarum, Lundee,
1805, p. 35.
1816. Comatula multiradiata, Lamarck (pars), Histoire Naturelle des Animaux sans Vertebres,
Paris, 1816, t. ii. p. 533.
1834. Comatula multiradiata, de Blainville (pars), Manuel dActinologie, Paris, 1834, p. 249.
1843. Asterias multiradiata, Miiller, Archiv f. Naturgesch., 1843, Jahrg. ix. Bd. i. p. 133.
1849. Comatula (Alecto) multiradiata, Miiller (pars), Abhandl. d. k. Akad. d. Wiss. Berlin,
Jahrg. 1847 [1849], p. 261.
1862. Actinometra multiradiata, Dujardin and Hupe, Hist. Nat. des Zoophytes, Echinodermes,
Paris, 1862, p. 210.
1879. Actinometra multiradiata, P. H. Carpenter, Trans. Linn. Soc. Lond. (Zool.), ser. 2, 1879,
vol. ii. p. 27.
1882. Actinometra multiradiata, P. H. Carpenter (pars), Journ. Linn. Soc. Lond. (Zool.),
1882, vol. xvi. p. 521.
1S82. Actinometra multiradiata, Bell, Proc. Zool. Soc. Lond., 1882, p. 533.
1882. Actinometra multiradiata, P. H. Carpenter (pars), Ibid., p. 747.
Centro-dorsal a thick circular disk, often hollowed in the centre, and bearing fifteen
to twenty stout marginal cirri of twenty-two to twenty-six joints. The basal ones are
very broad, the sixth and seventh longer than wide, and from the tenth onwards the
joints bear dorsal spines.
The ends of the basal rays are more or less visible. The first radials are almost
entirely concealed and sometimes parts of the second, which are imperfectly united
laterally. Three distichals, the third axillary with a syzygy ; two palmars, the second
axillary with a syzygy. Post-palmars, resembling the palmars, are but rarely present.
Eighteen to twenty-four arms, of about one hundred and twenty to one hundred
and fifty short overlapping joints, which are triangular at the arm-bases, but become
discoidal towards the middle of the arm ; their distal margins are very spinose.
A syzygy in the second brachial ; the next between the fifteenth and fortieth, with
others at intervals of four to nine joints.
REPORT ON THE CRINOIDEA. 323
The second distichals, first palmars (when present), and first brachials bear long-
tapering pinnules ; the first one reaching 18 mm., while the others are rather smaller.
That of the third brachial is considerably so, and the next three pinnules are of
decreasing size. The lowest pinnules have a well-marked comb, which may extend out
to the twentieth or twenty-fifth brachial. The basal joints of the lower pinnules may
be somewhat carinate, and in the following pinnules the edges of the joints project
laterally.
Mouth radial ; disk naked, or with scattered calcareous nodules.
Colour in spirit, — blackish-brown ; the disk sometimes mottled with white.
Disk 19 mm.; spread reaching 25 cm.
Locality.— Station 186, September 8, 1884 ; Prince of Wales Channel ; lat. 10° 30' N.,
long. 142° 18' E.; 8 fathoms ; coral mud. One specimen.
Other Localities. — Indian Seas (Linnaeus) ; Australian Seas (Peron and Lesueur) ;
Sumatra ; Bohol ; China Sea ; Kagoshima Bay, Japan.
Remarks. — The type of this species is a dry and somewhat mutilated Actinometra
in the Retzian collection at Lund, on which Linnaeus seems to have based his brief
description of Asterias multiradiata} He also referred to it the Caput-Medusse
cinereum and the Caput-Medusse brunnum of Linck ; but the exact specific relations of
these two forms must remain uncertain, as Linck's figures are not sufficiently clear for
the characters of their arm-divisions to be made out.
Retzius gave a more detailed description of the original type of Asterias multi-
radiata in 1783,2 stating the number of arms as thirty to forty, and that of the cirrus-
joints as twenty-three. He noticed it again in 1805 ;3 while in 1816 Lamarck
established the species Comatida multiradiata, i under which he placed Asterias
multiradiata, Linn., with a (?) appended. He described it as having fifty to sixty,
or even more arms, and referred to the Indian seas as its locality. Some years later
Goldfuss 5 applied the name Comatida multiradiata, Lamarck, to a many-armed
specimen, the distichal and palmar series of which each consisted of three joints, with the
axillary a syzygy. Midler,6 regarding this form as " die zuerst genau beschriebene,"
proposed in 1841 to retain the specific name multiradiata for it alone, and on the
basis of Troschel's examination of the Paris collection, he published a description of
Comatida multiradiata, Lamarck, under the name of Alecto multiftda. He distinguished
this type from that of Goldfuss by its palmar and post-palmar series each consisting of
but two joints, with the axillary not a syzygy. He went to Sweden, however, in the
1 Systema Nature, ed. 10, Holrnia;, 1758, t. ii. p. 663.
2 K. Svensk. Vetensk. Akad. Handl., Ar 1783, t. iv. p. 241.
» Dissertatio, sistens species cognitas Asteriarum, Lundoe, 1805, p. 35.
4 Hist. Nat. des Anim. sans vertebres, Paris, 1816, t. ii. p. 533.
6 Petrefacta Germanise, t. i. p. 202, pi. Isi. fig. 2. ° Monateber. d. h. irreuss. Akad. d. Wiss. Berlin, 1841, p. 188.
324 THE VOYAGE OF H.M.S. CHALLENGER.
same year, and examined at Lund the original of Asterias midtiradiata, Linn. This
he found to have a pinnule on the first and a syzygy in the second joint above the
distichal and palmar axillaries, i.e., there are two palmars, with the axillary a syzygy.
He gave a careful description of this form,1 to which, after his visit to Paris he added
some details derived from his personal examination of some examples collected by Peron
and Lesueur and by Quoy and Gaimard. His final diagnosis was headed Oomatula
(Alecto) multiradiata, Nobis ; 2 though, as we have already seen, he had referred the
Retzian specimen to the type of his new genus Actinometra. Dujardin and Hupe
described it under the latter name,3 entirely on the basis of Miiller's diagnosis of it ;
but they made no mention of the specimens obtained by Peron and Lesueur and by
Quoy and Gaimard, which resemble the Retzian individual in having syzygies in all the
axillaries.
I have already separated off one of these forms as Actinometra peroni,4 owing to
its palmar series consisting of three joints, instead of only two as in the Retzian type,
which has no post-palmars and not more than twenty -five cirrus-joints. One of Peron's
specimens presents the same characters as Asterias midtiradiata, and I have since met
with a considerable number of similar individuals. But the spirit-specimen brought
from the Moluccas by Quoy and Gaimard, which was referred by Midler and afterwards
by myself5 to Actinometra midtiradiata, must, I think, be separated from this species
on account of its larger number of cirrus-joints, and more numerous arms, owing to the
presence of post-palmar series.
Two examples of it were obtained by the Challenger at Banda, and wUl be
described immediately as Actinometra sentosa (PL LX VI. fig. 4).
I have had some doubts as to the projmety of separating Actinometra coppingeri
from Actinometra midtiradiata, the chief difference between the two being the absence
of palmars in the former and their presence in the latter. The character seems to be a
fairly constant one, however, as the two forms have not hitherto been found associated
together in one locality. Actinometra coppingeri is known from East Australia,
Singapore, Amboina, Banda, the China Sea, and Samboangan ; while palmars occur in
three examples of Actinometra midtiradiata from Bohol, another Philippine locality,
in two from Japan, in one from Sumatra, and in one from Torres Strait. It is a
generally more robust form than Actinometra coppingeri, with the lower brachials
relatively shorter and more overlapping ; while the spines on the cirri are of a much
more definite character than in that species. The second syzygy also is much further
from the calyx than in Actinometra coppingeri, especially in the Philippine examples of
Actinometra midtiradiata and in the Retzian type, in which last it may not occur till
the thirty -ninth brachial.
1 Archivf. Naturgesch., 1843, Jahrg. ix. Bel. i. p. 133. a Abhandl. d. k. Akad. d. Wiss. Berlin, 1847 [1849], p. 261.
3 Op. cit., p. 210. * Notes from the Leyden Museum, 1881, vol. iii. p. 214.
5 Journ. Linn. Soc. Lond. (Zool.), 1882, vol. xvi. p. 523.
REPORT ON THE CRINOIDEA. 325
The Hamburg Museum contains the fragmental remains of a dry specimen from
Sumatra which seems to belong to Actinometra multiradiata. There are only twenty
to twenty-five cirrus-joints, but a single post-palmar series is present in one ray. This,
however, is the only example which I have seen that has a post-palmar axillary as in
Actinometra sentosa, with the cirri of Actinometra multiradiata; and the additional
axillary is not improbably due to regeneration, as is so often the case.
Two very fine examples of this type, with somewhat smoother arms than usual, were
dredged by Dr. Doderlein in Japan. Apart from their large size, they are also remark-
able for the peculiar mottled appearance of the disk, which is naked, and without the
calcareous concretions that occur in the examples from further south and in the original
type of the species.
The basal star seems to be pretty well developed in Actinometra multiradiata.
Nearly all the specimens of it which I have seen show more or less indication of the
star between the centro-dorsal and the radials. None of them have any non-tentaculi-
ferous arms, and there appears to be no great difference in length between those coming
off from opposite poles of the disk.
Except for the presence of palmar axillaries, the three individuals from Bohol
correspond very well with Midler's description of Comatula Jimbriata ; and they were
referred to that species by Professor Semper, who found them to be the hosts of
Myzostoma lobatum, von Graff.1 I have since found one Myzostoma in the pharynx of
one of these individuals, its edge being just visible through the mouth.
4. Actinometra sentosa,, n. sp. (PI. LXVI. figs. 4-6).
1849. Comatula (Alecto) multiradiata, Miiller (pars), Abhandl. d. k. Akad. d. Wiss. Berlin,
Jahrg. 1847 [1849], p. 261.
1882. Actinometra multiradiata, P. H. Carpenter (jmrs), Journ. Linn. Soc. Lond. (Zool.), 1882,
vol. xvi. p. 521.
1882. Actinometra multiradiata, P. H. Carpenter (pars), Proc. Zool. Soc. Lond., 1882, p. 747.
Specific formula — a. 3. 2. (p.p'. °r)-^-
Centro-dorsal a thick disk, sometimes almost columnar, with the dorsal pole partially
hollowed, and bearing twenty to thirty moderately stout marginal cirri. These have
twenty-six to forty joints, of which the fifth is usually longer than wide, and the next
two or three the longest, least markedly so in the older cirri ; the later joints are nearly
square and somewhat compressed laterally, small spines appearing near their distal edges,
which increase in distinctness up to the penultimate joint.
First radials visible, least so in the larger specimens. The second partly united
1 See von Graff, Das Genus Myzostoma, Leipzig, 1877, p. 19, and also Zool. Chall. Exp., part xxvii. p. 57, 1884.
326 THE VOYAGE OF H.M.S. CHALLENGER.
laterally ; the rays and their subdivisions are well separated from one another. Three
distichals, the axillary a syzygy. Palmar and post-palmar series of two joints, the
axillary with a syzygy. Forty to sixty-five arms, of one hundred and twenty to one
hundred and fifty joints, the first few nearly oblong ; the following ones overlapping and
shortly triangular, with coarsely spinous distal edges. From about the fortieth onwards,
the joints become more oblong, as the arms narrow, and their terminal joints are squarer.
The anterior arms may be slightly the longer.
A syzygy in the second brachial ; the next from the fifteenth to thirtieth, usually
about the twentieth, with others at intervals of four to eight, usually five or six, joints.
The pinnules on the second distichals are nearly 30 mm. long, and moderately stout
at the base, but soon become more slender. The following pinnules are on the first
joints after each axillary, and the length decreases to those of the fifth and sixth
brachials which are not specially small. Their successors increase again slowly. The
lowest pinnules have a large terminal comb, which may extend out to the fifteenth
brachial ; and the edges of the pinnule joints are fringed with spines.
Mouth radial or nearly so ; disk naked or with a few calcareous nodules.
Colour in spirit, — blackish-brown.
Disk 15 mm.; spread 25 cm.
Locality. — Banda ; two specimens.
Other Localities. — Moluccas (Quoy and Gaimard).
Remarks. — This fine species cannot wTell be confounded with any other Actinometra,
the only form which at all approaches it being Actinometra multiradiata, in which,
however, there are normally no post-palmars, while the cirri do not have more than
twenty-six joints.
I have only seen three specimens of Actinometra sentosa, one which was brought
from the Moluccas to the Paris Museum by Quoy and Gaimard, and the two dredged at
Banda by the Challenger. The Paris specimen was referred by Midler1 to the type of
Asterias multiradiata, Linn., his final diagnosis of the species differing but little from
his previous description of the Retzian type, except that he gave the number of cirrus-
joints as twenty to thirty instead of simply twenty-four ; while he described forty to
fifty arms, instead of thirty to forty, the number assigned by Ketzius. The latter
change involved the presence of post-palmar axillaries, to which, however, Muller made
no reference.
I was at first inclined to follow Midler's example, and to describe the two Challenger
individuals under the name Actinometra multiradiata ; 2 but I have since examined a
greater variety of specimens, and have come to the conclusion that the larger number
1 Abhandl. d. h. Akad. d. Wiss. Berlin, Jahrg. 1847 [1849], p. 261.
3 Journ. Linn. Soc. Loud. (Zool.), 1882, vol. xvi. p. 521.
REPORT ON THE CRTNOIDEA. 327
of cirrus-joints and the additional axdlary together constitute a good specific character.
Palmars occur on every ray of each of the two Challenger specimens, so that the number
of arms reaches twelve or sixteen to each ray ; while in Actinometra multiradiata there
are not usually more than six. Some of the cirri in both individuals have thirty joints
or more, though the number may fall to twenty-six in cirri that are apparently mature ;
and on the other hand there may be as many as forty joints. The Paris specimen has
about thirty.
5. Actinometra lineata, n. sp. (PI. V. figs. 2, a-e ; PI. LX. fig. 3).
1879. Antedon sp., Rathbun, Trans. Connect. Acad., 1879, vol. v. p. 157.
1880. Actinometra lineata, P. H. Carpenter, Journ. Linn. Soc. Lond. (Zool.), 1882, vol. xv.
p. 213, pi. xii. figs. 27, a, b.
1882. Actinometra lineata, P. H. Carpenter, Proc. Zool. Soc. Lond., 1882, p. 747.
Specific formula — a.3.2.[(p).&r.].— .
Centro-dorsal discoidal, bearing twenty to thirty marginal cirri. These have eleven
to seventeen joints, usually not more than fourteen, several of which are longer than wide,
the later joints overlapping dorsally.
The first radials are usually concealed, together with more or less of the second,
which may or may not be united laterally. Three distichals, the axillary with a syzygy,
and sometimes two palmars, the axillary with a syzygy. The perisome between the rays
is occasionally plated as far as the distichal axillary.
Eighteen to thirty-four arms, the lower joints triangular and overlapping, but little
wider than long ; the middle joints more quadrate, and the later ones elongated.
A syzygy in the second brachial, and the next between the ninth and twelfth ; others
at intervals of one to five, usually three or four, joints.
The distichal pinnule reaches nearly 15 mm. long, with a large terminal comb. The
next pinnule is but little smaller ; but the size decreases considerably after the pinnule
on the second brachial, till the third or fourth on the same side. The following pinnules
increase slowly in length, becoming very long and slender in the terminal third of the
arm. The first five or six brachial pinnules are sometimes webbed by perisome for about
one-third of their length and have a small comb, which does not usually extend further,
though it may occur as far out as the eighteenth brachial. The basal joints of the lower
pinnules are sometimes slightly carinate.
Mouth variable in position ; a few of the hinder arms may be non-tentaculiferous.
Disk naked, or bearing a few scattered grains.
Colour in spirit, — reddish or yellowish -brown, with a dark purple medio-dorsal line.
Disk 15 mm.; spread 16 cm.
Locality. — Bahia ; 7 to 20 fathoms. Eight specimens.
328 THE VOYAGE OF H.M.S. CHALLENGER.
Other Localities. — Coast of Brazil; also the "Blake," 1878-79, Station 285, off
Barbados ; 13 to 40 fathoms ; and possibly Station 155, off Montserrat, 88 fathoms.
Remarks. — This Atlantic species may be readily distinguished from its allies of the
Eastern seas by the greater relative length and the more quadrate shape of the arm-
joints, the edges of which are by no means so spiny as in Actinometra Jimbriata,
Actinometra multiradiata, and their allies. The relative shortness of the syzygial
interval and the frequent plating of the interradial perisome are distinctive characters
of minor value.
The position of the mouth in this type seems to be a somewhat variable one. So far
as I have been able to make out, it is radial in the Caribbean variety, but interradial in
the Brazilian form. Neither of the Caribbean individuals that I have seen has any
palmar series and they are sometimes absent in those from Bahia.
The calyx of Actinometra lineata is not unlike that of Actinometra maculata. Iu
both alike the radials fail to cover the centro-dorsal entirely; while their angles are
everted so as to appear beyond its edge (PI. V. figs, la, lb, 2a, 2b); but the latter
character is more marked in the Atlantic species (PI. V. fig. 2d). Figs. 2c and 2b on
PI. V. show the upper and side views of a centro-dorsal, from which three radials have
been removed, so as to expose the rosette and a portion of the basal star.
Closely allied to this species is a very remarkable Actinometra, which was dredged
by the " Blake " in the Caribbean Sea. At first sight it greatly resembles a large
example of Actinometra lineata; but the palmar series are represented by single
axillary joints, and the post-palmars may be of the same character, or there may be two
joints united by syzygy. The second brachial is generally a syzygy on the outer arms
of each ray, and sometimes also on the adradial arm, which is on the inner side of each
distichium. But the other arms generally have the first two joints united by syzygy,
— (o)
so that the specific formula comes to be — a.3.1.1. br . y^. I really cannot tell what to
2 (l)
make of this remarkable form, and should much like to see some more examples of it.
For the present at any rate it may remain in the neighbourhood of Actinometra lineata.
The Comatula with an excentric mouth which was described by Eathbun 1 as
Antedon, sp., from some locality either on the the coast of Pernambuco or of Parahyba
do Norte, is I think identical with Actinometra lineata. But the question is a little
difficult to decide, as he makes no reference to the presence or absence of syzygies in the
distichal and palmar axillaries ; and the position of the first brachial pinnule is not
described very clearly.
1 Trans. Connect. Acad., 1879, vol. v. p. 157.
REPORT ON THE CRINOIDEA. 329
8. The Parvidrra-gcowp.
Tridistichate species, with a pinnule on the second brachial and a syzygy in the
third.
Remarks. — The tridistichate species of Actinometra which have the first arm-syzygy
in the third brachial, make up nearly half the whole number of the species of this genus
which are considered in this Report ; and so far as I can judge from the undescribed
material which I have examined, this proportion is not likely to be greatly affected by
future work. In the genus Antedon, on the other hand, the number of tridistichate
forms is quite small, both the ten-armed and the bidistichate groups containing a large
number of species.
The Parvicirra-group is more widely distributed than any other in the genus
Actinometra; though it does not occur in the Caribbean Sea, as the Fimbriata- and
Echinoptera-groups do. It is represented on the Peruvian coast and at Tahiti, is
abundant at Samoa, Tonga, and Fiji, and extends throughout the Eastern Archipelago to
Japan on the north and the Nicobar Islands on the west, being also represented by one
species on the southern coast of Australia. Actinometra parvicirra itself occurs at
Natal and Simon's Bay ; but I do not know for certain of any Atlantic representative of
the group, though there is possibly one on the Brazilian coast.
A striking feature in some members of this group is the tendency to the development
of two-jointed palmar series, either generally, as in Actinometra divaricata (PI. LXIII.
fig. 6), or on the outer parts of each ray only, as in Actinometra belli, Actinometra
duplex, and Actinometra nobilis (PI. LXIV. figs. 1, 3 ; PI. LXV. fig. 1). In Actinometra
multifida and Actinometra variabilis the palmar and all subsequent divisions are two-
jointed ; but in Actinometra alternans and Actinometra divaricata there are three-
jointed post-palmars, followed in the former case by two joints again, and in the latter
by a three-jointed series (PI. LXIII. fig. 6). On the other hand in Actinometra regalis
(PI. LXVIII. fig. 2), Actinometra bennetti, &c, there are three, or even four, three-
jointed series above the radial axillary, which is a very rare condition in Antedon.
These large multibrachiate species are all confined to the littoral fauna ; but an
example of Actinometra parvicirra with about thirty arms was obtained, together
with the multibrachiate Actinometra typica, from a depth of at least 210 fathoms at
Station 174.
The species of the Parvicirra -group may be classified as follows : —
(ZOOL. CHALL. EXP. — PART LX. 1888.) OoO 42
330
THE VOYAGE OF H.M.S. CHALLENGER.
A. Three distichals, not succeeded by palniars.
I. Ten to twenty cirri, generally with less than fifteen joints.
(a) Arm-joints triangular till some distance from the disk, and con-
siderably wider than long, .....
(b) Arm-joints relatively long, becoming quadrate about the fifteenth,
II. Thirty or more cirri of fifteen to twenty joints.
(a) Arm-joints short; lower joints of distichal pinnules not specially
marked, .......
(b) Arm -joints of moderate length; lower joints of distichal pinnules
rather large and carinate, .....
B. Palmar series developed above the distichals.
I. Two palmars, the axillary not a syzygy.
(a) Post-palmars like palmars.
1. Ten cirri, ......
2. Twenty cirri ; one or two further divisions, like palmars, .
(6) Post-palmar series of three joints, the axillary with a syzygy.
1. Fifty cirri on a hemispherical centro-dorsal,
2. Centro-dorsal reduced, with few or no cirri.
a. Further division like palmars; centro-dorsal stellate,
/S. Further division like post-palmars.
(i.) Fifteen to twenty small cirri, .
(ii.) Centro-dorsal stellate, without cirri.
Rays well separated ; mouth radial, .
Rays closely united, and the interradial peri-
some plated ; mouth interradial, .
II. Palmar series at outside of ray two-jointed, without a syzygy ; the inner
series three-jointed, with a syzygy.
(a) Post-palmars of two joints, the axillary not a syzygy ; pinnule
joints carinate, ......
(6) Post-palmars of three joints, the axillary with a syzygy.
1. Fifteen cirri ; rays quite free, ....
2. No functional cirri ; rays closely united, .
III. Three palmars, the axillary with a syzygy.
a. No further division. Centro-dorsal bears functional cirri.
1. Ten to twenty -five cirri, generally with less than fifteen
joints, ......
2. Thirty or more cirri, of fifteen to twenty joints.
(a) Lower pinnules not specially large.
Arm-joints short ; lower joints of distichal pin-
nules not carinate, ....
Arm-joints of moderate length ; lower joints of
distichal pinnules rather large and carinate, .
(b) Distichal and palmar pinnules very large and stout,
b. Post-palmar series present.
1. No functional cirri ; post-palmars of two joints, the
axillary not a syzygy, ....
2. Cirrus-sockets not entirely obliterated ; post-palmars like
palmars.
(a) Ten to thirty cirri.
(i.) Ten to twenty cirrus-joints.
a. No further division,
1. parvicirra, Mull., sp.
2. quadrata, n. sp.
3. trichoptera, MiilL, sp.
japonim, MiilL, sp.
multifield, MiilL, sp.
variabilis, Bell.
grandicalyx, Carpenter.
alternans, Carpenter.
briareus, Bell, sp.
4. divaricata, n. sp.
magnifica, Carpenter, MS.
5. belli, n. sp.
6. duplex, n. sp.
7. nobilis, n. sp.
1. parvicirra, Mull., sp.
3. trichoptera, MiilL, sp.
japonica, Mull., sp.
robustipinna, Carpenter.
8. littoralis, n. sp.
1. parvicirra, MiilL, sp.
REPORT ON THE CRINOIDEA. 331
(3. A fourth post-radial series of three joints,
the axillary with a syzygy.
First radials largely visible ; arm-joints
of moderate length ; pinnules on
fourth and fifth brachials short, . 9. regalis, n. sp.
First radials mostly concealed; short
arm-joints; pinnules on fourth and
fifth brachials not specially short, . schlegeli, Carpenter,
(ii.) Thirty cirrus-joints ; first radials mostly con-
cealed, ..... peroni, Carpenter.
(b) Forty to fifty cirri of twenty-five joints, . . bennetti, Mull., sp.
1. Actinometra parvicirra, Mull., sp. (PI. LXL; PI. LXVII. figs. 3, 4).
Specific formula — a.3.[3(3)].—.
Remarks. — The synonymy and diagnosis of this remarkable type will be given on a
subsequent page, when the species with palmar series are considered. The simpler forms
of it, with not more than twenty arms, are known from the following localities : —
H.M.S. Challenger: — Simon's Bay; Ternate; Ban da; Admiralty Islands; Prince of
Wales Channel ; Samboangan.
Other Localities. — Cape of Good Hope ; Nicobar Islands ; Timor ; North Borneo ;
Solor ; Ceram ; Bohol ; Port Molle ; Moreton Bay, Fiji ; Peru.
2. Actinometra quadrata, n. sp. (PI. LXII. fig. 1).
Specific formula — a. 3.—.
Description of an Individual. — Centro-dorsal a small thin disk, bearing a single row
of ten marginal cirri, with eleven joints, of which the fourth and fifth are slightly the
longest.
The first radials are largely visible, and the second partly united laterally, the rays
being quite free. Three distichals, the axillary with a syzygy.
Sixteen long and slender arms, consisting of about one hundred and twenty tolerably
smooth joints ; the lower joints triangular and relatively long, soon becoming distinctly
quadrate, then more square, and finally elongated. Syzygies in the third, tenth, and
fourteenth brachials, and then at intervals of two to four joints.
The second distichal bears a pinnule about 8 mm. long, and that on the second
brachial is but little shorter ; but the next pair are considerably so, and the size decreases
to about the sixth brachial, and then increases again, the terminal pinnules becoming
very slender and reaching 12 mm. A terminal comb on the pinnules of the first eight
brachials, and then irregularly till the twentieth.
Mouth interradial, and disk naked ; all the arms are grooved.
332 THE VOYAGE OF H.M.S. CHALLENGER.
Colour in spirit — greyish-green.
Disk 8 mm. ; spread 22 cm.
Locality. — Tongatabu Eeefs.
Remarks. — This is a very elegant species which may be distinguished from Actino-
metra parvicirra by the characters of its arm-joints. The lower joints lose their
triangular shape very soon and become unequally quadrate; the two sides gradually
become more equal untd the outline is nearly square, and finally the joints become
almost cylindrical with slightly oblique ends. The relative length of the lower joints
varies in some of the arms ; that selected by the artist for representation having rather
shorter joints than its fellows.
The small size of the cirri, and their fewness in numbers, will prevent this species
from being confounded with Actinometra trichoptera. Some specimens from the Nicobar
Islands in the museums at Copenhagen and Vienna should perhaps be referred to it on
account of the length of their arm -joints.
3. Actinometra trichoptera (Valenciennes), Mull., sp. (PI. LXIII. figs. 1-5).
be
Specific formula — a. 3.(3)—.
Remarks. — This species, like Actinometra parvicirra, may or may not have palmar
series, and will therefore be considered later. It was obtained by the Challenger at Port
Jackson.
4. Actinometra divaricata, n. sp. (PL LXIII. figs. 6-8).
Specific formula — a.3.2.3.3.— .
Description of an Individual. — Centro-dorsal stellate, without traces of cirri, and a
little below the level of the radial pentagon, the inner sides of which are somewhat cut
away. The second radials are relatively long and incompletely united laterally ; the
rays are quite free and may divide five times.
Three distichals, the axillary with a syzygy ; two palmars without a syzygy ; the
first and second post-palmar divisions, when present, each of three joints, the axillary
with a syzygy.
Arms very numerous, eighteen or twenty to the ray, and all grooved ; but the hinder
arms are only faintly so and are very narrow and short, with one hundred to one hundred
and twenty slightly overlapping joints ; the anterior arms have rather more. The lower
joints are shortly triangular, becoming more oblong, and finally nearly square.
Syzygies in the third, twelfth, and sixteenth brachials, and then at intervals of three
or four joints.
REPORT ON THE CRINOIDEA. 333
The pinnules on the second distichal and post-palmar joints are about equally long,
reaching 12 mm.; but the following pinnules are considerably shorter, diminishing to
that of the third brachial which is the smallest ; after which the length increases
slightly, but the pinnules are always comparatively short. The lower pinnules have a
terminal comb as far the fourth or fifth brachial, and it is continued at intervals to the
eleventh or twelfth.
Mouth radial ; disk naked, except for a few granules near the anal tube.
Colour in spirit, — dark blackish-brown.
Disk 30 mm.; spread 18 cm.
Locality. — Banda; 17 fathoms.
Bemarks. — This species is remarkable for the relatively small size of the arms and
pinnules as compared with that of the disk. Its nearest allies are the Antedon briareus
of Bell and the Philippine species to which I have referred in Part I. as Actinometra
magnified.1 The latter has nearly twice the spread of Actinometra divaricata, with a
relatively smaller disk ; while the rays are in close lateral contact as far as the distichal
axillary, above which the perisome is strongly plated.
Antedon briareus is really an Actinometra, as is shown by the absence of sacculi, and
the presence of a terminal comb on the lower pinnules, two points which seem to have
escaped Bell's notice. According to his description of the species,2 the post-palmar series
resemble the palmars in consisting of but two joints, the axillary without a syzygy.
This would indicate an alliance with Actinometra multifield. Bell's figure shows, however,
that about two-thirds of the post-palmars have three joints, the axillary with a syzygy,
and also that there are four 'cases of a further division, which he does not mention at all.
In one case these second post-palmars consist of two joints, the axillary without a syzygy ;
but the remainder consist of three joints, the axillary with a syzygy, just as in Actino-
metra divaricata and Actinometra magnifica. Pending the discovery of other examples
of this species, therefore, its formula must be — a.3.2.3.3.—, and not — A.2.3.(2).— , as was
assigned to it by Bell.3 Its centro-dorsal is evidently undergoing reduction to the
Phanogenia-condition, but some poorly developed cirri still remain attached to it ; while
in Actinometra alternans, Actinometra divaricata (PI. LXIII. fig. 6), and Actinometra
magnifica it is stellate, with few or no traces of any cirri at all.
We now come to a group of species, which in one respect stand altogether alone in
the whole family of Cornatulge. In each case there are three distichals, the axillary with
a syzygy ; but the two secondary arms borne on each distichal axillary are not alike
when they divide again. That on the outside of the ray has but two palmar joints, the
1 Zool. Chall. Exp., part xxxii. p. 57, pi. lvi. fig. 7. - " Alert " Report, p. 163, pi. xiv. ' Ibid., p. 155.
334 THE VOYAGE OF H.M.S. CHALLENGER.
axillary without a syzygy, while the inner palmar series resembles the distichal one in
consisting of three joints, the axillary with a syzygy. This is shown very well in the
single example of Actinometra duplex, which has no very great number of arms
(PI. LXIV. fig. 3) ; and the fact that this arrangement is not merely an accidental one is
shown by its occurrence in three individuals of Actinometra belli (PL LXIV. fig. 1), and
in six of Actinometra nobilis (PI. LXV. fig. 1), both species having four post-radial
axillaries, and therefore a large number of arms.
5. Actinometra belli, n. sp. (PI. LXIV. figs. 1, 2).
Specific formula — a. 3. -A — \ 2.2.
ab
b-
Centro-dorsal a moderately thick circular disk, hollowed in the centre, and bearing
about fifteen marginal cirri. These are fairly stout, of fifteen to twenty joints, a few of
which are rather longer than wide.
The first radials are partly visible, and the second incompletely united; the rays
are quite separate from one another, but the intervening perisome is regularly plated
as far as the palmar axillary. The rays may divide five times, giving sixty-five
to seventy arms.
Three distichals, the axillary with a syzygy ; palmar series two-jointed without a
syzygy on the outside of the ray, but three-jointed with a syzygy on the inside.
The first and second post-palmar series, when present, are also two-jointed. The
anterior arms are long and slowly tapering, with one hundred and twenty to one hundred
and fifty overlapping joints, which are shortly triangular at the base, becoming quadrate
about the middle, and slightly elongated near the tip. The posterior arms are shorter and
taper more quickly, with only eighty to one hundred joints. A syzygy in the third
brachial ; the next about the tenth or twelfth, with others at intervals of three to six joints.
The distichal pinnule is moderately stout and reaches 20 mm. in length ; the palmar
pinnule on the inside of the ray, and that of the second brachial are nearly as long ;
but that of the third brachial is only half their length and much more slender, while
the next pair are the smallest on the arm. The terminal pinnules are long and slender
on the anterior arms but shorter on the posterior ones. The basal segments of the
genital pinnules have sharp dorsal keels, which are less distinct in the first few pinnules
than in those immediately following. In the anterior arms they are lost after about the
fiftieth brachial, but are traceable to near the end of the posterior arms. The lowest
pinnules have a well-marked comb, which becomes gradually smaller and is lost about
the fifteenth brachial.
Mouth interradial ; the disk bears a variable number of small granules, especially
round the ambulacra ; several of the hinder arms are ungrooved.
REPORT ON THE CRINOIDEA. 335
Colour in spirit, — darkish-brown, with a dark medio-dorsal line ; the pinnules some-
times tipped with green.
Disk 30 mm. ; spread 21 em.
Locality. — Station 186, September 9, 1874; Prince of Wales Channel; lat. 10° 30'
N., long. 142° 18' E. ; 8 fathoms; coral mud. Three specimens.
Remarks. — This fine species, which I have dedicated to the energetic curator of the
National Collection of Echinoderms, is readily distinguished from the two others which
have a similar arrangement of the palmar series, by the fact that its first and second
post-palmar series are only two-jointed ; so that on the outside of the ray there is no
pinnule between that of the second distichal and that on the second joint of the free arm.
The difference in the lengths of the anterior and posterior arms is very considerable ;
and in each of the three individuals all the arms of the D ray are unprovided with
ambulacra, which may also be the case on the adjacent arms of the C and E rays as well.
This is well shown in fig. 2 on PI. LXIV. The C ambulacrum only supplies the anterior
half of the corresponding ray,1 so that all the C2 arms are grooveless. The same is the
case with all the arms of D and of E2, and even with some of those on E1} as the groove
which supplies them suddenly ceases before reaching the level of the palmar axillary.
One rather striking character of this species is the strong carination of the third and
following joints in the pinnules of the tenth brachial and its successors ; and another
peculiarity is the very definite nature of the perisomic plating between the rays. The
two radial axillaries are separated by a well-marked polygonal piece which rests on the
truncated angles of the second radials, and corresponds exactly with the first interradial
piece of the Apiocrinidae.
One of the three individuals of this species was presented by Sir Wyville Thomson to
the Natural History Museum at Stockholm, where I found it in August, 1886. The other
two have yielded seven examples of Myzostoma. I could, however, find none of these
parasites when I first searched for them, and again when I examined the species for
descriptive purposes. But they seem to have become detached at a later period after the
type had been drawn, and they are therefore not yet described.
6. Actinometra duplex, n. sp. (PI. LXIV. fig. 3).
Specific formula— a.3.|/^)3.(3).^- .
Description of an Individual. — Centro-dorsal a rounded and slightly convex disk,
bearing some fifteen marginal cirri which have fourteen to seventeen tolerably uniform
joints. Three radials visible, the second partly united laterally ; the rays are quite free
1 See woodcut, fig. 6 on page 274.
336 THE VOYAGE OF H.M.S. CHALLENGER.
and may divide four, or rarely five, times. Three distichals, the axillary with a syzygy ;
palmars, when present, two-jointed without a syzygy on the outside of the ray, but three-
jointed with a syzygy on the inside. The first and second post-palmars, when present,
also three-jointed, the axillary with a syzygy.
Forty-five arms ; the anterior ones of a hundred and twenty slightly overlapping,
triangular joints which gradually become quadrate ; the hinder arms shorter, with only half
as many joints. Syzygies in the third, twelfth, and sixteenth brachials, and then at
intervals of three joints.
The distichal pinnule is relatively long and stout, reaching 13 mm., and that on the
second post -palmar, which is but little smaller, is nearly twice the length of that on the
second brachial. Those of the next three joints are still smaller, after which the size
again increases, but the terminal pinnules are not specially long. The first few pinnules
have a well-marked comb, which becomes gradually weaker and is lost after the tenth
brachial. Mouth interradial; disk naked; some of the hinder arms have very faint
grooves and others none at all.
Colour in spirit. — light reddish-brown.
Disk 15 mm.; spread 21 cm.
Locality. — Band a, 17 fathoms. One specimen.
Remarks. — This is an elegant little species, which differs altogether from Actinometra
belli in having its post-palmar series like the distichals instead of being only two-jointed
(PL LXIV. figs. 1, 3). They occur on all the rays but one, and in the anterior half of
the ray regeneration has taken place to such an extent that there is a fifth post-radial
axillary. As this is probably not the normal condition I have put brackets round the
figure which indicates it in the specific formula.
The great difference in length between the anterior and posterior arms of this type is
very striking, the more so as most of the hinder arms have grooves, though only faint
ones. The five hinder arms of the E ray are, however, altogether devoid of ambulacra, as
the groove which should supply them suddenly stops quite short on the disk at the base
of the distichium ; and the right or western curve of the horse-shoe passes by them
altogether, on its way to the posterior or D ray.
7. Actinometra nobilis, n. sp. (PI. LXV.).
2
Specific formula — a. 3. g-.( — J. 3. 3. -y.
Centro-dorsal a thin disk, with about ten marginal cirri in immature individuals ;
more or less stellate and rather below the level of the radial pentagon in the adult.
Second and third radials short and closely united laterally. The two first distichals of
REPORT ON THE CRINOIDEA. 337
each ray are also closely united laterally ; those of adjacent rays are sometimes united
all round the calyx, and sometimes separated by a strong interradial plating which
extends to about the level of the palmar axillaries.
The rays may divide five times ; three distichals, the axillary with a syzygy ; palmars
two-jointed without a syzygy on the outside of the ray, but three-jointed with a syzygy
on the inside. First and second post-palmars three-jointed, the axillary with a syzygy.
Eighty to one hundred arms ; the anterior ones long and slowly tapering, of one
hundred and fifty to two hundred slightly overlapping joints, which remain almost
triangular till near the end ; the posterior arms tapering rapidly, with eighty to one
hundred more quadrate joints. A syzygy in the third brachial ; the next from the tenth
to the seventeenth, with others at intervals of three to five joints.
The distichal pinnule is moderately stout, reaching 30 mm. in length. Those on the
second joints of the following arm-divisions gradually decrease in size, but that of the
second brachial is only half as long as its predecessor, and the next two pairs of pinnules
are not much smaller. The terminal pinnules are much longer in the anterior than in
the posterior arms. The terminal comb of the lower pinnules is variable, being sometimes
small and ceasing about the tenth brachial, and sometimes much larger, extending out to
the fortieth joint.
Mouth interradial ; the ventral surface of the disk is usually naked, except for a Uttle
plating round the peristome. The hinder arms are mostly without ambulacra, and in
one case, at least, there are ungrooved arms on each ray.
Colour in spirit, — dull green, either alone or mottled with purple, brown, and white.
Disk 50 mm.; spread 30 cm.
Localities. — Station 208, January 17, 1875 ; Philippine Islands; lat. 11° 37' N.,long.
123° 31' E.; 18 fathoms; blue mud. One specimen.
Samboangan ; 10 fathoms. Five specimens.
Remarks. — This is a large and finely developed species, which differs from Actino-
metra duplex in the normal presence of a fifth post-radial axillary, and in the large
amount of perisomic plating between the rays and their subdivisions. It is further
distinguished by the characters of the centro-dorsal, which does not bear comparatively
stout cirri, as in Actinometra duplex (PI. LXIV. fig. 3), but almost reaches the condition
seen in Actinometra typica (PL EVIL fig. 1). The details of the process, which are
illustrated on PI. LXV. figs. 2-7, were explained on p. 15.
The five specimens obtained at Samboangan are generally very similar in their
characters ; but that from Station 208 is more uniformly coloured and has a much larger
terminal comb, which extends to the fortieth brachial instead of ceasing about the tenth
as in the Samboangan form, to which I referred as Actinometra dissimilis on pp. 110,
111, of Part L; for I did not then consider them as specifically identical with the type
(ZOOL. CHALL. EXP. PART LX. — 1888.) OoO 43
338 THE VOYAGE OF H.M.S. CHALLENGER.
from Station 208. Both in the latter individual and in one of those from Samboangan
the disk bears cysts of Myzostoma platypus, which open into the ambulacral grooves, as
seen in PI. LXV. fig. 8.
1. Actinometra 'parvicirra, Mull., sp. (PL LXI. ; PI. LXVII. figs. 3, 4).
Specific formula — a. 3. [3. (3)]—.
1841. Alecto parvicirra, Miiller, Monatsber. d. k. preuss. Akad. d. Wiss. Berlin, 1841, p. 185.
1841. Alecto timorensis, Miiller, Ibid., p. 186.
1843. Alecto Wahlbergii, Miiller, Arcliiv f. Naturgesch., 1843, Jahrg. ix. Bd. i. p. 131.
1849. Comatula (Actinometra) Wahlbergii, Miiller, Abhandl. d. k. Akad. d. Wiss. Berlin, Jahrg.
1847 [1849], p. 256.
1849. Comatula (Alecto) parvicirra, Miiller, Ibid., p. 260.
1849. Comatula timorensis, Miiller, Ibid., p. 263.
1862. Comatula parvicirra, Dujardin and Hupe, Hist. Nat. des Zoophytes, Echinodermes,
Paris, 1862, p. 206.
1862. Comatula timorensis, Dujardin and Hup6, Ibid., p. 206.
1862. Actinometra Wahlbergii, Dujardin and Hupe, Ibid., p. 211.
1875. Comatula mertensi, Grube, 53e Jahresber. der Schlesisch. Gesellsch. f. Vaterl. Cult.,
1875, p. 74.
1876. Actinometra (Comatula) armata (Semper, MS.), P. H. Carpenter, Journ. Anat. aud
Physiol., 1876, vol. x. p. 582.
1877. Actinometra polymorpha, P. H. Carpenter, Journ. Linn. Soe. Lond. (Zool.), 1877, vol. xiii.
p. 443.
1879. Actinometra parvicirra, P. H. Carpenter, Trans. Linn. Soc. Lond. (Zool.), ser. 2, 1877
[1879], p. 27.
1879. Actinometra Wahlberghii, P. H. Carpenter, Ibid., p. 27.
1879. Comatula timorensis, P. H. Carpenter, Ibid., p. 29.
1879. Actinometra polymorpha, P. H. Carpenter, Ibid., p. 51.
1881. Actinometra parvicirra, P. H. Carpenter, Notes from the Leyden Museum, 1881, vol.
iii. p. 204.
1882. Actinometra parvicirra, P. H. Carpenter, Journ. Linn. Soe. Lond. (Zool.), 1882, vol.
xvi. p. 519.
1882. Actinometra meijeri, P. H. Carpenter, Ibid., p. 525.
1882. Antedon mei-tensi, Bell, Proc. Zool. Soc. Lond., 1882, p. 533.
1882. Actinometra parvicirra, Bell, Ibid., p. 535.
1882. Actinometra wahlbergi, Bell, Ibid., p. 535.
1882. Actinometra annulata, Bell, Ibid., p. 535.
1882. Actinometra annulata, P. H. Carpenter, Ibid., p. 747.
1882. Actinometra parvicirra, P. H. Carpenter, Ibid., p. 747.
1882. Actinometra wahlbergi, P. H. Carpenter, Ibid., p. 747.
1882. Actinometra meyeri, P. H. Carpenter, Ibid., p. 747.
1884. Actinometra parvicirra, Bell, Rep. Zool. ColL H.M.S. "Alert," Lond., 1884, p. 168.
1887. Actinometra parvicirra, Bell, Trans. Dublin Soc, 1887, vol. iii. ser. ii. p. 645.
... Actinometra mutabilis, Liitken, MS., (pars), Godeffroy Museum.
Centro-dorsal discoidal and sometimes greatly reduced, but always with some
traces of marginal cirrus-sockets. Sometimes only three cirri visible, generally ten or
REPORT ON THE CRINOIDEA. 339
twelve, and occasionally twenty or twenty-five. They have ten to eighteen joints, a
few of which are longer than wide, and the later ones may have small dorsal spines.
First radials generally visible, sometimes largely so ; the second are partly united
laterally, but the rays are free and may divide two, three, and occasionally four times ;
each division of three joints, the axillary with a syzygy. Thirteen to forty-four arms of
generally overlapping triangular joints, which become gradually quadrate and elongated
at the ends. The arms are often dimorphic, the tentaculiferous anterior ones tapering
slowly with one hundred and twenty to one hundred and fifty joints, while the hinder
arms are ungrooved and taper rapidly with only fifty to ninety joints. A syzygy in
the third brachial, the next from the eighth to twelfth, usually in the tenth, and others at
intervals of one to ten, usually three or four joints.
The distichal pinnule may reach 15 mm., and the palmar pinnule, when present, is
usually about the same length ; but that on the second brachial is shorter, and those of
the fourth and fifth brachials are the smallest, their successors increasing in size. The
lower joints of these genital pinnules often overlap considerably and are sometimes
carinate. The terminal pinnules of the anterior arms are generally long and slender, but
those of the posterior arms are shorter and stouter, often with brownish ovoid bodies on
the dorsal aspect of several joints. The terminal comb varies much in size, being some-
times quite small and inconspicuous. It may not extend beyond the pinnule of the
third brachial, or occur on all the pinnules to the twelfth, and at intervals to the twenty-
fifth brachial, occasionally even to neax the end of the arm.
Mouth interradial or nearly so. Disk naked or bearing a few scattered granules
round the anal tube ; occasionally covered by a continuous pavement of plates. The
hinder arms and sometimes also the corresponding part of the disk are often ungrooved
and non-tentaculiferous.
Colour in spirit, — greenish-grey, or brown in various shades and more or less mixed
with white ; sometimes there is a medio-dorsal line, either white or dark.
Disk 20 mm. ; spread reaching 22 cm.
Localities. — Simon's Bay. One specimen.
Station 174 (b, c, or d), August 3, 1874; near Kandavu, Fiji; lat. (about) 19° 6'
S., long, (about) 178° 18' E. ; 255, 210, or 610 fathoms1 ; coral mud ; bottom temperature
at 610 fathoms, 39° F. One specimen.
Station 186, September 3, 1874 ; Prince of Wales Channel ; lat. 10° GO' S., long.
142° 18' E. ; 8 fathoms; coral mud. Five specimens.
Banda ; 17 fathoms. Two specimens.
Ternate. One specimen.
Admiralty Islands ; 16 to 20 fathoms. One specimen.
Samboangan ; 10 fathoms. Seven specimens.
1 The exact Station, and consequently the exact depth, is not recorded.
340 THE VOYAGE OF H.M.S. CHALLENGER.
Other Localities.— Cape of Good Hope; Port Natal; Ceylon; Nicobar Islands (?) ;
Australian Seas (Peron and Lesueur) ; Timor ; Solor ; North Borneo ; Sooloo ; China Sea ;
Yedo ; Zebu ; Bohol ; Ubay ; Cabulan ; Batjan ; Ceram ; H.M.S. " Alert," Warrior Reef,
Torres Strait, and Port Molle ; Kingsmills Islands ; Moreton Bay, Fiji ; Vavao ; Peru.
History. — -This name was given by Midler to an individual from some unknown
locality which was found by Troschel in the Paris Museum. It had twenty-seven arms,
owing to the presence of both distichal and palmar series, and twenty cirri of twelve
joints. Midler's description of it in his final memoir1 does not differ essentially from
that which was drawn up for him by Troschel in 1841 ;2 but he added to it a
more detailed diagnosis, based on his own observation, of a specimen from Vavao which
he was inclined to refer to the same type.
Although on two occasions I have searched carefully through the large Comatula-
collection in the Paris Museum, a privdege for which I am indebted to the kindness of
Professor Perrier, I have been unable to identify the original type of Midler's species.
The number of arms, twenty-seven, mentioned by him, is larger than that in some
individuals from the voyage of Peron and Lesueur which certainly belong to this species,
though I do not think that they can be the type of it as I formerly suggested. But I
can find no reference to them in any of Midler's writings, though he must certainly have
seen them when at Paris ; while they must also have been known to Lamarck, who
founded other species on Comatulae obtained by Peron and Lesueur.
Although, however, Midler's first type specimen seems to have disappeared, the second
one, that from Vavao, is in excellent condition. It was obtained by Hombron and
Jacquinot in 1841, during the voyage of the "Astrolabe," and is fortunately not dry,
but preserved in spirits. Had Midler been able to visit the Paris Museum himself in
1840, he would probably have recognised the identity of the form which he called Alecto
parvicirra with that which he found in the Leyden Museum under the name of Comatula
timorensis. The two species were described on successive pages of the Berlin Monats-
bericht for 1841, but I cannot regard them as different; and though the diagnosis of
Comatula timorensis is better than that of Comatula parvicirra, which precedes it, I
have preferred to retain the latter name, not on account of its one-page claim to priority,
but because it expresses a definite character of this widely distributed type, and does not
connect it with any particular locality.
Two years after making his first communication on the subject of Comatula-s'pecies,
Midler described a twenty-armed form from Natal in the Stockholm Museum under the
name "Alecto Wahlbergii."3 It has no palmar series, and further differs in several minor
1 Abhandl. d. k. Akad. d. Wiss. Berlin, Jahrg. 1847 [1849], p. 256.
2 Monatsber. d. k. preuss. Akad. d. Wiss. Berlin, 1841, p. 185.
3 Archivf. Naturgesch., 1843, Jahrg. ix. Bd. i. p. 131.
REPORT ON THE CRINOIDEA. 341
points from the types of Comatula parvicirra. and Comatula timorensis, so that I was
for a long time inclined to regard it as specifically distinct ; but I have at last been
obliged to abandon this view, and now consider the type as another variety of Actino-
metra parvicirra.
Grube's description of his Comatula mertensi 1 only differs from those of Comatula
parvicirra and Comatula timorensis in one essential point. He states that there are
but " 2 Radialia, das Axillare mit Syzygium." Were this really the case, his type would
be most closely allied to Actinometra distincta of the Typica-growp. But, having been
enabled by the kindness of Professor Schneider, Grube's successor at Breslau, to examine
the types of this species for myself, I can state positively that there are three radials
with a bifascial articulation between the second and third, as in most Comatulse ; while
in all other respects the characters of the type are those of Actinometra parvicirra, and
Grube's name is therefore reduced to the rank of a synonym.
During his residence in the Philippine Islands, Professor Semper collected several
examples of an Actinometra with thirteen to thirty-nine arms, on which, believing it to
be new to science, he bestowed the MS. name armata. This name was employed by
myself in a couple of anatomical papers,2 though I subsequently found reason to replace
it by polymorpha? when giving a detailed description of the type, which did not appear
to me to be absolutely identical with the Vavao variety of Actinometra parvicirra.
Further experience, however, has convinced me that the two forms cannot be separated
specifically, and I must also refer to the same variable type the dry specimen in the
Hamburg Museum which I have noticed as Actinometra meyeri.4 The same may be said
of the Actinometra annulata of Bell,5 in whose diagnosis I can find no single point of
specific value by which this type can be distinguished from the Actinometra polymarpha
which I had described some years previously, and had subsequently referred to Actino-
metra parvicirra, Miill., sp.;6 while as Bell gave no hint of his views respecting
the relationship of his new species, his reasons for establishing it are somewhat
obscure.
Some of the specimens which have been distributed by the Godeffroy Museum under
the name Actinometra mutabilis, Liitken, MS., must also be referred to Actinometra
parvicirra, e.g., No. 6146, from Moreton Bay, Fiji. There is a tridistichate individual
from the Nicobar Islands which I found under this name in the Copenhagen Museum, and
I subsequently saw a sinidar form at Vienna. The arm-joints are rather long in both
cases, and without making a renewed examination of the specimens I should not like to
1 53e Jahresber. der Schlesisch. Gesellsch.f. Vaterl. Cult, 1875, p. 74.
2 Journ. ofAnat. and Phys., 1876, vol. x. p. 582 ; vol. xi. p. 91.
3 Trans. Linn. Soc. Land. (Zool.), ser. 2, 1877, [1879], p. 50.
1 Journ. Linn. Soc. Lond. (Zool.), 1882, vol. xvi. p. 525.
6 Proc. Zool. Soc. Lond., 1882, p. 535.
0 Notes from the Ley den Museum, 1881, vol. iii. p. 204.
342 THE VOYAGE OF H.M.S. CHALLENGER.
speak positively as to their nature. But they must certainly belong either to Actino-
metra quadrata or to Actinometra parvicirra.
Remarks.— This latter variable and much be-named species is a somewhat isolated
one. It is separated from Actinometra trichoptera and Actinometra japonica by its
smaller number of cirri ; while Actinometra regalis and its allies have many more arms
which are united more or less completely by interradial plating (PI. LXVIII. fig. 2).
The only type which approaches Actinometra parvicirra at all closely is Actinometra
quadrata, which seems to be distinguished from it by the shape of its middle and later
arm-joints (PI. LXII. fig. 1). There are, however, one or two forms among those collected
at Samboangan by the Challenger which appear to approach Actinometra quadrata, and
it may be that the latter name will have to be abandoned.
When describing the series of specimens which I called Actinometra polymorpha, I
put down the number of arms as ranging from thirteen to thirty-nine ; and among all
the many specimens of this type which I have examined during the last twelve years I
have found but one in which those limits have been exceeded. Bell, indeed, described
his single individual of Actinometra annulata as having forty arms, which would mean
that it has the full complement of ten distichal and twenty palmar axillaries ; but I have
found on examination that this is not the case in reality. Post-palmar axillaries occur in
but few individuals that I have examined ; and even when they are present I have only
once found the number of arms to exceed thirty-nine, owing to the absence of palmars
and even of distichal axillaries in other parts of the rays, as is well shown in PI. LXVII.
fig. 3. In this individual three of the primary arms do not divide at all, i.e., there are
only seven instead of ten distichal axillaries, and the deficiency of arms arising from their
absence is partly compensated by the presence of three post-palmar axillaries. The same
is true, though in a less degree, of two more Challenger specimens from Samboangan
(PI. LXI. fig. 5), and also of one from Batjan which I have seen in the Berlin Museum ;
but in one example from Banda which I must provisionally refer to this type, all the
distichal and palmar axillaries are present, together with four post-palmars in addition.
There are five more individuals in the Challenger collection which have distichals and
palmars but no post-palmars, and eight more in which palmars are not present at all, a
condition which also occurs in three of Semper's eleven examples, one of which has only
thirteen arms. So great a variation in the number of arms as this is certainly unusual, but
I have found myself quite unable to draw any fixed line of separation, often as I have
attempted it. It does seem, however, as if forty were the usual limit of the number
of arms in this species, even though post-palmars may sometimes be present ; and I
am inclined to lay more stress upon this as a character of systematic value than upon the
presence or absence of palmar or post-palmar axillaries.
The potential dimorphism in the characters of the arms of Actinometra is very
well shown in this species. It presents itself in two large individuals dredged by the
REPORT ON THE CRTNOIDEA. 343
Challenger at Station 186, and in at least six of the seven from Samboangan. The usual
rule is that the tentaculiferous anterior arms have about twice as many joints as the
ungrooved hinder arms, which terminate definitely in a miniature axillary joint, bearing
a couple of pinnules ; while the anterior arms always seem to end in a growing point as
is the case with all the arms of Antedon.
The problematical ovoid bodies, which occasionally appear as brown spots in the
centre of the dorsal surface of some of the segments of the pinnules on the ungrooved
arms, occur in single individuals of this species from Station 186, Banda, and
Samboangan, and also in five out of the eleven examples obtained by Semper in the
Philippines. 1 am at present quite unable to throw any light upon their character,
though I hope that the researches of Dr. 0. Hamann, in whose skilled hands I have
placed several of the pinnules containing them, may add considerably to our knowledge
of their nature and structure. They are not peculiar to Actinometra parvicirra, as they
also present themselves in Actinometra elongata from Banda, and in the Brazilian
Actin ometra me ridiona lis.
The number of cirri which occur in Actinometra parvicirra seems to vary consider-
ably, though the number of joints remains fairly constant at ten to sixteen. In three
of the Philippine specimens the centro-dorsal is reduced to a thin disk bearing three or
four moderately developed cirri, with indications of other sockets which have been more
or less completely obliterated (PI. LXI. figs. 1,5); while in the individual from Station
174 the centro-dorsal is very irregularly shaped, and bears quite rudimentary cirri with
imperfect sockets for others (PI. LXI. fig. 3). Another Samboangan specimen has a
larger number of cirri, but they are all small and rudimentary on a very thin centro-dorsal
(PL LXI. fig. 4). As a general rule there are ten or a dozen cirri which are not unfre-
quently disposed in pairs, two at each angle, with a few others in intermediate positions
(PI. LXI. figs. 2, 6 ; PI. LXVII. fig. 3). But I have seen individuals, both from the
Philippines and from the Cape of Good Hope, with as many as twenty -five sockets on the
centro-dorsal, which almost entirely conceals the first radials, though of course they may
not all have borne cirri simultaneous^. There is also a good deal of variation in the
development of the spines on the later cirrus-joints, and in the characters of the terminal
comb on the lower pinnules. Three modifications of this comb in different individuals
from Samboangan are shown in figs. 8-10 on PL LXI. In the original of fig. 8, the comb
is so small that it might easily escape notice ; but the other two pinnules are more normal
in character. The number of pinnules which bear a comb is also very variable. I have
seen specimens both from Africa and from the Philippines, in which there is no comb
after the third brachial ; while in others from both localities it may be found on the
pinnule of the fifteenth brachial, and in some of the Philippine specimens the later
pinnules of the arms may have small combs. In like manner I have seen individuals
from the Cape and from the Philippines in which the basal joints of the genital pinnules
344 THE VOYAGE OF H.M.S. CHALLENGER.
are considerably produced towards the dorsal side ; but in other individuals from these
and from other localities this character is entirely absent. That obtained by the
Challenger at Simon's Bay seems to have just liberated its ova, as a small group of them
is collected on the distal side of each genital pinnule, in the angle between it and the
arm (PL LXI. fig. 7).
In nearly all the examples of this species which I have seen the mouth is very
distinctly interradial, as is well shown in Midler's diagram of Comatula icahlbergii,1 and
in my own figures of Actinometra polymorpha.2 In one or two cases, however, the
A ambulacrum is somewhat displaced forwards, though never so much so as to cause
the mouth to become radial.
The disk is generally naked, but the neighbourhood of the anal tube sometimes bears
scattered granules ; while in one individual from Torres Strait there is a tolerably close
pavement of minute scale-like plates over the whole disk. The perisome of the arms
and pinnules in this individual is considerably reduced, and the genital glands are but
poorly developed, though in another from the same station, which was presented by Sir
AVyville Thomson to the Stockholm Museum, the perisome is much more substantial
and the genital pinnules, especially in the posterior arms, are much swollen, so that the
two forms differ greatly in their external appearance.
Actinometra parvicirra, as described above, is a somewhat comprehensive type,
embracing as it does three of Midler's species, together with four others which have been
regarded as distinct at various times ; and its distribution therefore is considerably
extensive. It occurs to a distance of about 35° on either side of the equator, and has a
range in longitude of some 260° from the Cape of Good Hope to Peru. Long since
known from Natal, Timor, and from the Friendly Islands, it has subsequently been
discovered at numerous intermediate localities, such as Ceylon, the Moluccas, Phdippines,
Japan, Fiji, and East Australia ; and I quite expect that it will be eventually found in
the Atlantic, more especially as the species of Actinometra characteristic of that ocean
seems also to occur in the Arafura Sea, while Antedon carinata of the Indian Ocean and
East Pacific is a common species in the West Atlantic.
1 Abhandl. d. k. Akad. d. Wiss. Berlin, Jahrg. 1847 [1849], p. 245.
2 Trans. Linn. Soc. Lond. (Zool.), ser. 2, 1877 [1879], pi. i. figs. 6-10.
REPORT ON THE CRTNOIDEA. 345
3. Actinometra trichoptera (Valenciennes), Miill., sp. (PI. LXIII. figs. 1-5).
be
Spec ific formula — a. 3 . ( 3 ) . — .
Comatula trichoptera, Valenciennes, MS.
1846. Comatula trichoptera, Miiller, Monatsber. d. k. preuss. Akad. d. Wiss. ^Berlin, 1846, p. 178.
1849. Comatula trichoptera, Miiller, Abliandl. d. k. Akad. d. Wiss. Berlin, Jahrg. 1847 [1849],
p. 257.
1862. Comatula trichoptera, Dujardin and Hupe\ Hist. Nat. des Zoophytes, fichinodermes,
Paris, 1862, p. 205.
1879. Actinometra trichoptera, P. H. Carpenter, Trans. Linn. Soc. Lond. (Zool.), ser. 2, 1877
[1879], p. 27.
1882. Actinometra trichoptera, Bell, Proc. Zool. Soc. Lond., 1882, p. 535.
1882. Actinometra trichoptera, P. H. Carpenter, Ibid., p. 747.
Centro-dorsal a relatively wide disk, bearing some thirty or more marginal cirri.
These have about sixteen joints, a few of which are longer than wide, the penultimate
with but little trace of au opposing spine.
First radials scarcely visible, and the second but partially united laterally ; the rays
quite free and the axillary angle rather sharp. Three distichals and sometimes three
palmars, the axillary with a syzygy.
Fifteen to twenty-two arms, of slightly overlapping joints, the lower ones relatively
short and triangular, gradually becoming longer and more quadrate. Syzygies in the third
and in the tenth or twelfth brachials, and then at intervals of three or four joints.
The distichal pinnule is about 9 mm. long, and that on the second brachial but little
shorter. The next two or three diminish rapidly in length, but become swollen for the
genital glands and lose their terminal comb. The lower joints of the earlier pinnules
sometimes overlap rather sharply and have spinose edges.
Mouth interradial ; disk naked.
Colour in spirit, — light yellowish -brown, mottled with grey or darkish brown.
Disk 9 mm.; spread 12 cm.
Locality. — Port Jackson ; 10 to 12 fathoms. One specimen.
Other Localities. — Port Philip ; King George's Sound.
Remarks.- — The types of this species were brought to Paris from King George's Sound
by Quoy and Gaimard, and received from Valenciennes the MS. name trichoptera, which
was adopted by Miiller when he afterwards described them. Its range was extended to
Port Jackson by the Challenger, and the British Museum has since obtained examples of
it from Port Philip, so that it may be assumed to be common along the whole southern
coast of Australia. But I have never met with any form like it from the tropical seas.
Actinometra robustipinna from the Moluccas resembles it in the presence of a large
number of cirri, but is readily distinguished by the great size of its first three pinnules.
On the other hand there is a considerable resemblance between Actinometra trichoptera
(ZOOL. CHALL. EXP.— PABT LX.— 1888.) OoO 44
346 THE VOYAGE OF H.M.S. CHALLENGER.
and Miiller's other species, Actinometra japonica, to which I would now refer — possibly
as a varietal form — the individual which I called Actinometra morsei when asked by
von Graff to name the host of Myzostoma nigrescens.
' This is a little specimen without palmars, which are also absent in most of the examples
of Actinometra trichoptera that I have seen. It has rather shorter axillaries than the
type of Actinometra japonica, and less developed spines on the terminal cirrus-joints,
both of which are points of resemblance to Actinometra trichoptera. It seems, however,
to have longer arm -joints than the Australian species, and shows the carination of the
large basal joints of the distal pinnules, which in Actinometra japonica extends further
out on the arm. This does not appear in Actinometra trichoptera, and for the present,
therefore, I should be inclined to regard the two species as distinct, though it is by no
means improbable that other intermediate forms may eventually be discovered.
8. Actinometra littoralis, n. sp. (PI. LXVII. figs. 1, 2).
Specific formula — a.3.3.(2).— .
Description of an Individual. — Centro-dorsal a very thin pentagonal disk with
slightly incurved sides, rather above the level of the radial circlet and separated from
it by faint clefts. Cirri all lost. Three radials visible ; the second almost completely
united laterally, but the axillaries free. The rays may divide four times. Three
distichals and three palmars, the axillary with a syzygy ; post-palmars, when present,
of two joints only, the axillary without a syzygy. Thirty-eight arms, which are all
grooved, but dimorphic. The anterior of one hundred and fifty, and the posterior of
one hundred segments, which are triangular at the base, gradually becoming more
quadrate and slightly elongated towards the end. Syzygies in the third and tenth or
eleventh brachials ; others at intervals of three or four joints.
The palmar pinnule is nearly as long as the distichal one, which reaches 12"5 mm.:
but that of the second brachial is much shorter. The next pair are the smallest, their
successors increasing again. The terminal pinnules are much longer and more slender in
the anterior than in the posterior arms. The lowest joints of the proximal pinnules are
rather wide and overlap slightly with spinose margins. The proximal pinnules have a
well-defined comb which disappears by the fourth or fifth brachial.
Mouth interradial ; all the arms grooved ; disk naked.
Colour in spirit, — deep blackish-brown.
Disk 20 mm.; spread 20 cm.
Locality. — Banda ; 17 fathoms. One specimen.
Remarks. — But one individual of this species having been obtained, I am unable to
state its characters as definitely as I could wish. In the normal arrangement of the arm-
REPORT ON THE CRINOIDEA. 347
divisions there are three distichals and three palmars, just as in Actinometra parvicirra ;
but sometimes there are only two palmars ; and in one case this arrangement is followed
by a post-palmar series of the same character, which does not seem to be due to regenera-
tion (PI. LXVII. fig. 1) ; but as it may not occur in other individuals, I have enclosed
the sign for it within brackets in the specific formula.
Apart, however, from the possible presence of the two-jointed post-palmar series,
Actinometra littoralis differs from Actinometra parvicirra and Actinometra trichoptera
in the more complete reduction of its centro-dorsal. This is not quite lowered to the
level of the radial pentagon, from which it is separated by commencing clefts, a condition
not reached by any specimen that I have seen which in other respects presents the
general characters of Actinometra parvicirra. A minor point of distinction between
the two species is afforded by the overlap and the very spiny margins of the lower
pinnule-joints in Actinometra littoralis ; while the terminal comb disappears earlier than
is usually the case in Actinometra parvicirra, though it is well-developed on the proximal
pinnules (PL LXVII. fig. 2).
9. Actinometra regalis, n. sp. (PI. LXVIIL).
Specific formula — a.3. 3. 3.3.—.
Centro-dorsal a rudely circular disc, much hollowed in the centre, and bearing fifteen
to twenty marginal cirri, of fifteen nearly equal segments ; the penultimate without an
opposing spine.
Three radials visible, the second closely united laterally ; axillaries short, widely
triangular, and in contact laterally. The rays may divide five times but do not spread
much, as the first joints beyond each division are closely united laterally. The rays and
their divisions are united by interradial plating to the level of the distichal axillaries or
slightly beyond it. Each division of three joints, the third of which is axillary with a
syzygy-
Arms very numerous, thirteen to twenty-four on a ray. They have moderately long,
triangular, and much-overlapping joints, which soon become quadrate and are nearly
square at the ends ; the anterior arms have one hundred and sixty, and the posterior
only sixty or seventy joints. Syzygies in the third and the tenth or twelfth brachials,
and then at intervals of two to six, usually three or four joints.
The distichal and palmar pinnules are of about equal size, reaching 15 mm.; the
length diminishes to that on the second brachial, which is considerably shorter, and
those of the fourth and fifth brachials are much more so, after which the length increases
again. Later pinnules of the anterior arms not specially long ; terminal comb to the
tenth brachial.
348 THE VOYAGE OF H.M.S. CHALLENGED.
Mouth interraclial or nearly so ; disk naked. or slightly plated.
Colour in spirit, — deep brown, the pinnules tipped with yellow-green.
Disk 27 mm.; spread 22 cm.
Locality. — Banda ; 17 fathoms. Two specimens.
Remarks. — These two individuals seem to be different from that which I found in
the Leyden Museum and described under the name Actinometra schlegelii.1 They show
much more of the first radials,. which are almost entirely concealed in the Lej^den species,
and have relatively longer arm-joints. This character is best marked in the middle and
outer parts of the arms, those of Actinometra schlegelii being much wider than long ;
while in Actinometra regalis the joints are more equally quadrate, and the overlap of the
lower joints is more marked. In this species too the pinnules of the fourth to sixth
brachials are quite small, which is not the case in Actinometra schlegelii. The number of
cirrus-joints in the latter type is not known ; but Actinometra regalis has less than
twenty, being thus distinguished from Actinometra peroni with its very long cirri of
thirty joints ; while in Actinometra bennetti there are fifty cirri of twenty-five joints.
Genus 6. Promachocrinus, P. H. Carpenter, 1879.
1879. Promachocrinus, P. H. Carpenter, Proo. Boy. Soc, 1879, vol. xxviii. p. 385.
1880. Promachocrinus, P. H. Carpenter, Journ. Linn. Soc. Lond. (Zool.), 1880, vol. xv. p. 214.
Definition. — Centro-dorsal hemispherical or conical, bearing numerous closely-set
cirri. Ten radials with high distal faces which have large muscle-plates. Mouth central;
ambulacra symmetrically distributed and not provided with any definite skeleton.
Sacculi well developed.
Remarks. — The principal distinctive character of this remarkable genus, which is
only known from the dredgings of the Challenger, is the presence of ten radials in the
calyx instead of the usual five (PL I. figs. 1, a, b, c). In all other respects there are no
essential differences between Promachocrinus and Antedon. The species of the latter
genus to which Promachocrinus is most allied are those of the Eschrichti-gv oup, in
which the radials have high articular faces with large muscle-plates (PI. I. figs. 1, 6, 8, a).
The latter character also presents itself in Antedon accela, Antedon basicurva, and their
allies (PL II. figs. 1-5, a); but all these forms have a well-defined ambulacral skeleton
which is altogether absent in Promachocrinus.
One of the three species of this genus was obtained at a depth of 500 fathoms off the
Meangis Islands (Station 214). Unfortunately, however, it is only represented by one
individual in a most mutilated condition (PL LXIX.' figs. 9, 10). But each of the other
two species occurred at two localities in the southern sea. The type-species, Promacho-
1 Notes from the Leyden Museum, 1881, vol. iii. p. 210.
REPORT ON THE CRINOIDEA. 349
crinus kerguelensis, was obtained in various shallow-water dredgings round the coast of
Kerguelen Island, and was also found at 75 fathoms near Heard Island. In its general
facies it has a singular resemblance to Antedon eschrichti and its allies; while Promacho-
crinus abyssorum, from 1600 and 1800 fathoms (Stations 147, 158) is more like the
circumpolar and abyssal members of the Tenetta-group among the species of Antedon.
Two of these were associated with it at Station 147, while at Station 158 there was also
obtained the remarkable genus Thaumatocrinus.
Although there are ten radials in the calyx of Promachocrinus, the symmetry of the
basals is only pentamerous. Five of the radials are essentially like those of Antedon,
with a smooth dorsal surface and two openings on the inner face, between which is the
shallow groove lodging the radial axial furrow. This seems to have been converted into
a canal by the radial process of a rosette, just as in Antedon and Actinometra (PI. I.
fig. 8c ; PI. III. figs. 4c, 5b) ; but I was unfortunately unable to obtain this rosette entire,
for the central portions of it broke away from the peripheral part which remained firmly
attached to the radials (PI. I. fig. lc).
In many of the five-rayed Comatulae the interradial angles of the rosette become
connected with the five elements of the basal star, which are developed in the synostosis
between the centro-dorsal and the radials as I have explained elsewhere ; ' and these
basal rays lie beneath the sutures between the five primary radials (PI. I. fig. 6c; PI. II.
figs. 1-5, c; PL III. figs, lc, 3a, 36, 4c; PI. IV. fig. 3c; PL V. figs, lc, 5c/). In
Promachocrinus, however, with its ten radials (or at any rate in Promachocrinus
kerguelensis), there is a basal ray beneath the middle of every alternate radial (PL I.
figs. 1, a, c). Its inner end is broad and flattened, and extended laterally into two
processes which meet those of the adjacent basal rays beneath the dorsal surface of the
intervening primary radials (PL I. fig. lc). When these ten radials are separated from one
another the basal rays come away with the " interradial radials " to which they are attached
(PL I. figs. 2, a, b), and their impressions are left upon the inner ends of the dorsal surface
of the true primary radials with which they were in contact (PL I. figs. 1, 3, c).
The isolated centro-dorsal of Promachocrinus is indistinguishable from that of
Antedon. Its ventral surface is marked by five grooves lodging the basal rays (PL I.
figs, lc, 5). But there are. only five large radial areas without any indication whatever
that each of these lodges portions of two additional radials, as well as its true or primary
one. In the large centro-dorsal of Promachocrinus kerguelensis the five interradial
pillars within the central cavity are very distinct, as is also the case in Antedun
antarctica (PL I. figs. 1, 6, d).
In one of the three species of Promachocrinus the rays divide so as to produce twenty
arms ; but they remain simple in the other two species, just as in Eudiocrinus and
Thaumatocrinus. In both alike the first pinnule is on the second joint above the
1 Trans. Linn. Soc. Lond. (Zool.), 1879, ser. 2, vol. ii. pp. 95-100.
350 THE VOYAGE OF H.M.S. CHALLENGER.
primary radial ; but there seems to be no constancy as to the side on which it appears,
some arms having it on the right, and others on the left side.
The three species of the genus may be classified as follows : —
I. Twenty arms, . . . . . . . . . 1. kerguelensis, n. sp.
II. Ten arms only.
A. Centro-dorsal small, with but few cirri ; elongated arm-joints, . . 2. abyssorum, n. sp.
B. Centro-dorsal large, bearing numerous cirri ; arm-joints not specially long, . 3. naresi, n. sp.
1. Promachocrinus kerguelensis, n. sp. (PI. I. figs. 1, a-d; PI. LXX.).
1879. Promachocrinus kerguelensis, P. H. Carpenter, Proc. Roy. Soc, 1879, vol. xxviii. p. 385.
1880. Promachocrinus kerguelensis, P. H. Carpenter, Journ. Linn. Soc. Loud. (Zool.), 1880,
vol. xv. pL xii. fig. 28.
Centro-dorsal conical and thickly covered almost to the apex with eighty or more cirri.
These may reach to 40 mm. in length, and consist of thirty-five to forty tolerably uniform
joints, which are mostly rather longer than wide. The later joints may overlap slightly,
but the penultimate is small, with little or no trace of an opposing spine.
First radials barely visible; the second short, nearly oblong, and but slightly joined
laterally ; axillaries widely rhombic. The first brachial is scarcely incised by the second,
which is irregularly quadrate. The next few joints are quadrate, and their succcessors
triangular, wdder than long, and slightly overlapping. A syzygy in the third brachial ;
the next in the seventh or eighth, with others at intervals of two to four joints. The
first two pinnules on each side are tolerably equal, slender, and flagellate, and reach over
20 mm. in length.
The lowest pinnules have the most slender joints, those of their successors increasing
in stoutness, but diminishing in number. The two lowest joints of the middle and later
pinnules are somewhat flattened, with their apposed edges incurved.
Mouth central and anus marginal ; disk naked ; genital glands long and slender ;
sacculi abundant on the pinnules.
Colour in spirit, — light yellowish-brown, or greyish-white with dark red spots.
Disk 16 mm.; spread 18 cm.
Localities. — Kerguelen Island; 10 to 100 fathoms. One specimen.
Station 149c, January 19, 1874 ; Balfour Bay ; 20 to 60 fathoms. Two specimens.
Station 149d, January 20, 1874; Royal Sound; lat. 49° 28' S., long. 70° 13' E.;
28 fathoms. One specimen.
Station 149e, January 21, 1874 ; off Greenland Harbour ; 30 fathoms. One specimen.
Station 149h, January 29, 1874; off Cumberland Bay; 127 fathoms. Two young
specimens.
The bottom deposit at all these stations is volcanic mud.
Station 151, February 7, 1874; off Heard Island; lat. 52° 59' 30" S., long.
73° 33' 30" E.; 75 fathoms ; volcanic mud. One young specimen.
REPORT ON THE CRINOIDEA. 351
Remarks.— There is a most remarkable general resemblance between this species and
tlie two allied forms Antedon eschrichti and Antedon antarctica, which last was found
associated with it off Heard Island. The characters of the cirri, arm-joints, pinnules,
and even of the genital glands are very closely similar in the two types ; so that if
nothing were known of Promachocrinus kerguelensis but some fragments of its arms,
one would unhesitatingly refer them to an Antedon of the Eschrichti-grouTp.
Three of the seven specimens obtained at Kerguelen appear to be fully developed,
while two are premature, and two more, those from Christmas Harbour, are quite young
(PI. LXX. fig. 2). The cirri of these last exhibit a very striking dimorphism. Most of
them belong to the " small mature " type, while there are others with a much larger
number of immature joints. The latter type is the one which chiefly presents itself in
the adult.
The first radials of these young individuals are much more distinctly visible than is
the case in the adult ; while the second radials and first brachials are more deeply
incised, and the arm-joints relatively longer with a more distinct overlap. The Heard
Island specimen is considerably older than those from Christmas Harbour, but also has a
large number of the " small mature "' cirri.
The anal tube of this species, in the five examples of which I have examined the
disk, is near the margin and not close up to the central mouth, as is usually the case in
Antedon.
2. Promachocrinus abyssorum, n. sp. (PL I. figs. 4, 5 ; PI. LXIX. figs. 5-7).
1879. Promachocrinus abyssorum, P. H. Carpenter, Proc. Roy. Soc, 1879, vol. xxviii. p. 385.
Centro-dorsal small and rounded, nearly covered by the sockets of about twenty cirri,
which seem to have had very long lower joints.
Radials partially visible ; the first brachial of moderate length, and somewhat incised
by the second, which bears a pinnule. The next few joints are nearly oblong, and their
successors quadrate, gradually becoming much elongated. There is a syzygy in the
fourth or fifth brachial, and others at intervals of one to five joints.
The first few pinnules on each side are tolerably equal, slender, and flagellate,
reaching 8 mm. in length. The lowest pinnules have the smallest joints, those of their
successors becoming both longer and stouter.
Mouth central; disk naked. Genital glands short and stout. Sacculi fairly
abundant on the pinnules.
Colour in spirit, — white.
Disk 6 mm.; spread perhaps 10 cm.
Localities.— Station 147, December 30, 1873; lat. 46° 16' S., long. 48° 27' EL;
1600 fathoms ; Diatom ooze ; bottom temperature, 34°-2 F. Three specimens.
352 THE VOYAGE OF H.M.S. CHALLENGER.
Station 158, March 7, 1874 ; south-west of Melbourne ; lat, 50° l'S.,long. 123° 4' E.;
1800 fathoms; Globigerina ooze; bottom temperature, 33°"5 F. One specimen.
Remarks. — The general characters of this little species are altogether those of an
abyssal Antedon belonging to the Tenella-gvovq). It is remarkable for the shape of its
genital glands, which are short and thick (PL LXIX. figs. 7 a, b), instead of being long
and fusiform as in Promachocrinus kerguelensis and in Antedon eschrichti.
3. Promachocrinus naresi, n. sp. (PL LXIX. figs. 8-10).
1879. Promachocrinus naresii, P. H. Carpenter, Proc. Eoy. Soc, 1879, vol. xxviii. p. 385.
Description of an Individual. — Centro-dorsal hemispherical, 9 mm. in diameter, and
covered except at the dorsal pole by the sockets of some forty cirri.
Radials just visible ; the first brachial rather short and but little incised by the
second, which bears a pinnule. The next few joints are nearly oblong or quadrate, with
somewhat tubercular junctions and pinnules on their shorter sides ; the later joints
become more distinctly triangular and quadrate towards the ends. A syzygy in the
fourth brachial, and others at intervals of four to twelve joints. Sacculi abundant on
the pinnules.
Colour in spirit, — brownish-white.
Locality. — Station 214, February 10, 1875 ; off the Meangis Islands; lat. 4° 33' N.,
long. 127° 6' E.; 500 fathoms ; blue mud ; bottom temperature, 41°-8 F. One mutilated
individual.
Remarks. — This specimen is unfortunately so mutilated that a complete description
of it is impossible. But it is obviously not identical with the other ten-armed species
just described, as it has a larger centro-dorsal with more numerous cirri, and the
arm-joints relatively shorter and more triangular (PL LXIX. figs. 5, 8). The general
characters of the arms and pinnules, so far as can be judged from the fragments of them
which are preserved, are essentially those of Promachocrinus kerguelensis.
REPORT ON THE CRINOIDEA. 353
VII. BATHYMETRICAL DISTRIBUTION AND STATION LIST.
Station List of the Comatdl^e which have been obtained by the various British
Expeditions for Deep-Sea Exploration between the Years 1868 and 1882.
This list also contains the generic names of those Stalked Crinoids which were
dredged at Stations where Comatuke occurred. The specific details respecting them
will be found in Chapter xiii. of Part I.
H.M.S. "Lightning," 18G8.
Station 13. Lat. 59° 5' N, long. 7° 29' W.; 189 fathoms ; bottom temperature, 49°-3 F.
Antedon phalangium. Also in the Minch ; off Cape Mondego and
Cape Sagres ; the Seine Bank, and Mediterranean.
H.M.S. " Porcupine," 1869.
The Minch, 60 to 88 fathoms ; and off Loch Scavaig, Skye.
Antedon phalangium. Also at Station 13 (1868); off Cape
Mondego and Cape Sagres ; the Seine Bank, and
Mediterranean.
Station 51. Lat. 60° 6' N, long. 8° 14' W.; 440 fathoms; bottom temperature, 42° F.
Antedon tenella. Also at Stations 54, 55, 74 ; and 1870, Station
17a. Also the "Triton," 1882, Stations 2, 5.
Station 54. Lat. 59° 56' N, long. 6° 27' W. ; 363 fathoms; bottom temperature,
31°-4 F.
Antedon tenella. Also at Stations 51, 55, 74; and 1870,
Station 17a. Also the "Triton," 1S82, Stations
2, 5.
Station 55. Lat. 60° 4' N, long. 6° 19' W.; 605 fathoms ; bottom temperature, 29°8 F.
Antedon tenella. Also at Stations 51, 54, 74 ; and 1870, Station
17a. Also the "Triton," 1882, Stations 2, 5.
(ZOOL. CHALL. ESP — PART LX. 1888.) OoO 45
354 THE VOYAGE OF H.M.S. CHALLENGER.
Station 57. Lat. 60° 14' N., long. 6° 17' W.; 632 fathoms; bottom temperature,
30°-5 F.
Antedon eschrichti. Also the "Valorous," 1875, Station 1 ; the
"Alert," 1875, Franklin-Pierce Bay; the
Challenger, Station 48 ; and the "Triton," 1882
Station 4.
Station 74. Lat. 60° 39' N., long. 3° 9' W.; 203 fathoms; bottom temperature,
47°-6F.
Antedon tenella. Also at Stations 51, 54, 55 ; and 1870, Station
17a ; also the "Triton," 1882, Stations 2, 5.
Stations not recorded.
Antedon rosacea. Also off the coast of Tunis.
Antedon hystrix. Also the " Triton," 1882, Station 4.
H.M.S. "Porcupine," 1870.
Station 13. Off Cape Mondego ; lat. 40° 16' N., long. 9° 37' W.; 220 fathoms ; bottom
temperature, 52° F.
Antedon phalangium. Also in the Minch ; off Cape Sagres ; the
Seine Bank, and the Mediterranean.
Station 17a. Lat. 39° 39' N., long. 9° 39' W. ; 740 fathoms; bottom temperature,
49°'3 F.
Antedon lusitanica.
Antedon tenella. Also 1869, Stations 51, 54, 55, 74 ; also the
" Triton," 1882, Stations 2, 5.
Station 25. July 27, 1870 ; near Cape St. Vincent; lat. 37° 11' N., long. 9° 7' W.;
374 fathoms ; rock ; bottom temperature, 53°"5 F.
Actinometra pidchella. Also at Station 31, and by the Challenger
at St. Paul's Rocks; perhaps also at Station 192.
Abundant in the Caribbean Sea.
Station 31. Lat. 35° 56' N., long. 7° 6' W.; 477 fathoms; clay; bottom temperature,
50o>5 F.
Actinometra pulchella. Also at Station 25, and at St. Paul's Eocks.
REPORT ON THE CRINOIDEA. 355
Off Cape Sagres ; 45 fathoms.
Antedon phalangium. Also in the Mineh ; off Cape Mondego ;
the Seine Bank, and Mediterranean.
Off Carthagena ; 80 fathoms.
Antedon phalangium. Also in the Atlantic.
Bay of Benzert ; 50 to 100 fathoms.
Antedon phalangium.
Antedon rosacea (young). Also in the Fasroe Channel.
Skerki Bank ; 30 to 120 fathoms.
Antedon phalangium.
Antedon rosacea (young). •
H.M.S. " Valorous," 1875.
Station 1. July 22, 1875 ; off Hare Island in Davis Strait ; lat. 70° 30' N., long. 54° 41'
W. ; 175 fathoms ; sand and mud.
Antedon eschrichti. Also the "Porcupine," 1869, Station 57;
the "Alert," 1875, Franklin-Pierce Bay; the
Challenger, Station 48; and the "Triton," 1882,
Station 4.
Station 6. August 10, 1875 ; lat. 64° 5' N., long. 56° 47' W.; 410 fathoms; sand and
mud ; bottom temperature, 34°"6 F.
Antedon quadrata. Also the "Alert," 1875, Discovery Bay and
Franklin-Pierce Bay ; the Challenger, Station 48 ;
and the "Triton," 1882, Stations 4, 6.
H.M.S. " Alert," 1875.
Franklin-Pierce Bay ; lat. 79° 25' N.
Antedon eschrichti. Also the " Porcupine," 1869, Station 57 ; the
" Valorous," 1875, Station 1 ; the Challenger,
Station 48 ; and the "Triton," 1882, Station 4.
Antedon quadrata. Also at Discovery Bay ; also the " Valorous,"
1875, Station 6 ; the Challenger, Station 48 ; and
the "Triton," 1882, Stations 4, 6.
356 THE VOYAGE OF H.M.S. CHALLENGER.
Discovery Bay; lat. 81° 41' N.; 25 fathoms; hard bottom.
Antedon prolixa.
Antedon quadrata. Also at Franklin-Pierce Bay ; the "Valorous,"
1875, Station 6 ; the Challenger, Station 48 ; and
the "Triton," 1882, Stations 4, 6.
H.M.S. " Knight Errant," 1880.
On the plateau N.N.W. of North Bona; lat. 59° 12' N., long. 5° 57' W.; 53 fathoms;
rough ground.
Antedon rosacea. Also in the Mediterranean.
H.M.S. "Triton," 1882.
Station 2. Lat. 59° 37' 30" N., long. 6° 19' 0" W.; 530 fathoms; mud; bottom
temperature, 46° '2 F.
Antedon tenella. Also at Station 5. Also the "Porcupine," 1869,
Stations 51, 54, 55, 74 ; and 1870, Station 17a.
Station 3. August 8, 1882 ; on the Feeroe Banks ; lat. 60° 39' 30" N., long. 9° 6' 0"
W.; 87 fathoms; sand and shells ; bottom temperature,
49° F.
Antedon petasus.
Station 4. Lat, 60° 22' 40" N. and 60° 31' 15" N., long. 8° 21' 0" W. and 8° 14' 0"
W.; 327 to 430 fathoms ; stones, mud ; bottom tempera-
ture, 31°-5 to 30° F.
Antedon eschrichti. Also the "Porcupine," 1869, Station 57;
the "Valorous," 1875, Station 1; the "Alert,"
1875, Franklin-Pierce Bay; and the Challenger,
Station 48.
Antedon hystrix. Also the "Porcupine," 1869.
Antedon quadrata. Also at Station 6. Also the " Valorous," 1875,
Station 6 ; also the " Alert," 1875, Discovery Bay
and Franklin-Pierce Bay ; and the Challenger,
Station 48.
REPORT ON THE CRINOIDEA. 357
Station 5. Lat. 60° 11' 25" N. and 60° 20' 15" N., lonff. 8" 15' 0" W. and 8° 8' 0"
W.; 433 to 285 fathoms ; hard ground, stones ; bottom
temperature, 43°-5 to 40°-8 F.
Antedon tenella. Also at Station 2. Also the " Porcupine," 1869,
Stations 51, 54, 55, 74 • and 1870, Station 17a.
Station 6. Lat. 60° 9' 0" N., long. 7° 26' 30" W.; 466 fathoms; stones; bottom
temperature, 29°-5 F.
Antedon quadrata. Also at Station 4. Also the " Valorous,"
1875, Station 6; also the " Alert," 1875, Discovery
Bay and Franklin-Pierce Bay ; and the Challenger,
Station 48.
H.M.S. Challenger, 1873-1876.
Station 48. May 8, 1873 ; on the Le Have Bank; lat. 43° 4' N., long. 64° 5' W.;
51 fathoms; rock.
Antedon eschrichti. Also the "Porcupine," 1869, Station 57;
the "Valorous," 1875, Station 1; the "Alert,"
1875, Franklin-Pierce Bay; the "Triton," 1882,
Station 4.
Antedon quadrata. Also the " Valorous," 1 875, Station 6 ; the
"Alert," 1875, Discovery Bay and Franklin-
Pierce Bay; and the "Triton," 1882, Stations
4,6.
St. Paul's Rocks. Lat. 0° 55' 36" N., long. 29° 22' 32" W.
Actinometra pulchella. Possibly at Station 192. Also the
"Porcupine," 1870, Stations 25, 31. Abundant
in the Caribbean Sea.
Station 122. September 10, 1873 ; off Barra Grande ; lat. 9° 5' S., long. 34° 50' W.;
350 fathoms ; red mud.
Atelecrinus balanoides. Also at several Stations in the
Caribbean Sea.
Pentacrinus maclearanus.
358 THE VOYAGE OF H.M.S. CHALLENGER.
Bahia ; 7 to 20 fathoms.
Antedon carinata. Also in the Pacific, Eastern Archipelago, and
Indian Ocean.
Antedon diibeni.
Actinometra lineata.
Actinometra meridionalis. Abundant in the Caribbean Sea.
Station 135e. October 18, 1873; near Tristan da Cunha ; lat. 37° 21' 0" S., long.
12° 22' 30" W.; 1000 fathoms; hard ground, shells,
gravel.
Three Pentacrinoid larvae.
Station 135g. October 18, 1873; off Tristan da Cunha; lat. 37° 10' 50" S.,'long.
12° 18' 30" W.; 550 fathoms; hard ground.
Antedon multispina. Also at Station 344.
Simon's Bay, Cape of Good Hope.
Actinometra parvicirra. Also at Stations 174, 186, Banda,
Ternate, Admiralty Islands, and Samboangan.
Off Marion Island ; 50 to 75 fathoms.
Antedon exigua. Also at Station 145.
Station 145. December 27, 1873; off Marion Island; lat. 46° 43' 0" S., long.
38° 4' 30" E.; 140 fathoms ; volcanic sand.
Antedon exigua. Also in 50 to 75 fathoms.
Antedon hirsuta.
Station 147. December 30; between Marion Island and the Crozets, 1873; lat,
46° 16' S., long. 48° 27' E.; 1600 fathoms; Diatom
ooze ; bottom temperature, 34°-2 F.
Antedon abyssorum.
Antedon bispinosa.
Antedon remota.
Promachocrinus abyssorum. Also at Station 158.
Bathycrinus aldrichianus.
Hyocrinus bethellianus.
REPORT ON THE CRINOIDEA. 359
/Kerguelen Island ; 10 to 100 fathoms.
Station 149c. January 19, 1874 ; Balfour Bay ; 20 to GO fathoms.
Station 149d. January 20, 1874; Royal Sound; lat. 49° 28' S., long. 70° 13' E. ;
/ 28 fathoms.
Station 149e. January 21, 1874; off Greenland Harbour; 30 fathoms.
Station 149h. January 29, 1874; off Cumberland Bay; 127 fathoms.
\ The bottom deposit at all these Stations is volcanic mud.
Promachocrinus herguelensis. Also at Station 151.
Station 150. February 2, 1874 ; near Heard Island; lat. 52° 4' S., long. 71° 22' E.;
150 fathoms; coarse gravel; bottom temperature, 3 5° "2 F.
Antedon australis.
Station 151. February 7, 1874; off Heard Island; lat. 52° 59' 30" S., long.
73° 33' 30" E.; 75 fathoms; volcanic mud.
Antedon antarctica.
Promachocrinus Jcerguelensis. Also at Stations 149c, D, E, h.
Station 158. March 7, 1874; south-west of Melbourne; lat. 50° 1' S., long.
123° 4' E.; 1800 fathoms; Globigerina ooze; bottom
temperature, 33°-5 F.
Promachocrinus abyssorum. Also at Station 147.
Thaumatocrinus renovatus.
Station 160. March 13, 1874; south-west of Melbourne; lat, 42° 42' S., long.
134° 10' E.; 2600 fathoms; red clay ; bottom tempera-
ture, 33°-9 F.
Antedon abyssicola. Also at Station 244.
Port Jackson ; 8 to 10 fathoms.
A ctinometra trichopta -a .
Port Jackson ; 30 to 35 fathoms.
Antedon macronema.
300 THE VOYAGE OF H.M.S. CHALLENGER.
Station 164. June 12, 1874; off Port Jackson; lat. 34° 8' S., long. 152° 0' E.;
950 fathoms ; green mud ; bottom temperature, 36°-5 F.
Antedon spinicirra.
Eudiocrinus semperi. Also at Station 169.
Station 169. July 10, 1874 ; north-east of New Zealand ; lat. 37° 34' S., long.
177° 22' W.; 700 fathoms ; blue mud ; bottom tempera-
ture, 40° F.
Antedon alternata. Also at Stations 170a, 218, and 236.
Eudiocrinus semperi. Also at Station 164.
Station 170a. July 14, 1874 ; near the Kermadec Islands ; lat. 29° 45' S., long.
178° 11' W.; 630 fathoms; volcanic mud; bottom
temperature, 39°'5 F.
Antedon alternata. Also at Stations 169, 218, and 236.
Antedon basicurva.
Antedon breviradia. Also at Station 175.
Antedon echinata.
Antedon insequalis. Also at Station 174 (b, c, or d).
Antedon incerta.
Antedon incisa. Also at Station 174 (b, c, or d).
Pentacrinus naresianus.
Metacrinus (two species).
Tongatabu Reefs.
Antedon regalis.
Actinometra quadrata.
Station 174 (b, c, or d). August 3, 1874 ; near Kandavu, Fiji ; lat. (about) 19° 6' S.,
long, (about) 178° 18' E.; 225, 610, or 210 fathoms;
coral mud ; bottom temperature (at 610 fathoms), 39° F.
Antedon inasqnalis. Also at Station 170a.
Antedon incisa. Also at Station 170a.
Antedon occulta.
Antedon similis.
Antedon tuberculata.
REPORT ON THE CRINOIDEA. 361
Actinometra parvicirra. Also at Station 186, Simon's Bay, Banda,
Ternate, Admiralty Islands, and Samboangan.
Actinometra stelligera.
Actinometra typica.
Atelecrinus ivyvillii.
Station 175. August 12, 1874; near Kandavu, Fiji; lat. 19° 2' S., long. 177° 10' E.;
1350 fathoms ; Globigerina ooze ; bottom temperature,
36° F.
Anteclon acutiradia.
Antedon breviradia. Also at Station 170a.
Doubtful.
Antedon basicurva. Also at Station 170a.
Antedon insequalis. Also at Stations 170a, 174 (b, c, or d).
Pentacrinus naresianus.
Cape York. September 7, 1874 ; Channel between Albany Island and Somerset.
Actinometra paucicirra. ' Also at Station 187, and the Arrou
Islands.
Actinometra pectinata. Also at Samboangan.
Actinometra Solaris. Also at Station 187.
Station 186. September 8, 1874; Prince of Wales Channel; lat. 10° 30' S., long.
142° 18' E.; 8 fathoms ; coral mud.
Antedon microdiscus.
Antedon variipinna. Also at the Arrou Islands.
Actinometra belli.
Actinometra maculata.
Actinometra multiradiata.
Actinometra p>arvicirra. Also at Simon's Bay, Bauda, Ternate,
Admiralty Islands, Samboangan, and Station 174.
Actinometra valida.
Station 187. September 9, 1874 ; off Booby Island ; lat, 10° 36' S., long. 141° 55' K;
6 fathoms ; coral mud.
Antedon midtiradiata.
Actinometra paucicirra. Also at Cape York, and at the Arrou
Islands.
Actinometra Solaris. Also at Station 186.
(ZOOL. OH ALL. EXP. — PART LX. — 1888.) OoO 46
362
THE VOYAGE OF H.M.S. CHALLENGER.
Station 190. September 12, 1874 ; in the ArafuraSea ; lat. 8° 56' S., long. 136° 5' E.;
49 fathoms ; green mud.
Antedon denticulata.
Antedon elegans.
Station 192. September 26, 1874; near the Ki Islands; lat. 5° 49' 15" S., long.
132° 14' 15" E.; 140 fathoms; blue mud.
A ntedon angustiradia.
Antedon compressa. Also at Station 201.
Antedon discoidea.
Antedon Jlexilis.
Antedon longicirra.
Antedon manca.
Antedon parvipinna.
Antedon patida.
Antedon pusilla.
Antedon quinquecostata.
Antedon robusta.
Actinometra. pulchella ?
Metacrinus (five species).
Arrou Islands.
Antedon variipinna. Also at Station 186.
Actinometra pancicirra. Also at Station 187 and Cape York.
Banda ; 1 7 fathoms.
Actinometra coppingeri. Also at Samboangau.
Actinometra divaricata.
Actinometra duplex.
Actinometra elongata.
Actinometra Jimbriata. Also at Station 208.
Actinometra littoralis.
Actinometra multibrachiata.
Actinometra parvicirra. Also at Simon's Bay, Ternate,
Admiralty Islands, Samboangan, and Stations
174, 186.
Actinometra regalis.
Actinometra sentosa.
REPORT ON THE CRINOIDEA. 363
Tern ate, Shore.
Actinometra parvicirra. Also at Simon's Bay, Baucla, Admiralty
Islands, Samboangan, and Stations 174, 186.
Station 201. October 26, 1874; oft' Mindanao, Philippine Islands; lat. 7° 3' N.»
long. 121° 48' E.; 82 fathoms; stones, gravel.
Antedon balanoides.
Antedon compressa. Also at Station 192.
Station 203. October 31, 1874; off Panay ; lat 11° 6' N., long. 123° 9' E.;
20 fathoms ; mud.
Antedon milberti. Also at Station 212.
Station 205. November 13, 1874; off Luzon ; lat. 16° 42' N., long. 119° 22' E.;
1050 fathoms ; blue mud ; bottom temperature, 37° F.
Eudiocrinus varians.
Station 208. January 17, 1875 ; Philippine Islands ; lat, 11° 37' N., long. 123° 31' E.;
18 fathoms ; blue mud.
Antedon informis.
Antedon marginata.
Antedon parvicirra.
Actinometra Jimbriata. Also at Banda.
Actinometra nobilis. Also at Samboangan.
Zebu Reef, Philippines.
Antedon conjungens.
A ntedon disciformis.
Station 210. January 25, 1875 ; off the Panglao and Siquijor Islands; lat. 9° 26' N.,
long. 123° 45' E.; 375 fathoms ; blue mud ; bottom
temperature, 540,1 F.
Antedon distincta.
Antedon tuberosa.
Pentacrinus (two species ?).
Metacrinus (one species ?).
364 THE VOYAGE OF H.M.S. CHALLENGER.
Station 212. January 30, 1875; off Samboangan ; lat. 6° 54' N., long. 122° 18' E.;
1 0 fathoms ; sand.
Antedon anceps.
Antedon clemens.
Antedon milberti Also at Station 203.
Antedon quinduplicava.
Samboangan, Philippines.
Actinometra coppingeri. Also at Banda.
Actinometra distincta.
Actinometra nobilis. Also at Station 208.
Actinometra parvicirra. Also at Simon's Bay, Banda, Ternate,
Admiralty Islands, and Stations 174, 186.
Actinometra pectinata. Also at Cape York. '
Actinometra rotalaria.
Station 214. February 10, 1875 ; off the Meangis Islands ; lat. 4° 33' N., long. 127° 6'
E.; 500 fathoms; blue mud; bottom temperature,
41°-8 F.
Antedon accela.
Antedon actdeata.
Antedon angusticalyx.
Antedon gracilis.
Antedon Isevis.
Antedon valida.
Promachocrinus naresi.
Pentacrinus (two species).
Metacrinus (four species).
Station 218. March 1, 1875 ; north of Papua ; lat. 2° 33' S., long. 144° 4' E.; 1070
fathoms ; blue mud ; bottom temperature, 36°*4 F.
Antedon alternata. Also Stations 169, 170a, and 236.
Station 219. March 10, 1875 ; north of the Admiralty Islands ; lat. 1° 54' 0" S., long.
136° 49' 40" E.; 150 fathoms ; coral mud.
Antedon tenuicirra.
REPORT ON THE CRINOIDEA. 365
Admiralty Islands ; 16 to 20 fathoms.
Actinometra parvicirra. Also at Simon's Bay, Banda, Ternate,
Samboangan, and Stations 184, 186.
Actinometra simplex.
Station 232. May 12, 1875; off Japan; lat. 35° 11' N., long. 139° 28' E.; 345
fathoms ; green mud ; bottom temperature, 410,1 F.
Antedon latipinna.
Station 235. June 4, 1875 ; south of Japan; lat. 34° 7' N., long. 138° 0' E.; 565
fathoms ; green mud ; bottom temperature, 38°-l F.
Fjudiocrinus japonicus.
Pentacrinus (?) mollis.
Station 236. June 5, 1875 ; south of Japan ; lat. 34° 58' N., long. 139° 29' E.; 775
fathoms ; green mud ; bottom temperature, 37°-6 F.
Antedon alternate/,. Also at Stations 169, 170a, and 218.
Station 244. June 28, 1875; east of Japan; lat. 35° 22' N., long. 169° 53' E.;
2900 fathoms ; red clay ; bottom temperature, 35°-3 F.
Antedon abyssicola. Also at Station 160.
Station 308. January 5, 1876; off Tom Bay, Patagonia; lat. 50° 8' 30" S., long.
74° 41' 0" W.; 175 fathoms ; blue mud.
Antedon rhomboidea.
Station 320. February 14, 1876 ; off Monte Video ; lat. 37° 17' S., long. 53° 52' W.;
600 fathoms ; green sand ; bottom temperature, 37°-2 F.
Antedon angustipinna.
Antedon lineata.
Antedon longipinna.
Station 344. April 3, 1876 ; Ascension Island ; lat. 7° 54' 20" S., long. 14° 28' 20" W.;
420 fathoms ; volcanic sand.
Antedon multispina. Also at Station 135g.
Antedon porrecta.
366
THE VOYAGE OF H.M.S. CHALLENGER.
BATHYMETEICAL TABLES.
A Roman numeral opposite the name of a species shows that it also occurs in one
of the other Tables.
Table I.
Antedon acuticirra.
adeonse.
anceps.
articulata.
bidens.
bimaculata.
bipartipinna.
brevicuneata.
carinata, II.-V.
carpenteri.
clemens.
conjungens.
disciformis.
diibeni.
elegans, II.
elongata.
eschrichti, II. -VII.
jlagellata.
gyges.
imparipinna.
impinnata.
indica.
informis.
Isevicirra.
Isevipinna.
Imvissima.
loveni.
ludovici.
macronema, II.
marginata.
microdiscus.
-Species found at depths down to 20 fathoms.
Antedon milberti.
milleri.
multiradiata.
palmata.
parvicirra.
perspinosa.
petasus, II., III.
philiberti.
pinniformis.
protecta.
pumila.
quinduplicava.
regalis.
regime,
reynaudi.
rosacea, II., III.
savignyi.
serripinna.
spicata.
tessellata.
variipinna, II.
Actinometra alternans.
belli.
bcnnetti.
borneensis.
brachiolata.
briareus.
coppingeri.
cumingi.
distincta.
divaricata.
REPORT ON THE CRINOIDEA.
367
Actinometra duplex.
Actinometra paucicirra.
echinoptera.
pectinata.
elongata.
peroni.
Jimbriata.
quadrata.
grandicalyx.
regalis.
japonica.
robustipinna
lineata, II.
rotalaria.
littoralis.
rubighiosa.
maculata.
schlegeli.
magnijica.
seniosa.
meridionalis, II.
-V.
simplex.
multibrachiata.
Solaris.
multifida.
stelligera, V.
midtiradiata.
trichoptera.
nigra.
typica, V.
nobilis.
variabilis.
novse-guinese.
valida.
parvicirra, V.
Totals. — Antedon, 52 specie
s ; Actinometra, 45 species
Table II
Antedon armata.
carinata, I.-V.
dcnticidata.
elegans, I.
eschrichti, I.-VII
macronema, I.
magellanica.
'petasus, I. -III.
Species found at depths between 20 and 50 fathoms.
Antedon prolixa, III. -VII.
quadrata, III. -VI.
rosacea, I. -III.
variipinna, I.
Actinometra lineata, I.
meridionalis, I.-V.
Eudiocrinus in divisus.
Promachocrinus kerguelensis, III., IV.
phalangium, III.-V.
Totals. — Antedon, 13 species ; Actinometra, 2 species.
Table III. — Species found at depths between 50 and 100 fathoms.
Antedon antarctica.
balanoides.
carinata, I.-V.
compressa, IV.
dejecta, IV, V.
Antedon duplex, IV., V.
eschrichti, I.-VII.
hageni, IV., V.
petasus, I., II.
phalangium, I I.-V.
3(58
THE VOYAGE OF H.M.S. CHALLENGER.
Antedon prolixa, II.— VII.
quadrata, II. -VI.
rosacea, L, II.
spinifera, IV., V.
tenella, IV. -VII.
Actinometra blakei, IV., V.
discoidea, IV.
meridionalis, I.-V.
pidchella, IV.-VII.
Promachocrinus herguelensis, II. -IV.
Totals. — Antedon, 15 species; Actinometra, 4 species.
Table IV. — Species found at depths between 100 and 200 fathoms.
A ntedon angustiradia.
australis.
barentsi.
carinata, I.-V.
compressa, III.
defecta, III.-V.
discoidea.
duplex, III.-V.
eschrichti, I.-VII.
exigua.
flexilis.
granulifera.
hageni, III.-V.
hirsuta.
longicirra.
■manca.
parvipinna.
Totals. — An tedon.
Antedon patula.
phalangium, II.— V.
pourtalesi, V.
prolixa, II.-VII.
pusilla.
quadrata, II.-VI.
quinquecostata.
robusta.
rhomboidea.
spinifera, IJI.-V.
tenella, III. -VII.
tenuicirra.
Actinometra blakei, III.-V.
discoidea, III.
meridionalis, I.-V.
pulchella, III.— VII.
Promachocrinus herguelensis, II., III.
29 species ; Actinometra, 4 species.
Table V. — Species found at depths between 200 and 350 fathoms.
Antedon brevipinna.
carinata, I. -IV.
defecta, III., IV.
duplex, III., IV.
eschrichti, I.-VII.
hageni, III., IV.
insequalis 1 ?
incisa 1 ?
Antedon latipinna.
occulta.2
phalangium, II.-IV.
pourtalesi, IV.
quadrata, II. -VI.
similis.2
spinifera, III., IV.
tenella, III. -VII.
1 Most probably obtained from 610 fathoms at Station 174.
2 Most probably obtained from 210 or 255 fathoms at Station 174.
REPORT ON THE CRINOIDEA.
;<;<i
A n tedon tubereulata. 1 Actinometra pulchella, III. -V 1 1 .
Actinometra blakei, III., IV. stelligera,1 I.
meridionalis, I.-IV. typica,1 I.
parvicirra,1 I. Atelecrinus balanoides, VI.
Totals. — Antedon, 15 species; Actinometra, 6 species.
Table VI. — Species found at depths between 350 and 500 fathoms.
Antedon porrecta.
prolixa, II.-VII.
quadrata, II.-IV.
tenella, III. -VII.
tuberosa.
valida.
.Actinometra pulchella, III. -VII.
Atelecrinus balanoides, V.
cubensis.
Eudiocrinus atlanticus.
Promach ocrinus naresi.
Antedon accela.
aculeata.
angusticalyx.
colurnnaris.
cubensis.
distincta.
eschrichti, I.-VII.
gracilis.
hystrix.
leevis.
inultispina, VII.
Totals. — Antedon, 17 species; Actinometra, 1 species.
Table VII. — Species found at depths between 500 and 800 fathoms.
Antedon alternata, VIII.
angustipinna.
basicurva.
breviradia, IX.
echinata.
eschrichti, I. -VI.
insequalis.
incerta,
incisa.
lineata.
longipinna.
lusitanica.
multispina, VI.
Totals. -
Antedon occulta2 1
prolixa, II.-VI.
similis2 ?
tenella, III.-VI.
tubereulata2 %
Actinometra pulchella, III.-VI.
parvicirra2 \
stelligcra2,1.
typica2 ?
A telecrinus wyvillii.
Eudiocrinus japonicus.
semperi, VIII.
-Antedon, 15 species; Actinometra, 1 species.
1 Most probably obtained from 210 or 255 fathoms at Station 174.
2 Only if obtained from 630 fathoms at Station 174, which is improbable.
(ZOOL. CHALL. EXP. — PART LX. 1888.) OcO 47
370
THE VOYAGE OF H.M.S. CHALLENGER.
Table VIII.— Species found at depths between 800 and 1100 fathoms.
Antedon alternata, VII.
spinicirra.
Pentacrinoid larvae of Antedon, sp.
Eudiocrinus varians.
serwperi, VII.
Table IX. — Species found at depths between 1100 and 1500 fathoms.
Antedon acutiradia.
basicurva1 ?
Antedon breviradia, VII.
iniequalis %
Table X. — Species found at depths between 1500 and 1800 fathoms.
A n tedon abyssorum .
bispinosa.
remota.
Promaehocrinus abyssorum.
Thaumatocrinus renovatus.
A.
B.
Table XL — Species found at 2600 and 2900 fathoms.
Antedon abyssicola.
Summary.
I. 97 species found at depths down to 20 fathoms.
II. 17 species found between 20 and 50 fathoms.
III. 20
5 5 5
50 and 100
IV. 34
55 J
100 and 200
V. 22
55 5
200 and 350
VI. 22
55 5
350 and 500
VII. 19
5 5 5
500 and 800
VIII. 5
5 5 J
800 and 1100
IX. 2
J5 )
1100 and 1500
X. 5
55 J
1500 and 1800
XL 1
55 )
2600 and 2900
I. 97 species found at depths down to 20 fathoms.
I. -II. 9 of these descend to 50 fathoms.
I.— III. 5 of which reach 100 fathoms.
L, V. 3 also occur at 210 or 255 fathoms.
Only if obtained from 1350 fathoms at Station 175.
REPORT ON THE CRINOIDEA.
371
I.-V.
I.-VII.
II. -IV.
II.-V.
II.-VII.
III.-IV.
III.-V.
III.-VII.
IV. -V.
V.-VI.
VI.-VII
VII.-VIII.
VII. -IX.
XL
c.
III.
IV.
V.
VI.
VII.
VIII.
IX.
X.
XL
3 of these 5 descend to 350 fathoms.
1 of which descends to 632 fathoms.
4 species descend from 20 to 200 fathoms.
2 of these descend to 220 fathoms.
1 of which descends to 743 fathoms.
9 species descend from 50 to 200 fathoms.
5 of these descend to 350 fathoms.
2 of which descend to 800 fathoms.
1 species descends from 124 to 262 fathoms.
1
1
2
1
1
291 to 422
420 to 550
550 to 1100
630 to 1350
2600 to 2900
I. 86 species only found at depths down to 20 fathoms.
II. 4 species only found at depths of 20 to 50 fathoms.
2
17
5
15
11
2
1
5
1
50 to 100
100 to 200
200 to 350
350 to 500
500 to 800
800 to 1100
1100 to 1500
1500 to 1800
2600 to 2900
An analysis of Summaries B and C shows tbat of twenty-eight Comatula-species
which occur in the abyssal zone, twenty-two are peculiar to it. Seventeen of these
twenty-two belong to the genus Antedon, seven of them to the Tenelhi-group, and the
remainder to the Basicurva-, Spinifera-, and Granulif era-groups, all of which have
flattened rays and plated ambulacra. Furthermore, the only continental species of
Antedmi which extends downwards into the abyssal zone also has plated ambulacra ;
while two of the three littoral species found in the abyssal zone belong to the Tenella-
group, the third being Antedon eschrichti, which is so widely distributed in the
northern circumpolar region.
572
THE VOYAGE OF H.M.S. CHALLENGER.
Table showing the Species of Comatulee which occur in the Abyssal Zone.
Genus.
Species confined to the Abyssal Zone.
Continental Species
occurring in the
Abyssal Zone.
Littoral Species
occurring in the
Abyssal Zone.
Thaumatocrinus,
Atelecrinus,
Eudiocrinus,
Promachocrinus,
Actinometra, .
renovatus.
wyvillii.
C sem/peri.
(_ varians.
abyssorum.
japonicus ?
pulcltella.
Antedon,
A. Species with
Plated Ambulacra.
B. Species of the
TfeweZZa-group.
A. mvltispina.
.
acidirctih'a.
basicurva.
bispinosa.
breviradia.
eckinata.
insqualis.
incerta.
incisa.
lusitanica.
spinieirra.
abyssicola.
abyssorum.
alternata.
angnstipinna.
lineata.
longipinna.
remota.
eschricl
B. prolixa
B. tenella.
iti.
The characteristic abyssal species of Antedon thus belong to two very distinct types. —
(1) That with the bases of the rays flattened laterally and plated ambulacra. It is
rarely represented above 100 fathoms and ranges downward to 1600 fathoms. Eight of
the ten abyssal species are simple forms with but ten arms. This type is represented
in the fossil state by Antedon costata (the Solanocrinus costatus of Goldfuss) from
the White Jura (e) of Southern Germany, as has been already indicated on p. 101.
(2) Delicate ten-armed species allied to Antedon tenella of the Subarctic region, which
has a range in depth of 50 to 740 fathoms. The only Antedon found below 1600
REPORT ON THE CRINOIDEA. 373
fathoms belongs to this group ; and it was obtained at two localities, one in the Southern
Ocean (2600 fathoms), and one in the North Pacific (2900 fathoms).
The only species of Actinometra which extends downwards into the abyssal zone is
common among the Caribbean Islands, and also occurs in the continental region of the
East Atlantic.
Eudiocrinus atlanticus may possibly extend into the abyssal zone, but we have no
definite information on this subject as yet (see p. 79) ; while Eudiocrinus japonicus from
563 fathoms in the North Pacific may possibly also occur as a continental species in Japanese
seas, for Dr. Hilgendorf thinks that his specimen was dredged from 300 or 400 fathoms.
The following list, containing the names of one hundred and twenty species of
Antedon and forty-eight species of Actinometra, embodies the result of our present
knowledge of the Coniatulidas. But many species are still awaiting description, and the
geographical range of others will be greatly extended when the large collections in many
European museums have undergone a critical revision.
On the other hand it is more than probable that some of the names in the following
list will eventually be reduced to the rank of synonyms. Thus, for example, I strongly
suspect that Actinometra meridionalis is identical with the Comatula echinoptera of
Midler, while I have no doubt whatever that some of the following are not good species,
Antedon diibcni, Antedon hageni, Antedon milleri, Antedon petasus, and Antedon
rosacea. But the time has not yet come for a discussion of their mutual relations.
A List of the known Living Species of Comatul.e, showing their
Bathymetrical and Geographical Distribution. ■
Explanation of the Letters used.
A. Species discovered by the " Blake," and other U. S. ships.
B. Previously known species collected by the " Blake," &c.
C. Species discovered by the Challenger.
D. Previously known species collected by the Challenger.
E. Species discovered by the Arctic Expedition, 1875-76.
F. Previously known species collected by the Arctic Expedition.
G. Species discovered by H. M.S. "Alert," 1878-82.
H. Previously known species collected by the " Alert."'
K. Species discovered in the Philippine Islands by Professor Semper.
L. Previously known species collected by Professor Semper.
N. Previously known species collected by the " Voriugen " (Norwegian).
374
THE VOYAGE OF H.M.S. CHALLENGER.
P. Species discovered by the " Porcupine."
Q. Previously known species collected by the " Lightning," " Porcupine," " Knight
Errant," and " Triton."
S. Species discovered by the " Talisman " and " Travailleur " (French).
T. Previously known species collected by the "Talisman" and " Travailleur."
V. Species discovered by the " Varna " (Dutch).
W. Previously known species collected by the " Willem Barents " and " Varna."
X. Previously known species collected by the "Vettor Pisani" (Italian).
Y. Species discovered by the "Tegetthoff" (Austro-Hungarian).
How
obtained.
Eange in
Depth.
Principal Localities.
Fathoms.
Thaumatocrinus, Carp., .
renovatus, Carp.,
C.
1800
Southern Ocean, Station 158.
Atelecrinus, Carp., .
...
balanoides, Carp., .
B. C.
291-422
South-West Atlantic, Station 122; Caribbean
Islands.
cubensis, Pourt., sp.,
A.
450
Near Havana.
■wyvillii, Carp.,
C.
610
Pacific— near Fiji, Station 174c.
Eudiocrinus, Carp., .
atlanticus, Perr., .
s.
486
East Atlantic — Bay of Biscay.
indivisus, Semp., sp.,
K.
30
Pandanon, Philippine Islands.
japonicus, Carp., .
C.
300-565
Pacific — South of Japan, Station 235.
semperi, Carp.,
C.
700-950
South Pacific — off Port Jackson, and north-
east of New Zealand, Stations 164, 169.
f
varians, Carp.,
c.
1050
Pacific — off Luzon, Station 205.
Promachocrinus, Carp.,
abyssorum, Carp., .
c.
1600-1800
Southern Ocean, Stations 147, 158.
kerguelensis, Carp.,
c.
28-120
Near Kerguelen ; and off Heard Island, Station
151.
naresi, Carp.,
c.
500
Pacific — near the Meangis Islands, Station 214.
REPORT ON THE CRINOIDEA.
375
Antedon, de Frem.
Series I.
Elegans-group.
elegans, Bell,
microdiscus, Bell, .
multiradiata, Carp.,
Series II.
1. Basicurva-group
aculeata, Carp.,
acutiradia, Carp., .
basicurva, Carp.,
bispinosa, Carp.,
brevipinna, Pourt.,
breviradia, Carp., .
dentiadata, Carp., .
duplex, Carp., MS.,
echinaia, Carp.,
flexilis, Carp.,
gracilis, Carp.,
incerta, Carp.,
incisa, Carp.,
latipinna, Carp.,
Iniitjkirra, Carp.,
lusitanica, Carp.,
multispina, Carp.,
How
obtained.
D. H. K.
D. H.
C.
C.
C.
C.
C.
A.
C.
C.
A.
C.
C.
C.
C.
C.
C.
C.
P.
C.
Range in
Depth.
Fathoms.
12-49
G-12
Principal Localities.
500
1350
630
1600
270
630-1350
49
88-262
630
140
500
630
610-630
345
140
740
420-550
Torres Strait ; Port Molle ; Arafura Sea,
Station 190; Mergui ; Philippines.
Port Molle and jSTicol Bay, Australia ; Torres
Strait, Station 186.
Torres Strait, Station 187.
Pacific — near the Meangis Islands, Station 214.
Pacific — near Fiji, Station 175.
Pacific — near the Kermadecs, Station 170a.
Southern Ocean, Station 147.
Straits of Florida.
Pacific — near the Kermadecs, Station 170a;
near Fiji, Station 175.
Arafura Sea, Station 190.
Caribbean Islands ; Straits of Florida.
Pacific — near the Kermadecs, Station 170a.
Ki Islands, Station 192.
Pacific — near the Meangis Islands, Station 214.
Pacific — near the Kermadecs, Station 170a.
Pacific — near the Kermadecs, Station 1 70a ;
mar Fiji ; Station 17 4.
Pacific — off Japan, Station 232.
Ki Islands, Station 192.
East Atlantic — off Portugal.
South Atlantic — off Tristan da Cunha, Station
135g ; near Ascension, Station 344.
THE VOYAGE OF H.M.S. CHALLENGER.
Antedon, de Frera., eontd.
parvipinna, Carp., .
How
obtained.
Range in
Depth.
Principal Localities.
C.
Fathoms.
140
Ki Islands, Station 192.
pusilla, Carp.,
C.
140
Ki Islands, Station 192.
spinicirra, Carp., .
C.
950
South Pacific — near Port Jackson, Station 164.
tuberosa, Carp.,
c.
375
Pacific — off the Panglao and Siquijor Islands,
Station 210.
valkla, Carp.,
c.
500
Pacific — near the Meangis Islands, Station 214.
2. Acoela-group.
accela, Carp.,
c.
500
Pacific — near the Meangis Islands, Station 214.
discoidea, Carp.,
c.
140
Ki Islands, Station 192.
3. Eschrichti-group.
antardica, Carp., .
c.
75
Southern Ocean — near Heard Island, Station
151.
australis, Carp.,
c.
150
Southern Ocean — near Heard Island, Station
150.
barentsi, Carp.,
V.
132
Kara Sea.
eschrichti, Mull., sp.,
D. F. Q. W.
20-632
Circumpolar and North Atlantic, Station 48.
magellanica, Bell, .
G. X.
30
Straits of Magellan.
quadrata, Carp.,
D.F.Q.W.Y.
25-410
Smith's Sound, Davis Strait, Barents Sea, Kara
Sea ; and North Atlantic, Station 48.
rhomboidea, Carp.,
c.
175
Straits of Magellan, Station 30S.
4. Tenella-group.
abyssicola, Carp., .
C.
2600-2900
Southern Ocean — South-West of Melbourne,
Station 160; Pacific — East of Japan, Station
244.
abyssorum, Carp., .
C.
1600
Southern Ocean, Station 147.
altemata, Carp.,
C.
630-1070
Pacific — North-East of New Zealand, Station
169; near the Kermadecs, Station 170a ;
North of Papua, Station 218; South of
Japan, Station 236.
angustipinna, Carp.,
C.
600
West Atlantic — off Monte Video, Station 320.
armata, Pourt.,
A.
35
Florida Straits.
REPORT ON THE CRINOIDEA.
377
Antedon, de Frem., contd.
columnaris, Carp.,
cubensis, Pourt.,
diibeni, Bohlsche,
exigua, Carp.,
hageni, Pourt.,
hirsuta, Carp.,
hystrix, Carp.,
Isevis, Carp., .
lineata, Carp.,
longipinna, Carp.,
milleri, Miill., sp.,
petasus, Dub. and Kor., sp.,
phalangium, Miill., sp.,
prolixa, Sladen,
remota, Carp.,
rosacea, Linck, sp.,
tenetta, Retz., sp., .
tenuiciira, Carp., .
5. Milberti-group.
anceps, Carp.,
carinata, Carp.,
carpenteri, Eell,
informis, Carp.,
How
obtained.
Range in
Depth.
Principal Localities.
A.
A.
D.
C.
A.
C.
P.
C
C
c
Q
Q. T.
N. W. Y.
C.
Q. X.
B. Q. W.
C.
B. D. X.
G.
C.
Fathoms.
422
450
20
50-175
82-242
140
320-430
500
600
600
20-100
30-220
25-743
1600
100
50-740
150
10
7-278
7
18
Off St. Lucia.
Florida Straits.
Bahia ; Rio Janeiro.
Southern Ocean — off Marion Island, and Station
145.
Caribbean Islands and Straits of Florida.
Southern Ocean — near Marion Island, Station
145.
Fseroe Channel.
Pacific — near the Meangis Islands, Station 214.
West Atlantic— off Monte Video, Station 320.
West Atlantic— oS Monte Video, Station 320.
Milford Haven.
North-East Atlantic.
East Atlantic — Hebrides to Gibraltar and
Mediterranean.
Smith's Sound; Kara Sea; North-East Atlantic.
Southern Ocean, Station 147.
East Atlantic — Hebrides to Madeira and Medi-
terranean.
Barents Sea ; Kara Sea ; Scandinavia ; Fseroe
Channel ; North Atlantic — off Coasts of
Portugal and New England.
Pacific — North of Admiralty Islands, Station
219.
Pacific — off Samboangan, Station 212.
Brazil; Venezuela; St Lucia; Chile; Java (?);
Ceylon ; Seychelles ; Red Sea ; Zanzibar ;
Mauritius ; Madagascar ; St. Helena.
Queensland.
Philippine Islands, Station 208.
(ZOOL. CHALL. EXP. — PAET LX. — 1888.)
Ooo 48
378
THE VOYAGE OF H.M.S. CHALLENGER.
Anteclon, de Frtkn., contd.
Ixvissima, Grube, sp.,
loveni, Bell, .
milberti, Miill., sp.,
parvicirra, Carp., .
perspinosa, Carp., .
[liiniiformis, Carp.,
pumila, Bell, .
serripinna, Carp., .
tessellata, Miill., sp.,
variipinna, Carp., .
Unclassified species.
adeonx, Lam., sp.,
halanoides, Carp., .
bidetis, Bell, .
defecta, Carp., MS.,
impinnata, Carp., MS., .
Ixvipinna, Carp., .
Series III.
6. Spinifera-group.
brevipinna, Pourt.,
compressa, Carp., .
duplex, Carp., MS.,
fleseilis, Carp.,
lusitanica, Carp., .
macronema, Miill., sp., .
How
obtained.
Bange in
Depth.
G.
D. H.
C.
H.
G.
C. H.
Fathoms.
3-4
3-20
18
8-36
82
10
77-242
15
A.
270
C.
82-140
A.
88-262
C.
140
P.
740
D.
35
Principal Localities.
Borneo.
Queensland.
Pacific — off Panay, Station 203 ; off Sam-
boangan, Station 212. Queensland; Torres
Strait ; Ceram ; Borneo ; Mergui.
Philippine Islands, Station 208.
Jobie.
New Guinea ; North-West Australia.
Port Jackson ; Port Stephens ; Nelson's Bay.
New Guinea.
"Indien."
Torres Strait, Station 186, and Arrou Islands.
Arafura Sea ; Borneo ; Canton.
Queensland.
Pacific — off Mindanao, Station 201.
Torres Strait.
Caribbean Islands.
Mauritius.
Canton.
Straits of Florida.
Ki Islands, Station 192 j off Mindanao, Station
201.
Caribbean Islands ; Straits of Florida.
Ki Islands, Station 192.
East Atlantic — off Portugal.
King George's Sound ; Port Jackson ; Port
Stephens.
REPORT ON THE CRINOIDEA.
379
Antedon, de Fr&n., contd.
patula, Carp.,
How
obtained.
Range in
Depth.
C.
Fathoms.
140
pourtalem, Carp., MS., .
A.
124-262
quinquecostata, Carp.,
C.
140
robiwta, Carp.,
C.
140
spinifera, Carp.,
B.
80-297
7. Palmata-group.
xquipinna, Carp.
articulata, Miill., sp.,
H.
bimaculata, Carp.,
brevieuneata, Carp.,
clemens, Carp.,
C.
10
conjungens, Carp., .
C.
disciformis, Carp.,
C.
elongata, Mull., sp.,
flagellata, Mull., sp.,
gyges, Bell, ....
G.
imparipinna, Carp.,
...
indica, Smith, sp., .
Ixvicirra, Carp.,
manca, Carp.,
C.
140
marginata, Carp., .
c.
18
occulta, Carp.,
c.
210 or 255
palmata, Miill., sp.,
X.
protects, Liitk., MS.,
...
regalis, Carp.,
c.
regime, Bell, ....
G.
similis, Carp.,
c.
210 or 255
Principal Localities.
Ki Islands, Station 192.
Caribbean Islands.
Ki Islands, Station 192.
Ki Islands, Station 192.
Caribbean Islands.
Moluccas ; Queensland.
Amboina.
Amboina.
Pacific — off Samboangan, Station 212.
Zebu Reefs.
Zebu Reefs.
New Guinea.
?
Torres Strait.
?
Rodriguez.
Arrou Islands.
Ki Islands, Station 192.
Philippine Islands, Station 208.
Pacific — near Fiji, Station 174.
Red Sea ; Ceylon.
Fiji; Tonga.
Tongatabu Reefs.
Queensland.
Pacific — near Fiji, Station 174.
380
THE VOYAGE OF H.M.S. CHALLENGER.
Antedon, de Freni., contd.
spicata, Carp.,
How
obtained.
Eange in
Depth.
Principal Localities.
Fathoms.
Banda Sea ; Ugi.
tuherculata, Carp., .
C.
210 or 255
Pacific — near Fiji, Station 174.
Series IV.
8. Granulifera-group.
angusticahjx, Carp.,
C.
500
Pacific — off the Meangis Islands, Station 214.
distinda, Carp.,
C.
01 0
Pacific — off the Panglao and Siquijor Islands,
Station 210.
granulifera, Pourt.,
A.
101-120
Caribbean Islands.
ineequalis, Carp., .
C.
610-630
Pacific — near the Kermadecs, Station 170a ;
near Fiji, Station 174.
multispina, Carp., .
C.
420-550
South Atlantic — off Tristan da Cunha, Station
135g ; near Ascension, Station 344.
porreda, Carp.,
c.
420
South Atlantic — near Ascension, Station 344.
9. Savignyi-group.
acuticirra, Carp., .
1
anceps, Carp.,
c.
10
Pacific — off Samboangan, Station 212.
angustiradia, Carp.,
c.
140
Ki Islands, Station 192.
hipartipinna, Carp.,
Hong Kong.
ludnvici, Carp.,
Hong Kong.
philiberti, Miill., sp.,
Java.
quinduplicava, Carp.,
c.
10
Pacific — off Samboangan, Station 212.
reynaudi, Miill., sp.,
Ceylon.
savignyi, Miill., sp.,
Red Sea ; Kurrachee.
variipinna, Carp., .
C. H.
8-36
Torres Strait, Station 186, and Arrou Islands.
Arafura Sea; Borneo; Canton.
Actinometra, Miill.
Series I.
1. Solaris-group.
brachiolata, Lam., sp., .
Australia.
REPORT ON THE CRINOIDEA.
381
Actinometra, Mull., contd.
pedinata, Retz., sp.,
How
obtained
Range in
Depth.
Principal Localities.
D. H. L.
Fathoms.
8-12
Java; Singapore; Moluccas; Celebes; Arafura
Sea ; Torres Strait ; Queensland ; Bohol ;
Samboangan.
Solaris, Lam., sp., .
2. Paucicirra-group.
D. II.
6-12
Singapore ; Hong Kong ; Torres Strait, Station
187 ; Queensland.
pauddrra, Bell,
3. Typica-group.
D. H.
3-12
Arrou Islands; Torres Strait, Station 187;
Queensland.
distinda, Carp.,
C.
10
Samboangan.
multibradiiata, Carp., .
C.
17
Banda.
novx-guinex, Mull., sp., .
Eidouma, New Guinea.
typica, Loven, sp., .
D.
210 or 255
Pacific — near Fiji Station 174. Malacca;
Jobie; Zebu; Kingsniills Islands.
Series II.
4. Bchinoptera-group.
blakei, Carp., MS.,
A.
62-262
Caribbean Islands.
cumingi, Miill., sp.,
H.
Malacca ; Queensland.
echinoptera, Miill., sp., .
...
1
meridionalis, Pourt., sp.,
A. D.
7-262
Florida Straits ; Caribbean Islands ; Brazil.
puldiella, Pourt., sp.,
B. D. P. T.
73-830
Caribbean Islands ; St. Paul's Rocks ; East
Atlantic — near Gibraltar, and off Rochefort.
Ki Islands, Station 192 (1).
rubiginosa, Pourt., sp., .
A.
9-17
Bahamas ; Florida Straits.
Series III.
5. Stelligera-group.
maculata, Carp.,
C.
8
Torres Strait, Station 186.
nigra, Carp., MS., .
K.
Philippines.
puldiella, Pourt., sp.,
B. D. P. T.
73-830
Caribbean Islands ; St. Paul's Rocks ; East
Atlantic — near Gibraltar, and off Rochefort.
Ki Islands, Station 192 (?).
stelligera, Carp.,
D.
210 or 255
Pacific — near Fiji, Station 174. Tonga; Samoa;
Reef of Atagor.
382
THE VOYAGE OF H.M.S. CHALLENGER.
Actinometra, Miill., contd.
6. Valida-group.
How
obtained.
Range in
Depth.
Principal Localities.
Fathoms.
elongata, Carp.,
C.
17
Banda.
rotalaria, Lam., sp.,
D.
10
Samboangan ; Australia.
simjilex, Carp.,
C.
16-25
Admiralty Islands.
valida, Carp.,
C.
8
Torres Strait, Station 186.
Series IV.
7. Fimbriata-group.
borneensis, Grube, .
North Borneo.
mp'pingeri, Bell,
C. H.
10-17
Banda and Samboangan. Singapore; Arnboina;
China Sea; Queensland.
diseoidea, Carp.,
A.
8S-118
Caribbean Islands.
firnbriata, Lam., sp.,
D.
17-18
Banda and Philippine Islands. Sunda Strait ;
Java ; Nicobar Islands ; Madagascar (?).
lineata, Carp.,
B.C.
7-40
Brazil ; Barbados.
multiradiata, Linn., sp.,
D. L.
8
Torres Strait, Station 186. Philippines; China
Sea ; Japan.
sentosa, Carp.,
D.
17
Moluccas.
8. Parvicirra-group.
altemans, Carp.,
H.
12-20
Queensland.
belli, Carp., ....
C.
8
Torres Strait, Station 186.
bemictti, Bohlsche, .
Loyalty Islands ; Pelew Islands ; Sooloo Sea ;
Singapore.
briareus, Bell, sp., .
G.
Queensland.
divaricata, Carp., .
C.
17
Banda.
duplex, Carp.,
C.
17
Banda.
grandkalyx, Carp.,
Canton.
japcmica, Miill., sp.,
•
Japan.
littoralis, Carp.,
C.
17
Banda.
magnified, Carp., MS., .
K.
Philippines.
REPORT ON THE CRINOIDEA.
383
Actinometra, Mull., contd.
multifida, Miill., sp.,
How
obtained.
Range in
Depth.
Principal Localities.
H.
Fathoms.
Queensland ; Torres Strait.
ndbilis, Carp.,
C.
10-18
Philippine Islands — Samboangan, and Station
208.
parvicirra, Miill., sp.,
D. H. L.
8-210 or
255
Pacific — near Fiji; Station 174. Torres Strait,
Station 186. Borneo; Moluccas; Admiralty
Islands ; Philippines ; Ceylon ; Nicobar
Islands; Natal; Cape of Good Hope; China
Sea ; Japan ; Kingsmills Islands ; Fiji ;
Friendly Islands ; Peru.
peroni, Carp.,
...
Ceram.
quadrata, Carp.,
C.
Tongatabu.
regalis, Carp.,
C.
17
Banda.
robustipinna, Carp.,
Moluccas.
scldegeli, Carp.,
East Indies (?).
trklmptera, Miill., sp., .
D.
10-12
Port Jackson ; Port Thilip ; King George's
Sound.
variabilis, Bell,
G.
Torres Strait.
Analysis of the above List.
Genus.
Number of
Living
Species.
" Porcupine."
" Challenger."
"Alert."
New
Species.
Species
previously
known.
New-
Species.
Species
previously
known.
New
Species.
Species
previously
known.
Thaumatocrinus, .
Atelecrinus, .
Eadiocrinus, .
Promaclwcrinus, .
Antedon,
Actinometra, .
1
3
5
3
120
48
2
1
G
1
2
3
3
64
15
8
14
7
2
7
8
180
3
6
88
22
9
15
3S4 THE VOYAGE OF H.M.S. CHALLENGER.
Two facts shown in the above Table are worth notice : —
1. Kepresentatives of each of the six living genera of Comatulse were obtained by
the Challenger, two of thern being new to science.
2. Five-ninths of the recognised living species of Comatulse have been discovered by
British ships. This proportion will be largely modified, however, when the "Blake"
collection and those of the Continental Museums have been properly worked up.
SUPPLEMENTAL BIBLIOGRAPHY OF THE NEOCRINOIDEA.
The following list, which has been compiled upon the same principles as that given
in Part I., carries the Bibliography of the Neocrinoidea down to the end of April
188S. It also contains a few titles which had escaped notice when the first list was
compiled.
Agassiz, A., The Affinities of Crinoids. American Naturalist, 1872, vol. vi. pp. 305-306.
[Anonymous], Notes and Observations on Injured or Diseased Crinoids. By a Corresponding Member.
Second Paper. Proc Nat. Hist. Soc. Glasgow, 1878, vol. iii. pp. 333-339.
Notes and Observations of Additional Structures on Crinoid Stems. By a Corresponding Member.
Third Paper. Ibid., 1878-1880 [1881], vol. iv. pp. 73-77.
Barrett, L., On two Species of Echinodermata new to the Fauna of Great Britain. Ann. ami Mag. Nat. Hist.,
1857, ser. 2, vol. xix. pp. 32, 33.
Barrois, J., Sur l'embryogenie de la Comatule (C. mediterranea). Comptes rendus, 1886, t. cii.
pp. 1176-1177.
Des homologies des larves de Comatules. Comptes rendus, 1886, t. ciii. pp. 892, 893.
Recherches sur le developpement de la Comatule (C. mediterranea). Bee. zool. Suisse, 1888, t. iv.
pp. 545, pis. xxv.-xxxi.
Bell, F. J., Notes on a Collection of Echinodermata from Australia. Proc. Linn. Soc. N.S.W., 1884 [1885],
vol. ix. pp. 496-507.
■ The Echinoderm Fauna of the Island of Ceylon. Trans. Dublin Soc., 1887, pp. 643-657,
pis. xxxix., xl.
Bury, H., The early Stages in the Development of Antedon rosacea. Rep. Brit. Assoc, 1887, pp. 735, 736 ;
and Proc. Boy. Soc, 1888, vol. xliii. pp. 297-299.
Carpenter, P. H., On some Points in the Morphology of the Echinodernis, and more especially of the
Crinoids. Ann. and Mag. Nat. Hist., 1885, ser. 5, vol. xvi. pp. 100-119.
On the Geographical and Bathymetrical Distribution of the Crinoidea. Bep. Brit. Assoc, 1884,
pp. 758-760.
An Encysting Myzostoma in Milford Haven. Nature, 1885, vol. xxxii. p. 391.
The new British Myzostoma. Nature, 1885, vol. xxxiii. p. 8.
On the Variations in the Form of the Cirri in certain Comatuhe. Trans. Linn. Soc Land. (Zool.),
1886, ser. 2, vol. ii. pp. 475-480, pi. lvii.
The Comatulae of the ""Willeni Barents" Expeditions, 1880-1884. Bijdragen tot de Dierkunde,
1886, Afl. 13, pp. 1-12, pi. i.
Fossil Crinoids. Ann. and Mag. Nat. Hist., 1886, ser. 5, vol. xvii. pp. 276-289; Ibid., vol. xviii.
pp. 406-412.
The Morphology of Antedon rosacea. Ann. and Mag. Nat. Hist., 1887, ser. 5, vol. xix.
pp. 19-41.
(zool. chall. exp. — part lx. — 188S.) Ooo 49
386 THE VOYAGE OF H.M.S. CHALLENGER.
Carpenter, P. H., Notes on Echinoderm Morphology, No. X. On the supposed presence of Symbiotic Algae in
Antedon rosacea. Quart. Journ. Micr. Sci., 1887, vol. xxvii. pp. 379-391.
The Generic Position of Solanocrinus. Ann. and Mag. Nat. Hist., 1887, ser. 5, vol. xix. pp. 81-88.
Professor Perrier's Historical Criticisms. Zool. Anzeiger, 1887, Jahrg. x. pp. 57-62, and 84-88.
The supposed Myzostoma-cysts in Antedon rosacea. Nature, 1887, vol. xxxv. p. 535.
Zoologische Bijdragen tot de Kennis der Karazee. 11. Report on the Coinatuke. Bijdragen tot de
Dierkunde, 14 Aflevering, pp. 39-49, 1 pL
Further Remarks upon Professor Perrier's Historical Errors. Zool. Anzeiger, 1887, Jahrg. x. pp.
262-265.
On Crinoids and Blastoids. Proc. Geol. Assoc, 1887, vol. x. pp. 1-10.
Dendt, A., Description of a Twelve-armed Comatula from the Firth of Clyde. Proc. Roy. Physic. Soc.
Edin,, 1886, vol. ix. pp. 180-182, pi. x.
On the Regeneration of the Visceral Mass in Antedon rosaceus. Stud. Biol. Lab. Owens College,
1886, vol. i. pp. 299-312.
Deshayes, G. P., Rapport sur une Encrine vivant donnoe an Museum par M. Schramm, inspecteur des douanes
a la Guadeloupe. Nouv. Arch. Mus. Paris, 1870, t. vi. Bull., pp. 3-6.
Eck, H, Bemerkungen tiber einige Encrinus-A.vt<a.\. Zeitschr. d. deutsch. geol. Gesellsch., Jahrg. 1887,
pp. 540-558.
Etheridge, R., jun., and Carpenter, P. H, Catalogue of the Blastoidea in the Geological Department of the
British Museum (Natural History), with an account of the Morphology and Systematic Position of the
Group, and a Revision of the Genera and Species. London, 1886, pp. 1-322, pis. i.-xx.
Fischer, F., Echinodermen von Jan Mayen. Die Osterreichische Polarstation Jan Mayen. Bd. iii., Vienna,
1886, pp. 29-38.
Fritsch, C. Freiuerr von, Uber Encrinus Carnalli, Beyr. Zeitschr. f. Naturwiss., 1887, Bd. Ix. pp. 83, 84.
Graff, L. von, Ueber einige Deformitiiten an fossilen Crinoiden. Palxontographica, 1885, Bd. xxxi. pp.
185-191, Taf. xvi.
Report on the Myzostomida. Supplement. Zool. Chall. Exp., part Ixi., 1887, pp. 1-16, pis. i.-iv.
Gray, J. E., Notes on Holopusand Pentacrinus. Ann. and Mag. Nat. Hist., 1871, ser. 4, vol. viii. pp. 394-396.
Hall, J., On the occurrence of an internal convoluted plate within the body of certain species of Crinoidea.
Proc. Boston Soc. Nat. Hist, 1864-66, vol. x. pp. 33-34.
Hamann, 0., Die Wandernden Urkeimzellen unci ihre Reifungsstatten bei den Echinodermen. Zeitschr. f.
wiss. Zool,, 1887, Bd. xlvi. pp. 80-98, Taf. xi.
Hartog, M. M., On the True Nature and Function of the Madreporic System in Echinodermata. Rep. Brit.
Assoc, 1887, p. 736.
The True Nature of the " Madreporic System " of Echinodermata, with Remarks on Nephridia. Ann.
and Mag. Nat. Hist, 1887, ser. 5, vol. xx. pp. 321-326.
Herdman, "W. A., Report upon the Crinoidea, Asteroidea, Echinoidea, and Holothuroidea of the L. M. B. C.
District. First Report on the Fauna of Liverpool Bay, Liverpool, 1886, Svo; also, Proc. Lit. Phil. Soc.
Liverpool, 1886, vol. xl., Appendix, pp. 131-139.
Jeffreys, E. Gwyn, On a Pentacrinus (P. Wyville-Thomsoni) from the Coasts of Spain and Portugal. Rep.
Brit Assoc, 1870, p. 119.
Koenen, A. von, Beitrag zur Kenntniss der Crinoiden des Muschelkalks. Abhamdl, d. Kgl, Gesellsch. d, Wiss.
zu Gottingen, 1887, Bd. xxxiv., 44 pp., 1 Taf.
Ueber Muschelkalk Encriniten. Neues Jahrb. f. Mineral., 18S7, Bd. ii. pp. 86-88.
LacazeJJuthiers, H. de, Note sur une station d'une Encrine vivante (Pentacrinus Europaeus) sur les cotes de
France. Comptes rendus, 1869, t. lxix. pp. 1253-1256.
Levinsen, G. M. R, Kara-Havets Echinodermata. Dijmphna-Togtets zoologisk-botaniske Udbytte ;
Kjobenhavn, 1887, Svo, pp. 383-418, Tab. xxxiv., xxxv. ; also separately, Copenhagen, 1886, 8vo.
Loriol, P. de, Note sur le genre Millericrinus. Assoc Franc-, pour V avancement des Sciences. Compte rendu de
la 13me Session, 1884, part ii. pp. 247-252.
REPORT ON THE CRINOIDEA. 387
Loriol, P. de, Coup d'ceil d'ensemble sur les Crino'ides recueillis dans les couches Jurassiques de la France.
Ibid., 14me Session. Grenoble, 1885, part ii. pp. 3G4-371.
Paleontologie Franchise. Terrain Jurassique, Crino'ides, t. xi. lfre Partie, Paris, 1882-1884, 627 pp.
pis. i.-cxxi. 2me Partie, Paris, 1885-1888, pp. 1-352, pis. cxxi.-ccvii. (unfinished).
Notes sur quelques Eehinoderines fossiles des environs de la Rochelle. Ann. de la Soc. des Sci. Nat.
de la Rochelle, 1887, vol. xxiii. pp. 1-12, pis. i.-iii.
Marshall, A. M., [On the Nervous System of Antedon]. Rep. Brit. Assoc, 1884, pp. 256-258.
Peach, C. W., On the Occurrence of the " Rosy Feather Star " (Comatula [Antedon] rosacea) on the Eastern
Shores of Scotland, especially on that of Caithness. Proc. Roy. Physic. Soc. Edin., 1864, vol. iii. pp.
81-83.
Perhier, E., Note sur FAnatornie de la Comatule (Comatula [Antedon] rosacea, de Blainville). Comptes rendus,
1873, t. lxxvi. pp. 718-720.
Sur le developpement de l'appareil vasculaire et de l'appareil genital des Comatules. Comptes rendus,
1885, t. c. pp. 431-434.
Resume des Recherches sur l'organogenie et l'anatomie des Comatules. Zool. Anzeiger, 1885, Jahrg.
viii. pp. 261-269.
Les Encrines vivantes. Revue Scientifique, 1885, t. xxxv. pp. 690-693.
Les Explorations sous-marines. Paris, 1886, 352 pp.
Memoire sur l'Organisation et le Developpement de la Comatule de la Mediterrante (Antedon rosacea,
Linck.). Nouv. Arch,. Mus. Paris, 1886-87, t. ix. pp. 1-300, pis. i.-x.
Reponse a M. Herbert Carpenter. Zool- Anzeiger, 1887, Jahrg. ix. pp. 145-147.
Preyer, W., Ueber die Bewegungen der Seesterne. Zweite Hiilfte. Mittheil. d. Zool. Stat, zu Neapel, 1887,
Ed. vii. pp. 191-233, Taf. vii.
Rathbun, R., Notice of a Collection of Stalked Crinoids made by the Steamer Albatross in the Gulf of Mexico
and Caribbean Sea, 1884 and 1885. Proc. U. S. Nat. Mas., 1885, pp. 628-635.
Rope, J., Note on the Cause and Nature of the Enlargement of some Crinoidal Columns. Geol. Mag., 1869,
vol. vi. pp. 351-353.
Vogt, C, and Yung, E., Traite dAuatoniie Comparee Pratique. Crino'ides in Livr. 7, 8. Paris, 1886,
pp. 518-572.
Wachsmuth, C, and Springer, F., Revision of the Palseocrinoidea, Part III., First Section. Proc. Acad.
Nat. Sci. Philad., 1885, pp. 225-360, pis. iv.-ix. Second Section. Ibid., 1886, pp. 64-227. (Also
published separately, with index.)
The Summit Plates in Blastoids, Crinoids, and Cystids, and their Morphological Relations. Proc.
Acad. Nat. Sci. Philad., 1887, pp. 82-114, pi. iv.
Wagner, R., Die Encriniten des Unteren Wellenkalkes von Jena. Jenaische Zeitschr., 1886, Bd. xx. N.F.
xiii. pp. 1-32, T. i., ii.
Ueber Encrinus Wagneri, Ben. aus dem unteren Muschelkalk von Jena. Zeitschr. d. deutsch. geol.
Gesellsch., Jahrg. 1887 [1888], pp. 822-828.
Walther, J., Untersuchungen liber den Bau der Crinoiden mit besonderer Beriicksichligung der Formen aus
dem Solenhofener Schiefer und dem Kelheimer Diceraskalk. Palseoulographica, 1886, Bd. xxxii. pp.
155-200, Taf. xxiii.-xxvi.
INDEX TO AUTHORS QUOTED.
Agassiz, A., 170, 267, 301.
Agassiz, L., 2, 64, 86, 89, 172, 266.
Alder, J., 170.
Allman, G. J., 19.
Barrett, L., 56, 57, 87, 151, 158, 160.
Bell, F. J., 43, 44, 46-53, 55, 56, 59, 60, 88, 90, 95,
97-99, 137, 138, 149, 159, 160, 170, 176, 181,
193, 194, 199, 200, 206, 212, 256, 258-261,
2.64, 265, 267, 279, 282-285, 288, 290-293,
296, 301, 304, 313, 317, 320-322. 333, 338,
341, 345.
Blainville, H. M. J. de, 63, 86, 89, 199, 266, 286,
288, 313, 317, 322.
Bohlsche, W., 3, 42, 53, 87, 181, 183, 267, 271.
Bronn, H. G., 64, 65, 86, 87, 267.
Carpenter, P. H., 2-5, 8, 10, 14, 16-20, 22, 42, 49-51,
53, 57, 59, 60, 66, 68, 70, 72-74, 78, 80-82,
87, 88, 90, 109, 137-139, 142, 149, 151-153,
158, 159, 162, 165, 170, 181, 194, 199, 211,
212, 229, 256, 257, 262, 264, 272, 275, 279,
281, 282, 285, 287, 288, 290-293, 296, 301,
304, 305, 312, 313, 317, 320, 322, 324-327,
338, 341, 345, 348-352.
Carpenter, W. B., 6, 11, 12, 16-21, 23, 77, 80, 87, 90,
169, 182, 261.
Cams, J. V., 88, 90.
Claus, C, 88, 267.
Cuvier, G, 86, 266.
Dendy, A., 90, 110.
Diiben and Koren, 41, 57, 86, 170, 172, 173, 175.
Dujardin, F, and Hupe, H., 3, 41, 42, 65, 87, 89, 91,
138, 158, 170, 194, 197, 199, 201, 212, 267,
270, 279, 284, 288, 313, 317, 322, 324, 338,
345.
Duncan, P. M., and Sladen, W. P., 57, 88, 90, 138,
140, 149, 151, 153, 174.
Etlieridge, E., and Carpenter, P. H., 27, 28.
Filhol, H., 35, 306.
Fischer, F., 139, 149, 170, 174-178.
Fleming, J., 86, 89.
Fontannes, F., 87.
Forbes, E., 86, S7.
Fraas, O., 87, 89.
Freniinville, C. P. de, 1, 2, 64, 85, 88-91, 199, 201.
Geinitz, H. B., 87.
Goldfuss, G. A., 2, 4, 5, 63, 64, 86, 89, 172, 323.
Graff, L. von, 53, 56, 96, 198, 262, 325.
Gray, J. E., 86, 89.
Greeff, E., 17, 18, 90, 181.
Griffith, E., 199.
Grube, E., 42, 53, 194, 197, 267, 271, 317, 321, 322,
341.
Gunther, A., 284.
Hagenow, F. von, 86.
Hamann, O., 182, 343.
Herdman, W. A., 90.
Herklots, J. A, 278, 288.
Hoffmann, C. K., 138.
Jickeli, C. R, 17, IS.
Kcenen, A. von, 28.
Lamarck, J., 1-3, 41, 52, 59, 85, 88, 89, 91, 172, 199,
201, 266, 268-270, 279, 286, 288, 313, 317,
322, 323.
Leach, W. R, 1, 2, 85, 88, 89, 91, 199, 200.
Levinsen, G. M. E., 88, 139, 142, 143, 149, 153, 155,
168, 169.
Linck, J. H., 1, 85, 89, 286.
Linnanis, C, 1, 41, 85, 89, 169, 171, 172, 266, 284,
286, 322.
Llhuyd, R, 1, 89.
Loriol, P. de, 37, 65, 73, 76, 87.
Loven, S, 4, 14, 15, 87, 89, 97, 13S, 267, 272, 276,
296-298.
Ludwig, H., 17, 18, 28, 88, 90, 159, 164, 199, 202,
267, 301.
Lundgren, B., 87.
390
THE VOYAGE OF H.M.S. CHALLENGER.
Liitken, C. F., 3, 42, 53, 87, 90, 91, 138, 170, 199,
201, 267, 271, 272, 285, 296, 297, 315, 338.
M' Andrew, R., and Barrett, L., 158, 160.
Marenzeller, E. von, 57, 87, 90, 149, 151, 153, 170,
174, 175.
Marion, A. F., 90, 159-163.
Marshall, A. M., 17, 18, 20, 90.
Michelotti, G., 87.
Miller, J. S., 2, 85, 89, 182, 318.
Miiller, J., 2-4, 17, 41-43, 49, 52, 56, 59, 64, 80, 86,
89, 138, 140, 158, 160, 170, 172, 173, 193-196,
199,201,212,252,266-271,278-281,284,286-
288, 298, 313, 314, 317, 318, 322-326, 338,
340, 345.
Minister, G. von, 86, 89.
Nansen, F., 167, 176.
Norman, A. M., 2, 3, 42, 87, 89-91, 158, 170.
d'Orbigny, A, 2, 23, 63, 64, 86, 87.
Pausanias, 91.
Pennant, T., 85.
Perrier, E., 4, 73, 74, 76-81, 88, 90.
Pliilippi, R. A., 86.
Pictet, F. J., 87.
Pourtales, L. F. de, 5, 42, 53, 57, 59, 60, 68, 70, 72,
87, 90, 199, 201, 202, 211, 239, 247, 248, 267,
300, 301, 304, 305.
Quenstedt, Aug., 87, 88, 93, 101, 138, 140, 267.
Rathbun, R., 90, 199, 201, 202, 267, 301, 327, 328.
Retzius, A. J., 41, 57, 85, 169, 171-173, 266, 268,
279, 284-287, 322, 323, 326.
Sars, M., 87, 142, 168-170, 177, 178.
Say, T., 57, 85, 169.
Schlotheim, E. F. von, 85, 89.
Schluter, C, 70, 74, 87, 90.
Semper, C, 3, 4, 65, 73, 74, 78, 79, 83, 304, 338, 341.
Sladen, W. P., 151-153, 155, 166, 167, 174.
Smith, A. E., 210, 232.
Stebbing, T. R. R., 90.
Stinipson, W., 138, 140.
Stuxberg, A., 88, 139.
Teuscher, R., 17, 18.
Thompson, J. V., 85, 86, 90.
Thomson, C. W., 6, 19, 87, 90, 138, 142, 144, 160,
161, 170.
Troschel, F. H., 279, 287, 314, 323, 340.
d'Urban, W. S. M., 138, 149, 170.
Valenciennes, A., 196, 345.
Verrill, A. E., 138, 170, 172, 181, 194, 197, 199, 201,
301.
Vogt, C, and Yung, E., 17-21, 90, 92, 309.
Wachsmuth, C, and Springer, F., 9-13.
Wagner, R., 27.
Walker, D., 138.
Walther, J., 52, 88, 90, 93, 101, 135.
Wright, E. P., 87, 90.
Zittel, K., 65, 88.
GENERAL INDEX.
The figures in dark type indicate the page on which the genus or species is first described.
Synonyms are distinguished by an asterisk *.
AcffiLA-group, 34, 99, 131.
Actinocrinus, 1.
Actinometea, 1-5, 7, 13-16, 23-27, 30, 31, 34-39,
41-44, 47, 52, 57-62, 64-66, 74-79,
89, 91, 134, 141, 171, 203, 208, 228,
241, 251, 266-277, 292, 297, 298,
301-303, 309, 311, 315, 317,329,
333, 342, 344, 349, 372, 373, 383 ;
bathymetrical distribution of, 35,
36; centro-dorsal of, 7, 13-16, 38,
39, 78, 276, 287, 290, 293, 302,
307; disk of, 268-270, 272-275,
290, 294, 297, 319; geographical
distribution of, 35, 36, 283, 284,
329, 344; radials of, 9, 12-16, 18,
19, 21-28, 37-39; terminal comb
of, 4, 42, 92, 271, 276, 343; Series I.,
277; Series II., 300; Series III.,
302 ; Series IV., 315.
*affinis, 59, 285, 287.
*alata, 262, 305.
*albonotata, 59, 288, 290.
altemam, 46, 48, 50, 58, 61, 329,
330, 333, 366, 382.
*annuluta, 60, 338, 341, 342.
*armata, 338, 341.
*aruensis, 292.
belli, 59, 61, 274, 275, 329, 330, 334,
336, 361, 366, 382 (PI. Ixiv. figs.
1,2).
bennetti, 59, 61, 329, 331, 348, 366,
382.
Uakei, 58, 301, 368, 369, 381.
borneensis, 58, 317, 321, 366, 382.
ACTIXOMETRA-
brachiolata, 57, 59, 278, 283, 366, 380.
*brasiliensis, 302.
briareus, 48, 58, 330, 366, 382.
clieltonetisis, 26, 39.
coiijringeri, 45, 58, 301, 317, 319, 320,
324, 362, 364, 366, 382 (PI. Ix.
figs. 1, 2).
cumingi, 58, 301, 366, 381.
discoidea, 58, 316, 317, 368, 382.
*J/."siuiiliK, 60, 337.
disttncta, 57, 294, 295, 341, 364, 366,
381 (PI. Iv. fig. 1).
divaricata, 15, 59, 78, 316, 329, 330,
332, 333, 362, 366, 382 (PI. lxiii.
figs. 6-8).
duplex, 59, 329, 330, 334, 335, 337,
362, 367, 382 (PI. lxvi. fig. 3).
echinoptera, 58, 302, 367, 381.
elongaia, 45, 58, 61, 208, 273, 275,
303, 311-313, 343, 362, 367, 382,
(PI. Ivii. figs. 2-4).
fimbriata, 45, 46, 58, 61, 273, 316,
317, 319, 320, 322, 328, 362, 363,
367, 382.
f/isea, 306, 307.
grandicalyx, 59, 330, 367, 382.
*hamata, 278, 288.
*imperiali8, 59, 268, 270, 278, 279,
288.
^intermedia, 59, 278, 282, 283.
intricata, 310.
japonica, 43, 58, 59, 330, 342, 346,
367, 382.
392
THE VOYAGE OF H.M.S. CHALLENGER.
AcTtNOMETRA
*jukesi, 59, 291, 292.
lineata, 8, 20, 22, 24, 26, 45, 58, 274,
316, 317, 327, 328, 358, 367, 382
(PL v. fig. 2 ; PL Ix. fig. 3).
littoralis, 15, 59, 330, 346, 347, 362,
367, 382 (PL Lxvii. figs. 1, 2).
lov&ni, 16, 26, 38.
metadata, 20-22, 24, 58, 293, 302,
303, 304, 306, 307, 309, 328, 361,
367, 381 (PL v. fig. 1 ; PL lv.
fig. 2).
magnified, 58, 61, 330, 333, 367, 382.
meridionalis, 22, 26, 36, 45, 46, 5S,
275, 300-302, 343, 358, 367-369,
373, 381 (PL iv. fig. 4 ; PL lvi.
figs. 1, 2).
*mertensi, 60, 338, 341.
*meyeri, 60, 338, 341.
*morsei, 346.
nmltibrachiata, 27, 57, 93, 294, 295,
299, 362, 367, 381 (PL lvi. figs. 3, 4).
multifida, 44, 48, 58, 329, 330, 333,
367, 382.
multiradiata, 52, 58, 246, 316, 317,
319, 322, 324, 326-328, 361, 367,
382 (PL lxvi. figs. 1-3, 8).
mutabilis, 53, 338, 341.
nigra, 58, 304, 309, 367, 381.
nobilis, 15, 16, 59-61, 77, 265, 275,
329, 330, 334, 336, 363, 364, 367,
383 (PL lxv.).
novx-guineas, 57, 295, 298, 299, 367,
381.
parvicirra, 15, 27, 36, 46, 48, 50-
52, 58-60, 209, 262, 265, 275,
284, 310, 312, .313, 316, 321, 329,
332, 338, 341-344, 347, 358, 361-
365, 367, 369, 383 (PL lxi.;
PL lxvii. figs. 3, 4).
paucicirra, 9, 13-15, 22, 24, 26, 27,
43, 44, 49, 50, 57, 59, 60, 77, 93,
94, 99, 276, 277, 283, 287, 290,
291-294, 296, 303, 361, 367, 381
(PL iv. fig. 6; PL v. fig. 3; PL
liv.).
pectinata, 41, 44, 57, 59, 93, 274-
276, 278-284, 285, 287, 291, 309,
314, 361, 364, 381 (PL liii. figs.
15-22).
ACTINOMETRA
peroni, 59, 324, 331, 348, 367, 383.
*polymorpha, 60, 338, 341-344.
pulchella, 22, 26, 27, 35, 36, 45,
50, 53, 58, 59, 241, 262, 300, 301-
307, 354, 357, 362, 368, 369, 372,
373, 381 (PL iv. fig. 5; PL lii.
figs. 1, 2).
^purpurea, 278.
quadrata, 45, 58, 311, 319, 330, 331,
342, 360, 367, 383 (PL lxii. fig. 1).
regalis, 45, 59, 274, 275, 329, 331,
342, 347, 365, 367, 383 (PL Ixviii.).
*robusta, 59, 264, 278, 282, 283, 288-290.
rdbustipinna, 59, 345, 367, 383.
* rosea, 278.
rotatoria, 58, 303, 310, 311-313, 314,
315, 364, 367, 382 (PL lis. fig. 2).
rubiginosa, 58, 60, 300, 301, 367, 381.
scldegeli, 59, 331, 347, 367, 383.
sentosa, 43, 45, 46, 58, 246, 317, 324,
325, 326, 365, 367, 382 (PL lxvi.
figs. 4-7).
simplex, 58, 275, 302, 303, 310-312,
314, 365, 367, 382 (PL lix. fig. 1).
Solaris, 24, 36, 39, 57, 59, 60, 93, 94,
275-283, 285, 287, 288, 291, 294,
303, 309, 361, 367, 381 (PL v. fig. 4;
PL liii figs. 1-14).
*stellata, 296, 297.
stelligera, 13, 20-22, 24, 45, 50, 58,
302-304, 306-308, 309, 361, 367,
369, 381 (PL v. fig. 5; PL lviii.).
*steiearti, 321.
*strota, 59, 278, 282, 288, 290.
Henax, 308.
*timorensis, 53, 60, 338.
trachygaster, 53, 310.
friehoptera, 35, 58, 59, 284, 330,
332, 342, 345-347, 359, 367, 383
(PL lxiii. figs. 1-5).
typica, 9, 13-16, 44, 57, 60, 93, 277,
283, 287, 294-296, 297-300, 337,
361, 367, 369, 381 (PL lvii. fig. 1).
valida, 58, 208, 303, 310-311, 314,
315, 361, 367, 382 (PL lix. fig. 3).
variabilis, 51, 52, 58, 60, 61, 330, 383.
*wahlbergi, 60, 271, 338.
wurtembergica, 26, 39.
Agassizocrinus, 1.
REPORT ON THE CRINOIDEA.
393
*Alecto, 1-3, 41, 42, 64, 85-89," 26G, 267, 269-
271.
brachiolata, 42.
carinata, 199-201.
*dentata, 170.
echinoptera, 42.
eschrichtii, 138.
fimbriata, 317.
*glacialis, 138.
japonica, 42.
meridionalis, 301.
milberti, 194.
multifield, 323.
multiradiata, 269, 322.
parvicirra, 338, 340.
phalangium, 158.
^purpurea, 59, 278, 283, 284.
rotalaria, 271, 313.
*sarsii, 57, 170.
soZam, 270, 279.
Himorensis, 42, 338.
*wahlbergii, 271, 338, 340.
Alectro dentata, 57, 169, 172, 173.
Allagecrinus, 27.
Allionia, 87, 89.
Ambulacra, 61, 101, 104, 134, 214.
Anilocra, 99.
Antedon, 1-8, 13, 21, 23-27, 29-39, 42-44, 47-49,
52-56, 60-62, 64-66, 68-70, 74-79,
85-88, 89-94, 96, 97, 99, 110, 161,
164, 171, 201, 203, 204, 206, 208,
214, 216, 227, 229-231, 238, 239,
241, 243, 251, 255, 257, 264, 272,
273, 276, 277, 290, 296, 302, 303,
315, 316, 319, 327, 329, 343, 348,
349, 351, 371-373, 383; batbymetrical
distribution of, 31-35, 157, 193, 210,
211, 239; centro-dorsal of, 7-10, 13,
38, 39, 112, 214; disk of, 231, 265,
266 ; geographical distribution of, 29-
35, 136, 157, 193, 210, 224, 239;
pinnules of, 91, 92, 123, 137, 231;
radials of, 23-26, 38, 39, 112, 123,
125, 133, 143, 146, 192, 203, 204,
216, 229, 243, 245, 263; Series I.,
94; Series II., 99; Series III., 208;
Series IV., 238.
abgssieola, 30, 31, 33, 54, 92, 158, 180,
191, 359, 365, 370, 372, 376 (PL xxxiii.
figs. 1, 2).
(ZOOL, CHALL. EXP. PART LX. — 1888.)
Antedon —
abyssorum, 54, 158, 185, 190, 191, 358,
370, 372, 376 (PI. xxix. figs. 10-13).
accela, 19, 23, 54, 61, 107, 127, 132, 133,
156, 164, 184, 243, 348, 364, 369, 376
(PI. ii. fig. 3; PI. xvi.).
aculeafa, 5', 103, 128, 364, 369, 375
(PL xxiii. fig. 3).
acuticirra, 55, 116, 253, 366, 380.
acuHradia, 32, 33, 54, 102, 105, 112, 113,
115, 246, 361, 370, 372, 375 (PL xi.
figs. 3, 4).
adeonse, 54, 206, 366, 378.
&quimargi?iafa, 26, 38.
xquipimia, 55, 225, 227, 379.
*alata, 59, 304.
alternata, 27, 33, 54, 61, 158, 179, 180,
190, 191, 360, 364, 369, 370, 372, 370
(PL xviii. figs. 1-3 ; PL xxxii. figs. 5-9).
alticeps, 38.
anceps, 54, 55, 110, 193, 194, 197-199,
205, 240, 252, 254, 255, 263, 364, 3GG,
377, 380 (PL xxxv. figs. 1-3).
angusticcdyx, 21, 45, 55, 75, 118, 134, 218,
240-242, 243, 245, 246, 248, 249, 364,
369, 380 (PL ii. fig. 4; PL 1. figs. 1, 2).
angustlpinna, 54, 91, 158, 185, 189, 190,
192, 365, 369, 372, 376 (PL xxix.
figs. 1-4).
angustiradia, 55, 211, 252, 253, 362, 368,
380 (PL xlv. fig. 4).
antarctica, 9, 10, 22, 24, 33, 54, 75, 138,
140, 141, 144 145-147, 155, 157, 349,
351, 359, 367, 376 (PL i. fig. 6;
PL xxv.).
armata, 54, 207, 376.
articulate, 50, 54, 55, 224, 226, 238, 366,
379.
australis, 33, 54, 61, 136-138, 141, 146-
149, 155, 157, 359, 368, 376 (PL xxvi.
figs. 4, 5 ; PL xxvii. figs. 14-20).
balanoides, 13, 54, 206, 207, 363, 367,
378 (PL xxxiii. figs. 6, 7).
barentsi, 54, 136, 137, 138, 156, 368,
376.
basieurva, 32, 54, 75, 100, 102, 120-126,
129, 133, 198, 223, 243, 246, 248, 251,
348, 360, 361, 369, 370, 375 (PL ii.
fig. 2 ; PL xxi. fig. 3 ; PL xxii. figs. 3, 4).
*bicolor, 199.
Ooo50
394
THE VOYAGE OF H.M.S. CHALLENGER.
Antedon —
bidens, 54, 206, 366, 378.
*bidentata, 97, 262.
Umaculata, 54, 226, 366, 379.
bipartipinna, 55, 253, 366, 3S0.
bispinosa, 33, 54, 102, 114, 115, 191, 252,
358, 370, 372, 375 (PI. xx. figs. 3, 4).
*braziliensis, 199, 201.
brevicuneata, 54, 223, 226, 235-237, 366,
379.
brevipinna, 54, 207, 208, 211, 212, 376,
378.
breviradia, 20, 32, 33, 54, 75, 101, 102,
108, 110, 112, 113, 115-117, 123, 124,
211, 246, 360, 361, 369, 370, 372, 375
(PI. iii. figs. 4, 5 ; PI. xi. fig. 5 ; PI. xix.;
PI. xx. figs. 1, 2).
*briareus, 48, 60, 333.
carinata, 8, 13, 22, 24-26, 32, 34, 36, 54,
62, 75, 77, 192, 193, 194, 199-205,
290, 306, 344, 358, 366-368, 377
(PI. iii. figs. 1-3 ; PI. xxxiv.).
carpenter i, 54, 193, 366, 377.
*celtica, 57, 149, 151, 158, 160, 161.
clemens, 54, 224, 225, 229-231, 364, 366,
379 (PI. xxxix. fig. 5).
colunmaris, 54, 207, 369, 377
compressa, 54, 212, 222, 362, 363, 367,
368, 378 (PL xli.).
conjungens, 55, 60, 224, 225, 233-235,
237, 363, 367, 379 (PI. xlv. fig. 1).
costata, 94, 135, 214, 372.
*crenidata, 57, 256-261.
cube7isis, 54, 68, 207, 369, 377.
decameros, 38.
*dvcipiens, 57, 256, 258-260, 262.
defecta, 54, 206, 207, 367, 368, 378.
*de?itata, 170.
denticulata, 34, 54, 101, 103, 130, 131,
211, 362, 367, 375 (PL xxii. figs. 1, 2).
disciformis, 8-10, 13, 25, 45, 54, 224,
225, 227, 228, 230, 231, 363, 366, 379
(PL iv. fig. 2 ; PL xxxix. fig. 4).
discoidea, 54, 132, 134, 135, 362, 368, 376.
distincta, 45, 55, 118, 127, 178, 240, 241,
243, 246, 247, 248, 363, 369, 380
(PL li. fig. 1).
diibeni, 54, 157, 158, 181-183, 192, 199,
201, 205, 358, 366, 373, 377 (PL xxxvii.
figs. 1-3).
Aktedon —
*dubia, 110, 197, 258-261.
duplex, 54, 207, 208, 211, 212, 217, 367,
368, 375, 378.
eehinata, 54, 119, 360, 369, 372, 375
(PL xxi. figs. 4, 5).
elegans, 23, 31, 48, 52, 53, 55„56, 90, 94,
96, 97, 130, 264-2G6, 276, 362, 366,
367, 375 (PL viii.).
dongafa, 35, 54, 224, 226, 366, 379.
eschrichti, 3, 10, 13, 19, 21-23, 25, 33,
34, 39, 41, 45, 47, 54, 60, 62, 75, 77,
80, 92, 105, 118, 13G, 137, 138-147,
149-157, 163, 166-169, 176, 177, 196,
204, 290, 297, 349, 351, 352, 354-357,
366-369, 371, 372, 376 (PL i. fig. 8;
PL xxiv. figs. 4-14).
exigua, 54, 157, 158, 178, 180, 181, 358,
368, 377 (PL xxxii. figs. 1-4).
riagellata, 55, 214, 223, 224, 226, 366,
379.
flexilis, 54, 103, 110, 127, 128, 208, 209,
211, 212, 217, 218, 220-223, 362, 368,
375, 378 (PL xlii.).
*fluduans, 90, 94, 96, 97, 130, 264-265.
gorgonia, 88, 89, 199, 201.
gracilis, 54, 76, 91, 102, 107, 108, 118,
184, 364, 369, 375 (PL xii. figs. 3-5;
PL xv. figs. 1-4).
granulifera, 55, 57, 239-241, 243, 248,
252, 368, 380.
ijreppini, 38.
gresslyi, 214.
gyges, 49, 50, 55, 224, 225, 366, 379.
hageni, 22, 54, 207, 367, 368, 373, 377.
hirsuta, 54, 157, 158, 188, 219, 358, 368,
377 (PL xxxi. fig. 5).
hystriz, 54, 118, 119, 136, 143, 147, 156,
157, 162, 164, 165-169, 173, 250, 354,
356, 369, 377 (PL xxvii. figs. 21, 22 ;
PL xxviii. figs. 4, 5).
imparipinna, 54, 225, 366, 379.
impinnata, 54, 206, 366, 378.
inxqualis, 20, 21, 32, 45, 55, 57, 61, 122,
240, 241, 243,244-246, 248, 249,316,
360, 361, 368, 370, 372, 380 (PL ii.
fig. 5 ; PL li. fig. 2).
incerta, 27, 54, 61, 91, 101, 102, 104-106,
107, 108, 134, 249, 251, 360, 369, 372,
375 (PL xviii. figs. 4, 5).
REPORT ON THE CRINOIDEA.
395
Antedon-
incisa, 54, 75, 102, 122-124, 125-127,
129, 133, 156, 218, 243, 248, 360, 368,
369, 372, 375 (PL ii. fig. 1 ; PI. xxi.
figs. 1, 2).
incurva, 38.
indica, 35, 54, 210, 225, 232, 233, 366,
379.
informis, 54, 194, 205, 206, 224, 227,
230, 363, 366, 377 (PI. xxxiii. fig. 3).
infracretacea, 26, 39.
*insignis, 56.
irregularis, 48, 57, 256, 258, 261.
italica, 38.
*jacquinoti, 194.
Ixvicirra, 54, 225, 366, 379.
Isevipinna, 54, 206, 366, 378.
Isevis, 54, 158, 187-189, 364, 369, 377
(PI. xxxi. fig. 6).
Ixvissima, 54, 194, 366, 378.
latipinna, 54, 102, 116, 365, 368, 375
(PI. x. fig. 3).
lineata, 54, 158, 183, 184, 365, 369, 372,
377 (PL xiii. figs. 4, 5).
longicirra, 22, 23, 54, 102, 103-105, 136,
362, 368, 375 (PL xvii.).
longipinna, 54, 91, 157, 185, 365, 369,
372, 377 (PL xxx. figs. 1-3).
lov'eni, 54, 56, 194, 206, 366, 378.
ludovici, 55, 253, 366, 380.
lusitanica, 32-34, 47, 52, 54, 91, 102, 105,
108, 109-112, 118, 164,208-210, 212,
217, 252, 354, 369, 372, 375, 378 (PI.
xxxix. figs. 1-3).
macronema, 13, 22-26, 44, 54, 61, 75, 76,
210-212, 214, 284, 359, 366, 367, 378
(PL iv. fig. 3 ; PL xxxviii. figs. 4, 5).
magellanica, 30, 54, 137, 138, 149, 367,
376.
mama, 54, 206, 225, 226, 228, 362, 368,
379 (PL xliv. figs. 2, 3).
margitiata, 54, 224, 225, 227, 228, 230,
231, 233, 363, 366, 379 (PL xl.).
marmorata, 202.
*mediterraneus, 158, 160.
meridionalis, 300, 301.
microdiscus, 53, 56, 94, 96, 97, 264, 265,
361, 367, 375 (PL xxxvii. figs. 4-6).
milberti, 54, 56, 192-194, 284, 363, 364,
367, 378 (PL xxxv. figs. 4-6).
Antedon-
milleri, 54, 366, 373, 377.
multiradiata, 31, 53, 56, 90, 94, 96-98,
265, 276, 361, 367, 375 (PL ix.).
multispina, 33, 34, 45, 54, 55, 91, 102,
110, 117-120, 169, 177, 178, 240, 241,
248-251, 358, 365, 369, 372, 375, 380
(PL xiii. figs. 1-3 ; PL xiv. figs. 5-7 ;
PL lxix. figs. 1-4).
*notata, 187.
occulta, 50, 55, 61, 91, 208-210, 224, 226,
236, 360, 368, 369, 379 (PL xlviii. figs.
1,2; PL xKx. figs. 3, 4).
orhignyi, 38.
palmata, 35, 44, 54, 55, 208, 213, 224,
226, 366, 379.
parvicirra, 54, 192-194, 204, 224, 363,
366, 378 (PL xxxvi. figs. 7, 8).
parvispina, 54, 61, 103, 127, 128, 362,
368, 376 (PL xv. fig. 9).
patula, 54, 61, 212, 219-223, 362, 368,
379 (PL xliii.).
perforata, 8.
l)erspi?wsa, 54, 193, 366, 378.
petasus, 54, 158, 181, 357, 367, 368, 373,
377.
phalangium, 13, 22, 23, 34, 47, 54, 56,
61, 76, 77, 80, 137, 143, 151, 156-
158, 159-167, 169, 175-178, 181, 229,
353-355, 367, 368, 377 (PL xxvii. figs.
23-29; PL xxviii. figs. 1-3).
philiberti, 55, 253, 366, 380.
picteti, 26, 39.
pinniformis, 54, 193, 224, 366, 378.
porrecta, 45, 46, 55, 117, 177, 178, 241,
246, 250, 251, 316, 365, 369, 380
(PL Hi. figs. 3-5).
pourtalesi, 54, 208, 209, 211, 212, 368,
379.
prolixa, 54, 136, 147, 156, 157, 162, 166-
168, 173-178, 356, 367-369, 372, 377.
protecta, 53, 55, 91, 225, 234, 237, 366, 379.
*pukhella, 59, 304, 305.
pumila, 54, 56, 193, 206, 366, 378.
pudlla, 54, 101, 103, 130, 131, 362, 368,
376 (PL xxiii. fig. 1).
quadrata, 33, 34, 54, 57, 61, 136-138,
140-143, 147, 149-156, 168, 169,
187, 355-357, 367-369, 376 (PL xxvi.
figs. 1-3; PL xxviii. figs. 1-13).
396
THE VOYAGE OF H.M.S. CHALLENGER.
Antedon-
quinduplicava, 9, 10, 13, 55, 203, 253,
262-264, 364, 366, 380 (PI. iv. fig. 1 ;
PI. xlvii. figs. 4, 5).
quinquecostata, 8, 13, 47, 54, 75, 92,
208, 209, 211, 215-218, 306, 362,
368, 379 (PI. iii fig. 6; PI. xxxviii.
figs. 1-3).
regalis, 31, 55, 224, 226, 237, 360, 366,
379 (PI. xlvi.).
regime, 48, 55, 225, 366, 379.
remota, 27, 54, 158, 184-186, 188, 191,
358, 370, 372, 377 (PI. xxix. figs.
5-9).
retzii, 8, 9.
reymudi, 43, 55, 252, 255, 366, 380.
rhodanica, 40.
rhomboidea, 30, 54, 137, 138, 141, 148,
149, 365, 368, 376 (PI. xii. figs. 1, 2 ;
PI. xxiv. figs. 1-3).
robusta, 54, 212, 220, 221, 223, 362,
368, 379 (PI. xliv. fig. 1).
rosacea, 3, 7, 16, 19, 20, 22, 23, 27, 31-
34, 54, 77, 80, 89, 90, 110, 119,
123, 127, 137, 143, 158, 160-164,
168, 169, 171, 177, 178, 181-183,
192, 205, 229, 231, 240, 261, 286,
287, 309, 354-356, 366-368, 373,
377.
*rubiginosa, 60.
rugosa, 8.
*sarsii, 53, 142, 170, 172-175, 177.
savignyi, 55, 252, 253, 255, 263, 366, 380.
scrobicidata, 26, 38, 214.
semiglobosa, 8.
sempinna, 54, 192, 198, 366, 378.
similis, 55, 20S, 210, 211, 223, 224, 226,
235-237, 360, 368, 369, 379 (PI. xlvii.
figs. 1-3).
spieaia, 55, 225, 232, 233, 366, 3S0.
spinieirra, 54, 102, 112-114, 360, 370,
372, 376.
spinifera, 50, 55, 208-211, 216, 217, 306,
368, 379.
striata, 8.
tendla, 7, 22, 32, 33, 54, 57, 118, 119,
136, 142, 157, 162, 168, 169, 173-
180, 196, 353, 354, 356, 357, 368,
369, 372, 377 (PI. xiv. fig. 4 ; PI. xxxi.
figs. 1-4).
Antedon —
tenuicirra, 54, 158, 186-188, 192, 364,
368, 377 (PL xxx. figs. 4-8; PL xxxiii.
figs. 4, 5).
tessellata, 54, 193, 366, 378.
tessoni, 38.
tourtix, 8.
tubercuJata, 45, 55, 208, 210, 224, 225,
231, 232-234, 360, 369, 380.
tuberosa, 31, 54, 103, 126-129, 133, 178,
363, 369, 376 (PI. xiv. fig. 9; PI. xxiii.
fig. 2).
valida, 54, 61, 76, 91, 101, 102, 104-
108, 112, 116, 123, 141, 19S, 216,
249, '251, 364, 369, 376 (PL xv.
figs. 5-8).
variipinna, 45-47, 54, 55, 57, 61, 97,
193, 197-199, 205, 224, 240, 252, 253,
255, 256-263, 361, 362, 367, 378, 380
(PL xxxvi. figs. 1-6 ; PL xlviii. figs. 3-
5 ; PL xlix. figs. 1, 2).
Apiocrinidse, 10-12, 119, 335.
Apiocrinus, 11, 12, 27, 101.
Articulata, 64.
Asterencrinidea, 63.
*Asterias, 1, 85, 88, 269.
multiradiata, 89, 172, 173, 268-271, 322-
324.
pectinata, 59, 171-173, 268, 269, 271, 279,
280, 284, 286, 287.
tenella, 169, 171-173, 286.
*ASTEEIATITES, 85.
Asterites liberi, 63.
*Astkocoma, 86, 266.
Astylidea, 64.
Atelecrinus, 5, 13, 19, 23, 31, 35, 66, 68-71,
91, 207, 273, 383 ; basals of, 68, 69;
bathy metrical distribution of, 31, 70,
73; centro-dorsal of, 13, 69; disk
of, 70, 273; geographical distribu-
tion of, 31, 70.
balanoides, 69, 70-72, 357, 369, 374
(PL vi. figs. 6, 7).
cubensis, 69-73, 369, 374.
' wyvillii, 35, 69, 70, 72, 361, 369, 372,
374 (PL vi. figs. 4, 5).
Axillaries, 41-48, 50, 51, 53.
Barycrinds, 8.
Basals, 6, 10-13, 19-23, 38, 69, 214.
of Atelecrinus, 68, 69.
REPORT ON THE CRINOIDEA.
oy<
Basal bridge, 21.
grooves, 10.
rays, 14, 22, 23, 349.
star, 10, 21, 22, 204, 302, 307, 308, 349. ]
BASicuEVA-group, 33, 99, 100-102, 131, 133, 137,
157, 162, 192, 197, 198, 209, 211, 213, 215,
216, 221, 223, 239, 249, 266, 371.
Bathycrinus, 164, 191, 358.
Bathymetrical range of Coniatula?, 31-36.
of Actinometra, 35, 36.
olAntedon, 31-35, 157, 193, 210, 211, 239.
of Atelecrinus, 31, 70, 73.
of Eudiocrinus, 31, 79.
of Promachocrinus, 31.
of Thaumatocrinus, 31.
Blastoidea, 28.
Bourgueticrinida?, 8, 9, 65.
BOURGDETICRINUS, 65, 119.
ooliticus, 37.
* Caput-Medusse, 85, 323.
Central capsule, 17-21.
Centro-dorsal, 6-20, 22-24, 37-40, 349.
of Actinometra, 7, 13-16, 38, 39, 78, 277, 287,
290, 293,. 302, 307.
of Antedon, 7-10, 13, 38, 39, 112, 123, 125, 145,
161, 214, 216, 229, 243, 245, 263.
of Atelecrinus, 13, 69.
of Eudiocrinus, 13.
of Promachocrinus, 13, 349.
of Tliiolliericrinus, 65.
Chambered organ, 16-20.
Cirri, 6, 7, 11-15, 38, 39, 43, 47, 76-78, 82, 143,
147, 161, 162, 280, 343, 351.
Cirrus-sockets, 7, 13-15, 78.
Comaster, 2, 4, 5, 41, 64, 69, 87, 89, 266, 267.
*Comatula, 1-3, 5, 10-12, 21, 27, 30, 37, 41, 42, 44,
50, 63, 64, 69, 73, 77, 79, 85-89, 91,
171, 172, 175, 191, 263, 265, 266, 267,
269-272, 286, 287.
adeonse, 42.
*barbata, 287.
bracliiolata, 42, 172, 278.
*brevicirra, 314, 318.
carinata, 89, 199, 201.
*celtica, 57, 158.
coccodistoma, 320.
echinoptera, 42, 301, 373.
eschrichti, 138.
fimbriata, 41, 182, 317, 318, 325
flagellata, 42.
CoMATULA —
*hamata, 59, 278.
*jacquinoti, 56, 194-196.
japonka, 42, 43.
Ixvissima, 194, 197.
macronema, 43, 212.
*mediterranea, 169, 172.
meridionalis, 301.
*mertensi, 60, 338.
milberii, 56, 194-196.
multiradiata, 41, 269, 31S, 322-325.
palmata, 43.
parvicirra, 340, 341
pectinata, 287.
plialangium, 158.
*purpurea, 284, 285.
reynaudi, 43.
*rosea, 59, 278.
rotatoria, 41, 269, 271, 313, 314.
*sarsii, 170.
Solaris, 268-270, 279, 284, 286, 288.
tenella, 172.
tessellata, 266.
*timorensis, 42, 338, 340, 341.
triclwptera, 345.
*wahlbergi, 269, 271, 344.
woodwardii, 57, 158.
Comatuke, 1, 2, 4-7, 9-13, 19-23, 25-45, 47, 49-51,
53, 61, 64-66, 76-79, 89, 91, 137, 141,
198, 209, 250, 263, 319, 333; bathy-
metrical and geographical distribution of, 29-
36.
Comatulida;, 2, 27, 43, 63-66.
*Comatulina, 23, 87.
Comatulinse, 64.
*Comafulithes, 85.
*Comaturella, 86, 87.
Costata, 64, 65.
CtJPRESSOCMNUS, 8, 9.
Cyathocrinidae, 65.
Decacnemos, 2, 85, 86, 89.
*barbata, 171, 286.
rosacea, 111, 286.
*Decameros, 2, 86, 87, 89.
*Democrinus, 80.
Disk, of Actinometra, 268-270, 272-275, 290, 294
297, 319, 335, 336.
of Antedon, 231, 265, 266.
of Atelecrinus, 70, 273.
Distichals, 43-46, 48-53.
398
THE VOYAGE OF H.M.S. CHALLENGER.
Dorsoeentral, G.
Echini, genital plates of, 20.
EcHiNOPTERA-group, 300, 301-303, 329.
Edriocrinus, 1.
ELEGANS-group, 266.
Encrinds, 101.
gracilis, 27.
lilii/ormis, 28.
EscHRicim-group, 33, 99, 136-138, 156, 157, 349,
351.
Eudiocrinus, 4, 5, 13, 30, 31, 37, 39, 06, 70, 73-
81, 91, 92, 229, 273, 349, 383;
bathynietrical and geographical dis-
tribution of, 31, 79; centro-dorsal
of, 13; radials of, 75.
atlanticus, 4, 31, 74, 76-81, 369, 373,
374.
hyseh/i, 37, 75.
indivisus, 76, 79-81, 83, 367, 374.
japonicus, 79-83, 84, 85, 365, 367,
369, 372-374 (PI. vii. figs. 1, 2).
semperi, 25, 75, 79-82, 83, 85, 360,
369, 370, 372, 374 (PL iii. fig. 7;
PL vi. figs. 1-3).
varians, 76, 79-81, 83, 230, 362, 370,
372, 374 (PL vii. figs. 3-7).
Eugeniacrinidse, 65.
Eugeniacrinus, 2, 65, 192.
FiMBRiATA-group, 310, 316, 329.
Formulation, methods of, 43-52.
*Ganysieda, 86, 89.
Genital plates of Echini, 20.
♦Geocoma, 2, 87, 89.
Geographical range of Comatulse, 29-36.
of Actinometra, 35, 36, 283, 284, 329, 344.
of Antedon, 29-35, 136, 157, 193, 210, 224,
239.
of Atelecrinus, 31, 70.
of Eudiocrinus, 31, 79.
of Promachocrinus, 31.
of Thaumatocrinus, 31.
Geological range of Comatulse, 39.
*Glenotremites, 2, 64, 86, 87, 89.
paradoxus, 37.
GRANULiFERA-group, 239, 241, 249, 266, 371.
guettardicrinus, 11, 12.
*Hertha, 2, 86, 89.
*Hibernula, 86, 89.
Holopus, 1, 101.
Hyocrinus, 108, 164, 168, 184, 191, 358.
*Hyponome, 87, 89, 97.
*Kallispongia, 87, 90.
Marsupites, 2, 64, 65.
Metacrinus, 27, 92, 230, 231, 360, 362-364.
angulatus, 168.
moseleyi, 230.
rotundus, 230.
MiLBERTi-group, 99, 192, 202, 209, 210, 223,
227.
MlLLERICRINUS, 1, 12.
pratti, 211.
Muscle-plates of radials, 24-26, 75, 77.
Muscular bundles, 76, 77.
Myzostoma, 135, 178, 198, 236, 245, 262, 335.
alatum, 159, 160, 165.
areolatum, 301.
brevipes, 211.
carinatum, 206.
carpenteri, 171, 178.
coriaceum, 56.
cornutum, 180, 181.
dental um, 262.
excisum, 206.
filiferum, 262.
fimbriatum, 144.
gigas, 144, 156, 204.
inflator, 254.
laibatum, 325.
murrayi, 111, 211.
nigrescens, 346.
platypus, 338.
pulvinar, 159, 165.
quadricaudatmn, 96.
quadriferum, 262.
tenuispinum, 124, 125, 243, 245, 246.
willemoesii, 124, 245, 246.
Neocrinoidea, 1, 11, 27, 52.
*Ophiocrinus, 4, 65, 73, 74.
Ophiolebes scorteus, 189, 219.
Ophiomusium, 218.
*Ophiurites, 85.
Ovoid Bodies, 275, 312, 343.
Palseocrinoidea, 5, 9-11, 27, 52, 66.
Palmars, 43-46, 48-52.
PALMATA-group, 209, 210, 222, 223, 224, 250, 284,
310.
PARViciRRA-group, 316, 321, 329.
PAUCICIRRA-grOlip, 290.
Pentacrinidse, 11, 12, 23, 56, 61, 101, 104, 106,
119, 136, 211, 290, 319.
REPORT ON THE CRINOIDEA.
399
Pentacrinoid larva, 7, 19, 23, 32, 65, G9, 118, 119,
127, 142, 143, 1G3, 1GS, 169, 177, 178, 250,
319, 358, 370.
Pentacrinus, 1, 2, 6, 7, 12, 16, 23, 25, 27, 37, 64, 85,
86, 89, 92, 268, 297, 298, 365.
alternicirrus, 69, 93.
asteriscus, 37.
*euro}Jxus, 90.
maclearanus, 227, 357.
mollis, 365.
miilleri, 27, 72.
naresianus, 32, 246, 360, 361.
wyville-thomsoni, 69, 72, 93, 164.
*Phanogenia, 14-16, 24, 267, 272, 276, 297, 312,
333.
typica, 296.
Phylloceinus, 192.
*Phytocbinus, 86, 90.
Pinnules, 61, 69.
of Antedon, 91, 92, 123, 137, 231.
Plicatocrinus, 27.
Post-palmars, 48-50, 52.
Promachoceinus, 5, 10, 13, 22, 27, 30, 31, 49, 65,
66, 74, 91, 191, 348, 349, 374,
383 ; bathymetrical and geo-
graphical distribution of, 31 ;
centro-dorsal of, 13, 349 ; radials
of, 349.
abyssorum, 31, 349-351, 358, 359,
370, 372, 374 (PI. i. figs. 4, 5 ;
PI. lxix. figs. 5-7).
kerguelensis, 9, 25, 349, 350-352,
359, 367, 368, 374 (PI. i. figs.
1-3 ; PI. lxx.).
naresi, 350, 352, 364, 369, 374
(PI. lxix. figs. 8-10).
*Pterocoha, 86, 87, 89.
Radials (first), 9, 12-16, IS, 19, 21-28, 37-39.
of Adinometra, 24-26, 38, 39, 287, 290, 293,
302, 309, 328.
Radials of Antedon, 23-26, 38, 39, 112, 123, 125, 133,
143, 146, 192, 203, 204, 216, 229, 243, 245,
263.
of Eudiocrinus, 75.
of Promachocrinus, 349.
Radial axial canals, 8, 16-18.
Radial pentagon, 6, 8, 14-16, 21.
Radial pits, 9.
Radial spaces of stem, 8.
Rhizocrinus, 28, 65, 80, 119, 164, 297, 298.
Rosette, 6, S, 12, 16, 18-22, 31, 38, 75, 143, 216,
349.
Saccocoma, 64, 172.
Sacculi, 76, 79, 81, 83, 91, 92, 104, 105, 107, 134,
218.
SAViGNYi-group, 239, 250, 252, 255.
*Solacrinus, 86.
*Solanocrinites, 86, 87.
*Solanoceinus, 2, 64, 86-90.
costatus, 90, 93, 101, 211, 372.
gracilis, 90, 93, 94.
imperialis, 90, 93, 94.
SoLAEis-group, 274, 278, 291, 293, 301, 302.
SpiNiFEEA-group, 209-211, 213, 215, 223, 239, 266,
310, 371.
STELLiGERA-group, 303, 304, 310.
Syzygy, 41-46, 48, 50, 80, 90, 92, 93, 240, 264,
265, 297, 298.
TENELLA-group, 33, 99, 156, 192, 349, 352, 371.
Terminal comb of lower pinnules in Adinometra,
4, 42, 92, 271, 276, 343.
Tessellata, 64.
Thadmatocrinus, 5, 19, 23, 30, 65, 66, 68-70, 91,
191, 349, 383.
renovatus, 66, 67, 359, 370, 372,
374.
Thiolliericrinus, 37, 39, 65, 66.
TYPiCA-group, 294, 315, 341.
Under basals, 10-13.
VALiDA-group, 303, 310.
EXPLANATION OF THE PLATES.
A reference is given after each figure to a certain page of the text. In the case of
figures which illustrate entire specimens, the reference given is usually to the page in the
systematic part of the Eeport on which the species is first described. But with those
figures which represent structural details, reference is given to the page containing the
explanation which the figure was designed to illustrate. This is sometimes in the
morphological and sometimes in the systematic part of the Eeport. In a few cases there
is no special reference to a figure in the text ; and the number of the page following its
explanation is either that of the specific diagnosis, or that of a page containing a
description of structural peculiarities which is more or less illustrated by the figure in
question.
ERRATA.
Plate I. — The figure of the centro-dorsal of Antcdon antarctica should be lettered " 6d '
instead of " 6a."
Plate III. — Instead of " Fig. 4, Antedon radiospina," and " Pig. 5, Antedon eversa," read " Figs.
4, 5, Antedon breviradia."
Instead of " Fig. 7, Antedon (Ophiocrimis) semperi," read " Fig. 7, Eudiocrinus
semperi."
Plate IV. — Fig. 3, Instead of " Antedon macrocnema," read Antedon macronema."
Fig. 4, Instead of " Actinometra brasilicnsis," read " Actinometra meridionalis."
Fig. 6, Instead of " Actinometra aruensis," read " Actinometra paucicirra."
Plate V. — Fig. 3, Instead of " Actinometra juhesii" read " Actinometra paucicirra."
Fig. 36 is wrongly lettered " 55."
Fig. 4, Instead of " Actinometra strata," read " Actinometra Solaris. "
Plate VIII. — Instead of " Antedon fluctuans, sp. n.," read " Antedon eke/ans, Bell."
Plate XXXIII. — Figs. 4, 5, Instead of "Antedon notata, sp. n." read "Antedon tenuicirra, sp. u."
Plate XXXVI. — Figs. 1-6, Instead of "Antcdon dubia, sp. n.," read "Antedon mriipinna, Carpenter."
(ZOOL. CHALL. EXP. PART LX. — 1888.) OoO 51
PLATE I.
(ZOOL. CHALL. EXP. — PART LX. — 1888.) — OoO.
PLATE I.
Figs. 1-3. Promachocrinus kerguelensis, n. sp., x
Fig. 1. a. The calyx from the side, .....
b. The radial circlet from above, ....
c. The radial circlet from beneath, ....
(The central portion of the rosette is broken away.)
d. The centro-dorsal from above, ....
Fig. 2. a. Internal aspect of an "interradial radial," with a portion of
the basal star attached to it, .
b. Lateral view of the same, ....
Fig. 3. a. Lateral view of an ordinary radial, .
b. Internal aspect of the same,
c. Dorsal aspect of the same, with the impression of the basal
star at its inner end, ....
Figs. 4, 5, Promachocrinus abyssorum, n. sp.,
Fig. 4. Lateral view of an ordinary radial,
Fig. 5. The centro-dorsal from above, ....
Diam.
Page
6
348
.
348
349
349
349
349
349
349
351
349
Figs. 6, 7. Antedon Antarctica, n. sp.
Fig. 6. a. The calyx from the side, ....
b. The calyx from above, ....
c. Dorsal aspect of the radial pentagon and basal star,
d.1 Ventral aspect of the centro-dorsal,
Fig. 7. An isolated basal, .....
a. Ventral aspect, ......
b. Dorsal aspect, .....
Fig. 8. Antedon eschrichti, Mull., sp.
Fig. 8. a. The calyx from the side, ....
a. The calyx from above, ....
c. Dorsal aspect of the radial pentagon and basal star,
d. Ventral aspect of the centro-dorsal,
X
6
146
146
■
146
146
X
15
22
22
X
3
143
X
3
143
X
3
143
X
3
10
Wrongly lettered 6a.
TJieVqya.de of H M S' Challenger
ComaIula£.PJ 1 -
128
'"S^"
,r
-, A"— ~ -
f
-»
VI
'•'..;."
6<>
8*
a"
8C
can dtl Alith
l_3. PROMACHOCRINUS KERCUEIENSIS
4_ 5 ABYSSORUM
6 7 ANTEDON ANTARCTICA.
8 ESCHRICHTII
Harfhart.imp.
PLATE II.
PLATE II.
Fig. 1. Antedon incisa, n. sp., .
Fig. 1. a. The calyx from the side, .
b, Ventral aspect of the ceutro-dorsal,
c, Dorsal aspect of the radial pentagon and basal star,
d, The calyx from above, .
Fi°-. 2. Antedon basicurva, n. sp.,
Fig. 2. «. The calyx from the side, .
b: Ventral aspect of the centro-dorsal,
c. Dorsal aspect of the radial pentagon and basal star,
d, The calyx from above, .
Fig. 3. Antedon ac<ela, n. sp., .
Fig. 3. a. The calyx from the side, .
b. Ventral aspect of the centro-dorsal, .
c. Dorsal aspect of the radial pentagon and basal star,
d. The calyx from above, .
Fig. 4. Antedon angdsticalyx, n. sp.
Fi<*. 4. a. The calyx from the side, .
b. Ventral aspect of the centro-dorsal,
c. Dorsal aspect of the radial pentagon and basal star,
d. The calyx from above, .
Fig. 5. Antedon in^equalis, n. sp.,
Fig. 5. a. The calyx from the side, .
b. Ventral aspect of the centro-dorsal,
c. Dorsal aspect of the radial pentagon and basal star,
d. The calyx from above, .
Diam.
Page
. x 6
125
m .
10
.
125
•
125
. x 6
123
, .
10
m .
123
•
123
. x 6
7
.
7
,
23
•
24
. x 6
243
,
10
,
21
•
243
. x 6
245
,
10
,
21
.
245
The Voyage of HM.S Challenger"
Coma-tulae PI II.
3*
3b
^
4*
CBerie.au del etlith
1 ANTEOON INCISA
2 „ BASICURVA
3 ANTEDON ACCLLA
4. „ ANGUSTICALYX
5 ANT INAEQUALIS
PLATE III.
(ZOOL. CHALL. EXP. — PART LX. — 1888.) — OoO.
PLATE III.
Figs. 1-3. Antedon carinata, Lam., sp.
Fig. 1. a. The calyx from the side,
b. Ventral aspect of the centro-dorsal,
c. Dorsal aspect of the radial pentagon and
removal of one basal,
d. The calyx from above,
Fig. 2. An isolated basal,
a. Dorsal aspect.
h. Ventral aspect.
Fig. 3. a. Ventral aspect of the calyx after removal of two radials,
b. Side view of the same, showing the relations of the basals,
Diam.
Page
X
6
13
. •
,
203
basal star, after
203
.
•
24
x
15
22
22
Figs. 4, 5. Antedon breviradia, n. sp.
Fig. 4. a. The calyx from above, .
b. The calyx from the side, . . . •
c. Dorsal aspect of the radial pentagon and basal star,
Fig. 5. A younger individual.
a. The calyx from the side, ....
b. Dorsal aspect of the radial pentagon and basal star,
c. The calyx from above, ....
Fig. 6. Antedon quinquecostata, n. sp.
Fig. 6. a. Ventral aspect of the centro-dorsal, .
b. Dorsal aspect of the radial pentagon and basal star,
c. The calyx from above, ....
d. The calyx from the side, ....
Fig. 7. Eudiocrinus semperi, n. sp.,
Fig. 7. a. The calyx from the side, ....
b. Dorsal aspect of the radial pentagon,
c. Ventral aspect of the same,
X
6
112
112
•
112
X
6
112
X
12
112
X
12
112
216
9
216
13
75
75
75
The Voyage of HMS'thallenger"
Comalulae.Pl.il! .
CBprjRa.ii. del etlith .
Ha/iharl imp
.0. ANTEDON CARINATA 5. ANTEDON EVERSA
4- „ RADIOSPINA. 6 „ QU I N QU EC DSTATA
7 ANTEDON (Ophiocrmus) SEMPERI
PLATE IV.
PLATE IV.
Fig. 1. Antedon qotnduplicava, n. sp.,
Fig. 1. a. The isolated centro-dorsal, side view,
b. Dorsal aspect of the same, ....
c. Dorsal aspect of the radial pentagon, after removal of the
rosette, ......
d. Ventral aspect of the centro-dorsal,
Fig. 2. Antedon disciforms, n. sp.,
Fig. 2. a. The calyx from the side, ....
b. The calyx from above, ....
c. Dorsal aspect of the radial pentagon,
d. Ventral aspect of the centro-dorsal, .
Fig. 3. Antedon macronema, Mull., sp.,
Fig. 3. a. The calyx from the side, ....
b. The calyx from above, ....
c. Dorsal aspect of the radial pentagon and basal star,
d. Ventral aspect of the centro-dorsal,
Diam. Page
8
263
262
9
9
13
229
9
9
25
24
6
10
Fig. 4. ACTINOMETRA MERIDIONALIS, Pourt., sp.,
Fig. 4. a. The calyx from the side, ....
b. (a) Ventral and (yS) dorsal aspects of an isolated basal,
c. The calyx from above, ....
X
6
26
X
15
22
X
6
26
Fig. 5. ACTINOMETRA PULCHELLA, Pourt., sp.
Fig. 5. a. (a) Ventral and (/3) dorsal aspects of an isolated basal,
b. Dorsal aspect of an isolated radial with one basal attached,
c. Side view of the radial pentagon with the basals attached,
X
8
22
X
6
22
X
6
22
Fig. 6. ACTINOMETRA PAUCICIRRA, Bell.
Fig. 6. a. Ventral aspect of the young centro-dorsal, .
b. (a) Ventral and (/3) dorsal aspects of an isolated basal,
x
x
6
15
10
22
The Voyage of H U
•
jjtfW $^? j^~ ' /
l
\
■ ' /
— '
T> ' |
Ss».
3*
'
''. .•■
r.
**
x»w
-
C Bc^jcau del et hili
I ANTEDON QUINDUPLICAVA .
2. „ DISCIFORMIS
3 „ MACRONEMA.
A-. ACTINOMETRA BRASILIENSIS
5 „ PULCHELLA.
6, „ ARUENSIS.
Q |
PLATE V.
(ZOOL. CHAIX. EXP. — PART LX. — 1888.) — OoO.
PLATE V.
Fig. 1. ACTINOMETRA MACULATA, 11. sp.,
Fig. 1. a. The calyx from above, .
b. The calyx from the side, .
c. Dorsal aspect of the radial pentagon and basal star, .
d. Dorsal aspect of the centro-dorsal, .
Fie 2. ACTINOMETRA LINEATA, n. sp.,
Fig. 2. a. The calyx from above, .
b. The calyx from the side, .
c. Ventral aspect of the centro-dorsal with the rosette and two
radials in situ, .
d. Dorsal aspect of the centro-dorsal, .
e. Side view of the centro-dorsal, with the rosette and two
radials in situ, .
Fig. 3. ACTINOMETRA PAUCICIRRA, Bell,
Fig. 3. a. The calyx from above, .
b.1 The calyx from the side, .
c. Internal aspect of three united radials, after removal of the
centro-dorsal, .
Fig. 4. ACTINOMETRA SOLARIS, Lam., sp.,
Fig. 4. a. The calyx from above, .
b. The calyx from the side, .
c. Dorsal aspect of the radial pentagon and imperfect basal star,
Fig. 5. ACTINOMETRA STELLIGERA, n. sp.,
Fig. 5. a. The calyx from above, .
b. The calyx from the side, interradial view, .
c. The calyx from the side, radial view,
d. Dorsal aspect of the radial pentagon and basal star,
e. Ventral aspect of the centro-dorsal, with the rosette and
basal star in situ, .
Diam.
x 6
x G
Page
307
26
21
307
328
26
8
328
26
26
14
14
26
7
290
309
22
26
21
Wrongly lettered 5b.
. ' ulae PI V.
•W
CBr-r]ea,u del etlith.
I ACTINOMETRA MACULATA 3 ACTINOMETRA JUKESII.
2 „ LINEATA 4- „ STRATA
5. ACT STELLICERA
Ha.in. '■
PLATE VI.
PLATE VI.
Figs. 1-3. Eudiocrinus semperi, n. sp.,
Fig. 1. Side view, .......
Fig. 2. The disk, from above, ......
Fig. 3. Distal portion of an arm, .....
Figs. 4, 5. Atelecrinus wyvillii, n. sp.
Fig. 4. The disk, from above, . . . . . x 6 70
Fig. 5. Side view, . . . . • . . x 4 72
Figs. 6, 7. Atelecrinus balanoides, n. sp.
Fig. 6. The disk, from above, .
Diam.
Page
X 4
•
82
•
74
82
x 6 70
Fig. 7. Side view, ...... x 4 70
TkeVoya-ge of KM. S." Challenger "
Comatulae.EI VI.
West IHewmaiL 4 ( ° lilh.
1-3 EUDIOCRINUS SEMPERI, sp n
4. 5. ATELECRINUS WY V I LLI I , sp n g 7 ATELECRI N US BALANOI DES , sp. n
PLATE VII.
(ZOOL. CHALL. EXP. — PART LX. — 1888.) — Ooo.
PLATE VII.
Figs. 1, 2. Eudiocrinus japonicus, n. sp.,
Fig. 1. Side view,
Fig. 2. A cirrus,
Figs. 3-7. Eudiocrinus varians, n. sp.,
Fig. 3. Side view, interradial, . . •
Fig. 4. Side view, radial, .
Figs. 5, 6. The base of a short-jointed cirrus, from opposite Bides,
Fig. 7. The base of a long-jointed cirrus,
Diam.
Page
X 4
•
84
•
84
x 4
•
81
•
81
•
82
82
Mlenger!'
C om a Lu la :
!
I. 2. EUDIOCRINUS JAPONICUS.
3_6. EUDIOCRINUS VARIANS.:'
PLATE VIII.
PLATE VIII.
Diatn. Page
Antedon elegans, Bell, . . . . . x 3 94
The Voyage of H. M.S." Challenger
Comatulae . PI . VIII
Berjeau. &. HigkLey del et lith .
ANTEDON FLUCTUANS, Spn
Mmtem Bros m\p
PLATE IX.
(ZOOL. CIIALL. EXP. PART \,X. 1888.) OoO.
PLATE IX.
Antedon multiradiata, n. sp.
Fig. 1. Side view. .....
Fig. 2. The disk, from above, ....
Fig. 3. Distal portion of an arm,
Fig. 4. Ambulacral groove of an arm, .
Fig. 5. A pinnule from the middle of an arm, .
Fig. 6. One of the proximal pinnules, its end broken, .
Diam.
Page
. x 3
96
x 4
266
•
96
96
96
96
The Voyage of EM.S
■
Berjeau
ANTEDON MU LTIRADIATA, Spi
■
PLATE X.
PLATE X.
Figs. 1, 2. A.NTEDON DISCOIDEA, 11. Sp.
Fig. 1. Side view, .
Fig. 2. a. The pinnule of the fourth brachial, .
6. The pinnule of the eighth brachial, .
Diam.
Page
x 3
134
x 5
135
x (5
135
Fig. 3. AXTEDON LATIPINNA, 11. sp.
o
Fig. 3. Side, view,
x 4
I L6
TkeA ELMS. Challenger "
I tj LLulai PI X
BerjeaiiZtHig'hley del
■ , . ■
1,2. ANTEDON DISCOIDEA, sp.n.
3 ANTEDON LATIPINNA, sp.n
PLATE XL
(ZOOL. CHALL. EXP. — PART LX. — 1888.) — Ooo.
PLATE XL
Figs. 1, 2. Antedon spinicirra. n. sp.
Diam. Page
Fig. 1. Side view, .... . . x 4 112
Fig. 2. The disk, from above, . x 6 112
Figs. 3, 4. Antedon acutiradia, n. sp.
Fig. 3. Side view, ... . x 4 113
Fig. 4. Middle portion of an arm, . . x 4 113
Fig. 5. Antedon breviradia, n. sp.
Fig. 5. Side view, ..... . x 2 110
'['lie Voyage of H.M.S. QiaJLenqer '
Con ...
u BcHiglileyclel
I 2 ANTEDON S P I N I CI R R A , sp n .
3, 4-. ANTEDON ACUTIRADI A, sp.n.
5. ANTEDON B R E V I R A D I A, sp . n
PLATE XII.
PLATE XII.
Figs. 1, 2. Antedon rhomboidea, n. sp.
Diaru. Page
X 2 148
Fig. 1. Side view,
Fig. 2. A lower pinnule, . . x 3 148
Figs. 3-5. Antedon gracilis, n. sp.
Fig. 3. The disk, from above, ...
Fig. 4. Side view of a mutilated individual,
Fie;. 5. A cirrus,
X
6
107
X
4
107
X
3
107
Hie "Voyage ofH.M.S "Challenger
Con BLtulae.Pl.XII
I, 2. ANTEDON RHOMBOlDEA.sp n 3-5. ANTEDON G RAC I LI S , sp. n .
PLATE XIII.
(ZOOL. CHAiL. EXP. — PART LX. — 1888.) — OoO.
PLATE XIII.
Figs. 1-3. Antedon multispina, n. sp.
Fig. 1. Side view of a young specimen,
Fig. 2. Middle portion of an arm of the same, .
Fig. 3. The pinnule on the third brachial,
Diam.
Page
X
4
117
X
8
117
X
12
118
Figs. 4, 5. Antedon lineata, n. sp.
Fig. 4. Side view,
Fig. 5. (a) Dorsal and (b) ventral aspects of the pinnule on the twelfth
brachial, . . . . . . . x
183
184
i tie Voyage oi'H f.
Sa
5i>
Berjeau&HigWey delellith
L3. ANTEDON M U LT I S PI N A .
4.5.ANTEDON LINEATA.Sp.n
Hani
PLATE XIV.
PLATE XIV.
Fig. 1. Early Pentacrinoid of Antedon phalangium, Mull., sp.
Ficr. 2. Pentacrinoid of A ntedon sp., .
Fig. 3. Pentacrinoid of Antedon sp., .
Fio\ 4. Pentacrinoid of Antedon tenella, Retz., sp.,
Figs. 5-7. Pentacrinoids of Antedon multispina, n. sp.,
Fie. 8. Pentacrinoid of Antedon sp.. .
Ficv. 9. Pentacrinoid of Antedon tuberosa, n. sp.,
Diam.
Page
. X 20
163
. X 12
168
. x 12
168
. X 12
178
. x 12
118
. x 12
250
. x 12
127
The Voyage of H. MS "Challenger"
Comatulae.Pl XIV.
i
Bei^eau. & HigKley del el litK .
MiniBniBfL
PENTACRINOID LARV/E OF ANTEDON
PLATE XV.
(ZOOL. CHALL. EXP. PA11T LX. 1888.) — Ooo
PLATE XV.
Figs. 1-4. Antedon gracilis, n. sp.
Fig. 1. Side view of a young individual,
Fig. 2. Middle portion of an arm.
Fig. 3. Terminal portion of a young arm.
Fig. 4. Dorsal aspect of a lower pinnule,
Diam. Page
x H 109
x 4 108
x 6 108
x H 108
Fio-s. 5-8. Antedon valida, n. sp.
Fig. 5. Side view of a mutilated individual. . . - . x 2 104
Fig. 6. Side view of a calyx and arm bases with one ray removed, . x 101
Fio-. 7. Lower portion of an arm from the third brachial onwards, x 2 105
Fig. 8. A middle, pinnule of a young individual, . . >
6 105
Fig. 9. Antedon parvipinna, n. sp.
Fig. 9. Side view, . • . x 2 127
The Voyage of H.M.S!thallengep!
Comatulae. PI . XV.
& Hufl Jev del et lkK .
■
I-4.ANTED0N GRACI LIS ,Sp.n. 5-8.ANTED0N VALIDA,Spn.
9.ANTED0N PARVIPINNA, Sp r.
PLATE XVI.
PLATE XVI.
Antedon acoela, n. sp.
Fig. 1. Side view,
Fig. 2. Dorsal aspect of a genital pinnule,
Fig. 3. The pinnule of the second brachial,
Fio-. 4. A pinnule from the middle of the aim.
Fig. 5. A young individual,
X
1 )iam.
2
Page
132
X
6
133
X
6
132
X
6
134
X
•2
133
Comabula
i
ANTEDON ACOEI.A i i
■
PLATE XVII.
(ZOOL. CHALI. EXP. PART LX. 18S8.) — < >oc>.
PLATE XVII.
Diaiu. Page
Antedon longicikra, u. sp., . . . . - . x 2 103
itulae.Pl.XVn.
s
RRA, Sp.n,
Uiricrr. Bros ixnp.
PLATE XVIII.
Fig. 1 . Side view.
Fig. 2. A distal pinnule,
Fig. 3. A lower pinnule,
PLATE XVII 1.
Fio-s. i-:3. Antedon alternata. n. sp.
° Diam. Page
. X 6 179
. x 10 179
x 10 179
Figs. 4, 5. Antedon incerta, n. sp., . . x -
Fig. 4. Side view, showing the absence of the second radial in one ray. . . 106
Fig. 5. Another aspect of the same individual, ■•
101
The Voyage of H.M.S "Challenger"
Coma.ti.fl a,.: PI XVIII .
Berjeau&HigHey del.et libh
1-3 ANTEDON ALTERNATA
4.5. ANTEDO N INCERTA
I .. arii imi)
PLATE XIX.
(ZOOL. CHALL. EXP. P.AKT hX. — 1888.) — OoO.
PLATE XIX,
Antedon breviradia, n. sp.
Fig. 1 . Side view,
Fig. 2. A smaller specimen with cysts of Myzostonia murrayi,
Fig. 3. The pinnule of the sixth brachial,
Fig. 4. A later pinnule, . . . ■
Diam.
Page
X
n
110
X
2-i-
111
X
6
110
X
6
110
I
!
ANTEOON BREVIRADIA
PLATE XX.
PLATE XX.
Figs. 1, 2. Antedon breviradia, n. sp.
Diarn. Page
Fig. 1. Side view, . . . . . . . x 4 110
Fig. 2. A distal pinnule, . . . . . .X12 110
Figs. 3, 4. Antedon bispinosa, n. sp.
Fig. 3. Side view, . . . . . . . x 3 1 1 5
Fig. 4. The pinnule of the fourth brachial, . . . x 12 115
The Voyage of R.M.S "Challenger"
Comatulae PI XX
. .
1,2 ANTEDON BREVI RAD I A . Sp n
3,4. ANTEDON BISPINOSA .
Harthart imp
PLATE XXL
(ZOOL. CHALL. EXP. — PART LX. — 1888.) — Ooo.
PLATE XXI.
Figs. 1, 2. Antedon incisa, n. sp.
Diam. Page
Fig. 1. Side view, . . x 2 124
Fig. 2. (a) Inner and (b) outer views of the pinnule on the eighth
brachial, .... . x 10 123
Fig. 3. Antedon basicueva, n. sp.
Fig. 3. A young specimen, . . . x 2 123
Figs. 4, 5. Antedon echinata, n. sp.
Fig. 4. Side view, . . . . . . .x4119
Fig. 5. The pinnule on the second brachial, . . . x 10 120
I
: xxi.
Berteau A. Highley olel
. : . . '. '.. Uth
I.2.ANTED0N INCISA.nsp.
3ANTED0N BASICURVA.n sp 4ANTEDON E C H I N A T A . ■
PLATE XXII.
PLATE XXII.
Figs. 1, 2. Antedon denticulata, n. sp.
Diani. Page
Fig. 1. Side view, . x 4 130
Fig. 2. The pinnule on the sixth brachial, . . x 6 130
Figs. 3, 4. Antedon basicurva, n. sp.
Fig. 3. Side view, .... x 2 120
Fig. 4. The pinnule on the second brachial, . . . x 8 123
... ■' iHenger"
ComaUilae Pi '
I. 2 ANTEDON DENTICULATA
3.4. ANTEDON BASICURVA ! |
Ha-nhai
PLATE XXIII.
(ZOOL. CHALL. EXP. — PART IX. — 1888.) — OoO.
PLATE XXIII.
Fig. 1 Antedon pusilla, n. sp.,
Fig. 2. Antedon tuberosa, n. sp.,
Fig. 3. Antedon aculeata, n. sp.
Diam.
Page
X
5
131
X
3
126
X
3
128
The Voyage of H. M.S. "Challengers
. ... . pi.ixm.
B«rj»» &. ftghicy dd a 1th
Mii\te™ Bros .
I.ANTEDON PUSILLA, Sp.ru 2 ANTEDON TUBER0SA,5p n
3. ANTEDON ACU LEATA.Sp n
PLATE XXIV.
PLATE XXIV.
Diam. Page
Figs. 1-3. Antedon rhomboidea, n. sp.
Figs. 1-3. The pinnules on the second, fourth, and sixth brachials, . x 4 141
Figs. 4-14. Antedon eschrichti, Mull., sp.
Fig. 4. A mature cirrus, .....
Fig. 5. " Small mature " cirrus, ....
Fig. 6. Normal young cirrus, .....
Figs. 7-9. The pinnules on the second, fourth, and sixth brachials,
Fig. 10. A "Porcupine" specimen with regenerated arm,
Fig. 11. A Challenger specimen, ....
Fig. 12. Terminal portion of an arm, dorsal aspect,
Fig. 13. Distal portion of an arm in side view,
Fig. 14. Middle portion of an arm, dorsal aspect,
X
2
143
X
2
143
X
2
143
X
2
141
X
2
141
X
2
139
X
4
140
X
4
141
X
4
153
The Voyage of H.M.S."ChallerLgerv
Corn:. ... . . .'IXIV.
L2
14
BerjeaAi & Hafikley del et lit k
1-3 ANTEDON RHOMBOIDEA,Sp.n
l;' .
4-14. ANTEDON ESCHRI CHTl .MUll.sp.
PLATE XXV.
(ZOOL. CHALL. EXP. PAKT LX. — 1888.) — OoO.
PLATE XXV.
ANTEDON ANTARCTICA, n. sp.
Figs. 1-3. The pinnules on the second, fourth, and sixth brachials,
Fig. 4. Normal young cirrus, .
Fig. 5. Another, still younger, .
Fig. 6. A mature cirrus,
Fig. 7. " Small mature " cirrus,
Figs. 8, 9. Two radial axillaries,
Figs. 10-12. Side views of three individuals,
Diam.
Page
X
4
145
X
5
.145
X
5
145
X
5
145
X
5
145
X
5
145
X
2
144
The Voyage of H M.S." Challenger"
Comalulae PI. XXV
Berjosuj & Higkley cUletlitK.
ANTEDON ANTARCTICA, Sp.Ti
Mil* Om Bl tJS . imp .
PLATE XXVI.
PLATE XXVI.
Figs. 1-3. Antedon quadrata, n. sp.
Fig. 1. A young specimen (Challenger), .
Fig. 2. An older individual (Challenger), .
Fig. 3. Another, rather small (Fseroe Channel),
Figs. 4, 5. Antedon australis, n. sp.
Fig. 4. A mature individual, . . . . . x 3 146
Fig. 5. Another, much younger, . . . . x 4 148
X
Diam.
4
Page
152
X
2
149
X
3
152
The Voyage of H M S." Challenger.'"
Comatulae PI XXVI.
Berjeau ft. HigVdey del et LtK
Minttm Broa
1-3. ANTEDON QUADRATA , Sp.n. 4-, 5ANTED0N AUSTRALIS, Sp n.
PLATE XXVII.
(ZOOL. CHALL. EXP. — PART IX — 1888.) — OoO.
PLATE XXVII.
Figs. 1-13. Antedon quadrata, n. sp.
Fig. 1. Mature cirrus, . ....
Fig. 2. Another, immature, .......
Fig. 3. " Small mature " cirrus, ......
Fig. 4. Normal young cirrus, ......
Figs. 5-7. Terminal, distal, and middle portions of the arms, in their dorsal aspect,
Figs. 8-10. The pinnules on the second, fourth, and sixth brachials of a Challenger specimen,
Figs. 11-13. The same pinnules in an Arctic specimen, .
Diam.
Page
x 2
155
x 2
155
x 2
155
x 2
155
x 4
153
x 4
152
x 4
152
Figs. 14-20. Antedon australis, n. sp.
Figs. 14-16. The pinnules on the second, fourth, and sixth brachials,
Fig. 17. A very young cirrus, .....
Fig. 1 8. " Small mature " cirrus, .
Fig. 19. Premature cirrus, ......
Fig. 20. Mature cirrus, . .....
X
4
147
X
5
147
X
5
147
X
5
147
X
5
147
Fig. 21. Premature cirrus,
Fig. 22. Mature cirrus,
Figs. 21, 22. Antedon hystrix, n. sp.
166
166
Figs. 23-29. Antedon phalangium, Miill., sp.
Fig. 23. A long-jointed cirrus, ....
Fig. 24. A short-jointed cirrus,
Fig. 25. A long-jointed cirrus, immature,
Fig. 26. Terminal arm-joints of a specimen from the Minch, .
Fig. 27. The same part of a Mediterranean specimen,
Fig. 28. The later arm-joints of a specimen from the Minch, dorsal aspect,
Fig. 29. The same part of a Mediterranean specimen,
X
2
162
X
2
162
X
2
162
X
4
163
X
4
163
X
5
163
X
5
163
|e of H.M.S "ChsJleng
i
23
J-t
Eeneaji &. HicShley- de*. et li&
: ANTEDON QUADRATA.Sp.n.. 14- -20 ANTEDON AUSTRALIS, Sp.n.
2 .12. ANTEDON HYSTRIX ,Sp.n. 23-29. ANTEDON PHALANGI U M ,Miill.sp.
PLATE XXVIII.
PLATE XXVIII.
Figs. 1-3. Antedon pealangium, Miill., sp.
Fig. 1. Side view, radial, .....
Fig. 2. Side view, interradial, .....
Fig. 3. A youug specimen,
Figs. 4, 5. Antedon hystrix, n. sp.
Fig. 4. Side view, . . . . . . . x 3 165
Fig. 5. Another specimen, with an abnormal ray, . . . x 3 167
Diam.
Page
X
2
159
X
2
159
X
6
163
The Voyage of H.M.S."ChalleTLger:
Comatulae . PI . XXVIII
V
fc Hi£Hley lei et liiti
-3. ANTEDON PHALANGIUM, Mull.sp 4,5. ANTEDON HYSTRIX,
. i
PLATE XXIX.
(ZOOL. CHALL. EXP. — PABT LX. — 1888.) — OoO.
PLATE XXIX.
Figs. 1-4. Antedon angustipinna, n. sp.
Fig. 1. Side view, ...
Fig. 2. The pinnule on the second brachial,
Fig. 3. A genital pinnule,
Fig. 4. A distal pinnule,
Figs. 5-9. Antedon eemota, n. sp.
Fig. 5. Side view,
Fig. 6. A calyx with only two radials in one ray,
Fig. 7. The pinnule on the second brachial (incomplete),
Fig. 8. The pinnule on the third brachial (incomplete),
Fig. 9. A distal pinnule,
Figs. 10-13. Antedon abyssorum, n. sp.
Fig. 10. Side view, ....
Fio\ 11. The pinnule on the second brachial, .
Fig. 12. A genital pinnule, . .
Fig. 13. A distal pinnule,
Diam. Page
X 4 189
x 6 189
x 6 190
x 6 190
X
4
184
x
4
185
X
6
185
X
6
185
X
6
185
X
4
190
X
6
190
X
6
190
X
6
190
The Voyage of H.M.S." Challenger,"
Comatulae PI. XXIX.
13
10
Bej]ea.\i & Highley del et litK
M intern Bros \mp
1-4. ANTEDON ANGUSTI PI N N A , sp n 5-3. ANTEDON REMOTA.sp.n.
1013. ANTEDON A BYSSORU M . sp n
PLATE XXX.
PLATE XXX.
Figs. 1-3. Antedon longipinna, n. sp.
Fig. 1. Side view of a young individual,
Fig. 2. An older specimen,
Fig. 3. A cirrus, ...
Figs. 4-8. Antedon tenuicirra, n. sp.
Fig. 4. Side view, . • •
Fig. 5. The pinnule of the second brachial (incomplete),
Fig. 6. The pinnule of the fourth brachial (incomplete),
Fig. 7. A distal pinnule,
Fig. 8. A cirrus, ...
Diam.
x 8
x 4
Page
186
185
185
X
4
186
X
8
187
X
8
187
X
8
187
X
2
187
The Voyage of H.M. S "Challenger:'
Comatulae PI XXX
3erjea.u k H>#,W i»l et lift, Kfctem Bros imp
1-3. ANTEDON LON Gl PI N NA.sp.n . 4-8;ANTED0N TEN U I Cl R RA, spn.
PLATE XXXI.
(ZOOL. CHALL. EXP. PART LX. 1888.) OoO.
PLATE XXXI.
Figs. 1-4. Antedon tenella, Ketz., sp.
Diain. Page
Fig. 1. A " Porcupine " specimen, . . . . x 4 170
Fig. 2. Portion of an arm of an individual from the West Atlantic
("Blake"), . . . • • • . x 6 170
Fig. 3. Cirrus of a " Porcupine " specimen, . . . x 6 170
Fig. 4. Cirrus of a " Blake " specimen, . . . . . x 6 170
Fig. 5. Antedon hirstjta, n. sp.
Fig. 5. Side view, . . . . . . . x 4 188
Fig. 6. Antedon l^evis, n. sp.
Fig. 6. Side view, . . . . . . . x 4 187
The Voyage of H.M.S'.'ChallcngeT\
Comalulae . PI . XXL
Bcrjeau &. Highley deVetlith.
1-4. ANTEDON TENELLA.Ret-zmssp. S.ANTEDON HIRSUTA,nsp.
6, ANTEDON LAEVIS.n.sp.
5ro3.in\p.
PLATE XXXII.
PLATE XXXII.
Figs. 1-4. Antedon exigua, n. sp.
Fig. 1. Dorsal aspect of an arm,
Fig. 2. Side view of an arm and pinnules,
Fig. 3. A cirrus, .....
Fig>. 4. A mature individual, ....
X
Diam.
6
Page
178
X
6
179
X
6
179
X
4
178
Figs. 5-9. Antedon alternata, n. sp.
Fig. 5. Side view of a young arm and pinnules,
Fig. 6. An abnormal individual with four radials in one ray,
Fig. 7. Side view of an arm and pinnules,
Fig. 8. A mature individual, ....
Fig. 9. A cirrus, .....
X
6
180
X
4
180
X
6
180
X
4
179
X
8
180
The Voyage of H.M.S "Challenger."
Comatulae.Pl.Xm
Eerjeau. &. Hiutiey del et lith -
SCiLerai Bros rux.
I-4.ANTED0N EX lt,UA , Sp n.
5-9= ANTEDON ALTERNATA, Sp.n.
PLATE XXXIII.
(ZOOL. CHALL. BXP. — PART LX. — 1888.) — OoO.
PLATE XXXIII.
Figs. 1, 2. Antedon abyssicola, n. sp.
Diam. Page
Fig. 1. Individual from Station 244, . . . . x 6 191
The tenth and fourteenth brachials are wrongly drawn as syzygial joints.
Fig. 2. Individual from Station 160, . . . . . x 8 191
The sixth brachials are wrongly drawn as syzygial joints.
Fig. 3. Antedon informis, n. sp.
Fig. 3. Side view, . . . . . . . x 6 205
Figs. 4, 5. Antedon tenuicirra, n. sp.
Fig. 4. Dorsal aspect of a broken specimen, . . . . x 4 186
Fig. 5. Side view of an arm, . . . . . x 4 186
Figs. 6, 7. Antedon balanoides, n. sp.
Fig. 6. The calyx, from the side, . . . . . x 2 207
Fig. 7. Portion of an arm, ... . . x 2 207
f
The Voyage of H.M.S" Challenger"
Coxnatulae . PI .
Bcr^ea-Ti & HigWey del eL liLh .
I,2.ANTED0N ABYSS ICO LA , Sp.n. 3. ANTEDON INFORMIS, Sp.r,.
4,5.ANTED0N NOTATA,Sp.n 6,7. ANTEDON BALAN 01 DES, Sp n
Mmtern Bros imp
PLATE XXXIV.
PLATE XXXIV.
Antedon carinata, Lam., sp.
Diaru. Page
Fig. 1. Side view, . . . . . . . x 2 199
■■&■
203
Fig. 2. Calyx and arm-bases of an immature individual, . . x 3
Fig. 3. Dorsal aspect of the youngest individual obtained, . . x 4 204
Figs. 4-7. Portions of arms of different individuals, to show the varying
degree of carination of the joints, . . . x 4 203
The Voyage of H.M S." Challenge
Berjexo it HigUey deL.et Kllv.
■
ANTEDON C/ F NATA .Lam.sp,
PLATE XXXV.
(ZOOL. CHALL. EXP. PAET LX. — 1888.) OoO.
PLATE XXXV.
Figs. 1-3. Antedon anceps, n. sp.
Fig. 1. Dorsal aspect, ...•••
Fi2. 2. Portion of an arm from the third brachial,
Fig. 3. A cirrus,
Figs. 4-6. Antedon milberti, Mull., sp.
Fio-. 4. Side view, ..-•••
Fio-s. 5, 6. Dorsal and ventral views of an arm,
Diam.
Page
X 3
254
X 4
255
X 5
254
x 2 194
x 4 194
The Voyage c£ H.M.S "Challenger"
Comatulae .PI . XXIV.
Bcrjeau 4 ffitfhlq nc.\ et Klh.
Mintern Bros . imp .
I-3.ANTED0N ANCEPS.spn. 4-6,ANTED0N M I LBERTI . MiiU.,sp.
HATE XXXYI.
PLATE XXXVI.
Figs. 1-6. Antedon variipinna, Carpenter.
Fig. 1. Side view,
Figs. 2, 3. Ventral and dorsal views of an arm, .
Fig. 4. Dorsal aspect of the pinnule on the sixth brachial,
Fig. 5. Dorsal aspect of the pinnule on the fourth brachial,
Fig. 6. Dorsal aspect of the pinnule on the second brachial,
Diam.
Page
. x 3
259
. x 3
256
. x 6
260
. x 6
260
. x 6
260
Figs. 7, 8. Antedon paevicirra, n. sp.
Fig. 7. Dorsal aspect, ......
Fig. 8. Lower portion of an arm in side view, .
x 4
x 4
204
205
Lge of H."M S.Thallei
. .
Berjeaii 8s n:ghley del et LiV.
I-G ANTEDON DUBIA. Sp.n.
7,8. ANTEDC I >-•-■
■
PLATE XXXVII.
(ZOOL. CHALL. EXP. — PART LX. — 1888.) — OoO.
PLATE XXXVII.
Figs. 1-3. Antedon dubeni, Bohlsche.
Diarn.
Page
Fig. 1.
Side view of the Challenger specimen, .
X
6
181
Fig. 2.
Dorsal aspect of the type specimen,
X
6
181
Fig. 3.
The pinnules on its second and fourth brachials,
Figs. 4-6. Antedon microdiscus, Bell.
X
6
182
Fig. 4.
Side view, ......
X
3
97
Fig. 5.
A cirrus, ......
X
4
97
Fig. 6.
A palmar pinnule, .....
X
4
99
The Vqvage of H.M.S"ChaJlenger'.'
Comatuke. PI. XXXVTL
Beqaui k H^Mey del et bth
l-3,ANTED0N D U B E N I , BBKlscht
4-6,ANTED0N M I CRO D I S C US , Bell.
Mintern t.i
PLATE XXXVIII.
PLATE XXXVIII.
Figs. 1-3. Antedon qtjinquecostata, n. sp.
Fig. 1. Side view, ......
Fig. 2. A lower pinnule, .....
Fig. 3. A distal pinnule, .....
Figs. 4, 5. Antedon macronema, Mull., sp.
Fig. 4. Ventral aspect of the disk and arm-bases, . . . x 3
Fig. 5. Side view, ......
Diam.
Page
x 1|
215
x 7
216
x 7
216
214
212
The Voyage of H M : ! "( Jhallsnger.'1
iiilaePLXXXVH!
S Cow.ij-d, dpJ.othtb.
1—3. ANTEDOIM Q U I N Q U E C 0 S T AT A , ;;p n
■ ■
4,5. ANTEDON M A C R 0 N E M A . Mull,
PLATE XXXIX.
(ZOOL. CHALL. ESP. — FAKT LX. — 1888.) OoO.
PLATE XXXIX.
Figs. 1-3. Antedon lusitanica, n. sp.
Diam. Page
Fig. 1. A bidistichate individual, . . . . . x 3 109
Fig. 2. Ventral aspect of the calyx and arm-bases, . . x 3 109
Fig. 3. A ten-armed individual, . . . . x 3 109
Fig. 4. Antedon disciforms, n. sp.
Fig. 4. Side view, . . . . . . . x 1\ 228
Fig. 5. Antedon clemens, n. sp.
Fig. 5. Side view, . . .... x 2 229
Tlie Voyage oJ II M S bhalli a ■
.•idaePUDOaX.
Parker & Coward, deLet Yith.
■ ■
1—3. ANTEDON LUSITANICA, :;} n
4 ANTEDON DISCIFORMIS, sp. n. 5 . ANTEDON CLEMENS, ap n
PLATE XL.
PLATE XL.
Antedon maeginata, n. sp.
Diam. Page
Fig. 1. Dorsal aspect, . . . . . • .x2230
Fig. 2. The disk, from above, . . . • • . x 2 231
The Voyage of H.M.S.'bhalLenger"
lorn,
Parker &.< JcTWBxaLdal ecTtch
.
ANTEDON MARGINATA. ■
PLATE XLI.
(ZOOL. CHALL. EXP. — PART LX. — 1888.) OoO.
PLATE XLI.
Antedon compressa, n. sp.
Diam. Page
Fig. 1. Side view, . . . . . . . x 3 222
Fig. 2. The disk, from above, . . . . . . x 3 222
Fig. 3. A distal pinnule, . . . . . . x 4 222
Fig. 4. The pinnule on the fourth brachial, . . . x 4 222
The Voyage of H.M.S"Gha ei
a
v Coward del .ethQi-
.
ANTE DON C 0 M PR ES S A, sp.n
PLATE XLII.
PLATE XLII.
Diam. Page
Antedon FLEXILIS, 11. sp., . . - ... X 1¥ 217
Voyage of
..
Earkenr & Coward, dr I e
:■ . ■
ANT EDO N F LE.XI LI S, sp. n
PLATE XLIII.
(ZOOL. CHALL. EXP. — PART LX. — 1888.) — Ooo.
PLATE XLIII.
Diani. Page
Antedon patula, n. sp., . x 1?
x 11 219
The Voyage of H.J/1 "■ ■ enger!
:'..,.tulae.Pl '
Par-ker Sl Covrard 4;L et liLh
Weat,Nevnnitji X-Cfunp.
A NTE D 0 N PATULA,3pn
PLATE XLIV.
PLATE XLIV.
Fig. 1. Anteeon eobusta, n. sp.
Diam. Page
Fig. 1. Side view, . . . . . . . x 1^ 220
Figs. 2, 3. Anteeon manca, n. sp.
Fig. 2. Ventral aspect of the calyx and arm-bases, . . . x 3 227
Fig. 3. Dorsal aspecl^of the same, . . . . . x 3 226
I'he Voyage of H M S Challe] i
Q
Parker & Coward del etlirjj
M
ANTEDON ROBUSTA sp n
2. 3. ANTEDON MANCA sp. ti.
PLATE XLV.
(ZOOL. CHALL. EXP. — PART LX. 1888.) — OOO.
PLATE XLV.
Fig. 1. Antedon conjungens, n. sp.
Diani. Page
Fig. 1. Dorsal aspect, ... . . x 2 233
Figs. 2, 3. Antedon tuberculata, n. sp.
Fig. 2. Dorsal aspect, . . . . . . x 2| 232
Fig. 3. The lower pinnules, . . . ... .x4 233
Fig. 4. Antedon angustiradia, n. sp.
Fig. 4. Side view,
. x 3 253
The Voyage of H/M. S . Gh.aTLei] •
ComatulsLe .PI JXLV.
Parker 5c Cowaradel.etlifch
Ne-wmsn ftCimp
I . A IN T E D 0 N CONJUNGENS, 3p. n.
2,3. ANTE DO IN TUBERCU LATA Sp. n 4-. ANTEDOIM A IN G U S T I R A D I A , sp. n.
PLATE XLVI.
PLATE XLVI.
Diam. Page
Antedon regalis, n. sp., . . x 2| 237
Fig. 1. Dorsal aspect.
Fig. 2. The disk, from above.
The Voyage of H.M. ^'Challenger1'
Comatulat: PI XI, VI
Parier S: C<jwajTA,del etlith
'
ANTEDON REGALIS, ap. n.
PLATE XL VII.
(ZOOL. CHALL. EXP. PART LX. — 1888.) — OoO.
PLATE XLVII.
Figs. 1-3. Antedon similis, n. sp.
Diam. Page
Fig. 1. Dorsal aspect, . . . . . . . x 2 235
Fig. 2. The disk, from above, . . . . . . x 2| 235
Fig. 3. Diagram of the ray-divisions, . . . . x 2^ 235
Figs. 4, 5. Antedon quinduplicava, n. sp.
Fig. 4. Side view, . . . . . . . x 2| 262
Fig. 5. Diagram of the ray -divisions, . . . . x 3 262
The Voyage • ' .Tialleiiger"
Comatiilae.PlJLVIl
^rW fc Ccwairl dtl « hik
. ,,
1-3. ANT E. DO N SIMILIS, sp. n.
4, 5. ANTEDON p U I N DU PLICAVA, cp. ji.
PLATE XLVIII.
PLATE XLVIII.
Figs. 1, 2. Antedon occulta, n. sp.
Fig. 1. Dorsal aspect,
Diam. Page
x 2 236
Fig. 2. The base of an arm, with the pinnules of the second, fourth, and
sixth brachials, . . . • • . x 5 237
Figs. 3-5. Antedon variipinna, Carpenter.
Fig. 3. The base of an arm, with the pinnules of the third, fifth, and
seventh brachials, ......
Fig. 4. The disk, from above, .....
Fig. 5. Dorsal aspect, ......
X
5
260
X
3
256
X
3
256
• ■ of H.M.S .Challenger"
Comaxulae PI XLVII1
-
Parker & Coward, deL et lith-
I, 2.ANTED0N OCCU LTA . sp. n
3_5 ANTEDON VARIIPINNA, Carpenter.
PLATE XLIX.
(ZOOL. CHALL. EXP. — PART LX. — 1888.) — Ooo.
PLATE XLIX.
Figs. 1,2. Antedon variipinna, Carpenter.
Diam. Page
Fig. 1. Dorsal aspect, . . . . . . . x 3 256
Fig. 2. A lower pinnule, . . . . . . x 3 260
Figs. 3, 4. Antedon occulta, n. sp.
Fig. 3. Dorsal aspect, . . . . , . x 3 236
Fig. 4. The base of an arm, with the pinnules of the second, fourth, and
sixth brachials, . . . . . x 5 237
The Voyage of H.M.S tlhall :nger
Com,: I XL IX.
Pai-kig- ScCowa-rd, del.etlith
I, 2. ANTE DON VARII PINNA, Caj-penter.
W*at,Newmaji&C? imp
3,4-. ANTE DON OCCULTA, 3p.Il.
PLATE L.
PLATE L.
Figs. 1, 2. Antedon angusticalyx, n. sp.
Fig. 1. Side view, ......
Fig. 2. A genital pinnule, .
Diam. Page
242
x 21
243
Figs. 3-6. Antedon multispina, n. sp
Fig. 3. Calyx and arm-bases of a tridistichate individual,
Fig. 4. The disk, from above, ....
Fig. 5. Portion of an arm, in side view,
Fig. 6. A cirrus, .....
X
3
249
X
3
249
x
3
249
X
3
249
The Voyage oF H.M.S"ChaJLenger"
Con U iia.e i'l
■ .V. Cow axd, dele l .
I.2.ANTED0N A N G USTICALYX, sp. n.
Weat,KewmiwYk0o imp
3 6 A NT ED ON MULTISPINA.sp n
PLATE LI.
(ZOOL. CHALL. EXP. — PART LX. — 1888.) — OoO
PLATE LI.
Diam. Page
Fig. 1. Antedon distincta, n. sp., . . . . . x 2 247
Fig. 2. Antedon IN.EQUALIS, n. sp., . . . . x 2 244
'he Voyage of II. M I) GhaJlenfr r
Coma lulae.PJ.-LJ
Parknr &. Coward, del. etlith.
..■ . ,
I ANTEDON D I ST I N CT A, Bp.n.
2. ANTEDON I N/EQUALIS sp. n.
PLATE LIL
Diam. Page
x 3 301
PLATE LI I.
Figs. 1, 2. ACTINOMETRA PULCHELLA, Pourt., sp.
Fig. 1. A " Porcupine " specimen, ....
Fig. 2. A doubtful specimen from the Arafura Sea, . . . x 3 306
Figs. 3-5. Antedon pokrecta, n. sp.
Fig. 3. The calyx and arm-bases, from the side,
Fig. 4. The same, from above, .....
Fig. 5. Side view of an arm, .....
X
2
250
X
2
250
X
9i
•^2
250
, . ge of H.M.S ChaJlf
Gomatuiie PI LI1
; .riii,- ! ward I e !ith
I, 2. ACTINOMETRA PULCHEL.LA, Pourt, sp.
3_5. ANT EDO N PORRECTA, Sp. xi.
PLATE LTTL
(ZOOL. CHALL. EXP. PART LX. 1888.) Ooo.
PLATE LIII.
FigS. 1-14. ACTINOMETRA SOLAEIS, Lam., Sp.
Fig. 1. Dorsal aspect of a young individual,
Fig. 2. An older individual, .....
Figs. 3, 4. The first pair of pinnules, ....
Figs. 5, 6. The second pair of pinnules,
Figs. 7, 8. The third pair of pinnules, ....
Figs. 9, 10. The first pair of pinnules in another example,
Figs. 11, 12. Its second pair, .....
Figs. 13, 14. Its third pair, ......
Figs. 15-22. ACTINOMETRA PECTINATA, Retz., sp.
Fig. 15. Dorsal aspect. . - • ■ . x 1^ 285
Figs. 16, 17. The first pair of pinnules, . . . . x 3 281
Figs. 18, 19. The second pair of pinnules, . . . . x 3
Figs. 20, 21. The third pair of pinnules,
Fig. 22. The pinnule on the eighth brachial, . . • . x 3
1 >iam.
Page
X
3
290
X
H
288
X
3
281
X
3
282
X
3
288
X
3
282
X
3
28]
X
3
281
281
3 281
285
The "Voyage of H M S .Challengei "
Camatulae.Pl LIII
Parker Se Cowa.nl, dsl et li£h
1 — 14. ACTINOMETRA SOLARIS, l.m, sp.
15-22. ACTINOMETRA PECTINATA, Ret:
PLATE LIV.
PLATE LIV.
ACTINOMETEA PAUCICIRRA, Bell.
Diam. Page
Fig. 1. Dorsal aspect, . . . . . . . x 3 291
Fig. 2. Dorsal aspect of the calyx and one ray with a palmar series, . x 3 293
Figs. 3-9. Various stages in the modification of the centro-dorsal, x 3 14
Fig. 10. The youngest specimen obtained, . . . . x 3 293
The Voyage oi' H.MS.'Cliall.-i i
Comatul li ■ i I '.
raiku- fc. Coward, AbL«I lilX
WitttjBcwiBinJ ; -T' j.
ACTI N OM ETR A PAUCICIHRA.
PLATE LV.
(ZOOL. CHALL. EXP.— PART LX. 1888.) 000.
PLATE LV.
Diain. Page
Fig. 1. ACTINOMETRA DISTINCTA, n. sp., . . . . . X 3 295
Fig. 2. ACTINOMETRA MACULATA, n. sp., . . . . X 3 307
r/age af H.M. S .' Challenger"
Cama.tulae.H.LV
Parker & Coward, del etli'Ji
■
I, ACTIN0METRADISTINCTA8Pm 2, ACTI NOMETRA MACU LATA, 5M
PLATE LVL
PLATE LVI.
FigS. 1, 2. AcTINOMETRA MERIDIONALIS, Pourt., Sp.
Fig. 1. Dorsal aspect, ......
Fig. 2. Terminal comb of a lower pinnule,
FigS. 3, 4. ACTINOMETRA MULTIBRACHIATA, n. sp.
Fig. 3. Dorsal aspect of the calyx and two rays,
Diam. Page
x 4 301
x 20 276
x 2 299
Fig. 4. Terminal comb of a lower pinnule, . . . x 20 276
TheVoya.£e of H.M. S ." Challenger "
Comatulae. PI. LVI.
■'/,-•
,. , 'x;'-
Geo West & 6ons del bth et unp.
I, 2. ACTINOMETRA M E RID ION ALIS, Pourt., sp.
3, 4.. ACTINOMEITRA M ULTI BRACHIATA , sp n.
PLATE LVIL
(ZOOL. CHALL. EXP.— PAKT LX.— 1888.)— Ooo.
PLATE LVII.
Fig. 1. Actinometra typica, Loven, sp.
Diam. Page
Fig. 1. Dorsal aspect of the calyx and two rays, . . x 3 296
Figs. 2-4. Actinometra elongata, n. sp.
Fig. 2. Dorsal aspect, .....
Fig. 3. Ventral aspect of the disk,
Fig. 4. Pinnules with " ovoid bodies," .
X
4
311
X
4
273
X
5
275
The Voyage of H.M.S. Challenger '.'
Oomatulae. Pl.LVII.
Parlrar X, Coward, Asletlith
WeBt.Uewmaji&Cimp
I, ACTINOMETRA TYPICA, Loven.sp 2-4-, ACTINOMETRA ELONG ATA,spn
PLATE LVIII.
PLATE LVIII.
ACTINOMETEA STELLIGERA, n. sp.
Diam. Page
Fig. 1. Side view, . . . . . . x 2 308
Fig. 2. The lower pinnules, . . . . . . x 4 308
The Voyage of H.M.S ."C}l^tlleIlge^.,'
! ma.tulae i'l LVili
■
lOUUL&C^h:
ACTINOMETRA ST E LLi G E R A, sp.n.
PLATE LIX.
(ZOOL. CHALL. EXP.— PABT LX.— 1888.) — OoO.
PLATE LIX.
Fig. 1. ACTINOMETRA SIMPLEX, n. Sp., .
Fig. 2. ACTINOMETRA ROTALAEIA, Lam., Sp.,
Fig. 3. ACTINOMETRA VALIDA, n. Sp., .
Diam.
Page
x 3
312
x 3
313
x 2
314
\ a^eafH M.S.'bhalleri
.
WS.Evans.del
ParkEr & Cow:, i
WeBt,Tfewmaii&C?iinp
I. ACTINOMETRA SIMPLEX, sp. 71.
2. ACTINOMETRA ROTALARIA Lam.,sp. 3. ACTINOMETRA V A LI DA, gp. n.
PLATE LX.
PLATE LX.
Figs. 1, 2. ACTINOMETRA COPPINGEIU, Bell.
Diam. Page
Fig. 1. An individual from Saniboangan, . . . . x 3 320
Fig. 2. Another from Banda, . . . . - . x 3 320
Fig. 3. ACTINOMETRA LINEATA, n. Sp.
Fig. 3. . . • • • • • . x 3 327
N.B. — Some of the cirri have been represented by the artist with too many joints.
W.S.'Chal
, c.del-
-
1,2. ACTINOMETRA COPPINGERl. Bell. 3 ACTI NOMETRA LINEATA, sp n
PLATE LXI.
ZOOL,
. CHALU EXP. PART LX. 1888.)— Ooo.
PLATE LXI.
ACTINOMETRA PARVICIREA, Mull., Sp.
Fig. 1. A specimen with many arms and few cirri ; the second radial is
absent in one ray, .....
Fig. 2. A smaller individual, without palmars, ....
Fig. 3. The calyx of another example with stellate centro-dorsal bearing-
rudimentary cirri,
Fig. 4. Another specimen with more rounded centro-dorsal,
Fig. 5. A larger form with post -palmars,
Fig. 6. Another example from Simon's Bay,
Fig. 7. Ventral aspect of its arm with the ova attached,
Figs. 8-10. Modifications of the terminal comb on the lower pinnules, .
Diam. Page
X
2
343
X
6
343
X
3
343
X
4
343
X
3
338
X
3
343
X
4
344
X
10
343
The Voyage of H M". S . Challenger."
Cama.tulae Pl.LXI.
Qm** <X
West. ,Nevaaaji2tC? imp.
W.SJEvans.del
Parker & Coward, Ml1-
ACT1IM0METRA P A R V I C I R R A , M U L L, sp
PLATE LXII.
PLATE LXII.
Fig. 1. ACTINOMETRA QUADRATA, U. Sp.
Diam. Page
Fig. 1. Dorsal aspect. . . . . . . .x4331
Figs. 2-4. ACTINOMETRA FIMBRIATA, Lam., sp.
Fig. 2. Ventral aspect of an unusually symmetrical disk, . . x 3 319
[The arrow indicates the line of the antero-posterior axis.]
Fig. 3. Dorsal aspect, . . . . . . .x3317
Fig. 4. A more normal disk, . . . . . . x 3 319
The Voyage of HI S . Challenger"
ComatulM. Fl
Paxkwr &. Coward, dflL et lah.
WCR!.,Mt!lS ".■'■
I, ACTINOMETRA Q U A D R AT A , sp n. 2-4-, A C T I N 0 M E T R A FIMBRIATA Lam.sp
PLATE LXIIJ.
(ZOOL. CHALL. EXP. PART LX. — 1888.) OoO.
PLATE LXIII.
Figs. 1-5. ACTINOMETRA TKICHOPTERA, Milll., sp.
Fig. 1. Dorsal aspect, ......
Fig. 2. Portion of an arm, from the side,
Fig. 3. The same, from above, .....
Fig. 4. A cirrus, ......
Fig. 5. Terminal comb of a lower pinnule,
Figs. 6-8. ACTINOMETRA DIVARICATA, n. Sp.
Fig. 6. Dorsal aspect, . . . . . . .x2332
Fig. 7. Terminal comb of a lower pinnule, . . . . x 7 276
Fig. 8. Dorsal aspect of an arm, . . . . . x 2^ 332
Diam.
Page
X
6
345
X
4
345
X
4
345
X
6
345
X
15
276
The Voyage of H M S "Challenger."
Comatulae. PL LXffl.
\
Berjeau. &. HigWey del etlith.
Mint em. Bros .
-5. ACTINOMETRA TRICHOPTERA,Mull.sp. 6-8. ACTINDMETRA DIVARICATA, Sp.-n.
PLATE LXIV.
PLATE LXIV.
Figs. 1, 2. AcTINOMETRA BELLI, n. sp.
Diam. Page
Fig. 1. Dorsal aspect, . . . . . . x 2 334
Fig. 2. The disk, from above, . . . . . . x Lj 335
Fig. 3. ACTINOMETKA DUPLEX, 11. sp.
Fig. 3. Dorsal aspect, . . . . ... . x 3 335
The Voyage of H .M. S . Challenger"
Comatulae. Pl.LXIV.
Parker JcCcrwari^dc^etlL di.
W#8f .N^Tonatii5;<-?jmp
I, 2. A CT I N 0 M ETR A BELLI, sp n. 3. A C T I N 0 M E T R A D U P L E X , sp . n.
PLATE LXV.
(ZOOL. CHALL. EXP,— PART LX.— 1888.)— OoO
PLATE LXV.
ACTINOMETRA NOBILIS, n. sp.
Fig. 1. Dorsal aspect, ......
Figs. 2-6. Various stages in the modification of the centro-dorsal,
Fig. 7. Terminal comb of a lower pinnule,
Fig. 8. Portion of the disk, with cysts of Myzostoma platypus,
Diam.
Page
iat. size
336
x H
15
x 20
276
x 2
338
TheVoyage of H.M. S " Challenger
Comatulae. PI. U2T.
f>f'P\
-K^Jg^-
A
.^>
f-ft*
^<Mjy^
\:, <■?-'
.
:
Geo West * Sons del hth et tmp.
ACTINOMETRA NOBILIS, sp t,
PLATE LXVI.
PLATE LXVI.
FigS. 1-3, 8. ACTINOMETEA MULTIRADIATA, Linil., Sp.
Diam. Page
Fig. 1. Dorsal aspect, . . . . . . . x 2 '322
Fig. 2. A terminal pinnule, . . . . . . x 4 322
Fig. 3. A lower pinnule, . . . . . x 4 276
Fig. 8. Dorsal aspect of an arm, . . . . . x 2 322
Figs. 4-7. Actinometra sentosa, n. sp.
Fig. 4. Dorsal aspect, . . . . . . .x2325
Fig. 5. A lower pinnule, . . . . . x 4 276
Fig. 6. A terminal pinnule, . . . . . x 4 325
Fig. 7. Dorsal aspect of an arm, . . . . x 2 325
The Voyage of H.M.S"Challenge^,.,
Comatulae. PI. LXVI.
Berje^u & HiiWeu dej.et lilt
1-3,8. ACTINOMETRA MULTIRADIATA.Iinn.sp.
4-7 ACTINOMETRA SENTOSA,spn
Mintem ii
PLATE LXVIL
(ZOOL. CHALL. EXP. — PART LX. 1888.) — OoO.
PLATE LXVII.
Figs. 1, 2. ACTINOMETKA LITTOEALIS. n. sp.
Diam. Page
Fig. 1. Dorsal aspect, . . . . . . . x 2 346
Fig. 2. Terminal comb of a lower pinnule, . . . x 20 276
Figs. 3, 4. AcTINOMETRA PAEVICIRRA, Mill]., sp.
Fig. 3. Dorsal aspect, . . . . . . .x2338
Fig. 4. Terminal comb of a lower pinnule, . . . x 20 276
TheVoya£e afH.M.S " Challenger "
Comatulae. PI LXVn.
I, 2. ACTINOMETRA LITTORALIS, sp n.
3, 4.. ACTINOMETRA PARVICIRRA. Mull, sp
Geo West it 6ons del hih.etuup.
PLATE LXV11I.
PLATE LXVIII.
ACTINOMETRA REGALIS, 11. Sp.
Diam. Page
Fig. 1. The disk, from above, . . . . . . x 2 274
Fig. 2. Dorsal aspect, . . . . . . . x 2 347
Fig. 3. Terminal comb of a lower pinnule, . . . x 20 276
TheVoyage of H M S " Challenger
Comatulae. PI. LXVIII
GeoWest & 6one. del btii et imp
ACTINOMETRA REGALIS, sp n.
PLATE LUX,
(ZOOL. CHALL. EXP.— PART LX. — 1888.)— OoO.
PLATE LXIX.
FigS. 1-4. ANTEDON MULTTSPINA, U. Sp.
Diam. Page
Figs. 1, 2. Side views of a tridistichate individual, . . . x 4 249
Fig. 3. The disk, from above, . . . . . . x 4 249
Fig. 4. Dorsal aspect of an arm, . . . . x 4 117
Figs. 5-7. Promachocrinus abyssorum, n. sp.
Fig. 5. Side view, . . . . . . x 4 351
Fig. 6. Basal portion of a cirrus, . . . . x 4 351
Fig. 7. («, b) Arm fragment with genital pinnules, . . . x 4 351
Figs. 8-10. Promachocrinus naresi.
Fig. 8. Dorsal aspect of an arm, ....
Fig. 9. Dorsal aspect of the calyx, ....
Fig. 10. The calyx, from the side, ....
X
2
352
X
2
352
X
2
352
Tl is Voyage of H.M.S . bhaLLenger"
Comatulce . P1.LX1X.
10
Parker & Coward, del etHtJi.
Wrai.,Howr.' i V. ' ,, i
1-4. ANTE DON M U LT I S P I N A , sp n
5_7 PROMACHOCRINUS ABYSSORUM sp.n.8, 9, PROMACHOCRINUS NARESI,sp.n.
PLATE LXX.
PLATE LXX.
Promachocrinus kerguelensis, n. sp.
Diani. Page
Fig. 1. Side view, . . . x 2 350
Fig. 2. A young individual, . . . . . x 4 351
The Voyage of H.M.S'.'Challenger'
Comatulae.Pl. LXX.
Bcrjeavi &HigHey dfci et Ltk.
PROMACHOCRiNUS KERGUELENSIS, Spiv.
Mintern Bros, imp.
THE
VOYAGE OF H.M.S. CHALLENGER.
ZOOLOGY.
REPORT on the Seals collected during the Voyage of H.M.S. Challenger
in the Years 1873-76. By Sir William Turner, Knt., M.B., LL.D.,
F.R.SS. L. & E., Professor of Anatomy in the University of Edin-
burgh, Member of the General Medical Council.
In the first volume of the Reports of the Scientific Results of the Voyage of H.M.S.
Challenger (Zoology, vol. i., part iv., 1880) I gave an account of the Bones of the
Cetacea which had been collected by the expedition.
In this Report it is my intention to describe the Seals, and therefore to complete the
account of the Marine Mammals which were entrusted to me for examination and
description.
Specimens of Seals were procured by the expedition in the following localities : —
The Kerguelen Group of Islands.
Heard Island.
Messier Channel, off the west coast of Patagonia.
The Falkland Islands.
Juan Fernandez.
These specimens belong to the genera Macrorhinus, Leptonychotes, Otaria, and
Arctocephalus.
(zool. chall. exp. — part Lxvui. — 1887.) Yyy 1
CONTENTS.
page
PART I.— Description of Genera and Species, . ...
3
Macrorhinus, .........
3
Macrorhinus leoninus, .......
3
External Characters, ....
3
Skeleton, ... ....
5
Leptonychotes, ......
20
Leptonychotes weddelli, .....
20
Comparison of Skull with Stenorhynch us leptonyx,
20
Skeleton, ........
23
Otaria, ... ......
29
Otaria jubata, ........
29
External Characters, ... ...
29
Skull, ... . . .
29
Arctocephalus, ... .
36
Arctocephalus gazella, .......
36
Skull, . . . .
37
Arctocephalus atistralis, .......
39
External Characters, .......
38
SkulL
41
Comparison of Skeletons of Arctocephalus gazella and Arctocephalus australis,
43
Arctocephalus from Juan Fernandez, ......
52
PART II. — Classification of the Pinnipedia, ....
55
P h o c i d se,
55
PhocinaB,
57
Phoca, .
57
Halichosrus,
61
Ogmorhinina?,
62
Ogmorhinus,
62
Leptonychotes,
.
63
Ommatophoca, .
63
Monachus, .
64
Cystophorinse, .
65
CystopJwra,
65
Macrorhinus,
66
Trichechidse,
56,67
Trichechus (Odobxnus),
68
Otariidae,
56,70
Otaria, .
73
Eumetopias,
73
Arctocephalus, .
79
PART III. — Brain of the Elephant
Seal and Walrus, ....
89
Brain of Elephant Seal,
91
Brain of Walrus, .
102
Comparison of the Convolutions of the Seals aud Walrus with those of the Carnivora and
of Apes
i
and Man, .........
113
PART IV. — Viscera of Elephant Seal,
135
APPENDIX.— The Myology of the I
'innipedia. By Dr W. C. Strettell Miller,
139
PART I.
DESCRIPTION OF GENERA AND SPECIES.
Macrorhinus, F. Cuvier.
Macrorhine, F. Cuvier, Mini, du Museum, xi. p. 200, 1824.
In the Narrative of the Voyage of the Challenger it is stated that the Elephant Seal
was seen on Marion Island, in the South Atlantic : on Kerguelen Island and on Heard
Island -,1 whilst reference is made to the observations by previous voyagers of the occur-
rence of this Seal on the Crozet Islands, Tristan da Cunha, Inaccessible Island, and Juan
Fernandez, which places were also visited by the Challenger. Specimens were secured
and sent home both from Kerguelen Island and Heard Island. They were males and
females of the great Elephant Seal of the Southern Ocean, and are all I beHeve to be
referred to one species, Macrorhinus leoninus.
Macrorhinus leoninus (Linnaeus) (Pis. L, II., III., IV.).
Phoca leonina, Linnfeus, Syst. Nat., ed. 12, L p. 55, 1766.
Elephant Seal.
The specimens of this Seal consisted of (a) the carcase of a well-grown female, (b) the.
skin of another female containing bones of the limbs, (c) the complete skeleton of another
female, and (d) the skull of a young female from which the brain had been removed, all
killed at Christmas Harbour, Kerguelen, January 7, 1874 ; also (e) the skeleton of a well-
grown male, (/) the skull and some of the cervical vertebras of a large female shot at Betsy
Cove, Kerguelen, January 9, 1874, together with (g) a young skull, apparently of a
female, picked up on the same beach ; also (h) the skull of a large male picked up by
Professor Moseley on the shore of Heard Island, February 6, 1874.
External Characters. — I have been able to examine the external characters of the
Sea Elephant in the carcase of the well-grown female (a) and the skin of another female (b),
both killed on Kerguelen Island and preserved in salt. The carcase with the skin on had
been cut into halves immediately behind the pectoral limbs, and the abdominal viscera had
been removed, so as to allow it to be packed in a barrel. It is from this specimen that
the drawings in PI. I. have been made. When the halves were placed in contact in
1 Narr. C'hall. Exp., vol. i. pp. 294, 354, 373, 377.
4 THE VOYAGE OF H.M.S. CHALLENGER.
the plane of section the animal measured along the curve of the back, from the muzzle to
the tip of the tail, 6 feet 2 inches, and to the most distal point of the hind limb, when it was
elongated behind the tail, 7 feet 2 inches. The free part of the tail was only 2f inches long,
the girth immediately behind the pectoral limb was 3 feet 2 inches in the eviscerated carcase.
The other female, judging from the dimensions of the separated skin, had apparently been
somewhat bigger, as the length from muzzle to tip of hind limb was 7 feet 8 inches.
Between thirty and forty black bristles arranged in six rows on each upper lip, pro-
jected for the most part backwards, and the more posterior bristles were longer than the
anterior. The lips overhung the mouth, the slit of which was 2f inches long on each side.
The nostrils opened forwards on the face, but there was no proboscis. Immediately
above these orifices were three transverse wrinkles in the skin, from the upper of which
a single black bristle projected at each end, and by the depression of these wrinkles,
through the contraction of the facial muscles, that peculiar appearance of the face is
produced, which, in the Narrative of the Voyage, the animal is stated to assume when
disturbed. The palpebral fissure was if inch wide, and seven black bristles projected
from the skin, a short distance above the inner end of each of these fissures. A large
extensde third eyelid was situated at the inner canthus, which could be drawn outwards
so as to cover the front of the globe. The orifice of the external meatus, so small as
only to admit a fine probe, was situated 2^ inches behind the external canthus.
The pectoral limb was 1 2 inches long and 4 inches in greatest width ; five digits, each
with a long and strong convex nail on the dorsum, were individualised at the distal
border of the manus. A groove on the surface of the limb, between the pollex and index,
was short and shallow, but the surface grooves which differentiated the other digits
became deeper and more elongated as one passed from the second to the fifth digit, and
possessed a narrow intermediate web. Both surfaces of the limb were covered with hair,
and the nails projected beyond the distal border of the limb.
The left hind limb measured from the head of the thigh bone to the tip of the
innermost digit 25^ inches, and from the fold at the root of the tail to the tip of the
same digit 12^ inches ; but the corresponding measurements were not cpaite so long on
the right side. The pes was pentadactylous. The first and fifth toes were very
much longer and bigger than the three intermediate digits, so that the posterior free
border of the foot was deeply concave. A broad and deep web connected the toes
together, and only the tips of the three intermediate digits projected beyond the free
border of the web, but their outline was distinct on both the dorsal and plantar surfaces;
more especially on the latter. The web was longitudinally wrinkled, and permitted the
toes to be approximated or drawn widely asunder, so as to make the greatest width of
the foot 1 1 inches. Both the dorsal and plantar surfaces of the foot were haired up to
the free distal border of both web and toes, and no trace of nails was seen.
The vent was elongated from before backwards, and was situated a little in front of
REPORT ON THE SEALS. 5
the ventral aspect of the tail. Into it the urethra, vagina, and rectum opened. A pair
of nipples projected from the abdominal wall about ] 6 inches in front of the vent ; when
pressed on, each nipple receded into a depression in the integument,
The hair of the face and back was dark grey, dashed with a brown or yellow tint ;
down the sides and belly it was lighter and more yellow, with a dash of reddish-brown,
but the brownish tint was to some extent due to discoloration from the oil which had
escaped out of the blubber amongst the hair. In Dr. Gray's figure of a female Elephant
Seal in the Voyage of the "Erebus" and "Terror" (pi. ix.), the face and the sides and
belly are coloured a lighter yellow than was seen in my specimen. Mr. Eaton 1 does not
refer to the yellow colour of the hair, which he says in some specimens is uniformly
reddish-brown, in others is pale, blotched and spotted with darker grey.
Skeleton. — There does not appear to be on record any detailed description of the
skeleton of the Elephant Seal, or of the characters which differentiate the bones of the
male and female. F. Cuvier has given 2 a short description of the skull. Some measure-
ments both of the skull and other bones have been recorded by Mr. J. A. Allen in his
monograph on the North American Pinnipeds.3 Professor Flower has described some
characters of the cranium of a splendid specimen of a large male from the Falkland Islands,4
and in the Catalogue of the Skeletons of the Mammals in the Museum of the Eoyal
College of Surgeons of England5 he has given the length of the articulated skeleton as
4500 mm. from tip of nose to end of tail, and 4890 mm. to end of posterior digits. Dr.
St. George Mivart has also published6 short notes on the cranium. A more detailed
description is still, however, a desideratum. It is especially necessary to make a com-
parison of the male and female crania, as they differ from each other so much in size,
and to some extent in relative proportions, that a naturalist, in ignorance of the animals
from which they had been obtained, might easily regard them as belonging to different
species. As the collection contained crania in different stages of growth, some obser-
vations on the influence of age on the skull have also been made.
Skull. — The skulls which have been examined whilst writing the following description
consisted not only of those collected by the Challenger, both males and females (p. 3),
but of a fine male, which had been shot on Heard Island, presented to me for the
Anatomical Museum of the University of Edinburgh by my former pupil and assistant,
Professor J. Halbday Scott of Dunedin, N.Z.
In the following table I have stated the measurements of the three male skulls, and of
the large female (f) shot at Betsy Cove ; and to allow their relative size to be compared
with that of other recorded specimens, I have, in addition to a number of new measure-
ments, also adopted those employed by Professor Flower in his account of the skull of
1 Proc. Roy. Soc. Lond., vol. xxiii. p. 502, 1875, and Phil. Traiu., vol. clxviii. p. 96, 1879.
2 M4m. du Museum, t. xi. p. 200, pi. 14, 1824. 3 U.S. Geol. Survey, Washington, 1880.
4 Proc. Zool. Soc. Lond., January 4, 1881. 5 London, 1884. 6 Proc. Zool. Soc. Lond., May 19, 1885.
6
THE VOYAGE OF H.M.S. CHALLENGER.
the Elephant Seal obtained from the Falkland Islands, and, for convenience of reference,
have included in the table the measurements of his specimen.
From this table it will be seen that none of these skulls equalled in length the
Falkland Island specimen, or indeed two other large skulls referred to in Professor
Flower's table on p. 147 of his memoir, but that they closely approached in length the
skull in the Berlin Museum obtained in Kerguelen Island by the German Transit of Venus
expedition. In their other dimensions also they were much smaller than the skull from
the Falkland Islands above referred to, which is much the largest specimen that has yet
been measured. The very material difference between the dimensions of the male skulls
and the largest female will at once be recognised. It will be of importance therefore to
determine, if possible, whether this difference is sexual or merely due to a difference in
relative age. The female skull is, I believe, to be regarded as approaching adult, for the
occipito-sphenoidal synchondrosis was obliterated, except for a faint trace on the surface
of the bone on each side, and the junction between the pre- and post-sphenoids was only
indicated by a surface mark. But it should be stated that in the cervical vertebra?
which accompanied this skull the plate-like epiphyses of the bodies were not ankylosed.
In the female skull next in size to the above, the condylo-premaxillary length of which
was 274 mm., both the basi-cranial synchondroses were unossified, though the interval
between them was narrow.
Table I. — Crania of Elephant Seal.
Length from front of premaxilla to occipital condyle,
,, ,, to occipital crest,
Extreme interzygomatic width, ....
Extreme width between occipital crests, .
Greatest width at posterior edge of external meatus,
Greatest width of palate, .....
Width of maxilla across middle of rostrum,
Width between outer sides of base of upper canines,
, , , , upper lateral incisors,
,, „ lower canines,
Length of palate from mesial notch behind to incisor teeth,
Height of skull from basion to middle of occipital crest, .
Smallest inter-frontal width in plane of upper surface,
Length of nasals, ......
Greatest width of anterior nares, ....
Vertical diameter of mea-ethmoid at anterior nares.
From anteroinferior angle of mes-ethmoid to central incisor,
Greatest length of mandible, ....
Greatest width at condyles of lower jaw, .
rj
*
6
9
3
Challenger.
Prof. Scott.
Challenger.
Challenger.
Prof. Flower.
Heard
Heard
Kerguelen
Kerguelen
Falkland
Island, h.
Island.
Island, e.
Island. /.
Islands.
mm.
mm.
mm.
mm.
mm.
486
493
402
300
564
508
497
392
296
597
354
350
281
222
3S4
203
201
187
171
242
284
303
253
199
178
154
124
89
185
168
160
122
73
176
160
146
106
62
158
5S
43
31
60
83
64
38
93
250
248
176
128
272
167
160
141
112
65
71
52
38
55
65
55
43
98
96
82
48
80
85
64
48
158
145
127
85
350
326
253
191
375
318
236
195
352
In Dr. Scott's male the occipito-sphenoidal synchondrosis was obliterated mesially,
but on each side a gap about 1 mm. wide separated the two bones ; the synchondrosis
between the pre- and post-sphenoids was, however, open both mesially and laterally.
REPORT ON THE SEALS. 7
This skull, therefore, had not yet reached a stage in which the possibilities of additional
growth in length at the basis cranii were exhausted. In the Challenger specimens from
Heard and Kerguelen Islands both the occipito-sphenoidal and intra-sphenoidal synchon-
droseswere still unossified,so that if these animals had lived, their crania would undoubtedly
have increased in length and in other dimensions. It will be noticed from the measure-
ments given in Table I. that though the condylo-premaxillary length in Professor Scott's
specimen was more than the same dimension in the Challenger skull from Heard Island,
yet that the length to the extreme projection of the occipital crest was greater in the
latter cranium. The difference in absolute dimensions between the male and female
crania is clearly therefore to be regarded as a sexual differentiation.
Important sexual characters were seen also in the teeth in the two sexes. In both
2 — 2 1—1
the males and females the formula was — incisors 1_,, canines j~r> post-canines either
5_5 4_4
KZIK or JUi' ^though in one female there were six post-canines on one side of each
jaw. But the relative size of the teeth in the two sexes varied very materially, especially
in the canine and incisor teeth. In Table II. I have given the comparative dimensions
of some of the teeth in one of the Heard Island males and in the largest female from
Betsy Cove, Kerguelen ; all these measurements were taken in a straight line.
Table II. — Relative Size of Teeth.
3
?
Heard
Kerguelen
Island.
Island. /.
mm.
mm.
Length of upper canine,
153
,, its enamelled crown,
24
ie
Greatest transverse diameter at alveolar border,
36
12
Length of upper inner incisor, .
44
„ its enamelled crown,
5
7
Length of upper outer incisor, .
75
„ its enamelled crown,
7
8
Length of first post-canine,
39
„ its enamelled crown,
5
6
Length of last postcanine,
22
,, its enamelled crown,
4
4
Length of lower canine,
140
Greatest transverse diameter at alveolar border,
39
9
The incisor and canine teeth had conical crowns and elongated single fangs, which in
the canine teeth were fluted. The crowns of the post-canines were somewhat laterally
compressed, and many of them were marked by shallow, vertical grooves, which indicated
a division into two or even three imperfect lobes or cusps ; the fangs were all simple. In
the older crania the greatest circumference of the teeth was after they had emerged from
8 THE VOYAGE OF H.M.S. CHALLENGE!!.
the alveoli, and where in the living animal they would have been embraced by the gum.
Also in these older crania the entrance into the pulp cavity was obliterated, excepting in
the canines, but in the younger skulls the communication with the pulp cavity at the tip
of the fang was freely open in all the teeth. In both of the large males the canines were
not only worn down somewhat at the apex, but the lateral aspects, where the upper and
lower canines had rubbed against each other, were much flattened. In the Kerguelen
Island male (e) the teeth showed very slight signs of wear, so that this animal was far
from being adult,
When the skulls of the females were placed side by side with that of either of the
large males, other differences than that of relative size were observed. In the female
skull the summit from the occipital crest to the fronto-nasal suture was almost flat,
but sloped downwards and forwards on the nasal bones ; the occipital crests were only
faintly indicated ; the skull possessed great width in the occipital, parietal, and temporal
regions, and then suddenly narrowed in the frontal and interorbital regions. The
temporo-zygomatic fossa was capacious and continuous with the orbit, and the zygomatic
arch was massive and bulged outwards. In both the large males the temporo-zygomatic
fossse and arches were like those in the female, but on a larger scale. The summit of the
skull was not flat, but concavo-convex from behind forwards, the posterior concavity being
due to the elevation of the occipital crests and posterior border of the parietal bones.
The frontal bones were also somewhat depressed below the plane of the two parietals,
between the anterior borders of which they were received, but further forwards the
frontals were raised into a slight convexity in the interorbital region, and at their
anterior ends subsided into a hollow corresponding to the fronto-nasal suture. In the
Kerguelen Island young male (e) the occipital crests were much lower than in the other
males, and the summit of the cranium was less removed from the flattened form of the
skull seen in the female, and this flattening of the vertex was a character in all the young
skulls, both male and female. The frontal region in all the crania was constricted as
compared with the great breadth of the occipital, parietal, and temporal regions ; but in
the males the frontal width in front was proportionally more than in the female, owing
to the greater width of the anterior nares in the former sex. In the younger crania the
interfrontal width was not so constricted posteriorly as in the adult skulls. The
greatest width of the skull at the zygomata was at a point about midway between the
two ends of the arch.
The relative size of the orbits and temporo-zygomatic fossa was studied by com-
paring the diameter, measured from the anterior surface of the cranial box at or near
the fronto-parietal suture to the tip of the antorbital process, with the orbital diameter
from the tip of that process to the apex of the ascending or orbital process of the malar.
In the two older males the orbital diameter, as measured between the above two points,
was about two-thirds that of the entire distance ; in the younger male (e) the orbital
REPORT ON THE SEALS. 9
diameter was about four-fifths that of the entire distance, whilst in the female (/) and in
the youngest skulls, both male and female, the orbital diameter was almost equal to the
distance from the cranial box to the antorbital process, so that the orbital process of the
malar bone, was almost in the same transverse plane as the anterior wall of the cranial
box, instead of being considerably in front of it as in the older male crania. The
temporo- zygomatic fossa had therefore a greater relative antero-posterior diameter in
the adult males than in the female and in the younger skulls of both sexes, and this was
correlated with a greater elongation of the constricted part of the frontal region. The
zygomatic process of the temporal was bent abruptly upwards behind the orbital
process of the malar, as far as, or almost as far as its tip, and the two bones formed a
lofty process in this region of the face. The antero-inferior angle of the parietal bone
articulated with the alisphenoid.
The nasal bones in both sexes were relatively short, triangular, and with the apices
received between the anterior diverging borders of the frontals ; the base was forwards
and with a notch marking the interval between the two bones. The anterior edge of
the mes-ethmoid was vertical, and grooved for the reception of the nasal cartilage ; in the
males it came forward as far as the anterior border of the nasals above, but in the females
not quite so far forwards ; whilst below it was lodged in the bottom of the spoutdike
vomer, the anterior end of which projected horizontally for some distance beyond the
mes-ethmoid and the anterior border of the nasals. The premaxillary bones consisted
only of a horizontal part, which was prolonged far in front of the anterior nares, so that
in the males this bone had the extreme length of about 140 mm., and in the largest
female 69 mm. In the larger skulls each bone possessed a premaxillary tubercle above
the incisor teeth. The anterior end of the beak was broadly truncated in the males,
and the superior maxillse with their canines were almost in the same transverse plane
as the incisor teeth. In both sexes the upper surface of each premaxilla was almost
horizontal and fitted on to the inner surface and anterior end of the superior maxilla ;
it bifurcated posteriorly, the inner fork articulating with the outer side of the anterior
end of the spoutdike vomer, whdst the outer broader fork rested on the horizontal
portion of the superior maxilla. As the premaxilla did not possess an ascending jjart it
did not enter into the formation of the lateral boundary of the anterior nares.
The anterior nares were wide, and owing to the vertical direction of the mes-ethmoid
and their steep and almost vertical lateral boundaries, were in the vertical transverse
plane of the face almost on a level with the front of the zygomatic arch. They were
bounded above by the nasals, laterally by the nasal process of each superior maxilla,
and below by the vomer, superior maxdlas and premaxilla^, whilst the interval between
the mes-ethmoid and outer wall of each chamber was filled up by the highly subdivided
maxfilo-turbinal, which came forwards so as to be in the plane of the opening. In
Table I. the height and width of the anterior nares in the two sexes are given, from
(zool. cball. exp. — part Lxvui. — 1887.) Yyy 2
10 THE VOYAGE OF H.M.S. CHALLENGER.
which it will be seen that in the male the width is proportionally greater than the
height. In the male also the upper jaw is prolonged in front of the anterior nares,
both absolutely and relatively more than in the female, and the superior maxillae in the
male extend laterally beyond the prernaxillae, much more than in the female, which is
due partly to the much greater magnitude of the incisor and canine teeth in the male
sex, and partly to this region of the skull being associated with the development of a
proboscis in the male and not in the female. There is therefore a marked difference in
the two sexes between both the length and breadth of the pre-nasal part of the skull, and
between the adult and younger male crania in the same region.
As the nasal cartilages had been preserved in the Kerguelen Island male skull I
examined their arrangement and connections. Attached to the anterior border of the
two nasal bones was a triangular cartilaginous plate, 80 mm. long, the apex of which was
directed forwards. It prolonged the roof of the nose forwards in the plane of the nasal
bones, and had at one time evidently been divided into two lateral halves, as traces of a
median suture could be seen on its upper surface. By its under surface it was fused with
the septal cartilage, which was prolonged forwards in the mesial plane from the anterior
border of the mes-ethmoid for 12 mm. in front of the premaxillaries. Where it rested
in the vomer and on the prernaxillae it was broadened out into a base varying in width
from 30 to 40 mm. Attached to each lateral border of the roof cartilage of the nose was
a lateral cartilage, which passed outwards as far as the superior maxilla, where it formed
the side wall of the anterior nares. The two formed a pair of alar cartilages, and were
near their maxillary attachment fibrous in their structure.
The antorbital (maxillary) process was a well-marked triangle in both sexes, and
was situated immediately behind the anterior nares, whilst the infraorbital foramen was
somewhat in front of the nasal opening in the skull. The postorbitals were wanting,
but in one skull a stroug fibrous band stretched from the orbital process of the zygomatic
arch to the side of the frontal bone, and completed the ring of the orbit posteriorly.
The ascending processes of the superior maxillae, like the nasals, were received between
the two diverging frontals.
The hard palate was widest immediately behind the last molar tooth ; it was concave
anteriorly and mesially, though without much depth, and its outer edge behind the
dentary arcade was scarcely raised above the general plane of the surface. In one male
skull this edge extended 132 mm. from the socket of the 5th post-canine to the palato-
pterygoid suture, and in the largest female 69 mm. The palatal surfaces of the
prernaxillae were triangular, and the apex of each was received between the superior
maxillae ; an almost obliterated naso-palatine foramen was situated mesially between the
prernaxillae. The palato-maxillary suture was almost transverse, and placed some
distance behind the last molar tooth, though immediately behind the root of the malar
process of the superior maxilla ; behind it the palate diminished considerably in breadth,
REPORT ON THE SEALS. 11
and at its posterior border was doubly festooned, with a slight posterior process in the
region of the mesial palatal suture. The vertical plate of the palate bone extended
behind the posterior edge of the hard palate, and overlapped the outer surface of the
pterygoid. The hamular process was distinct and curved backwards and outwards. In
the older males the posterior border of the hard palate was in the same transverse plane
as the lower border of the articulation between the malar and the squamoso-zygomatic,
and a little in front therefore of the glenoid fossa ; but in the female and younger skulls
it was in a transverse plane, a little in front of this articulation.
The posterior edge of the nasal septum did not in either sex extend so far back as the
posterior nares, and consisted of the posterior border of the vomer, which sloped down-
wards and forwards, and of an ascending vomerine crest from the palate bone, articulating
with the vomer in front of the truncated border.
The alisphenoid canal was absent. The tympanic bulla was smooth and only
slightly elevated ; its general form was triangular, and prolonged into the greatly
elongated wall of the external meatus ; it was perforated at the postero-internal angle
of the base by the canal for the internal carotid artery, which looked almost directly
backwards and was quite distinct from the foramen lacerum posterius. When opened
into, the tympanic cavity was seen to consist of a chamber as big as a walnut, with which
both the external meatus and Eustachian tube communicated. At the posterior part of
the roof and immediately above the orifice of the meatus was a subordinate chamber of the
tympanum about the size of a hazel nut, and situated immediately to the outer side of the
petrous element ; it opened by a narrow fissure into the cranial cavity. As the tympanic
ossicles have already been so fully described by Mr. Doran1 and by Professor Flower,2 and
figured also by the former anatomist, it is unnecessary to redescribe them, as they correspond
so closely with the accounts which they have given. I need only state that the stapes
showed no trace of a division into crura. The optic foramina opened separately into the
cranial cavity, and between them was a mesial plate of bone continuous with a prominent
crista galli. The tentorium was partially ossified, although not so extensively as in some
seals. In the young skull, the cap of which had been sawn off for the removal of the
brain, the transverse diameter of the cranial cavity (148 mm.) was markedly greater
than the antero-posterior (127 mm.).
The occipital condyles closely approximated in front, and in the males were separated
by a narrow groove. In the females the condyles were more widely divergent than in
the males. In one female where the process of maceration was carefully watched, a
broad plate of unossified cartilage, continuous with the basis cranii, extended backwards
along the inner border of each condyle for 29 mm. from the basi-occipital, so that the
foramen magnum was greatly diminished in size, as compared with a fully macerated
specimen ; in two of the males the corresponding plate of cartilage had undergone a
1 Trans. Linn. Soc. Lond. (Zool.), ser. 2, vol. i., 1876. 2 Proc. Zool. Soc. Lond., January 4, 1881.
12 THE VOYAGE OF H.M.3. CHALLENGER.
partial ossification. A pair of foramina opened on the outer surface of the occipital bone
immediately behind the foramen magnum, and the canals continuous with them running
vertically upwards in the substance of the supra-occipital, opened again on the surface
below the occipital crest, the distance varying in different specimens ; the canals and
foramina probably transmitted veins, and may appropriately be named supra-occipital.
The basi-occipital was not perforated, and in the older skulls was marked by a transverse
ridge. A slight paroccipital process was present in the older males, but in the young male
and the females it was just visible. The mastoid was scarcely differentiated as a process.
The lower jaw was much more massive in the males than females, due in part to the
magnitude of the canine teeth and the size of the areas of attachment of the muscles
of mastication. In none of the specimens had fusion at the symphysis taken place. The
lower border of the body of the bone was slightly everted and terminated abruptly
behind in one of the males, but not in the other or in the females. At the posterior
border of the ascending ramus a subcondyloid i^rocess projected backwards a little
below the neck of the bone. The condyle was transversely elongated, the coronoid
process was low, and the sigmoid notch was shallow.
Spine.- — The description of the bones of the neck, trunk, and limbs has been based
upon the study of the skeleton of the well-grown male (e) from Betsy Cove, Kerguelen,
though the skeletons of the younger specimens have been also examined. In no specimen
were the plates ankylosed to the bodies, and the cartilaginous tips of the spinous, trans-
verse, and mam miliary processes were unossified.
The vertebral formula was C 7, D 15, L 5, S 3, Cd 10 = 40.
The cervical vertebrae, except the 7th, possessed a foramen at the root of the
transverse process ; in all except the axis this process was a massive bar of bone
projecting downwards and outwards, but not flattened into a plate except in the atlas.
Evidence of the presence of two tubercles at the end of this process was seen in all
except the axis and the 7th. The spinous process was feeble, except in the axis, where
it was massive. The articular processes were autero-posterior in direction, the anterior
pair looked upwards and inwards, the posterior pair downwards and outwards. The
bodies were elongated transversely. The atlas had plate-like transverse processes which
projected outwards and very slightly downwards ; the articular surfaces for the occipital
condyles were deeply concave, and separated from each other by a distinct interval ; the
lamina on each side was perforated by a foramen for the vertebral artery. The articular
surface for the odontoid was continuous with the posterior articular facets for the axis,
and they were covered by a common plate of cartilage. The axis had a well-marked
odontoid process and the bone showed the remains of the intervertebral disc between
this process and the body of the axis. A broad plate of cartilage covered the inferior
surface of the process, which was separated from the cartilage covering the anterior
articular surfaces by a narrow groove on each side. The surface of the odontoid for the
REPORT ON THE SEALS. 13
transverse ligament was covered by a much narrower plate of cartilage. The transverse
process was very short and pointed.
The dorsal vertebras had low spinous processes, those of the 2nd and 3rd being the
most prominent. They projected slightly backwards. The transverse processes were
thick and strong in the anterior and middle regions, but posteriorly they had almost dis-
appeared. Notwithstanding the relative disappearance of the transverse processes in the
last five dorsal vertebras, each possessed a large articular surface for a rib in the region
where the transverse should have been, so that throughout the series the vertebrae
possessed both an articular surface or surfaces on the side of the body for the head of a
rib and one for the tubercle. The last five dorsal vertebras had only a single facet on
each side of the body, which was placed at its anterior part. Anapophyses were very
faintly marked in the 11th to the 14th vertebras. Metapophyses were no more than very
slightly indicated in any. Only the more anterior and posterior dorsal vertebras were
keeled on the ventral surface of the body.
Each lumbar vertebra had a transverse process directed downwards, forwards, and
outwards. The spine was strong but low. The mammillary processes were short and
rounded and directed forwards and outwards. The anterior articular processes were
slightly concave and directed upwards and inwards, the posterior convex and directed
downwards and outwards. The bodies were elongated antero-posteriorly and faintly
keeled on the ventral surface.
The sacral vertebras were apparently three in number. The 1st was massive, 7 "2 cm.
in antero-posterior diameter, and 16 "5 cm. in transverse diameter at the base. Its
lateral articulation for the ilium was ear-shaped below, and rough above for the great
sacro-iliac ligament. This bone diminished rapidly in transverse diameter from the base
to the posterior surface. Its neural arch was complete in the larger animal, but the
laminae had not met in the young female. The 2nd sacral vertebra was 6-6 cm. in
antero-posterior, and 97 in its greatest transverse diameter. Its neural arch was com-
plete in both pelves. At first I thought that it had a slight articulation laterally with
the ilium, but a fresh examination leads me to say that it did not quite reach it. In
addition to the articulation between the bodies it articulated in front with the 1st sacral
by a pair of truncated processes springing from the pedicles, and situated ventrally to the
proper anterior articular processes, and behind it articulated with the 3rd sacral by a
corresponding pair of processes. The inferior and superior sacral foramina were situated
ventrally and dorsally to these processes. The 3rd sacral vertebra was smaller than the
2nd, and had in both pelves a complete neural arch. The epiphyses between the bodies
of the 1st and 2nd and the 2nd and 3rd sacral vertebras had fused with each other, but
had not ankylosed to the bodies of the vertebras to which they belonged.
I have referred ten vertebras to the caudal region. The first caudal had a neural
arch, the next one had a neural groove, the laminas not being united ; the rest consisted
14 THE VOYAGE OF H.M.S. CHALLENGER.
only of elongated bodies, diminishing in size to the end of the tail, the last being only
1*4 cm. in length.
Hibs. — There were fifteen pairs of ribs, nine of which articulated with the sides of
the sternum. The head of the 1st rib articulated with the side of the body of only the
1st dorsal vertebra, but the heads of the ribs from the 2nd to the 11th, both inclusive,
articulated with the sides of the bodies of two vertebrae. The heads of the four most
posterior ribs articulated each with the body of only a single vertebra. The 1st to the
11th ribs, both inclusive, possessed each a well-defined neck, with a distinct interval
between the head and the tubercle ; but in the last four ribs the neck was stunted and
the head and tubercle were more closely approximated, so that in the last rib they were
separated by an interval of only 8 mm. In a straight line the osseous part of the 1st
rib measured 145 mm., and the bony shaft gradually increased in size to the 8th rib,
which was 390 mm. in length, from which again the ribs diminished to the 15th, which
was 232 mm. long. The ribs as a rule were curved ; their shafts were thick and with
three surfaces, external, anterior and posterior. The two last ribs were almost straight.
The massive costal cartilages of the sternal ribs were either longer than, or closely
approximated in length to, the osseous division of their respective costal arches. The
cartilages of the asternal ribs were attenuated at their inner ends. In this animal the
ligamentum conjugale costarum originally described by Professors Mayer1 and Cleland'2
was seen to great advantage. As in the seal which Professor Cleland dissected, it con-
sisted of a strong ligamentous band attached on each side to a depression situated imme-
diately below the cartilage covering the undivided articular surface of the head of each
of the ribs which articulated with the bodies of two vertebras. It entered the spinal canal
in the plane of the intervertebral disc, immediately above which it was situated, and its
inferior surface as well as the superior surface of the disc was smooth and polished, and
was covered by a synovial membrane, continuous with that of the costo-vertebral joint
on each side. There can be no doubt therefore that this ligament plays upon the upper
surface of the disc in the respiratory movements of the ribs. It was kept in its place
by the superior common ligament which covered it.
Sternum. — This bone, formed of nine segments, was 960 mm. long. The 1st or
praesternal segment was 85 mm. in length, and consisted of a prsesternal cartilage,
broader posteriorly than anteriorly, where it terminated in a pointed apex, so that it had
somewhat of a triangular form. It extended forwards to the neck for 63 mm. in front of
the 1st pair of costal cartilages. The 2nd to the 8th segments were plates of bone, the
first, second and third of which were longer than broad, whilst the length and breadth of
the remainder were almost equal. These segments articulated with each other by movable
1 Muller'n Archivf. Anat. u. Physiol., vol. i. p. 273, 1834.
■ Edinburgh New Philosophical Journal, vol. viii., April 1859. See also J. B. Sutton, Journ. of Anat. and Phys.,
vol. xviii. p. 225, 1884, and Ligaments, their Nature and Morphology, London, 1887.
REPORT ON THE SEALS. 15
joints, and the margin of the segment next the joint was formed of unossificd cartilage.
The 9th or xiphisternal segment was situated behind the last pair of sternal ribs ; it was
290 mm. long, and whilst its anterior third was ossified, the remainder consisted of
cartilage, which widened at its free end into a leaf-like expansion. The first seven pairs
of costal cartilages articulated with the side of the sternum at the junction of its segments
with each other, the 8th pair was jointed to the side of the 8th segment and the 9th pair
at the junction of the 8th and xiphisternal segments. The 8th and 9th costal cartilages
also articulated with each other close to the sternum. The maximum sterno-vertebral
diameter of the cavity of the thorax was 430 mm. and the greatest transverse diameter
of the cavity was 370 mm.
Anterior Extremity. — The scapula did not have so well-marked a falciform shape as
is usual in the seals. The dorsum was divided into two fossae ; the postspinous was
deeply grooved immediately below and parallel to the spine, and its vertical diameter
was about three times more than its greatest antero-postcrior diameter. The pixespinous
fossa was almost triangular in shape, and its vertical diameter was about twice as great
as the antero-posterior. The area of the prsespmous was somewhat greater than that of
the postspinous fossa. The spine had no great prominence, and was without an acromion.
The coracoid was stunted, and in the younger skeletons was not fused with the body of
the scapula. The scapula was prolonged by a triangular suprascapular cartilage, and its
extreme breadth or vertical diameter, including this cartilage, was 34' 5 cm., whilst the
length or antero-posterior diameter was 19 "5 cm. The subscapular and prsespinous fossae
were smooth and only slightly concave.
The humerus was 26 cm. long, and had a strong deltoid ridge with a bicipital groove
internal to it. The upper and lower epijmyses were not fused with the shaft ; that of
the internal condyle was quite distinct from the radio-ulnar articular epiphysis, and that
for the head was separate from that for the inner tuberosity. There was no supra-
condyloid foramen and the shaft of the bone was not much twisted.
The radius, 24 cm. long, was rounded above and had the usual cup-shaped head, and
was flattened in its lower half as is usual in seals. The anterior border of the shaft was
strongly ridged for the tendon of insertion of the pronator teres. The ulna, 28 cm. long,
was expanded above at the olecranon and attenuated below. In both bones of the fore-
arm the epiphyses were not united to the shafts. The radius was anterior to the ulna,
and its cup was a more important factor in the elbow joint than the articular surface of
the ulna. The radius articulated at its lower end with the ulna, scapholunar, and cunei-
form ; the ulna articulated with the radius, cuneiform, and pisiform.
Manus. — The carpus possessed seven bones. The 2^isiform was a mere nodule and
articulated with both ulna and cuneiform. The cuneiform articulated with the ulna,
radius, scapholunar, unciform, and 5th metacarpal. The scapholunar was large and
articulated with radius; cuneiform, trapezium, trapezoid, os magnum, and unciform. The
16 THE VOYAGE OF H.M.S. CHALLENGER.
trapezium articulated with 1st and 2nd metacarpals, trapezoid, and scapholunar. The
trapezoid articulated with the trapezium, 2nd metacarpal, scapholunar, and os magnum.
The os magnum, was one of the smallest bones of the carpus, and articulated with the 2nd,
3rd, and 4th metacarpals, the trapezoid, scapholunar, and cuneiform. The unciform was
shut out from the inner border of the wrist by the approximation and articulation of the
5th metacarpal with the cuneiform ; it articulated with the 4th and 5th metacarpals, the os
magnum, scapholunar, and cuneiform. The carpal bones were roughened on their palmar
and dorsal surfaces for the attachment of ligaments, but there was an absence of the
ridges and processes which characterise the corresponding bones in the human carpus.
The digits were five in number, and both the entire digit and its metacarpal segment
diminished in length from the pollex to the minimus. The three segments of the pollex
were longer than the corresponding segments in any of the fingers. The so-called
metacarpal of the thumb and the phalanges generally possessed three centres of ossifica-
tion, one for the shaft and one each for a proximal and a distal epiphysis ; the ungual
phalanx, however, had only a proximal epiphysis. The metacarpals of the four fingers
had only a distal epiphysis, and if a proximal epiphysis had ever been present, it had
become fused with and indistinguishable from the shaft of the bone. The 1st metacarpal
articulated with the trapezium ; the 2nd with the trapezium, trapezoid, os magnum, and
3rd metacarpal ; the 3rd with the os magnum and 2nd and 4th metacarpals ; the 4th
with the os magnum, cuneiform, and 3rd and 5th metacarpals ; the 5th with the
cuneiform, unciform, and 4th metacarpal.
Pelvis. — The pelvis consisted of the sacrum and two innominate bones. The sacrum
has been described above. Each os innominatum articulated by the inner or sacro-pelvic
surface of the ilium with the area on the 1st sacral vertebra, which was partly auricular
and cartilaginous, and partly rough for the great sacro-iliac ligament. In the larger
animal, a male, the length of the bone was 320 mm., in a smaller specimen, a female (c),
215 mm. The acetabulum, though relatively deep, had only a feeble brim, and the non-
cartilaginous covered surface at the bottom was narrow and grooved. The ilium was
short, 98 mm., and its crest was 135 mm. long. Its dorsal surface was more than
twice as broad as the ventral surface. From the sacro-iliac joint the bone inclined almost
transversely outwards to the iliac crest, which was only a little anterior to the transverse
plane of the base of the sacrum. The os pubis, ischium, and obturator foramen were all
elongated, as is characteristic of the seals, and the diameter from the pectineal tubercle
to the pubic symphysis was 240 mm. The junction of the os pubis and ilium was
marked by a large pectineal tubercle for the insertion of the psoas parvus. In the larger
of the two pelves measured the ischium and os pubis were not fused with each
other at the pubic symphysis, but in the smaller female specimen the fusion was com-
plete. The ischial tuberosity was moderate. The epiphysial cartilages at the symphysial
end of both pubis and ischium, at the iliac crest, the ischial tuberosity, and the pectineal
REPORT ON THE SEALS. 17
tubercle were unossified, but the fusion of the three segments of the bone in the
acetabulum was complete. The interval between the two pubic bones at the symphysis
was of considerable width.
Posterior Extremity. — The femur was 173 mm. long, and was characteristically
flattened, its greatest width at the condyloid end being 100 mm. The head was smooth
and without a depression for the ligamentum teres. The trochanter major was well
marked, but there was neither trochanter minor nor trochanter tertius. The anterior
flattened surface of the shaft was divided by an oblique ridge, which separated the
crureus and vastus externus, and extended from the neck downwards and outwards
towards the outer condyle, and on the back of the shaft there was a faint linea
aspera. The trochlear surface for the patella was shallow and not continuous with the
articular surfaces of the condyles, from which it was separated by an intermediate rough
area, to which was attached a broad, strong, ligamentous band connected with the lower
end of the patella and the deep surface of the ligamentum patella. This band would
separate the patello- femoral joint from the femoro-tibial joints and was doubtless morpho-
logically the same as, though histologically different from, the ligamentum adiposum of
the human knee-joint. The condylar articular surfaces were feebly convex and separated
from each other by a roughened intercondylar fossa.. The epiphyses in one of the larger
femora were separable from the shaft, but in the other fusion had commenced.
The patella was 45 mm. in its long axis and 41 mm. transversely ; its trochlear
articular surface was feebly concave and not facetted. Its cutaneous surface was
roughened. At its upper end it was 26 mm. thick, and at its lower end only 12 mm.
The tibia was 340 mm. long, and the. fibula was 336 mm. They articulated with each
other above and below, and the shafts were separated in the upper three-fourths by an
interosseous interval of some width, but in the lower fourth they were closely approximated
and united by an intermediate ligament. Each bone had a malleolar prolongation at
the lower end, but that of the tibia was very short, and did not articulate with the
inner surface of the astragalus. The tibia had a broad surface superiorly, smooth on
each side for the femoral condyles, but rough between for the attachment of the semi-
lunar cartilages and crucial ligaments. The shaft of the tibia was almost straight and
possessed a ventral surface and ridge for the insertion of the gracilis, semitendinosus, and
semimembranosus tendons. Above this ridge was the surface of attachment of the
ligamentum patellae, fibulad to which the shaft was grooved for the tibialis anticus. The
posterior surface of the tibial shaft was grooved for the origin of the tibialis posticus, the
tendon of which also grooved the back of the lower end of the bone. The fibula was a
much more bulky bone than in the human leg, so as to give broader surfaces for the
origin of muscles ; two peroneal grooves marked the lower end of the shaft and the
external malleolus. The epiphyses at both ends of each leg bone were not fused with
the shafts.
(ZOOL. CHALL. EXP. PART LXVIII. — 1887.) ^JJ 3
18 THE VOYAGE OF H.M.S. CHALLENGER.
Pes. — The tarsus contained seven bones. The astragalus was the largest bone of
the foot, and articulated with the tibia by its superior surface, and with the fibula by its
external lateral surface, and its fibular surface was almost as large as the tibial ; also
with the os calcis by its inferior surface, which possessed two facets separated by a deep
groove for an interosseous ligament, and by its anterior surface or head with the scaphoid
bone ; it also had a posterior process which, though massive, did not projedt quite so
far back as the calcanear process of the os calcis, and was not grooved posteriorly.
The os calcis grooved for the peroneal tendons articulated with the astragalus and
cuboid ; its calcanear process was longer than, but not so massive as, the posterior process
of the astragalus, and possessed at its free end a separate epiphysis. The scaphoid had
the characteristic form of the bone and articulated with the astragalus, cuboid, and three
cuneiforms. Its tubercle for the tibialis posticus was massive, and it had also a pointed
plantar process. The cuboid had a plantar ridge and peroneal groove ; it articulated
with the 4th and 5th metatarsals, and by a very small surface with the 3rd, also with
the calcaneum, scaphoid, and ecto-cuneiform. The three cuneiforms varied much in
size, the ento- was the largest, the ecto- next in size, and the meso- so small as only to
be seen on the dorsum of the foot. The ento- articulated with the 1st and 2nd
metatarsals, the scaphoid, and meso-cuneiform. The meso- with the other cuneiforms,
the scaphoid, and the 2nd metatarsal. The ecto- with the 2nd and 3rd metatarsals,
the meso-euneiform, cuboid, and scaphoid.
There were five toes. The hallux and minimus, notwithstanding the difference in the
number of segments, were of almost equal length, although the hallux had slightly the
advantage. The 2nd and 4th toes, almost of equal length, reached to about the level of
the articulation of the terminal and penultimate phalanges of the hallux. The 3rd toe was
the shortest and ended almost opposite the joint between the 2nd and 3rd phalanges
of the 2nd toe. The segments of the hallux were longer than the corresponding segments
in the other toes. The 2nd metatarsal was in close relation to the outer side of the
1st, and the tarsal end passed behind that of the 1st, so as to articulate with nearly one-
half of its proximal end, the remainder being for the internal cuneiform ; this arrange-
ment gave to the tarsal end of the 2nd metatarsal a hook-like form ; it articulated with
all three cuneiforms and with the 1st and 3rd metatarsals. The 3rd metatarsal was the
shortest and articulated at its proximal end with the ecto-cuneiform, slightly with the
cuboid, and with the 2nd and 4th metatarsals. The 4th metatarsal was in close apposi-
tion with the 5th, and its tarsal end was hollowed on the outer side to allow the 5th
metatarsal to be lodged in it ; it articulated with the 3rd and 5th metatarsals and the
cuboid. The 5th metatarsal, though shorter than that of the hallux, was if anything
more massive ; its tarsal end articulated with the 4th metatarsal and cuboid, and was
somewhat elongated into a process on the outer side of the foot. The ossification of the
metatarsals and phalanges was on the same plan as that of the metacarpals and phalanges
REPORT ON THE SEALS. 19
in the manus. A fold of integument extended for some distance beyond the tip of each
ungual phalanx.
The vertebral column was 2580 mm. in length in the largest of the Challenger skeletons
(male e), measured with the intervertebral discs in place but shrivelled and dried, so that
during life the spine would have been somewhat longer; the extreme length of the skull
was 402 mm., giving 2982 mm. or 9 feet 9 inches from the front of the premaxilla to the
end of the tail. This dimension was very much shorter than that of the male measured
by Professor Flower (p. 5),1 or that which Professor Peters has measured,2 the length of
the spine of which was 3700 mm., and of spine with skull 4200 mm. or 13 feet 9 inches.
A spine with skull in the Museum at Cambridge, Mass., measured by Mr. J. A. Allen,
and said to be an adult male, was 4340 mm. or 14 feet 3 inches long, and in the same
animal the humerus was 335 mm., radius 310, femur 200, tibia 415 mm. in length. If
these measurements are compared with those of the corresponding bones in the young
male that I have described, it will be seen that they are very materially longer, so that
in all probability the Challenger animal had not attained much more than about two-
thirds of the growth of an adult male. Great differences in size exist between the adult
male and female Elephant Seals. Captain Scammon, whose observations were made on
the Californian Sea Elephant,3 states that the male is frequently triple the bulk of the
female — the oldest males average from 14 to 16 feet, whilst the largest he had ever seen
measured was 22 feet. Two females which he measured were 9 and 10 feet respectively.
Corresponding differences in magnitude may be seen in the skulls of the Southern
Elephant Seal measured in Table I.
The length-breadth indices of the skulls measured in Table I., calculated on the
relation of the condylo-premaxillary length to the interzygomatic width, were for the
three large male skulls respectively 72-8, 70'9, and 68, for the well-grown male (e)
69'9, and for the large female (/) 74. Calculated on the relation of the condylo-
premaxillary length to the width behind the external meatus the indices for the two
Heard Island males were 58 and 61 respectively, for the well-grown male (e) 62-9,
and for the large female (/') 66. The greater magnitude of the zygomatic index
expresses the greater breadth of the skull in that region.
1 Notwithstanding the dimensions of this animal the plates were not united to the bodies of the vertebras, nor the
epiphyses of the bones of the fore-arm and fore-leg to their respective shafts.
2 Monatsber. d. k. preuss. Akad. d. JViss. Berlin, 1875, p. 393.
3 The Marine Mammals of the North-West Coast of North America. San Francisco, 1874.
20 THE VOYAGE OF H.M.S. CHALLENGER.
Leptonychotes,1 Gill.
Lepionychotes, Gill, Arrangement fam. Mam., 1872.
On the 9th January 1874, a seal, regarded as a Sea Leopard, and believed to be a
female, was shot at Betsy Cove, Kerguelen. The skeleton was preserved and sent home.
It is referred to in the Narrative of the Voyage of the Challenger (vol. i. p. 355), and on
p. 373 it is stated that the sandy beach of Heard Island was strewn with bones of both
the Elephant Seal and the Sea Leopard, those of the former being the more abundant.
Leptonychotes weddelli (Lesson) (Plate V.).
Otaria Wedddlii, Lesson, Ferussac's Bull. d. Sci. Nat., vol. vii., 1826, pp. 437, 438.
Leptonyx weddelli, Gray, Ann. and Mag. Nat. Hist., vol. x., 1836.
False Leopard Seal, or Weddell's Seal.
The comparison of the skull of the animal shot at Betsy Cove with the drawings and
descriptions of the crania of the seals figured by Dr. Gray in the Zoology of the Voyage
of the "Erebus" and "Terror,"2 has satisfactorily shown that the specimen was not a true
Sea Leopard, such as is included under the generic names Stenorhynchus (Ogmorhinus)
and Lobodon, but that it was like the specimen of the seal from Santa Cruz, which was
named by Dr. Gray after Captain Weddell. Short notes on the characters of the
skull of Weddell's Seal have been given by Messrs. Allen and St. George Mivart in their
monographs already referred to, but the skeleton generally has not yet been described.
Skeleton. — The animal was not an adult, for the vertebral plates were not united to
their centra, and the epiphyses of the bones of the shafts of the limbs were not ankylosed.
Skull. — In drawing up the following description I have examined and compared the
skull of Weddell's Seal from Betsy Cove with two well-grown crania of Stenorhynchus
(Ogmorhinus) leptonyx, one of which was from Wellington Harbour, New Zealand,3 but
the locality from which the other and somewhat older specimen came is unknown.
In Table III. I have given the comparative measurements of these crania.
In none of the skulls was either of the basi-cranial synchondroses ossified, though the
interval between the bones was scarcely more than would admit the edge of a knife.
The dental formula of Stenorhynchus leptonyx was —
. 2-2 1-1 5-5
in-2^2 c-r=n p-c-5^5'
and of Weddell's Seal,
2-2 1-1 6-5
in.
2-2 "1-1 P-C'5-5
1 As the generic name Leptonyx, given to Weddell's Seal by Gray, has also been applied to one of the Mustelida?,
to one if not two genera of Birds, and to a genus of Gastropodous Molluscs, I have preferred to adopt the generic name
Leptonychotes employed by Gill and Allen.
2 Vol. i., Mammalia, London, 1814-1875.
3 This skull was presented to the Anatomical Museum of the University of Edinburgh by Sir James Hector, F.R.S.,
REPORT ON THE SEALS.
21
In Weddell's Seal the incisors were much smaller than in Stenorhynchus, but in Loth
species they were rather recurved, and the laterals both above and below were larger
than the centrals. In Weddell's Seal, however, as compared with Stenorhynchus, the
upper lateral incisors were proportionally bigger than the central, whilst the lower lateral
incisors were proportionally smaller than the central. The canines were also similarly
formed in both species, but considerably larger in Stenorhynchus. The post-canines,
however, showed important differences in the two species. In Stenorhynchus they had the
characteristic three large cusps so frequently described ; but in Weddell's Seal these teeth
were very much smaller and with a single prominent cusp, which represented the central
cusp of Stenorhynchus, though in the 3rd and 4th molars in both jaws a rudiment
of a posterior cusp was just visible, and a sharp-edged ridge or cingulum ran around the
inner side of the base. Except the first the post-canines were two-fanged. The difference
in size may be gathered from a comparison of the length of the second upper post-canine,
Table III. — Crania of Leptonychotes and Stenorhynchus.
Wellington
Challenger.
Harbour.
Weddell's
Stenorhynchus
Stenorhynchus
Seal.
leptonyx.
leptonyx.
mm.
mm.
mm.
Extreme condylo-premaxillary length,
237
323
321
,, interzygomatic width, ....
142
• 161
176
„ width behind external meatus,
157
165
172
Greatest width of palate, .....
57
68
68
Width between outer side of base of upper canines,
42
57
55
„ ,, of lower canines,
26
46
42
Length of palate in line of mesial suture to central incisor, .
87
114
Height of skull from basion to middle of occipital crest,
81
91
92
Smallest interfrontal width in plane of upper surface,
22
31
40
Length of nasals, ......
55
77
89
Greatest width of anterior nares, ....
29
31
33
Length of mandible, .....
147
232
241
Width between outer ends of condyles of mandible,
141
161
179
from the alveolar border to the tip of the cusp in both animals. In Stenorhynchus
it was 15 mm., in Weddell's Seal only 6 mm. In the latter specimen the first and last
post-canines both above and below, and on the left side above (where there were six
post-canines) the penultimate tooth also, were considerably smaller than the three inter-
mediate teeth, which were about ecpial in size, but in Stenorhynchus there was but little
difference in the relative magnitude of the five post-canine teeth both above and below.
In all the crania the extreme length was in the condylo-premaxillary diameter, for
the occipital crests, though present, were small. In one Stenorhynchus leptonyx measured
in the table the interzygomatic width was less than the greatest width of the skull,
but in the other the interzygomatic width slightly preponderated ; in this animal the
22 THE VOYAGE OF H.M.S. CHALLENGER.
widest part of the zygomatic arch was at its hinder end, and the arch diminished in
width when traced from behind forwards ; in Weddell's Seal the width of the arch was a
little greater at its mid-point than posteriorly. In Stenorhynchus the distance from the
antorbital process to the most anterior surface of the cranial box as compared with the
distance from the antorbital process to the orbital process of the molar was as 9 to 7 ; in
Weddell's Seal the former diameter very slightly exceeded the latter. In both specimens
of Stenovhynchus the total length of the skull both absolutely and relatively was greater
than the breadth as compared with the same dimensions in Weddell's Seal. The skull
was capacious in the parietal region, and comparatively flattened in all the crania, and
became greatly constricted in the frontal region ; this constriction was relatively longer
in Stenorhynchus than in Weddell's Seal. In Leptonychotes the antero-inferior angle
of the parietal articulated with the alisphenoid ; in one Stenorhynchus leptonyx they were
separated by an epipteric bone ; in the other they directly articulated.
In all the crania the nasal bones were elongated and ankylosed together posteriorly
and mesially. More than one-half of the length was received between the two divisions
of the frontal, where they formed a triangular area, with the apex backwards, whilst the
anterior part, lodged between the two superior maxillre, was quadrilateral in form. The
anterior edge of the mes-ethmoid was situated far back in the nasal chamber, and the
spout-like vomer, which contained the septal cartilage, sloped downwards and forwards
to the anterior nares. The ascending part of each premaxilla entered into the lateral
boundary of the anterior nares, but in Stenorhynchus leptonyx it did not quite reach the
nasal bone, whilst in Weddell's Seal it partially articulated with the anterior end of the
outer edge of the nasal. The lateral boundaries of the anterior nares sloped obliquely
downwards and forwards, so as to bring the floor of the opening close to the anterior end
of the rostrum. The interval between the vomer and the side wall of the nose was
occupied by a much subdivided maxillo-turbinal. In Stenorhynchus the antorbital process,
though small, was distinctly marked, but in Weddell's Seal it was only a faint tubercle ;
in the former there was an indication of a postorbital process, which was not visible
in Weddell's Seal. The antorbital process and infraorbital foramen in all these skulls
were in almost the same transverse plane, and considerably behind the opening of the
anterior nares. The ascending process of the superior maxilla was not received between
the diverging frontals.
In all the specimens the hard palate was widest opposite the last molar, and its
concavity was very slight. In Stenorhynchus it extended for 40 mm. from the last molar
to the palato-pterygoid suture, and in Weddell's Seal for 36 mm., and its border was not
raised above the general plane of the palate. The palatal surface of each premaxilla
was triangular and the naso-palatine canal was large enough to admit a stilet. The
palato-maxillary suture was transverse near the middle line and opposite the last molars,
but then sloped backwards and outwards and terminated immediately behind the malar
REPORT ON THE SEALS. 23
process of the superior maxilla. The posterior border of the hard palate was deeply
emarginate, and the posterior border of the vomer was visible between the two diverging
bones, though not to so great an extent in Weddell's Seal as in the other species. The
angle of junction of the two palate bones in the mesial line of the hard palate was in the
younger Stenorhynchus about opposite the last molar tooth, and in the older specimen
further back and almost on a line with the posterior border of the zygomatic process of
the superior maxilla. In Weddell's Seal again this angle was in a plane 1 1 mm. posterior
to the same process. In both specimens of Stenorhynchus two small triquetral bones
were situated at the antero-internal angle of the palato-maxillary suture, and in Dr.
Gray's figure of the skull procured during the voyage of the "Erebus" and "Terror"
a mesial triquetral bone is shown in the same region.
The alisphenoid canal was absent. The tympanic bulla was almost hemispherical and
smooth in Weddell's Seal, and its antero-internal angle was truncated ; in Stenorhynchus
a keel-like ridge, not very elevated, was prolonged from the postero-external to the antero-
internal angle, the latter of which was pointed. The carotid canal was separated from the
foramen lacerum posterius in all three specimens by a distinct plate of bone as in the
Elephant Seal. A deep fissure also separated the tympanic bulla from the mastoid part
of the bone, and in it the stylo-mastoid foramen opened. The two optic foramina had a
common opening into the cranial cavity in both Leptonychotcs and Stenorhynchus.
The hamular process was barely visible in Stenorhynchus leptonyx, but in Weddell's Seal
it was present and directed outwards.
The occipital condyles converged and met anteriorly in one skull of Stenorhynchus but
did not quite meet in the other, and in Weddell's Seal the cartilaginous covered surfaces
of the two were continuous. In Weddell's Seal the basi-occipital was thin and perforated
by a rounded hole, but in the other crania it was entire. A low par-occipital process was
present in Stenorhynchus, but was scarcely visible in Weddell's Seal. In both specimens
of Stenorhynchus the supra-occipital canals opened immediately within the posterior edge
of the foramen magnum ; in Weddell's Seal a single foramen only was present on the outer
surface of the bone close to the foramen magnum.
The lower jaw in Weddell's Seal was proportionally more slender than in Stenorhynchus,
-which was in part due to the smaller size of the teeth, requiring a shallower alveolar
border, and in part to the more limited surfaces for the attachment of the masticatory
muscles. The body of the bone was straight and smooth, and with no eversion of the
lower border. The mandible had scarcely any ascent behind to the condyle, and had no
angle ; the subcondyloid process was absent in Stenorhynchus leptonyx, but in Weddell's
Seal it was a faint incurved tubercle. In Weddell's Seal the mandible was much more
slender than the lower jaw of Ommatophoca rossi, or Ross's large-eyed seal, figured in
pi. viii. of the Voyage of the "Erebus" and "Terror."
Spine. — Vertebral formula, C 7, D 15, L 5, S 2, Cd 11 = 40. As the animal
24 THE VOYAGE OF H.M.S. CHALLENGER.
was immature the vertebrae had not assumed their adult characters, and various of
the processes were probably less strongly marked than would have been the case in
a mature animal. As previously stated, the epiphysial plates were not ankylosed to the
bodies.
In all the cervical vertebras, except the 7th, a vertebrarterial foramen was present
between the two roots of each transverse process, and in the atlas the neural arch was
also perforated on each side. The transverse process of the atlas was a broad plate pro-
jecting almost transversely outwards ; that of the axis was short and pointed ; those
of the 3rd, 4th, 5th, and 6th were more massive, and ended in two tubercles, that of
tbe 7th was a single bar of bone springing from the neural arch. In the configura-
tion of its transverse processes Weddell's Seal approximated closely to the Elephant
Seal and differed materially from the corresponding processes in Arctocephalus, in
which animal they were flattened into broad plates which projected almost vertically
downwards, though in the case of the atlas they were elongated downwards and out-
wards. The axis was the only cervical vertebra with a prominent spine ; its odontoid
process was 19 mm. high, and fused with the body of the axis. The ventral surface of
the bodies of the cervical vertebrae had a mesial keel.
The dorsal vertebras articulated with fifteen pairs of ribs ; the 1st with one and
a half, the 11th, 12th, 13th, 14th and 15th with only a single rib on each side, the
others with the halves of the heads of two ribs. When only a single rib articulated
with the side of the body, it was near its anterior part. The transverse processes were
prominent from the 1st to the 10th dorsal, behind which they diminished in size, and
were scarcely to be recognised in the 14th and 15th dorsal vertebrae. The spines were
feeble.
In the lumbar vertebras the transverse processes were elongated, and projected
forwards, outwards, and downwards. The spines were not very prominent. The body
was keeled on its ventral surface.
The sacrum was represented apparently by only two vertebras, though it is possible
that the more anterior of the two caudal vertebras which possessed a neural arch,
might in a mature animal be ankylosed with the sacrum. Of the two vertebras
which I have regarded as sacral, the first was much the larger, its breadth at the
base was 100 mm., and its antero -posterior diameter was 40 mm. It had abroad lateral
articidation with the ilium, 47 mm. in its longer diameter, whilst the corresponding
articulation of the second sacral was only 17 mm. in its longer diameter.
Each of the caudal vertebras, except the two most anterior, consisted of an elongated
body, without a neural arch, and they diminished in length from before backwards, the
terminal vertebra being only 12 mm. long.
Ribs. — Of the fifteen pairs of ribs, ten articulated with the sides of the sternum.
The capitular epiphysis was not in any bone ankylosed to the rest of the rib. The
REPORT ON THE SEALS. 25
cartilaginous division of the rib was long in relation to the osseous part, and in the 1st
rib it was as 87 mm. to 52 mm. The osseous parts of the ribs increased in length
from the 1st to the 8th, then they were almost equal in length to the 13th, whdst the
14th and 15th again were shorter. The last two ribs had no tubercles, and in the 13th
the tubercle was rudimentary.
Sternum. — This bone was narrow and elongated, 468 mm. long ; it consisted of
ten segments ; all the bony segments were quadrilateral in shape, except the 8th, which
was a flattened disc. The first bony segment was more elongated than the others.
The margins of articulation of the bony segments consisted of unossified cartilages, and
at least one pair of costal cartilages articulated with the side of the sternum where the
segments were jointed together ; but between the 8th and 9th bony segments both the
9th and 10th pairs of cartilages articulated with the bone. The most anterior or prse-
sternal segment was a slender mesial cartilage of almost uniform transverse diameter
throughout, and 70 mm. long ; it projected forwards into the neck, and the 1st pair of
costal cartilages was articulated at the junction of this praesternal cartilage with the 1st
osseous segment. The last or xiphisternal segment was prolonged behind the last pah*
of sternal ribs ; its most anterior half was an elongated bone 50 mm. long, which was
continuous behind with a broad platedike cartilaginous xiphisternum.
Anterior Extremity. — The scapula was falciform and 135 mm. in length. The
pras- and post-spinous fossae were almost of equal size. The upper two-thirds of the
spine formed so low a ridge as scarcely to be recognisable, the lower third, which was
32 mm. long, projected for 17 mm. from the dorsum of the bone. There was no
acromion and the coracoid was feeble.
The humerus had a prominent deltoid ridge, inner and outer tuberosities about
equal in size, bicipital groove deep, no supra-condyloid foramen, capitellum and trochlea
both distinct. It was 117 mm. lon^.
The ulna had a large olecranon ; its shaft was traversed by a strong anterior ridge
to which the internal lateral ligament of the elbow was attached ; its lower end was
somewhat rounded, and articulated with radius, cuneiform, and pisiform. It was
136 mm. long.
The radius had a cup-shaped head, below which was a feeble bicipital tuberosity.
The shaft was rounded above and flattened out at the lower end, which articulated
below with a large scapholunar bone, and at its inner border with the cuneiform and
ulna. Its length was 126 mm.
The manus was pentadactylous. Owing to the coalescence of the scaphoid and
lunare there were only seven carpal bones, which were rough both on the palmar and
dorsal surfaces for the attachment of ligaments, and which were devoid of ridges and
processes. The scapholunar articulated with radius, trapezium, trapezoid, os magnum,
and unciform. The cuneiform articulated with radius, ulna, pisiform, unciform, and 5th
(zool. chall. exp. —tart Lxvin. — 1887.) Yyy 4
26 THE VOYAGE OF H.M.S. CHALLENGER.
metacarpal. The pisiform was small, and articulated both with the cuneiform and
ulna. The trapezium articulated with scapholunar, trapezoid, 1st and 2nd metacarpals.
The trapezoid articulated with scapholunar, trapezium, os magnum, and 2nd metacarpal.
The os magnum, though small, articulated with the 2nd, 3rd, and 4th metacarpals,
and with the unciform, scapholunar, and trapezoid. The unciform was small and
did not reach the ulnar border of the wrist, so that the 5th metacarpal articulated
with both it and the cuneiform ; it also articulated with the scapholunar, os magnum,
cuneiform, and 4th metacarpal. The skin on the last phalanx had not been removed,,
and was covered with yellow hairs both on the dorsal and palmar aspects. Each
phalanx had an elongated dark -brown nail on its dorsum. The pollex was the longest
digit, and they gradually diminished in length to the minimus. Each of the three
segments of the pollex was longer than the corresponding segments in any of the other-
digits. Its first or so-called metacarpal segment had both a proximal and a distal
epijmysis, and in this respect it corresponded with all the phalanges, except the terminal,
which latter had only a proximal epiphysis. The metacarpals of the other digits had
each a distal epiphysis but no proximal. A parr of small sesamoids was situated on
the palmar aspect of each metacarpophalangeal joint.
Pelvis. — The innominate hones articulated with the 1st and slightly with the 2nd
sacral vertebra, and also with each other through the interposition of a cartilage at the
pubic symphysis. The ilium was 64 mm. long and its crest was truncated. Its
ventral surface was narrow and gave but little room for the attachment of an iliacus
muscle. The dorsal surface was four times the breadth of the ventral. From the
sacro-iliac joint the ilium passed almost transversely outwards to the crest which, as in
the Elephant Seal, was in nearly the same transverse plane as the base of the sacrum.
A pectineal tubercle marked the place of junction of the ilium and os pubis.
Although the ligamentum teres was absent a narrow and elongated noncartflaginous
covered area, bounded by a definite line, was at the bottom of the acetabulum. The
ischium, os pubis, and obturator foramen were all elongated. The diameter from the
pectineal tubercle to the pubic symphysis was 131 mm. A sharp pectineal line extended
from the tubercle to the pubic symphysis. An angular projection on the upper border
of the ischium marked the position of the ischial tuber, and between it and the acetabu-
lum, but nearer to the latter, was a ridge which probably represented the ischial spine.
Posterior Extremity. — The femur, 96 mm. long, was flattened, and with its
epiphyses not ankylosed. The head was smooth and without any depression for a
ligamentum teres. The great trochanter was large, and with a digital fossa. There
was no small or third trochanter. The condyloid end possessed a shallow trochlear
surface for a patella, which surface was not continuous with the articular areas on the
condyles. The two condyles were separated from each other by an intercondyloid fossa,
to which the crucial ligaments were attached.
REPORT ON THE SEALS. 27
The 'patella was 21 mm. long, and almost flat on both its articular and cutaneous
surfaces. The upper end of the bone was very slightly thicker than the lower.
The tibia had a broad upper end with two smooth surfaces for articulation with
the femoral condyles, and an intermediate rough part for the attachment of the crucial
ligaments and semilunar cartilages. Its shaft had three surfaces above but was antero-
posteriorly compressed below. The shaft had just below the condylar end a tubercle
for the attachment of the great patellar tendon, and externally an articular surface for
the fibula. About the middle of the ventral side of the shaft was a rough ridge for the
attachment of the gracilis tendon. The lower end of the tibia was prolonged into a
short malleolus, and articulated both with the fibula and the upper surface of the
astragalus ; it was grooved in front for the tendons of the tibialis anticus and long
extensor of the great toe ; whilst behind there was also a groove for the tendon of the
tibialis posticus. Its length was 201 mm.
The fibula, although about equal in length to the tibia, had only half its bulk. Its
upper end was relatively broad ; the lower end was prolonged into a malleolus, and
articulated with the tibia, the outer surface of the astragalus, and by the posterior part
of its tip with a small area on the os calcis external to the astragalo-calcaneal articula-
tion. The epiphyses were not ankylosed to the shafts of the two bones of the leg.
The interval between their shafts was wide in the middle.
The pes was pentadactylous and with dark yellowish -brown hair at the tips of the
toes both on the dorsal and plantar surfaces ; a small dark-brown nail, concealed amidst
the haii', was present on the dorsal aspect of the terminal phalanx of each toe. The
hallux and minimus, about equal in length, were much the longest digits, about
280 mm., they were rounded at the tip and the integument extended about 30 mm.
beyond the terminal phalanx. Digits 2 and 4, about equal to each other, though
2 was a little longer, reached a little beyond the line of articulation of the 2nd and
3rd phalanges of the thumb. Digit 3 was the shortest, and ended on a line with
the articulation of the 2nd and 3rd phalanges of digit 2. Each of the three segments
of the hallux was longer than the corresponding segment in the other digits. The
phalanges diminished in length from the 1st to the 3rd. The 3rd metatarsal was
the shortest. The 2nd metatarsal was equal in length to the 5th, and resembled in
shape the corresponding bone in the Elephant Seal ; it articulated behind with the
three cuneiforms, and the 1st and 3rd metatarsals. The 4th metatarsal was concave
on the external lateral surface at its proximal end where it articulated with the 5th
metatarsal. The first or so-called metatarsal segment of the hallux had both a
proximal and a distal epiphysis, a character which it shared along with all the
phalanges except the terminal, which latter had only a proximal epiphysis. The
metatarsal bone of each of the four outer toes had only a distal epiphysis. A pair of
sesamoid bones was situated on the plantar surface of each metatarso-phalangeal joint.
28 THE VOYAGE OF H.M.S. CHALLENGER.
The tarsalia were seven in number. The astragalus was a larger bone than the
os calcis. Its posterior process reached behind the corresponding process of the os
calcis, and formed the most projecting part of the heel ; it was grooved for the tendon
of probably the plantaris muscle. Its head passed in front of the same bone, and
articulated with the scaphoid and cuboid. Its inferior surface articulated with the
os calcis, and its superior and external lateral surface with the two bones of the leg.
The os calcis was attenuated behind into a calcaneal- process, and articulated with
the astragalus and fibula on its superior, and the cuboid on its anterior surface. The
cuboid possessed both a plantar tubercle and a deep peroneal groove, and articulated
with the os calcis, astragalus, scaphoid, ecto-cuneiform, and 4th and 5th metatarsals.
The scaphoid was shaped not unlike the human bone, and articulated with the
astragalus, cuboid, and three cuneiforms. Of the three cuneiform bones the ento- was
much the largest, and the meso- was so small as not to be visible on the plantar
surface. The ecto-cuneiform had a peroneal groove on its plantar surface, and it
articulated with the scaphoid, meso-cuneiform, cuboid, and 2nd and 3rd metatarsals.
The meso-cuneiform was only seen on the dorsum of the foot, and the ento-cuneiform
passed so far in front of it that the 2nd metatarsal had to be prolonged both backwards
and inwards in order to reach it ; it articulated with the other cuneiforms, the scaphoid,
and the 2nd metatarsal. The ento-cuneiform articulated with the meso-cuneiform,
scaphoid, and 1st and 2nd metatarsals.
The vertebral column of Lcptonychotcs measured, with the discs dried and in position,
1540 mm. or 5 feet, and as the skull was 237 mm. long, the length from the premaxillary
bone to the tip of the tail was 1777 mm. or 5 feet 9 inches. As the ossification of the
skeleton was so imperfect it is obvious that this seal in its adult condition must grow to
be a much longer animal than was the specimen above described.
The length-breadth indices of the skulls measured in Table III., calculated on the
relation of the condylo-premaxillary length to the interzygomatic width, were for
Weddeli's Seal 59*9, and for the crania of Stenorhynchus leptonyx 49 "8 and 54 "8 respec-
tively, but calculated on the width behind the external meatus this index was 66 for
Weddeli's Seal and 51 and 53 "5 for Stenorhynchus leptonyx. These figures show at a
glance how much wider in relation to the length the skull of Weddeli's Seal is than the
other two crania.
REPORT ON THE SEALS. 29
Otaria, Peron.
Otaria, Peron, Voyage aux Terres Australes, ii., 1816.
Under the generic name Otaria I include only those Eared Seals which possess a long
and deeply concave palate, truncated posteriorly, and extending back as far as, or nearly as
far as, the hamular processes of the pterygoids. One large adult skull, which possessed this
character of palate, was collected by the expedition. It was from an animal shot at Port
Stevens, West Falkland Islands, and was presented to Sir C. Wy ville Thomson by Mr. E. T.
Smith. The skin of another specimen, a young male, containing the skull, was presented
by Mr. Dean, of Stanley, Falkland Islands. In the Narrative of the Voyage it is stated
that along the coast of these islands many bones of seals and whales were scattered.
Otaria jubata (Forster).
Phoca jubata, Forster, 1755, and Schreber, Saugethiere, iii.
Lion Seal.
External Characters. — The young specimen of the Sea Lion obtained at Stanley was
probably from fifteen to twenty days old, as the skull which it contained is almost the
same size as one in the Royal College of Surgeons of England, which is said to be of that
age. The skin had been preserved in salt and was in good condition. It is unnecessary
for me to give a detailed description of the skin, as the external characters of the Sea
Lion have been so admirably described and figured by Dr. James Murie in his well-known
memoir on this animal.1 It may suffice if I state that from the muzzle to the tip of the
tail the length was 36 inches, and 41 inches to the tip of the pes, when the hind foot was
drawn backwards. The pinna of the ear was pointed at the tip and 16 mm. long. The
hair on the back was dark brown, almost indeed black, but the hair on the belly was
somewhat lighter, and with a slight reddish shade in the brown, and there was no under
fur. The dorsum of both manus and pes was haired as low clown as the nails, but the
skin of the palm and sole was hairless and much wrinkled.
Skull. — The skull of the young male closely resembled both in size and appearance
the specimen figured by Dr. Murie in pi. lxxvii. figs. 12, 13. The skull of the adult
was like that figured by him in the same plate, figs. 20, 21. It also was a male and of
full age, for the basi-cranial synchondroses were both ossified, and the teeth were worn.
This skull was accompanied by the hyoid apparatus and the atlas vertebra. After the
excellent description of the skull of Otaria jubata in both sexes and at different ages
which has been given by Dr. Murie, it might seem unnecessary again to describe the
skull of this animal ; but as one of the objects which I have in view in this Report is to
make a comparison between the skulls of different genera of seals so as clearly to bring out
1 Trans. Zool. Soc. Lond., vol. viii part ix.
30
THE VOYAGE OF H.M.S. CHALLENGER.
their differential characters, I have drawn up a short account of these specimens on the
same lines as with the other genera described. I have also compared these Falkland
Island crania with the adult male skull of a Sea Lion brought by Dr. R. 0. Cunningham,
from Laredo Bay, Magellan Strait,1 with another adult collected at Maldonado, Eiver
Plate, by the same naturalist, and with a third adult obtained in guano on the Chincha
Islands, off the coast of Peru, all of which specimens are in the Anatomical Museum of
the University of Edinburgh. The last-named skull was at one time in the collection of
Dr. M'Bain, and was described by him 2 as probably an example of the seal named by
von Tschudi and Peters, Otaria ullose.
The principal dimensions of the crania are given in the following table : —
Table IV. — Crania of Otaria.
Juv.
Adult.
Adult.
Adult.
Adult.
Stanley,
West
Laredo Bay,
Maldonado,
Chincha
Falkland
Falkland
Magellan
River Plate.
Islands.
Islands.
Island.
Strait.
mm.
mm.
mm.
mm.
mm.
Extreme condylo-premaxillary length,
161
365
365
255
252
From basion to optic foramen,
92
148
151
121
107
Extreme iuterzygomatio width,
97
235
226
154
Extreme width, immediately behind external
meatus, .....
90
217
210
125
Greatest width of palate,
34
71
62
47
43
Width between outer side of base of upper
canines, .....
33
116
106
49
Width between outer side of base of lower
canines, .....
26
91
Length of palate in line of mesial suture to
central incisor, ....
78
227
148
142
Height of skull from basion to middle of
occipital crest, ....
66
155
139
90
Smallest interfrontal width (at root of crest),
47
20
25
28
17
Length of nasals, ....
30
70
49
39
Greatest width of anterior nares,
24
41
49
34
27
Greatest width at postorbital processes,
54
145
116
85
52
Length of mandible,
100
279
• . •
Width between outer ends of condyles of
mandible, ....
91
213
From the above table it will be seen that the adult crania from West Falkland Island
and Laredo Bay were in all their dimensions considerably larger than the Maldonado
and Chincha Islands specimens, and as all four crania had the basi-cranial synchondroses
closed, the question arises — Are the smaller skulls a different species from the larger, or are
they the females and the larger specimens the males of the same species ? In addition to
these differences in size the two larger crania, more especially the West Falkland specimen,
possessed lofty occipital, sagittal, and interfrontal crests, the latter of which was grooved
1 Natural History of the Strait of Magellan, Edinburgh, 1871. 2 Journ. of Anat. and Phijs., vol. iii. p. 113, 1869.
REPORT ON THE SEALS. 31
on its summit antero-posteriorly ; also from each parietal, where it formed the anterior
wall of the cranial box, a strong tubercle projected forwards. In the two smaller adult skulls
the sagittal and occipital crests were very feeble, and the interfrontal was scarcely marked,
and although the frontal bone at the anterior wall of the cranial box bulged forwards, it
was not elevated into a tubercular process.
In the young Falkland Island skull, the crests were undeveloped, the summit of the
cranium was smooth and the frontal region was only slightly constricted behind the
postorbital processes, and its absolute width was about twice as great as the large male
crania. The length of the brain-cavity, measured from the basion to the optic foramen,
is given in Table IV., so that its proportional length to that of the entire cranium may be
estimated, and in the young skull the antero-posterior diameter of the cranial box was more
than one-half that of the entire skull, so that the more anterior part of the skull grows
in its progress to adult life at a much greater rate proportionally than the brain-cavity.
Qq -I 1
The dentition in all the specimens was as follows : — incisors ^tj » canines y^ ' post-
c &
canines -p — ? = 36. The individual teeth in the adults possessed the characters which have
so frequently been described in Otaria, so that I need not dwell upon them.
The teeth in the young male were so much smaller than the adult that they were
apparently the milk series. In the upper jaw only two incisors on each side, each with
an anterior and a posterior cusp, bad cut the gum ; in.3 being still concealed. Very
small canines had erupted, and the points of the 1st and 2nd post-canines could be seen.
In the lower jaw the incisors, canines, 1st and 4th post-canines had cut the gum.
As compared with the Elephant Seal, the zygomatic arches were much flatter, and the
greatest width was towards the posterior end. The zygomatic and temporal fossae were
not so capacious relatively to the size of the skull, but the frontal region in Otaria was
much more constricted immediately behind the orbits. In the two adult male skulls the
distance from the anterior surface of the cranial box (the tubercular process not being
included) to the antorbital process was about twice as great as the orbital diameter from
the antorbital process to the orbital process of the malar. In the smaller adult from
the Chincha Islands the orbital diameter was three-fourths that of the entire distance,
and in the Maldonado skull two-thirds. In the young male from the Falkland Islands
the two diameters were almost equal. It follows, therefore, that whilst in the young
animal the back of the orbit is in close relation to the front of the cranial box, in the
adult male it is separated from it by a wide interval, which marks the position of the
temporal muscle and acquires its magnitude in relation to the use of that muscle in the
masticatory process. The zygomatic process of the temporal did not curve upwards so
abruptly as in the Elephant Seal, and did not reach the tip of the orbital process of the
malar, which was much more stunted than in the Elephant Seal.
The nasal bones were relatively short and not ankylosed except in the Maldonado skull ;
32 THE VOYAGE OF H.M.S. CHALLENGER.
they diverged from each other posteriorly so as to admit between them in the middle line
an anterior or nasal prolongation of the frontal bone. The anterior border of the mes-
ethmoid was vertical, but did not quite reach the anterior nares. The anterior end of
the spout-like vomer terminated a little in front of the anterior border of the nasals.
The horizontal part of the premaxdla was relatively short, and gave origin to a nasal
tubercle»close to the floor of the anterior nares ; in one of the larger skulls the depth of
the bone from the anterior nares to the alveolar border was 45 mm., in one of the smaller
only 18 mm. The ascending part of the premaxilla mounted upwards and formed the
lateral boundary of the nares, and by its upper end articulated with somewhat more than
the anterior half of the outer border of the nasal bone. The superior maxilla articulated
with the rest of the outer border of the nasal, and completely shut out the frontal from
this border. A large, much divided maxillo-turbinal occupied the interval between the
mes-ethmoid and the outer wall of the nose, but it did not come quite so far forwards as
in the Elephant Seal. The plane of the anterior nares sloped downwards and forwards
from the nasal bones to the premaxilla, and the opening was well in front of both the
antorbital process and infraorbital foramen. Although the large males possessed massive
canines, yet the anterior end of the superior maxillae with their canines did not lie so
near to the transverse plane of the incisor teeth as in the Elephant Seal.
The postorbital processes were transverse in direction, much larger than the ant-
orbital in all the crania, but in the two large crania the antorbitals were several times
larger than in the smaller skulls. From Table IV. it will be seen that the skulls differed
greatly in width in this region, and this difference in relation to their almost equal length
was especially marked in the Maldonado and Chincha Island specimens. In the West
Falkland adult a strong fibrous band passed from the postorbital process to the zygoma,
and completed the orbital ring posteriorly.
The hard palate had the characteristically elongated form of the genus. In the larger
skulls the concavity was much deeper in proportion to the size of the specimens, and the
borders of the palate behind the molar teeth converged more closely together than in
the smaller crania. The distance from the last molar tooth to the posterior edge of
the hard palate was 101 mm. in the large West Falkland Island skull. The greatest
palatal width of the larger skulls was either between the canines or the more anterior
post-canine teeth, and in the smaller skulls immediately behind the last molar. The
premaxilla? were not so distinctly triangular as in the Elephant Seal, and each contained
a well-defined naso-palatine foramen. The most anterior part of the palato-maxillary
suture was triangular, and either just behind or opposite the last molar tooth. The
palatal surface of the palate bone formed nearly one-half of the length of the hard
palate, but in one of the larger crania the proportion varied on the two sides owing
to these bones not being symmetrical. The dentary border of the superior maxilla,
although continued behind the last molar, yet did not nearly reach the length of the
REPOKT ON THE SEALS. 33
posterior border of the hard palate. In the young Falkland Island male the hard
palate differed in some respects from the adult; it had scarcely any concavity in its
posterior part, but anteriorly it was somewhat hollowed out ; the anterior part of the
palato-maxillary suture was opposite the penultimate molars ; the length of the palatal
surface of the palate bone was equal to the length from the palato-maxillary suture to
the incisive canal ; the widest part of the hard palate was immediately behind the last
molars. The posterior edge of the hard palate and the posterior nares in the two large
skulls were in the same transverse plane as the anterior border of the glenoid fossa, but
in the smaller adults they were a little anterior to that plane, and more so in the
young skull from Stanley. The hamular processes were curved and projected down-
wards, inwards, and then outwards.
The great elongation of the palate in Otaria juhata is therefore due to the remarkable
antero-posterior diameter of the palatal plate of the palate bone, which completely con-
cealed both the sphenoidal articulation and the posterior border of the vomer, the latter
of which was falciform, and did not articulate with the palate, but passed forwards to
reach the vomerine crest of the superior maxilla. As the hard palate was covered by the
mucous membrane when the skull reached me, I took the opportunity of examining it
when softened prior to its removal. This membrane possessed numerous short papillae,
which, in the part of the palate situated between the molar teeth , were arranged in seven
low ridges, which were not quite transverse, but with a slight inclination backwards.
Between and in front of these rows similar dwarf papillae were scattered over tbe
mucous surface, but behind the last molars the membrane was smooth.
All the crania possessed alisphenoid canals and mastoid processes. In all, the
tympanic bulla had a process projecting vertically downwards from the inferior surface.
In the larger adults it was thick and truncated, in the smaller adults it formed a sharp
ridge ; in the young male, although the ridge did not project so much as in the smaller
adults, it was quite as thick. The tympanic cavity was opened into in the Maldonado
specimen, and consisted of a large chamber, dilated below, which suddenly narrowed as
it ascended to the outer side of the petrous-temporal. The carotid canal opened within
the boundary of the jugular foramen. The occipital condyles were not continuous
anteriorly in the adults but separated by a definite interval, and their inner borders in
front lay in a plane running almost directly from before backwards; in the young skull,
however, the condyles were continued into each other in front, and the cartilage was
prolonged from one to the other. No supra-occipital foramen was visible either at
the foramen magnum or below the occipital crest. The inferior surface of the basi-
occipital had in the four adult crania an elevated ridge running antero-posteriorly, and
there was no mesial perforation in any of the skulls. The carotid canal opened imme-
diately within the boundary of the jugular foramen. The par-occipitals were stunted.
The skulls from the Falkland Islands were the only specimens wh ich possessed a lower
(ZOOL. CHALL. EXP.— PART LXVIII. — 1887.) Y yj 5
34 THE VOYAGE OF H.M.S. CHALLENGER.
jaw. The angle was marked by a ridge-like tubercle wdiich projected backwards. A
strong, quadrangular, inflected subcondyloid process sprang from the posterior border of
the ascending ramus ; it was separated by a notch from the ridge-like angle, and by a
still deeper notch from the neck and condyle. The coronoid was broad, thin, and formed
an obtuse angle. The body was massive, with its lower border everted, and closely
corresponded in its characters to the description given by Dr. Murie. The mandible in
the young skull showed on a smaller scale the same character as the adult.
The differences between the larger and smaller adult crania, in addition to that of
size, may be summarised as follows : — In the smaller skulls the occipital and sagittal
crests were feeble ; no parietal tubercle ; the antorbital processes much smaller; the
interfrontal diameter relatively larger ; the front of the premaxilla was both absolutely
and relatively less deep, and its nasal tubercle was scarcely marked ; the breadth of the
palate was greater behind the molars ; the tympanic bulla was prolonged downwards
into a sharp ridge instead of a thick, truncated process ; the length of the brain-cavity
in the smaller crania, especially the Maldonado specimen, was, in proportion to that of
the entire skull, greater than in the larger specimens ; and the antero-posterior
diameter of the orbit bore a larger proportion to the distance between the front of the
cranial box and the antorbital process. These differences cannot be ascribed to age,
for the smaller skulls were as perfectly ossified as the larger crania. As nothing
is known of the sex of the animals from which the smaller crania were derived, it
cannot absolutely be stated that the differences were sexual only, though without doubt,
for the most part, they were such as are mainly occasioned by a more vigorous ossification
in the one skull than in the other, as we are in the habit of recognising in male crania
when compared with female. That important sexual differences do exist in the crania
of the Sea Lions has already been pointed out by Sir Richard Owen,1 Mr. J. A. Allen,2
and Dr. J. Murie,3 and both Drs. Gray 4 and Murie have dwelt on the changes in form
which the skull undergoes in passing from the young stage to that of adult life and old
age, and the specimens now before me show that the characters of the two smaller adults
in many respects approximated to those of the young male Falkland Island cranium.
The opinion of zoologists has greatly fluctuated regarding the number of species
which should be referred to the genus Otaria, even when that genus is restricted
according to the definition given on p. 29. A perusal of the numerous papers on the
Eared Seals by the late Dr. J. E. Gray will show how frequently he changed his views
on this subject. In a similar manner the late Professor Peters of Berlin from time to
time either added to or subtracted from the number of species. It is unnecessary to give
a resume of their various changes of opinion, as this has already been done by Mr. J. A.
1 Catalogue, Royal College of Surgeons.
2 Bull. Mus. Corwp. Zool, Cambridge, U.S., vol. ii. 1870-1871 ; and History of North American Pinnipeds, 1880.
3 Proc. Zool. Soc. Lond., 1869 ; Trans. Zool. Soc. Lond., vol. viii.
4 Proc. Zool. Soc. Lond., 1859 ; and Catalogue of Seals and Whales.
REPORT ON THE SEALS. 35
Allen in his important monograph above referred to. It may suffice to state that Dr.
Gray, in his latest writings1 on this subject, spoke of four species, viz., Otaria jubata,
Otaria minor, Otaria ulloas, and Otaria pygmsea; whilst Dr. Peters, in his last published
paper,2 recognised only one species, viz., the Otaria jubata of Forster. Peters regarded the
Otaria leonina of Fr. Cuvier, and the Otaria ullose of Tschudi, merely as " Localrassen,"
whilst he made no mention of the Otaria godeffroyi which he had described in May
1866 3 as a distinct species, so that he doubtless ultimately considered it also as only a
variety of Otaria jubata. The geographical distribution of Otaria jubata is said by
Peters to extend around the southern half of South America, from Rio de la Plata on
the east to Callao and the Chincha Islands on the west. Mr. Allen also favours the
view that Otaria jubata is the only species, and in addition to its habitat on the South
American continent, assigns it to the Galapagos Islands in the Pacific, from specimens
collected by the Hassler Expedition, whilst the crania collected by the Challenger, and
others previously procured, have established it to be a denizen of the Falkland Islands.
If it be correct to regard the genus Otaria as consisting only of the species jubata,
then it will follow that all the crania which I have described in this section, notwith-
standing their great difference in size, and to some extent in proportion, are to be
considered as of this species, and the small adult skulls would then be females, whilst
the larger crania would without doubt be males. Dr. Murie, in his monograph on the
Sea Lion, has given the following dimensions of crania which he has measured. In an
adult female the skull was 11 inches long (280 mm.), and 5-8 inches broad (148 mm.) ;
an old male was 12"8 inches long (325 mm.), and 7'5 inches broad (190 mm.), and a
very old male 14*3 inches long (363 mm.), and 9"4 inches broad (239 mm.). Mr. Allen
records the mean length of eight male skulls at 350 mm., and the mean breadth 223 mm.,
whilst the mean length of four females was 261 mm., and the mean breadth 143 mm.
These proportions closely approximate to the measurements of the adult male and female
crania which I have given in Table IV. Mr. Allen also states that there is a wonderful
disparity in size between the sexes in Otaria jubata, the weight of the adult males being
generally three to five times that of the adult females.
The length-breadth indices of the two large male skulls, calculated on the interzygo-
matic breadth, were respectively 64-3 and 61 -9, and on the width behind the external
meatus 59"4 and 57'5. The corresponding indices in the smaller adult from Maldonado
were 60"3 and 49, and in the young skull 60'2 and 55*9. The difference between the
interzygomatic width and the width behind the external meatus was most strongly
marked in the Maldonado cranium.4
1 Ann. and Mag. Nat. Hist., ser. 4, vol. xiii. p. 325 ; and Hand Atlas of Seals.
2 Monatsber. d. k. preuss. Akad. d. Wiss. Berlin, August 9, 1877, p. 506. 3 Op. cit, May 17, 1866, p. 266.
4 I have not thought it necessary to figure the skull of Otaria jubata, as Dr. Murie's memoir on this animal is so
completely illustrated.
36 THE VOYAGE OF H.M.S. CHALLENGER.
Arctocephalus, F. Cuvier.
Arctocephales, F. Cuvier, Mem. du Museum, xi., 1824.
In the Narrative of the Voyage it is stated that Fur-Seals frequented Nightingale
Island, one of the Tristan da Cunha group ; the Crozet Islands ; Kerguelen Island ; Juan
Fernandez ; the Messier Channel ; and Elizabeth Island in the Strait of Magellan.
Specimens of Eared Seals, which did not possess the elongated concave palate so
characteristic of the genus Otaria in the sense defined on p. 29, were procured from the
Kerguelen group of islands ; in Messier Channel on the west coast of South America, and
from Juan Fernandez. They consisted of the following specimens from Kerguelen : —
Two carcases of young Fur-Seals without the skin, procured from the " Emma Jane " at
Fullers Harbour, January 1874; two skeletons of Fur-Seals also at Fullers Harbour, which
were distinguished from each other as No. 1 and No. 2 (No. 2 having been killed on
Swaine Island). From the Messier Channel were obtained the skin and skeleton of a
male and the skin and skeleton of a female, also two skeletons of males shot on rocks in
January 1876. The specimen from Juan Fernandez was a skin containing the skeleton
of a very young animal.
Arctocephalus gazella, Peters (PI. VI.).
Arctocephalus gazella, Peters, Monatsber. d. k. preuss. Aknd. d. Wiss. Berlin, June 10, 1875, p. 393.
Kerguelen Island Fur-Seal.
This species of Fur-Seal was described by the late Professor W. Peters from two
specimens procured for the Berlin Museum by the German exploring ship " Gazelle,"
which visited Kerguelen Island shortly after the departure of the Challenger in 1874.
The larger of these two specimens was the skin of a male, but without the cranium,
whilst the smaller skin contained the head and trunk of a female, not quite adult.
Subsequently to his first description of this species, Peters ascertained ' that only
the female specimen had been obtained from Kerguelen ; whilst the male skin was either
from St. Paul or Amsterdam Island, and he named it Arctocephalus elegans.
The Fur-Seals collected by the Challenger at Kerguelen Island I have referred to
Arctocephalus gazella. At the request of the late Sir C. Wyville Thomson, I forwarded,
in November 1877, the skull of one of the larger specimens (No. 2) procured by the
Challenger to Professor Peters for examination. In returning the specimen to me in
April 1878, Professor Peters wrote that it was the male of Arctocephalus gazella, of
which he had previously seen only the skull of a female. The male, he said, showed
" all the peculiarities of the species, which consist, amongst others, in the extreme small-
ness of the tympanic bones, a part which is of so great an importance as it is peculiar for
each species."
1 Idem, May 18, 1876, p. 315.
REPORT ON THE SEALS. J7
Skeleton. — Neither of tbe two skeletons from Fullers Harbour had reached adult life,
as the epiphyses of the long bones and the plates of the vertebral bodies were not
ankylosed to their respective bones.
Skull. — In each of the larger crania the occipito-sphenoid synchondrosis was
ankylosed, but the joint between the two divisions of the sphenoid was yet open.
The principal dimensions of the crania are given in Table V. : —
Table V. — Crania of Kerguelen Island Fur-Seal.
No. 1.
No. 2.
mm.
mm.
Extreme condylo-premaxillary length, ....
212
211
From basion to optic foramen, ....
85
79
Extreme interzygomatic width, ....
131
120
Extreme width immediately behind external meatus,
120
109
Greatest width of palate, ....
39
35
Width between outer sides of base of upper canines,
45
42
Width between outer sides of base of upper lateral incisors,
24
25
Width between outer sides of base of lower canines,
30
32
Length of palate to incisor teeth,
92
91
From basion to middle of occipital crest,
77
69
Smallest inteifrontal width in plane of upper surface, .
33
30
Length of nasals, .....
29
Greatest width of anterior nares,
22
28
Vertical diameter of rues-ethmoid at anterior nares,
From anteroinferior angle of nies-ethmoid to central incisor,
Greatest length of mandible, ....
142
133
Greatest width at condyles of lower jaw,
124
93
Although the ossification of these two crania was so far advanced, yet they possessed
no sagittal or interfrontal crest, and there was only the faintest indication of a crest in
the occipital region. On the supposition that these crania were males, as was surmised
by Professor Peters from an examination of the smaller of the two specimens, it would
appear that in this species of Arctocephalus the development of cranial crests scarcely if
at all occurs.
The dentition was the same in both skulls, and the formula was as follows : — incisors
36. The first and second upper incisors possessed
canines
-, post-canines
2-2' 1-1"" 5-D
the customary anterior and posterior cusps. Both the upper and lower canines had only
a single large cusp. The greatest interval between the post-canines was between the 4th
and 5th and the 5th and 6th, the interval between the last named being the widest.
The 6th post-canine was the smallest, and both it and the 5th in the upper jaw had
two diverging fangs ; the remaining upper post-canines had only a single fang. The 5th
lower post-canine was two-fanged ; the remainder one-fanged.
38 THE VOYAGE OF H.M.S. CHALLENGER.
The zygomatic arches were widest behind, and the arch was somewhat flattened. The
malar bone formed a larger proportion of the arch than in Otaria. The antero-
posterior diameter of the orbital cavity, from the antorbital process to the ascending
process of the malar was in No. 2 T9xths and in No. 1 ^-Jths of the distance from the
cranial box to the antorbital process. The orbital process of the malar was pointed, and
the zygomatic process of the temporal only reached its base, and did not therefore turn
up behind it.
The nasal bones were separate, and received posteriorly a mesial process of the frontal
between them. The anterior border of the mes-ethmoid did not extend quite as far as
the anterior border of the nasals. The anterior end of the vomer reached to 22 mm. of
the tip of the premaxilla. The horizontal part of the premaxillse was short, so that the
anterior nares came close to the anterior end of the skull, and the premaxillse projected so
far in front of the superior maxillae that the upper canines were well behind the incisor
teeth. The ascending process of the premaxilla formed the lateral boundary of the
anterior nares and articulated with a little more than the anterior half of the outer
border of the nasal. The superior maxilla articulated with the nasal behind the pre-
maxilla, but a small part of the frontal also joined the outer border of the nasal behind
the superior maxilla. The maxillo-turbinals did not quite reach the anterior nares, the
plane of which sloped obliquely downwards and forwards to the incisive region, well in
front of both the antorbital process and the relatively large infraorbital canal. The post-
orbital processes were transverse and distinctly larger than the antorbital. As the left
process was broken off in each skull the width in this region could not be taken.
The hard palate was only slightly concave. Its posterior edge was transverse at and
on each side of the mesial suture ; it was so far in front of the glenoid fossa as to be in
the same transverse plane as the orbital process of the malar bone ; and the interval from
the posterior edge to the last molar tooth was only 20 mm. The most anterior part
of the palato-maxillary suture was opposite the 4th pair of post-canines, which were
distinct, and curved backwards, downwards, and outwards, and the posterior border
of the hard palate was 29 mm. iD front of the hamular pterygoids. The palatal plates
of the palate bones did not contribute so much to the formation of the hard
palate as the corresponding plates of the superior maxillae, the dentary border of
which latter bones extended almost as far back as the posterior edge of the hard palate.
The palatal surface of each premaxilla was quadrilateral in shape, and each contained a
large naso-palatinc canal. The posterior edge of the vomer sloped very obliquely forwards,
and was not seen at the posterior nares, which openings were not nearly so far back as
in Otaria, and permitted the junction between the pre- and post-sphenoids to be seen.
Both the crania had alisphenoid canals and mastoid processes. The tympanic bulla
was almost flattened and marked by only a low ridge. The occipital condyles were
separated by a wide interval in front, and their inner borders anteriorly were almost
REPORT ON THE SEALS.
39
parallel to an anteroposterior plane. The basi-oecipital was not perforated mesially.
Each skull had a single supra-occipital foramen immediately within the outer edge of the
foramen magnum. The carotid canal opened independently of the jugular foramen. The
par-occipitals were very slight.
The lower jaw, although much less massive than in Otaria, yet had a general
resemblance in form ; the subcondyloid process from the posterior border of the ascending
ramus was distinct and inflected, and the coronoid process was lelatively large. It also
had a tubercle which marked the angle, which was feeble as compared with Otaria.
Avctocephalus australis (Zimmermann) (Pis. VI., VII.).
Phoca australis, Zimmermann, Geogr. Geschichte, iii., 1783.
South American Fur-Seal.
The specimens from the Messier Channel have furnished me with material for studying
both the external characters and the osteology of the Fur-Seal which frequents the
southern part of South America.
External Characters. — The external characters were examined in the skins of
both the male and female specimens. The principal dimensions were as follows : —
Table VI. — Dimensions of South American Fur-Seal.
From snout to tip of tail in straight line,
From snout to tip of longest digit of pes,
Length of free part of tail,
Length of pectoral limb,
Greatest breadth of that limb,
Length of hind limb from root of tail,
Greatest breadth of that limb,
From root of pectoral limb to angle of mouth,
Male.
ft. in.
11
10*
3|
4
6|
5
0
Female.
ft. in.
3 10*
1 2
1 2
u
1 6"
The skin possessed two kinds of hairs, long and short. On the back of the neck and
chest the long over-hairs were from one to two inches long, the shaft of the hair being black
but tipped with grey or yellowish-grey so as to give a grizzled character. In the lumbar,
sacral, and caudal regions the black over-hairs were not more than an inch long, and their
tips were grey. On the under surface of the remarkably elongated neck the over-hairs
resembled in colour those on the back of the neck, but were not quite so long. On the
sides of the body they were like those in the lumbar region, but on the belly they were
blackish-brown, and without grey tips. The dorsal surface of both manus and pes was
40 THE VOYAGE OF H.M.S. CHALLENGER.
covered up to the nails with black hairs not tipped with grey, but beyond the nails and
on the inferior surface of both manus and pes the hairs were absent and the skin was
black and beautifully marked with ridges and furrows. The short under-hairs formed
the proper fur of the animal and constituted a thick undergrowth, concealed by the
long over-hairs, and only exposed when they were parted asunder. The fur was of a
brownish or reddish-brown colour.
In the manus rudimentary nails were present on the dorsum of the terminal phalanx
of each digit, but that on the minimus was so small as to be detected with difficulty.
The pollex was the longest digit and from it they diminished in length to the minimus.
The digits were all so closely enclosed in a common fold of skin that any widening of the
manus by the muscular efforts of the animal seemed impossible. The skin extended,
however, for some two inches beyond the terminal phalanx, being thickened in line with
the digits, but thinner and webbed between them. At the free posterior border of the
manus it was faintly indented and the position of each digit was marked by a slight
projecting fold of integument.
The pes had on the dorsal surface of each of the 2nd, 3rd, and 4th digits a strong
curved greyish-coloured nail ; on both the hallux and minimus the nail was feeble and
scarcely projected beyond the fold of skin at the nail-root. The toes varied but little
in length, digits 3 and 2 being slightly the longest, and the minimus being a trifle
shorter than the hallux. The hallux was the broadest digit, and next to it the minimus.
The toes were connected together by an intermediate web, haired on the dorsum but not
on the plantar surface, so that the animal could widen or diminish the transverse diameter
of the pes. This web, together with five thickened folds of skin, one corresponding to
each toe, was prolonged from 105 to 110 mm. beyond the nails and phalanges. The
thickened folds reached the free end of the foot, but the thinner web did not go so far,
so that the integument of the terminal border of the foot was deeply indented.
The snout was short and the tip of the nose was black and without hairs. From the
upper lip about twenty bristles, arranged in six rows, projected backwards and outwards.
As a rule they were white, though some of the smaller were greyish-black. A pair of
white bristles projected from the skin immediately above each eye. The external ear
was in the male situated 3 inches behind the outer canthus of the eye ; it was 1^ inch
long in the male and 1|- in the female. It was pointed at the tip, and whilst its dorsum
was haired the opposite surface was hairless, hollowed out into a concha and directed
forwards and outwards.
In the male the abdominal opening for the penis was 8 inches in front of the vent.
The female had two pairs of elongated nipples. The hinder or abdominal pair were
8^ inches in front of the vent, whilst the anterior or thoracic pair were 6 inches in front
of the abdominal and only a little behind the transverse plane of the posterior edge of
the pectoral fin.
REPORT ON THE SEALS.
41
Skeleton. — The four skeletons consisted of one female and three males. The female
(No. 2) skeleton was fully ossified. One male (No. 3) was fully ossified ; a second
(No. 1) had the epiphyses completely fused with the shafts of the long bones only at one
extremity, at the opposite a groove of demarcation was still visible ; the third (No. 4)
again was a much younger animal, and the epiphyses at both ends of the long bones were
separable from the shafts.
Skull. — Along with the specimens of Arctocephalus australis from the Messier
Channel I have examined two skulls from Tuesday Bay, Desolation Island, Strait of
Magellan, which were collected by Dr. R. 0. Cunningham when acting as naturalist on
H.M.S. "Nassau."1 In all the crania from the Messier Channel, the basi-occipito-
sphenoid joint was ossified, and in one specimen only (No. 4) was the intra-sphenoidal
joint uuossified. The Desolation Island specimens were aged crania and in all pro-
bability males.2 The dimensions of all these skulls are recorded in Table VII.
Table VII. — Crania of South American Fur-Seal.
6*.
6-
s.
?.
i.
6-
No. 1.
No. 3
No. 4.
No. 2.
Desolation
Island.
Desolation
Island.
mm.
mm.
mm.
mm.
mm.
mm.
Extreme condylo-premaxillary length,
231
233
206
202
241
245
From front of premaxilla to occipital crest,
225
228
195
179
228
236
From basion to optic foramen,
83
94
86
94
97
103
Extreme interzygomatic width
148
118
116
145
Extreme width immediately behind external meatus, .
129
138
104
107
142
125
Greatest width of palate, ......
38
31
26
33
33
Width between outer sides of base of upper canines, .
52
55
34
50
Width between outer sides of base of lateral incisors, .
26
25
24
19
29
Width between outer sides of base of lower canines,
37
35
28
20
Length of palate to incisor teeth, ....
101
86
85
106
108
From basion to middle of occipital crest,
79
"87
77
67
82
83
Smallest interfrontal width in plane of upper surface, .
31
29
25
25
Length of nasals, .......
36
30
30
Greatest width of anterior nares, ....
27
24
26
"31
Vertical diameter of mes-ethmoid at anterior nares,
28
From anteroinferior angle of mes-ethmoid to central
incisor
52
Greatest width at postorbital processes,
56
42
"37
Greatest length of mandible, .....
162
133
139
Greatest width at condyles of lower jaw,
119
111
105
The Desolation Island skulls and Nos. 1 and 3 from the Messier Channel possessed
occipital crests and sagittal crests extending more or less forward into the frontal region,
the greatest elevation of which was 10 mm. In No. 4, a younger male from the
Messier Channel, and in the adult female, these crests were scarcely developed at all.
1 These skulls were presented by Dr. Cunningham to the Anatomical Museum of the University of Edinburgh.
See his work already cited on p. 30 for an account of the seals in this locality.
s Some years ago I left these skulls from Desolation Island, for examination, with the late Dr. J. E. Gray, who
made some notes on them in the Ann. and Mag. Nat. Hist, vol. iv. ser. 4, p. 264, 1869. He referred them to the
species which he had described as " Euotaria nigrescens, the usual Fur-Seal of the Falkland Islands and other parts of the
«oast of South- West America," the same animal as is described in the text as Arctocephalus australis.
(zool. chall. exp. — part Lxviii.— 1887.) Yyy 6
4'2 THE VOYAGE OF H.M.S. CHALLENGER.
The dental formula in all the crania of Arctocephalus australis was as follows : —
3-3 . 1-1 . 6-6 oa
incisors o_9» canines -._-., post-camnes - — F = 36.
In the younger male the left upper canine had not erupted. The two last upper post-
canines were two-fanged, the 5th lower post-canine was also two-fanged. The 6th
upper post-canine, though smaller than the 5th, yet did not differ much from it in size.
In addition to the cingulum and large cusp the lower post-canines possessed a faint
anterior cusp, and the last two also a faint posterior cusp, but in the upper post-canines,
whilst the rudimentary anterior cusp was present, it was the exception to find a rudimentary
posterior cusp, though the last one or two frequently had one.
The zygomatic arches had generally the same form as in Arctocephalus gazella.
The proportion of the antero-posterior diameter of the orbit to the distance from the cranial
box to the antorbital process was in two adult males about fths, in the young male about
fths, and in the adult female about }-f ths. The orbital process of the malar was pointed,
and the zygomatic process of the temporal only reached its base.
In one Desolation Island skull the nasal bones were ankylosed with each other and
with the frontal ; in the remaining crania they were separated behind and received between
them a mesial prolongation of the frontal. The anterior edge of the mesethmoid
did not extend as far forwards as the anterior border of the nasals. The premaxilla
articulated by its ascending process with the anterior half of the outer side of the nasal.
The superior maxilla articulated with the same bone immediately behind the premaxilla,
but in one specimen allowed a slender process of the frontal to be intercalated between it
and the nasal, whilst the superior maxilla itself superiorly and posteriorly was received
into a recess in the anterior border of the frontal. The upper surface of the horizontal
part of the premaxilla was deeply grooved, and the ridges bounding this groove laterally
met in front to form a premaxillary tubercle, stronger than in Arctocephalus gazella,
which projected forwards in front of the plane of the incisor teeth. The postorbital
processes were stronger than the antorbital and in the male skulls were bent backwards at
the tip. The anterior nares sloped downwards and forwards to the premaxillse and were
well in front of both the antorbital process and the relatively large infraorbital foramen.
The hard palate was only slightly concave. Its posterior margin in most of the
specimens was truncated, but in two it was slightly emarginate. This margin was in a
line almost midway between the glenoid fossa and the posterior border of the malar
process of the superior maxilla, and almost opposite the orbital process of the malar bone ;
in the adult males it was from 25 to 30 mm. behind the last molar tooth, and about the
same distance in front of the hamular pterygoids. The hamular processes were distinct
and curved backwards, downwards, and outwards. In the female skull a strong fibrous
membrane had been preserved, prolonging the palate as far back as the hamular ptery-
goids, and it is probable that in all seals in which the hard palate is not itself continued
REPORT ON THE SEALS. 43
as far back, a similar membranous prolongation exists. The most anterior part of the
palato-maxillary suture was in one specimen opposite the 4th pair of post-canines, and in
another opposite the last molar ; a pair of small ossicles was situated in the middle of
this suture. The palate bones formed a smaller proportion of the hard palate than the
superior maxilla. The premaxillse had well-marked naso-palatine canals and in one
specimen a pair of ossicles was situated behind the maxillo-premaxillary suture. The
posterior edge of the vomer resembled that of Arctocephalus gazella, and the articulation
of the prae- and post-sphenoids was visible.
In the female and young male the tympanic bullae were swollen and marked by an
anterior and a posterior low ridge. In the adult males these ridges were stronger and
the posterior one projected downwards as a definite tubercle. Alisphenoid canals and
mastoid processes were present, The occipital condyles generally resembled those of
Arctocephalus gazella. The supra-occipital foramen was not visible. The carotid canal
opened independently of the jugular foramen. The basi-occipital was not perforated
mesially, and the paroccipitals were only distinct in the male skulls.
The lower jaw of Arctocephalus australis in its general form resembled that of
Arctocephalus gazella, but it was, especially in the adult males, more massive, and the
hollowing out of the ramus in the region of the masseter muscle was much deeper.
The hyoid consisted of a transversely elongated basi-hyal, which was articulated at
each extremity, both with a thyro-hyal and a kerato-hyal. The kerato-hyal though
not so long was thicker than the thyro-hyal. Both the epi-hyal and stylo-hyal were
ossified and jointed together, so that nine distinct bones formed the hyoid apparatus.
I do not consider it necessary to give a separate description of the spine and the
other bones of the trunk and of the limbs of the Kerouelen Island and South American
Fur-Seals, as they are in most respects so much alike that one description will, in a
great measure, suffice for both, and such differences as occur can easily be included in it.
As one of the Messier Channel specimens was a fully ossified male, the leading descrip-
tion has been written from Arctocephalus australis.
Spine. — The vertebral formula was C 7, D 15, L 5, S 3, Cd 11 =41.
The cervicals had, as a rule, a foramen at the root of each transverse process. The
7th cervical was, however, peculiar, for in the adult male from the Messier Channel each
of its transverse processes was perforated by a small foramen, but in the other three
skeletons from the same place there was no foramen. In both skeletons of Arctocephalus
gazella the right transverse process was perforated but not the left. In the atlas the
transverse process was broad, plate-like and elongated downwards and outwards ; in the
axis it was much shorter and styloid ; in the 3rd cervical the inferior lamella was
flattened out into a plate which increased in magnitude in the other cervical vertebrae
down to the 6th ; in the 7th this lamella was absent except in those specimens in
which the transverse process was perforated, when it was a thin, horizontal plate of
44 THE VOYAGE OF H.M.S. CHALLENGER.
bone. The spine of the atlas was rudimentary ; in the axis it was massive ; in the
3rd cervical it was short and it gradually increased in length to the 7th cervical.
A strong process, distinct from the transverse process, projected backwards for nearly
half an inch from the pedicle of the 7th cervical immediately below the articular process,
and indications of a simdar process were seen in both the 6th and 5th. The bodies of
the cervicals were mesially keeled on the ventral surface, and in the 7th the keel was
elongated into a plate-like hypapophysis, which was very projecting in the adult Messier
Channel male.
The dorsal vertebra} had relatively short transverse processes, which were the
longest in the more anterior dorsals, but diminished in length in the posterior dorsals
so as to be scarcely recognisable in the 14th and 15th. Anapophyses were present
in all the dorsals ; in the more posterior they were elongated and styloid, in the middle
and more anterior they were stunted and tubercular, and in series with the strong back-
ward-projecting process from the pedicle of the 7th cervical. Metapophyses projecting
forwards from the anterior articular processes were especially seen in the middle and pos-
terior dorsals. The spines projected backwards ; they were elongated and strong in the
anterior vertebra?, but gradually diminished in size from before backwards. The bodies of
the more anterior and more posterior dorsal vertebrae were keeled on the ventral surface.
The lumbar vertebrae had transverse processes which projected downwards, outwards,
and forwards; in the 1st lumbar they were very short, but they increased in length
to the 4th. The spines were short and hatchet-shaped. Anapophyses had almost
entirely disappeared ; metapophyses were present in all of these vertebras. The ventral
surface of the bodies was keeled.
Sacrum. — In the adult male from the Messier Channel the 1st and 2nd sacral
vertebrae were completely fused together, the lamina? of the 2nd were fused with those
of the 3rd, but the spines were not, and between their bodies the intervertebral disc was
partially present. The 3rd sacral vertebra was fused with the 1st caudal by the pair
of processes situated below the articular processes. The greatest breadth of the base
of the 1st sacral vertebra was 48 mm. and the length of its body was 32 mm. The
2nd and 3rd were smaller bones, especially in breadth ; they were all fully ossified
In Arctocephalus gazella fusion of the sacral vertebrae had not taken place.
The caudal vertebra? diminished in size from the 1st to the 11th. The 1st, 2nd,
and 3rd possessed laminae and spines, the 4th had a neural groove, but the rest consisted
only of the bodies, and the two terminal segments of the tail were only 6 and 4 mm.
long respectively.
Ribs. — There were fifteen pairs of ribs, of which nine articulated with the sides of
the sternum. The head of the 1st rib articulated only with the body of the 1st dorsal
vertebra, but from the 2nd rib backwards to the 12th, both inclusive, the head articulated
with the bodies of two vertebrae. The three most posterior ribs again articulated, each
REPORT ON THE SEALS. 45
only with a single vertebra. The osseous part of the 1st rib was about the same length
as its costal cartilage, but in the other ribs the bony part considerably exceeded the
cartilaginous. The 10th and succeeding pairs had more slender cartilages than those
which were anterior to them. In Arctocephalus gazella the arrangement was the
same, but the bones were more slender.
Sternum. — This bone consisted of eight segments, capable of being separated from
each other. The 1st was 125 mm. long, and 29 mm. in its widest part ; it was formed
of bone in its whole length, and consisted apparently of the pnesternal segment fused
with the most anterior segment of the meso-sternum, for it projected forwards to the
neck, and the 1st pair of ribs was articulated to it 90 mm. behind its anterior
end. It was an elongated bar of bone, and possessed three surfaces and a ventral
keel. The cartilages of the 2nd to the 7th ribs, both inclusive, were articulated at the
junctions of the segments of the meso-sternum. The 8th pair of cartilages were
articulated to the sides of the 7th segment, and the 9th pair at the junction of the
7th and xiphisternal segments. The segments of the meso-sternum were all elongated,
and the 7th was expanded in the plane of articulation of the 8th costal cartilages. The
xiphisternum projected behind the 9th pair of costal cartilages, it was bone in its more
anterior two-thirds, but the posterior third was a leaf-like plate of cartilage. The
sternum of Arctocephalus gazella closely resembled Arctocephalus australis.
Anterior Extremity. — The scapula in the Messier Channel adult male measured
205 mm. from glenoid fossa to vertebral border, and 253 mm. from the pointed posterior
to the rounded anterior angle. The spine was more distinct than in Macrorhinus
and Leptonychotes, especially than in the latter, and it ended below in a very feeble
acromion. The praespinous fossa was more than twice as large as the postspinous, and
was imperfectly divided into two almost equal areas by a ridge almost parallel to the
spine, and situated about halfway between the spine and the rounded auterior angle.
A strong ridge behind the spine, which was much more projecting than in Mac-
rorhinus and Leptonychotes, was for the origin of the third and fourth heads of the
triceps muscle. The axillary border was falciform and the coracoid process was very
feeble. The ventral surface was concave, but marked by ridges for the tendons of
origin of the subscapularis. In Arctocephalus gazella and the younger animals from
the Messier Channel a portion of the suprascapular cartilage was still unossihed.
The humerus had an extreme length of 185 mm. The deltoid ridge was strong
and rough, with its outer border everted, and terminated above in a strong external
tuberosity, which was separated from the inner tuberosity by a deep bicipital groove.
The outer condyloid ridge projected much more than the inner, but the inner condyloid
eminence wTas more prominent than the outer. The capitellum and trochlea formed a
continuous articular surface, and both the radial and olecranoid fossse were shallow.
The musculo-spiral groove was not very strongly marked, but the shaft was concave
46 THE VOYAGE OF H.M.S. CHALLENGE!;.
vertically on the outer side of the deltoid ridge, which was not the case in Macrorhinus.
In Arctocephalus gazella the humerus was 156 mm. long. The epiphysis of the head
was conjoined with that of the greater tuberosity, but that for the lesser tuberosity
was distinct. The epiphysis for the inner condyle was distinct from that for the
radio-ulnar articular surface. Neither species possessed a supracondyloid foramen.
The radius and ulna had the characteristic general shape and relative position
customary in the seals. The radius had a strong ridge on its anterior border for the
pronator teres, its lower end was grooved on the dorsal aspect for the extensor tendons,
and immediately below the neck was a low bicipital tuberosity. The olecranon and
upper half of the shaft of the ulna were distinctly grooved both on the dorsal and
palmar surfaces ; that on the dorsal surface was divided into two unequal parts by a
longitudinal ridge which commenced in a tubercle at the free edge of the olecranon.
Another tubercle was situated at the anterior end of the same border of the olecranon,
and a third at the posterior end. To these parts the triceps and dorsi-ejjitroehlear
muscles were attached. The interosseous interval was narrow. The radius articulated
at its lower end with the ulna, scapholunar, and cuneiform ; the ulna with the radius,
cuneiform, and pisiform. The radius in the adult male was 182 mm. long,, the ulna
224 mm. In Arctocephalus gazella they were similar in shape but smaller ; the
radius was 152 mm., the ulna 191 mm. long, and the epiphyses were not ankylosed.
Manus. — There were seven carpal bones. The scapholunar was large and trans-
versely elongated ; it articulated with the radius, cuneiform, and the four bones of the
distal row. The cuneiform was elongated in the dorsi-palmar diameter and articulated
with the radius, ulna, pisiform, scapholunar, unciform, and 5th metacarpal. The
pisiform was elongated for 21 mm., and projected inwards from the lower end of the
ulna, with which and the cuneiform it articulated. The trapezium, though much
smaller than the scapholunar, was next to it in size ; it articulated with it and with the
trapezoid and 1st and 2nd metacarpals. The trapezoid was wedged in between the
trapezium and magnum, and articulated with them and with the scajmolunar and 2nd
metacarpal. The os magnum was a comparatively small bone : it articulated with the
3rd metacarpal, trapezoid, unciform, and scapholunar. The unciform, as in the other
seals described, did not reach the inner border of the carpus, owing to the articu-
lation of the 5th metacarpal with the cuneiform ; it articulated with the 4th and 5th
metacarpal, cuneiform, scapholunar, and magnum. All the carpalia were rough on the
palmar and dorsal surfaces for the attachment of ligaments, and only the scapholunar
had a process, and that a low one, projecting from the inner side of its palmar surface.
There were five digits, which diminished in length from the pollex to the minimus.
The three segments of the pollex measured collectively 241 mm., they were not only
longer than the corresponding bones of the other digits but they were broader and
stronger. The metacarpal of the pollex, 110 mm. long, was flattened at its carpal end,
REPORT ON THE SEALS. 47
and articulated with the trapezium and second metacarpal. The 2nd metacarpal
articulated with the 1st and 3rd, and with the trapezium and trapezoid. The 3rd
metacarpal articulated with the 2nd and 4th and with the os magnum. The 4th
metacarpal articulated with the 3rd and 5th and with the unciform. The 5th meta-
carpal was only 56 mm. long, and was flattened, unlike the metacarpals of the other
fingers ; it articulated with the 4th metacarpal, the unciform, and cuneiform. The
metacarpal bone and phalanges of the minimus collectively measured 117 mm. The
nails were rudimentary, and the terminal phalanges ended abruptly and without an
ungual process. The skin, longitudinally wrinkled and without hairs, was prolonged
beyond the terminal phalanx, and in the pollex this cutaneous fold was 100 mm. long and
45 mm. broad.
In Arctocephalus gazella the carpalia corresponded in number, shape, and arrange-
ment to the bones in Arctocephalus australis, but the}7 were smaller. The bones of
the digits were also similar, but on a somewhat smaller scale. In both species the 2nd
phalanx of the minimus though wider was scarcely so long as the terminal phalanx.
Both in Arctocephalus gazella and the younger specimens of the Messier Channel
Arctocephalus, it was seen that the ossification of the phalanges and metacarpal bones
was on the same plan as in Macrorhinus and Leptonychotes. In Arctocepjhalus
gazella the length of the pollex and minimus was 178 and 89 mm. respectively.
Pelvis. — This division of the skeleton consisted of the three sacral vertebras and the
two innominate bones. The length of the os innominatum was 210 mm., that of the
ilium 85 mm., and of the ischio-pubic part 125 mm. The ilium was more elongated than
is usual in the seals ; its dorsal surface was three times broader than the ventral, which
was 1 1 mm. broad, and was bounded externally by a rough surface for the origin of the
rectus ; the inner surface was as usual articular for the side of the sacrum, but the crest
of the bone instead of being in almost the same transverse plane as the base of the
sacrum, as in Macrorhinus and Leptonychotes, projected forwards, so that it was 31 mm.
in front of the base of the sacrum. Between the crest and the base of the sacrum the
inner surface of the ilium was marked for the origin of a muscle, probably the multifidus
spina?. The acetabulum had a complete covering of cartilage immediately within the
brim, but at the bottom of the cup was a narrow rough depression which opened at the
back of the brim in a small cotyloid notch or foramen. The margin of the brim was
complete in bone, as the cotyloid notch was bridged by a bony bar which almost
converted the notch into a foramen for the passage of the vessels and nerves into the
joint. The os pubis and ischium were slender bars of bone, and the symphysis was
limited to the junction of the two pubic bones. The junction of the os pubis and ilium
was marked by a very prominent pectineal tubercle, and the pectineal line was sharp.
The ischium had neither definite tuberosity nor spine, and the obturator foramen was
elongated.
48 THE VOYAGE OF H.M.S. CHALLENGER.
Posterior Extremity. — The extreme length of the femur in the adult male from the
Messier Channel (jSTo. 3) was 112 mm. The head was smooth and without a fossa for
the ligamentum teres ; the neck was stunted, the great trochanter was well marked,
and with a shallow digital fossa for the obturator muscle ; the small trochanter was a
distinct tubercle, of no great size ; there was a faint posterior intertrochanteric line,
a very feeble anterior intertrochanteric line, but no trochanter tertius. The shaft was
broadened laterally, though not so much as in Leptonychotes and Macrorhinus ; its
inner lateral border was almost straight, its outer lateral border slightly concave,
differing therefore from the other two genera, in which, more especially in Leptonychotes,
these borders were markedly concave ; though the posterior surface was flat the
anterior was slightly convex. A rough ridge passed down the inner border of the bone
towards the posterior surface, which apparently represented a linea aspera. The external
tuberosity was more prominent than the internal, and they were both grooved ; the
outer groove being for the tendon of the popliteus. The patellar articular surface
was not hollowed from side to side, and was convex from above downwards ; it was
so large as to occupy almost the whole of the front of the lower end of the femur,
whilst in Macrorhinus and Leptonychotes it occupied only about the middle third of
that part of the bone. This surface was prolonged downwards and backwards so as to
be continuous with the internal condylar articular surface, but was separated from the
external condylar surface by a narrow groove ; the condylar surfaces were situated on
the back of the lower end of the bone, and were separated from each other by an
intercondylar fossa. The inner condylar surface was concave from side to side, whilst
the outer was convex. In Arctocephalus gazella the femur was only 93 mm. long, and
much more slender than in the Messier Channel seal, and this indeed was a character
which distinguished all the bones of the hind limb ; the trochanter minor was absent
even in the most fully ossified of the two skeletons, in which the head was united to
the neck of the bone. The patellar articular surface was continuous with the inner
condylar articular surface, but was separated from the outer by a narrow groove, to
which the adipose ligament was attached as well as to the intercondylar fossa. The
outer border of the shaft of the femur was more concave in Arctocephalus gazella than
in Arctocephalus australis.
The patella was 28 mm. long, and in both species of Arctocephalus had an articular
surface transversely elongated and slightly concave from above downwards ; its cutaneous
surface was elongated superiorly into a strong tubercle, whilst lower down the bone
was flattened, so that whilst the upper end of the bone was 18 mm. in antero-posterior
thickness, the lower end was only 10 mm.
The tibia was 235 mm. long in the Messier Channel adult male. Its superior surface
was divided into two articular facets with a rough groove between them, and the outer
facet was wider than the inner. The ligamentum patella? was attached to the front
REPORT ON THE SEALS. 49
of the bone immediately below the condylar surface. The shaft of the tibia was
divided into three surfaces in its upper part, but in the lower half the ventral and outer
surfaces were not so sharply differentiated from each other. The lower end of the
inner and posterior surfaces of the tibia had two longitudinal grooves, the inner for the
tibialis posticus being both wider and deeper than the outer for the flexor longus
liallucis. On its inner side a short malleolus projected downwards. The tibia articu-
lated below with the fibula and the upper surface of the astragalus. The tibia differed
from that of Leptonychotes in having a much less transverse diameter at the condylar
end, in not possessing a definite ridge on the shaft for the gracilis, in not being so
distinctly grooved at the lower end in front for the tibialis anticus, and in being more
deeply grooved behind for the tibialis posticus and flexor longus hallucis.
The fibula was a slender bone, 213 mm. long. Its upper end was fused with the
outer tuberosity of the tibia. The shaft was three-sided all the way down. A short
malleolus projected from the lower end which articulated by movable joints both with
the tibia and the external lateral surface of the astragalus, and was grooved externally
for the peronei ; it did not articulate with the os calcis, and the malleolus generally was
much less bulky than in Leptonychotes. In Arctocephalus gazclla the tibia was 180
mm. and the fibula 163 mm. long, and the epiphyses were not ankylosed.
Pes. — The tarsus contained eight bones. The astragalus possessed a trochlear
surface superiorly, which articulated with the lower end of the tibia ; internally it did
not articulate with the tibia, and externally it had a broad surface for the external
malleolus of the fibula, which looked forwards as well as outwards, but was not however
relatively so large as in Macrorhinus and Leptonychotes; anteriorly it had a convex
head for the scaphoid, immediately external to which was a narrow articular surface
for the cuboid ; inferiorly its articular surface was divided into two parts, separated
by an interosseous ligament, for the os calcis ; the posterior surface was narrow and
grooved, and not prolonged into a calcanear process. The extreme length of the
astragalus was 40 mm.
The os calcis was elongated behind into a strong calcanear process, which was
grooved posteriorly for the tendon of the plantaris. Its outer surface was also marked
by the peronei tendons, the position of which was expressed by a strong tubercle and
by two grooves ; superiorly it articulated with the astragalus, and anteriorly with the
cuboid. The extreme length of the os calcis was 54 mm.
The cuboid had both a plantar ridge and a peroneal groove. It articulated behind
with the os calcis, internally with the scaphoid and external cuneiform, anteriorly with
the 4th and 5 th metatarsals.
The scaphoid had the usual shape of the bone, but without a tubercle ; it articu-
lated behind with the astragalus, externally with the cuboid, internally with the ento-
scaphoid bone, anteriorly by three very distinct facets with the cuneiforms.
(zool. chall. exp. — part Lxvin. — 1888.) Yyy 7
50 THE VOYAGE OF H.M.S. CHALLENGER.
Of the cuneiforms the ento- was as usual the largest and the meso- the smallest.
The ento-cuneiform articulated behind with the scaphoid, internally with the eighth
tarsal bone, anteriorly with the 1st metatarsal, externally with the 2nd metatarsal and the
ccto-cuneiform. The meso-cuneiform was visible both on the dorsal and plantar surfaces
of the foot ; it articulated laterally with the other cuneiforms, behind with the scaphoid,
in front with the 2nd metatarsal. The ecto-cuneiform was also visible on both surfaces
of the foot; externally it articulated with the cuboid, internally with the meso-cuneiform
and 2nd metatarsal, behind with the scaphoid, and anteriorly with the 3rd and very
slightly with the 4th metatarsals.
The eighth bone of the tarsus, or entoscaphoid, was in the adult Arctocephalus
australis 21 mm. long by 14 mm. broad, situated internally, and articulating by distinct
facets with the inner surfaces of both the scaphoid and ento-cuneiform bones towards
the plantar aspect. It was present in the other skeletons from the Messier Channel, and
in both the skeletons of Arctocephalus gazetta. Obviously therefore it is a constant
bone in this genus.1 This additional bone represents, I believe, the tubercle of the
scaphoid which has remained as a separate ossicle. Occasionally in man the tubercle of
the scaphoid ossifies as a distinct ossicle in connection with the tendon of the tibialis
posticus, to which it seems to have the relation of a sesamoid bone, and some years ago
I described in an adult a specimen of this kind.2 Karl Bardeleben has also stated3 that
in the human embryo at the second month there is a special cartilage, corresponding to
the tuberosity of the scaphoid, which he regards as homologous with the scaphoid bone
of the carpus. In a youth of 1 5 years he had once seen it as a separate ossicle.
The toes were of almost equal length as regards their skeleton, but the integument
was prolonged in a variable extent from 105 to 110 mm. beyond the terminal phalanx.
Each toe had an elongated convex nad on the dorsum of the last phalanx, but the
integument was not haired. The three segments of the hallux measured collectively
237 mm. Each was longer than the corresponding segments in the other digits. The
1st metatarsal articulated by much the greater part of its proximal end with the ento-
cuneiform and very slightly with the 2nd metatarsal ; it was the longest and the most
1 Dr. Murie says that Otaria jubata possesses the normal number of tarsal bones, and he figures the seven tarsalia.
In a skeleton of the Grey Sea Lion of Australia, Eumetopias cinereus, I found the eighth tarsal bone occupying a position
similar to what I have described in Arctocephalus, and articulating with both scaphoid and ento-cuneiform. In a young
Arctocephalus gazella dissected by Dr. Miller, the entoscaphoid was still cartilaginous, and received a large part of the
tendon of the tibialis posticus. In Macrorhinus there was no separate entoscaphoid, and the tendon of the tibialis
posticus was inserted into a thick plate of cartilage continuous with the scaphoid bone in the region of the tubercle. In
both a young and adult Phoca vitulina dissected by Dr. Miller, the tendon of the tibialis posticus was inserted into the
tubercle of the scaphoid, but a strong slip passed distally from that tendon to end in an ossicle which in the adult was
13 mm. long and 8 mm. broad. This ossicle was situated internally to the ento-cuneiform, but had not a definite
faceted articulation either with that bone or with the scaphoid as was seen in Arctocephalus. In a young Walrus, the
ossicle was represented by a cartilaginous nodule intimately connected with the tendon of the tibialis posticus, and having
a distinct facet for articulation with the ento-cuneiform.
- Joum. of Anat. and Phys., October 1882, vol. xvii. p. 82.
3 Joum. of Anat. and Phys., July 1885, vol. xix. p. 510.
REPORT ON THE SEALS. 51
massive of all the bones of the toes. The 2nd metatarsal articulated with the 1st
and 3rd and with the three cuneiforms ; it was bent a little inwards, much less than
in the Elephant Seal, in order to pass slightly behind the 1st metatarsal. The 3rd
metatarsal articulated with the 2nd and 4 th and with the ecto-cuneiform ; it was the
shortest of the metatarsals. The 4th metatarsal articulated with the 3rd and 5th and
with the ecto-cuneiform and the cuboid, it was not hollowed out on the outer side of the
shaft as in Macrorhinus and Leptonychotes. The 5th metatarsal was next in length to
the first, but much less broad; it possessed at its tarsal end a peroneal tubercle and
articulated with the 4th metatarsal and the cuboid. The terminal phalanx of the 2nd,
3rd, and 4th toes was prolonged into a pointed ungual process on which the nail rested,
but the corresponding phalanx of the 1st and 5 th digits had no such process, and the
nails wTere smaller than in the other toes. A pair of sesamoids was situated on the
plantar surface of each metatarso-phalangeal joint. In the adult male all the epiphyses
were fused with the diaphyses, but in the young male the epiphyses were seen to have
a similar arrangement to those described in the metatarsals and phalanges of the
Elephant and Weddell's Seals.
The tarsalia in Arctocephalus gazella corresponded in number, form, and arrangement
to those of A rctocephalus australis. The bones of the digits were also similar, but more
slender, and the epiphyses as in the young male of the other species were not ankylosed.
Nearly twenty years ago the late Professor Alleu Thomson described : the ossifica-
tion of the digits in the common seal, Phoca vitulina. In the manus he said that the
1st metacarpal bone and all the digital phalanges, except the terminal phalanx, each
possessed only a proximal epiphysis, whilst in the four other metacarpal bones there were
only distal epiphyses. In the pes again, the 1st metatarsal bone had both a proximal
and a distal epiphysis like the phalanges generally, except that the terminal phalanx had
only a proximal epiphysis ; the four other metatarsals had each only a distal epiphysis.
In the year 1869, the late Mr. A. B. Stirling prepared and mounted, in the Anatomical
Museum of the University of Edinburgh, specimens to illustrate this method of ossification
of these bones. The description which has been given in this Report of the manus and
pes of Macrorhinus, Leptonychotes, and Arctocep>halus, shows that in them the bones
of the digits of the manus do not ossify in the same manner as in the manus of Phoca
vitulina, but that in all these three genera the digits both of the manus and pes ossify
after the same plan, which corresponds with that seen in the pes of the common seal.
The length of the spine of the adult male Fur-Seal from Messier Channel was 1490
mm., the dried intervertebral discs being included, and that of the skull of the same
animal was 233 mm., giving a total length of 1723 mm. or 5 ft. 6 in. The length of
the spine of the adult female, including the dried intervertebral discs but exclusive of
the six terminal caudal vertebrae which were missing, was 1100 mm., and the length of
1 Journ. of Anat. and Phys., November 1868, p. 140.
52
THE VOYAGE OF H.M.S. CHALLENGER.
the skull was 202 mm.; if 90 mm. be allowed for the missing vertebrae then the combined
leno-th of skull and spine would be 1392 mm. or 4 ft. 6 in. The length of the spine,
including the dried intervertebral discs, of the larger Kerguelen Island Fur-Seal was
1110 mm., that of the skull was 212 mm., giving a total length of 1322 mm.
or 4 ft. 3 in. Mr. H. E. Elliott in his account of the Northern Fur-Seal of the Pribylov
Islands (Callorhinus ttrsinus) states1 that the male, when fully mature and fat, weighs
on an average from 400 to 500 pounds, while the female gives a mean of from 80 to 85
pounds. Mr. Allen gives the length of the skeleton, including the skull of an adult
male ursinus, as 2040 mm. (6 ft. 7 in.); and of another as 1840 mm. (6 ft. ^ in.); whilst
one adult female measured 1370 mm. (4 ft. 5 in.); and another 1215 mm. (3 ft. 10 in.).
It would appear therefore that the Northern Fur-Seal, especially the male, is a
somewhat longer and probably more bulky animal than the Fur-Seal which frequents
the southern half of the South American continent.
The length-breadth indices of five male skulls of Arctocephalus australis,
calculated on the width behind the external meatus, were respectively 55 -8, 59, 50,
58-9, and 51, and of the female 52-9 ; but at the interzygomatic width the indices for
three males were respectively 63"5, 57, and 59, and for the female 57'4. The length-
breadth indices of the skulls from Kerguelen Island, calculated on the width behind the
external meatus, were respectively 56'6 and 51-6, and on the interzygomatic width
617 and 56-8 respectively.
Arctocephalus, sp. incerta.
Fur- Seal from Juan Fernandez.
This specimen from its size was obviously a very young pup. It was given alive to
Sir Wyville Thomson at Juan Fernandez, but died after being a few days on board ship.
The animal was preserved in strong spirit. Its principal dimensions are as follows : —
Table VIII. — Fur-Seal from Juan Fernandez.
Ft. In.
From snout to tip of tail,
1 7i
From snout to longest digit of pes,
1 10
Length of free part of tail,
11
Length of pectoral limb,
6f
Greatest breadth of that limb, .
.
2i
Length of hind limb from root of tail,
,
H
Greatest breadth of that limb, .
H
From root of pectoral limb to angle of mouth,
n
1 The Seal Islands of Alaska, a memoir issued by the Department of the Interior, Washington, 1881 ; and in his
buok, An Arctic Province, Alaska and the Seal Islands, London, 1886.
REPORT ON THE SEALS. 53
The skin of the body was covered with soft jet-black hairs, which were present also on
the dorsum both of the raanus and pes. The shape and general character of the manus
closely corresponded to the South American Fur-Seal. The pes also had generally the
same character, but the nails were in relation to the size of the foot stronger than in that
seal, especially the nails on the hallux and minimus. Some of the bristles in the upj^er
lip were dark brown, others black. The external ear was 1 inch long, pointed at the tip,
and with black hairs on the dorsum.
Mr. Elliott says that the pup of Ccdlorhinus ursinus at birth, and for three months
after, is of a jet-black colour both in the hair and nippers, save a tiny white j)atch at the
back of each forearm ; that it weighs at birth from 3 to 4 pounds, and is 12 to 14 inches
long. The jet-black colour of the hair in the foetus and at the time of birth seems to be a
character of the Fur-Seal. In Phoca vitulina it has been observed that the intra-uterine
hair is yellowish-white and woolly, and it is shed either in utero or immediately after
birth.1 Some years ago I showed that the intra-uterine hair of the foetus of Halichcerus
grypus 2 was yellowish-fawn colour and streaked with dark grey bands and spots, but
that it was neither woolly nor fur-like. The foetal hair is shed within about a month
after the animal is born.
The skull was 130 mm. long, 73 mm. in its zygomatic diameter, and 81 mm. at the
widest part of the cranial box. The orbits were immediately in front of the anterior part
of the cranial box, so that the orbital process of the malar was close to the anterior wall
of the cranium, and the zygomatic part of the temporal did not turn up behind the orbital
process of the malar. The frontal bone passed between the hinder ends of the nasals.
The ascending process of the premaxilla articulated with the anterior third of the outer
border of the nasal. The anterior and postorbital processes were small. The anterior
nares opened well in front of the antorbital process and infraorbital foramen. The basi-
occipital was not perforated. The occipital condyles were not continuous with each other,
and alisphenoid canals were present. Each tympanic bulla had a low antero-posterior
ridge.
The hard palate was emarginate posteriorly and ended in line with about the middle
of the zygomatic arch and the orbital process of the frontal bone. The posterior edge of
the vomer sloped very obliquely forwards and was not seen at the posterior nares, though
the joint between the prse- and post-sphenoids was well behind the posterior edge of the
palate. The skull showed the following dentition, and the teeth which had erupted or
3 3 , i i . 5 5
were just appearing were : — incisors \ — , canines - — , post-canines - — - = o2. The
canines and post-canines were small and unifanged. In the lower jaw the coronoid process
was expanded and there was a distinct quadrilateral and inflected subcondyloid tubercle.
1 Wright, Forhandl. vid de Skandin. Naturforsk. i Stockholm, 1842 (1843), abstract in Mailer's Archiv f. Anat.
u. Phys., 1844.
2 Memoir on the Placentation of the Seals, Trans. Roy. Soc. Edin., June 1875, vol. xxvii.
54 THE VOYAGE OF H.M.S. CHALLENGE!!.
Some years ago Dr. Philippi of Santiago obtained specimens of a Fur-Seal from Juan
Fernandez, which Professor Peters examined and named Arctocephalus philippii.1 He
regarded it as the same as the seal which had been named Arctocephalus (Otaria)
argentatus. It is not unlikely that the pup which I have described is the young of this
animal, though whether it should be regarded as a distinct species, or as of the same
species as Arctocephalus australis, is a matter which perhaps can scarcely as yet be said
to be definitely decided.
1 Monatiber. d. k. preuss. Alad. d. Wiss. Berlin, November 9, 1871 ; June 10, 1375; August 9, 1877.
PART II.
CLASSIFICATION OF THE PINNIPEDIA.
In the course of the study of the species of Seals described in the preceding part of
this Report, I have been led to examine and compare, so far as the material at my
disposal would admit, the skeletons of other species of Seals, belonging both to the
Earless and Eared families, and also that of the Walrus. This examination and compari-
son have enabled me to recognise several anatomical characters, which either have had
too little attention paid to them, or have been altogether overlooked, and which are of
undoubted value in assisting one to discriminate between the different genera and species.
I propose in this part of the Report to give a classification of the Pinnipedia, and to
introduce into it, along with the characters usually recognised, those additional ones
which seem to me to be of value in taxonomy.
Starting from the usually accepted position that the Pinnipedia are a suborder of
the Carnivora, which suborder is divided into three families, Phocidse, Trichechidas or
Odobaenidse, and Otariidse or Arctocephalidas,1 I shall first state briefly the distinguish-
ing characters of these families, and then subdivide them into their subfamilies, genera,
and species.
Phocida
Without pinna of ear and scrotum ; postorbital processes either wanting or
rudimentary ; no alisphenoid canal ; mastoid moderate and not entirely discontinuous
from the tympanic bulla ; nasal bones elongated backwards between the two halves of
the frontal. Inner wall of orbit complete or almost complete. Hard palate the widest
opposite or a little behind the last pair of molars and almost in line with the hinder
i See as authorities on the subdivision of the Pinnipedia, H. N. Turner, Proc. Zool. Soc. Land., 1848 ; Gill,
Proc. Essex Inst., vol. v., 1866, and Families of Mammalia, 1872 ; J. E. Gray, Cat. Seals and Whales, 1866 and
Supplement ; W. H. Flower, Proc. Zool. Soc. Lond., 'January 14, 1869, and article Mammalia in Ency. P.rit., 9th Ed.;
J. A. Allen, North American Pinnipeds, 1880 ; and St. George Mivart, Proc. Zool. Hoc. Lond., May 19, 1885.
56 THE VOYAGE OF H.M.S. CHALLENGER.
edge of the maxillary root of the zygoma, from which spot the palate diminishes in
transverse diameter both forwards and backwards ; posterior border either truncated or
emarginate. Articulation of vomer with floor of the nose variable in position. Carotid
canal distinct from and antero-external to the foramen lacerum posterius. Occipital
condyles converging anteriorly, their articular surfaces frequently continuous ; basi-
occipital flattened and usually perforated mesially. Anterior nares far from terminal.
Malar variable in its length. Neck short. Hind limbs directed backwards, not used in land
locomotion. Front limbs smaller than posterior, digits armed with strong terminal claws.
1-1 . 5-5
Palms and soles hairy. Dentition — incisors variable, canines f3j. post-canines g— g.
Mammas 2 or 4. Scapula with prse- and postspinous fossss almost equal in area.
Humerus with or without supra-condyloid foramen. Ilium with crest much everted and
in transverse plane of base of sacrum. Femur with small trochanter absent or rudi-
mentary. Astragalus with a long calcanear process.
Trichechid^e.
Without pinna of ear and scrotum ; postorbital processes either wanting or rudi-
mentary ; alisphenoid canal large ; mastoid massive but not entirely discontinuous from
the tympanic bulla ; posterior border of nasal articulating with anterior border of frontal
almost in the transverse plane of skull. Inner wall of orbit complete, but that of
zygomatic fossa defective. Lateral borders of hard palate almost parallel to each
other behind, though somewhat converging anteriorly ; posterior border truncated.
Vomer articulating with floor of nose well in front of hinder border of palate. Carotid
canal distinct from and in front of foramen lacerum posterius. Basi-occipital mesially
keeled and not perforated. Anterior nares terminal. Malar short. Neck longer than
in Phocidse. Palms and soles hairless ; manus and pes can be turned forwards and used
for land locomotion. Upper canines 1-1, elongated into great tusks. Mammas 4.
Scapula with prsespinous much larger than postspinous fossa. Humerus without a
supra-condyloid foramen. Ilium with crest everted and in front of base of sacrum.
Femur with rudimentary small trochanter. Astragalus without a calcanear process.
Otariid^e.
With pinna of ear and scrotum ; postorbital processes distinct ; alisphenoid canal
present ; mastoid quite discontinuous from tympanic bulla and prominent ; tympanic
bullae only slightly swollen ; anterior mesial process of frontal passing between the
diverging posterior ends of the nasals. Inner wall of the orbit very defective, so that
REPORT ON THE SEALS. 57
vomer can be seen through the opening. Hard palate either truncated or emarginate.
Malar very elongated and reaching back almost to glenoid fossa. Vomer articulating on
floor of nose only with crests of superior maxillte. Carotid canal opens either within
boundary of foramen lacerum posterius or immediately anterior to it. Basi-occipital
mesially keeled and seldom perforated. Anterior nares not quite terminal. Neck of
considerable length. Palms and soles hairless, used in land locomotion, and manus and
pes capable of being turned forwards. Three middle digits of pes short and feeble
compared with pollex and minimus, and with well-developed nails. Outer upper
incisor caniniform ; inner upper incisors with anterior and posterior cusps. Pernia-
• 3-3 1-1 5-5 6-6
nent dentition — m. 9-^, c. j^j, p- c. g^-g or - — ; = 34 or 36. Mammae 4. Scapula
with praespinous very much larger than postspinous fossa. Humerus without a supra-
condyloid foramen. Ilium with crest slightly everted and well in front of base of
sacrum. Femur with small trochanter usually present. Astragalus without a calcanear
process.
Phocidj:.
The family Phocidas has been divided into the following subfamilies — Phocinaa,
Ogmorhininaa, and Cystophorinae.1
PHOCIN.E.
Anterior nares oblique and in front of infraorbital foramen ; beak but little prolonged
in front of opening ; no postorbital process ; interorbital and interzygomatic parts of
frontal greatly compressed laterally ; horizontal part of premaxilla thin ; widest part of
hard palate behind molar teeth, and in line with hinder edge of maxillary root of zygoma,
from which spot the palate diminishes in transverse diameter both forwards and
backwards. Zygomatic process of maxilla not much prolonged back below malar.
Inner wall of orbit entire or almost entire. Tympanic bullae swollen. Pterygoids
3-3 1-1 5-5
vertical in direction. Dentition — in. ^—5, c. t~tt, p. c. g— r = 34. Mandible with both
a subcondyloid process and an angle. Nails strong in all the digits. Toes of hind
foot almost equal in length. Humerus with supracondyloid foramen.
This subfamily contains the genera Phoca and Halichcerus.
1 For a number of years I have lost no opportunities of collecting specimens of the crania of the Seals of the North
Atlantic for the Anatomical Museum of the University of Edinburgh, and from the courtesy of several gentlemen, more
especially the late Mr. Charles Edward Smith, Dr. A. J. M. Bentley, Dr. James Foulis, Captain M 'Donald, the Rev.
Dr. Joass, Dr. W. Stewart Campbell, Dr. W. Livesay, and Mr. Charles A. Anderson, the collection is now very
complete.
(ZOOL. CHALL. EXP. PART LXVIII. — 1888.) YjV 8
58 THE VOYAGE OF H.M.S. CHALLENGER.
Phoca.
Callocephale, F. Cuvier, Mem. du Museum, t. xi. p. 182, 1821
Infraorbital foramen opens into anterior part of floor of orbit, which slopes backwards
to become continuous with the thin posterior border of zygomatic root of maxilla.
Anterior nares not high, post-canines with more than a single cusp on the crown, mostly
three-cuspidate ; fangs two-rooted except in the first post-canine.
Phoca vitulina, Linnseus. Common Harbour Seal.1 North Atlantic Ocean.
Phoca vitulina, Linn., Syst. Nat., ed. x. p. 38.
Callocephalus vitulinus, Gray, Brit. Mus. Catal., p. 20, 1866.
Nasal bones elongated and attenuated between frontals, not ankylosed together ;
premaxilla forming side of anterior nares but usually not quite reaching the nasal ;
neither antorbital nor postorbital processes ; occipital ridge fairly marked ; sagittal
ridge indicated ; no ridge along line of sagittal suture. Hard palate with posterior
border acutely cleft, the sides of cleft forming with line drawn between hamular-
pterygoids almost an equilateral triangle; the apex of the cleft a little behind the
posterior edge of the maxillary root of the zygomatic arch and considerably in front of
the hamular-pterygoids ; posterior border of vomer visible _ in cleft and articulating with
vomerine crest ascending from the upper surface of the anterior part of the horizontal
plate of the palate bone. Zygomatic arches bulging, and their greatest width, which is
about the middle, considerably exceeding the greatest cranial width. Post-canines set
closely together and obliquely, so that the posterior part of the tooth in front is external
to the anterior part of the tooth immediately succeeding. Subcondyloid process of
posterior border of ramus of mandible short, inverted, distinct from the tubercle at the
angle of the bone ; lower border of horizontal ramus scarcely inverted ; coronoid
moderate.
Phoca grcenlandica, Fabricius. Harp Seal. North Atlantic2 and Arctic Oceans.
Phoca grcenlandica, Fabricius in Muller, Zocl Dan. Prodr., p. viii., 1776, and Fauna groen-
landica, p. 11, 1780.
Nasal bones elongated and attenuated between frontals, not ankylosed together.
Premaxilla forming side of anterior nares and articulating with about one-third of outer
border of nasal but not reaching its tip. Anterior nares oblique, but the lateral borders
more concave near the nasal bones than in Phoca vitulina, Phoca hispida, and Phoca
1 In writing the descriptions of the characters of the skulls of the several species of Seals I have as far as possible
selected those of adult males.
2 In November 1874 I recorded (Journ. Anat. and Phys., vol. ix. p. 163) the capture of a Seal on the coast of
Lancashire, which was identified by Mr. T. Gough and myself as Plwca grcenlandica.
REPORT ON THE SEALS. 59
barbata. Occipital crests well marked, no sagittal crest, ridges at upper border of
temporal fossa distinct. No definite ridge along line of sagittal suture. Hard palate
with posterior border truncated and immediately in front both of the hamular-pterygoids
and the malo-zygomatic articulations. Posterior edge of vomer immediately above this
border and at once articulating with vomerine crest of palate bones ; vertical diameter of
vomer short, that of posterior nares contracted, and about one-half the transverse
diameter. Tympanic bulla somewhat more swollen, the apex is more truncated and the
outer part of its under surface is rougher than in Phoca vitulina ; it is also prolonged
outwards into a much thicker wall for the external auditory meatus, the aperture of
which looks forwards. Greatest width of zygoma about middle of arch and inter-
zygomatic width exceeding greatest cranial width. Post-canine teeth not set closely
together, nor oblique. Subcondyloid process of mandible broadly triangular, inverted,
distinct from the tubercle at the angle ; lower border of body more incurved than in
Phoca vitulina, so that the transverse diameter between the angles is much less in
Phoca grcenlandica than in Phoca vitulina ; coronoid long and pointed.
Phoca hispida. Schreber. Einged-Seal or Floe-Rat. North Atlantic and Arctic Oceans.
Phoca hispida, Sclne'ber, Die Saugthiere, iii. p. 312, pi. lxxxvi., 1778.
„ fcetida, Fabricius in Miiller, Zool. Dan. Prodr., p. viii., 1776, and Fauna groenlandica,
p. 13, 1780.
,, annellata, Nilsson, Skand. Faun., i. p. 362, 1820.
The smallest species of the genus.
Interorbital part of frontal more constricted than in any other seal. Occipital
crests feeble, no sagittal crest, ridge at upper border of temporal fossa distinct ;
no definite ridge along line of sagittal suture. Nasal bones elongated and
attenuated between frontal, not ankylosed together. Premaxilla articulating with
about anterior fourth of outer border of nasal. Hard palate cleft at posterior
border though not so deep as in Phoca vitulina, the base of the triangle between
the hamulars being wider than the sides ; posterior border of vomer scarcely visible in
cleft, and running very obliquely forwards for some distance before articulating with
vomerine crest of palate bone. Transverse diameter of orbit more capacious than in
Phoca vitulina and Phoca groznlandica. Tympanic bulla and external meatus closely
resembling Phoca vitidina. Foramen lacerum posterius very capacious, basi-occipital
perforated mesially so that basi-occipital is largely unossified. Post-canines not crowded
and not obliquely set, frequently quadricuspidate, the second cusp being the largest.
Subcondyloid process of mandible not triangular, but elongated into a vertical ridge,
somewhat inverted and forming the posterior border of the bone, but separated by a
notch from the tubercle at the angle ; lower border of body incurved opposite the last
60
THE VOYAGE OF H.M.S. CHALLENGER.
molar tooth, behind which incurved part this border of the bone sweeps backwards and
outwards in a graceful curve ; coronoid long and pointed.
Some years ago, after an examination of the bones of the Seals which up to that
time had been found in brick clay in several localities in Scotland, I came to the
Fig. 1.— The ramus and part of the body of the mandible, in the upper figure from a fossil Seal found in
brick clay at Montrose, in the lower figure from an adult Phoca hispida.
conclusion1 that they belonged to Phoca hispida, and as the lower jaw is a very
characteristic bone, I figured it both in a recent and fossil specimen, and for con-
venience of reference now reproduce the figure.
Phoca barbata, Fabricius. Bearded Seal. North Atlantic and Arctic Oceans.
Phoca barbata, Fabricius in Miilier, Zool. Dan. Prodr., p. viii., 1776, and Fauna groenlandiea,
p. 15, 1780.
Erignathus barbatus, Gill, Proc. Essex Inst., vol. v., 1866.
The largest species of the genus.
Interorbital part of frontal bone less constricted than in the other species. Occipital
crests present, no sagittal crest, ridge at upper border of temporal fossa distinct; a
strong, curved ridge along line of sagittal suture, with a deep groove between the ridge
and the root of the zygoma. Nasals elongated backwards, but not so attenuated as in
preceding species, and not ankylosed together. Premaxilla articulating with tip and
for a short distance only with outer border of nasal. Superior maxilla having a much
1 I described and figured the characters of the lower jaw in Phoca hispida in my paper On Fossil Seals found in
Scotland, Journ. of Anat. and Phys., vol. iv. p. 268, May 1870. The figures are reproduced in the text.
REPORT ON THE SEALS. Gl
larger articulation with nasal proportionally than in the other species. Antorbital
processes are faint tubercles. Zygomatic arches comparatively flattened, their greatest
transverse diameter at posterior part of arch, and the transverse diameter of cranium
behind external meatus greater than interzygomatic diameter. Malar bone short, only
about equal in length to the same bone in Phoca hispida. Hard palate truncated behind
and reaching close to the hamulars ; posterior border of vomer almost entirely concealed
and extending very obliquely forwards to join the vomerine crest of the superior
maxilla. Tympanic bulla not so swollen as in the other species and somewhat rougher
on inferior surface. Foramen lacerum posterius small, basi-occipital not perforated
mesially, par-occipital s distinct. The floor of the orbit not so oblique as in the other
species of Phoca, but approximating more nearly to the vertical. Post-canines with
distinct intervals between, and not very strongly implanted. Subcondyloid process of
mandible large, triangular, deeply incurved, and continued by a ridge into the tubercle
at the angle of the jaw ; lower border of body a little incurved and concave in its
general outline. Coronoid process shorter and less pointed than in the other species,
and the sigmoid notch more shallow.
A comparison of the cranial characters of Phoca barbata with those of the three
other species of the genus Phoca, shows that it differs more from them than they do
from each other. These differences are especially seen in the lower jaw as just stated,
in the flattening of the zygomatic arches, so that the cranial breadth is greater than the
interzygomatic diameter, in the more vertical direction of the floor of the orbit, and in
the very pronounced ridge along the line of the squamous suture. It has been pointed
out by previous writers (Gill, Allen) that Phoca barbata differs from the other species
of Phoca in having a broader muzzle, in the middle digit of- the manus being the longest
instead of the digits slightly decreasing in length from 1st to 5th, and in possessing
four and not two mammae, and on these differences Gill has established for it the
genus Erignathus. To these external differences may now be added those cranial
differences which I have just described, so that from the point of view of those zoologists
who are in favour of the multiplication of genera, additional data are given for separating
it from the genus Phoca and calling it Erignathus barbatus.
I have not seen any specimens of the Seals which have been named Phoca caspica,
Phoca siberica, and Phoca equestris {Histriop)hocafasciata).
Halichcerus, Nilsson.
Infraorbital foramen does not open into anterior part of floor of orbit but below it,
for the floor of orbit does not form a continuous slope with the posterior border of the
zygomatic root of maxilla, but is separated from it by a deep vertical surface. Anterior
nares high and capacious. Post-canines each with a large, conical, simple cusp ;
62 THE VOYAGE OF H.M.S. CHALLENGER.
secondary cusps not present in upper post-canines except occasionally in 4th and 5th ;
in lower post-canines secondary cusps not unfrequently present. Teeth single-fanged
except the last lower and last two upper post-canines.1
Halichcerus grypus (Fabricius). Grey Seal. North Atlantic Ocean.
Phoca grypus, Fabr., Skr. Nat. Selsk., i. p. 167, pi. xiii. fig. 4, 1790.
Halichaerus griseus, Nilsson, Skand. Faun., i. p. 377, 1820.
Phoca gryphus, Fischer, Syn. Manila., p. 239, 1829.
The only species.2
Interorbital constriction of frontal somewhat swollen about the middle. Occipital
and sagittal crests present and not unfrequently a squamous ridge. Nasals wider than
in Phoca and not ankylosed together. Premaxilla not expanding at its upper end and
with only a limited articulation to outer border of nasal, not reaching its tip ; the two
premaxilke curve outwards from side of nasal, so that the widest part of anterior nares
is in the upper third and the opening generally is very capacious. Z}-gomatic arches
bulging, widest part of arch in the middle and much wider than the widest part of the
cranium. Hard palate with rounded arch at posterior border, the crown of the arch
considerably in front of both the hamulars and the malo-zygomatic joints ; posterior
border of vomer visible in concavity of arch, but soon joining vomerine crest of
palate. Horizontal part of premaxilla thicker than in Phoca and with distinct tubercle.
Tympanic bulla swollen, generally smooth, but with a ridge in outer half, which is
prolonged into thick wall of external auditory meatus. Foramen lacerum posterius
moderate. Basi-occipital usually not perforated mesially. Par-occipital short. Mandible
with a stunted, vertically elongated, subcondyloid process, scarcely inverted and quite
distinct from the tubercle at the angle ; lower border of body thickened and scarcely
inverted ; masseteric fossa very deep, coronoid broadly triangular.
1 Nehring (Sitzb. der Gesellsch. naturf. Freunde zu Berlin, October 17, 1882) gives an account of Halichamis grypus,
and refers to variations in the skull due to age and sex as well as individual modifications. He points out that the
roots may vary in number in the hinder post-canine teeth, and that the accessory cusps are variable in the lower post-
canines. He also states both from his own observations and those of Professor Gerstaecker that an accessory 6th upper
molar not unfreijuently occurs. Gerstaecker has seen it eight times in thirty-four crania, five times on one side only,
thrice on both sides. I may also refer to the skull of a young Halichoerus grypus which I described in the Journ. of Anat.
and Phys., vol. vii. p. 273, 1873, in which no teeth were developed except the canines.
2 This Seal is often regarded, in so far as its distribution in Scotland is concerned, as restricted to the west coast, but, in
addition to specimens from that side of the island, I have placed crania in the Anatomical Museum of the University
from animals killed at the mouth of the Tay, off Montrose, at Golspie in Sutherlandshire, and from the Shetland
Islands.
REPORT ON THE SEALS. 63
OgMORHININ.E.
Anterior nares in front of the infraorbital foramina ; beak moderately prolonged
in front of opening. Either a postorbital process or a mere rudiment of one ; premaxilla
may or may not articulate with nasal. Horizontal part of premaxilla moderate. Floor
of orbit sloping back to become continuous with the thin posterior border of zygomatic
root of maxilla ; infraorbital foramen opening into floor. Greatest width of palate behind
last molar and almost on line with posterior border of zygomatic root of maxilla.
Zygomatic process of maxilla prolonged back for some distance below malar. Inner wall
of orbit defective. Basi-occipital sometimes perforated mesially. Pterygoids horizontal
or almost horizontal in direction and with slit or foramen between upper border and base
• 2 — 2 1—1 5—5
of skull. Dentition — in. , c. , p. c. - —, = 32. Post-canines two-rooted except
2-2 1-1 F 5-5' 1
the first. First and fifth toes of hind foot longer than the rest.
This subfamily contains the genera Ogmorhinus, Leptonychotes, Ommatophoca,1 and
Monachus.
Ogmorhinus}
Stenorhinque, F. Cuvier, Mem. du Museum, t. xi. p. 190, 1824.
Ogmorhinus, Peters, Monatsber. d. k. preuss. Akad. d. Wiss. Berlin, p. 393, footnote, 1875.
Premaxilla does not reach the nasal. Cranial width may or may not be greater
than the interzygomatic width ; nasals ankylosing together early. Anterior nares oblique.
Hard palate emarginate posteriorly, transverse part of palato-maxillary suture opposite
penultimate post-canine ; posterior border of vomer distinctly visible in palatal cleft,
and articulating only slightly with vomerine crest of palate. Postorbital process rudi-
mentary or absent. Basi-occipital not perforated, par-qccipital process present. Wall of
auditory meatus short, foramen opens outwards ; groove between tympanic and mastoid
temporal.
1 Dr. J. E. Gray has given, in vol. i. of the Zoology of the Voyage of the " Erebus" and "Terror," figures of the
palatal aspect and the profile of the skull of the following seals : — Stenorhynchus leptonyx and Lobodon carcinophaga,
Leptonyx weddelli, Ommatophoca rossi, Macrorhinus leoninus, Eumetopias hookeri, Otaria julata, and the palate and teeth of
Arctocephalus hiatus.
2 I wish to express my obligations to Mr. William E. Hoyle of the Challenger Commission for having revised and
verified the references to the names given to the various species of Seals described in the text. In the course of this
revision, which was not made until after Part I, of the Report had been sent to press, he pointed out to me that the
names Stenorhynchus and Macrorhinus have both been applied to different animals. The name Stenorhynchus was given
to a Brachyurous Crustacean so far back as 1818 (Lamarck, Hist. Nat. des Anim. sans Vert), and is regularly in use at
the present time (see Report by E. J. Miers on the Brachyura, part xlix., Zool. Chall. Exp., vol. xvii.). Taking as a
precedent Gill's name Leptonychotes, as a modification of Leptonyx, it would have been better to have modified Stenorhynchus
into Stenorhynchotes, and thus to obtain a generic name which, whilst distinctive, would have been a less departure from
the name most commonly in use than the generic term Ogmorhinus proposed in 1875 by Peters. Macrorhinus was used
in 1825 by Latreille (Earn. Nat. du Regne Animal), to designate a genus of Coleoptera, whilst F. Cuvier in the
previous year had applied to the Elephant Seal the name " Macrorhine." Thus the name as applied to the Elephant
Seal has the priority, and it rests with the entomologists to change the name of the Beetle.
64 THE VOYAGE OF H.M.S. CHALLENGED.
Ogmorhinus leptonyx (de Blainville). Leopard Seal. Southern Ocean.
Phoca leptonyx, Blv., Journ. de Phys., t. xci. p. 298, 1820.
Stenorhynchus leptonyx, F. Cuvier, Diet. d. ScL Nat., t. xxxix. p. 549.
Gray, Zool. Voy. " Erebus" and " Terror," p. 16, 1875.
Tympanic triangular, with pointed apex placed antero-internally, not much swollen,
with ridge running from base to apex. Superior maxilla with extensive articulation
with side of nasal. Hard palate deeply emarginate posteriorly, pterygoids not much
everted. Hamular processes rudimentary. Post-canines large, with three pointed
cusps, the middle being the largest and somewhat recurved at tip, the tips of the
others being inclined towards the middle cusp ; all the post-canines almost equal in size.
Mandible with only a thickening in the border of the bone to represent the subcondyloid
process and with no. angle ; symphysis moderate, lower border of body scarcely incurved.
Skull elongated.
Ogmorhinus carcinophagus (Hombron and Jacquinot). The Crab-eating or Saw-
toothed Seal. Southern and Antarctic Oceans.
Phoca carcinophaga, Hombron and Jacquinot, Voy. Pole Sud, Atlas, Mamm., pis. x., xa.,
1842-1853.
Lobodon carcinophaga, Gray, Zool. Voy. "Erebus" and "Terror," p. 5, 1844.
Stfnorhynchus serridens, Owen, Ann. and Mag. Nat. Hist., vol. xii. p. 331, 1843.
Tympanic not so pointed at apex as in preceding species, swollen and without so
definite a ridge from base to apex. Superior maxilla with limited articulation with side
of nasal. Hard palate moderately concave posteriorly, posterior border of vomer
scarcely seen in the concavity. Hamular processes elongated. Post-canines much
longer than in Ogmorhinus leptonyx; one cusp recurved and especially elongated
and somewhat bulbous at the apex, with a small cusp anteriorly and one, two, or three
behind it. Mandible with a distinct angle and rudimentary subcondyloid process ;
symphysis greatly elongated, lower border of body incurved. Skull not so elongated as
in preceding species.
Leptonychotes, Gill.
Premaxilla reaching the side of the nasal. Cranial width greater than the inter-
zygomatic width , nasals ankylosing together early and the interfrontal part laterally much
constricted ; anterior nares oblique, superior maxilla with a moderate articulation with
the nasal. Hard palate emarginate, posterior border of vomer visible in cleft and
articulating with vomerine crest at anterior part of palate bone. No postorbital process ;
REPORT ON THE SEALS. (55
hamular process and pterygoid everted, basi-occipital perforated mesially. Tympanic
bulla not ridged and with antero-internal angle truncated. Mandible more slender than
in Ogmorhinus. Humerus without a supracondyloid foramen.
Leptonychotes weddelli (Lesson). Weddell's Seal. Southern Ocean.
Otaria Weddellii, Less., Ferussac, Bull. Sci. Nat., t. vii. p. 438, 1826..
Leptonyx Weddellii, Gray, Zool. Voy. "Erebus" and "Terror," p. 7, 1844.
Leptonychotes iceddelli, Gill, Proc. Essex Inst., vol. v., 1866.
Post-canines small, with one moderately prominent cusp, though in the 3rd and 4th
a rudiment of a posterior cusp is also visible ; the last post- canine distinctly smaller
than the rest. Mandible with a faint, incurved tubercle to represent the subcondyloid
process and with no angle ; symphysis moderate, the halves of body widely diverging.
Ommatophoca, Gray.
Nasals ankylosed together, greatly elongated and reaching to the back of the con-
stricted part of the skull ; premaxilla does not reach the nasal ; anterior nares almost in
the vertical plane and slightly in front of infraorbital foramina. Orbits very large and
the interzygomatic width greater than the cranial ; the lower border of zygoma reaches
below the plane of the dentary border of superior maxilla. Malars greatly elongated.
Marked protuberance on squamous temporal above and behind root of zygoma. Hard
palate slightly emarginate, the apex of the cleft a little behind the posterior border of
the maxillary root of the zygoma ; palatal plate of palate bone much smaller than that
of superior maxilla. Vomer visible in palatal cleft. Tympanic bulla not greatly
swollen, triangular and ridged on inferior surface, prolonged outwards into a strong
process which forms the floor of the external meatus, and may extend considerably
beyond it as a thick truncated process. Basi-occipital not perforated. Pterygoids
everted. Nails rudimentary on front and absent on hind feet.
Ommatophoca rossi, Gray. Ross's Seal. Antarctic Ocean.
Ommatophoca Rossii, Gray, Zool. Voy. "Erebus" and "Terror," p. 8, pis. vii., viii., 1844.
Teeth small ; post-canines three-cusped, the central cusp the longest and recurved.
Mandible with a moderate angle and low subcondyloid process ; coronoid short but
pointed, sigmoid notch shallow, symphysis moderate, lower border of body slightly
incurved.
The genus Ommatophoca is, so far as we know, represented only by a single species
living in the Antarctic seas, two skulls of which are in the Natural History department
(zool. chall. exp. — fart Lxviu.— 1888.) Yyy 9
66 THE VOYAGE OF H.M.S. CHALLENGER.
of the British Museum.1 The crania have characters which distinguish them from the
other genera of the same family and which approach in some degree to the characters of
Cystophora, so that Ommatophoca is a well-marked genus. It approximates to Cysto-
phora in the vertical direction of the anterior nares, in their relation to the infraorbital
foramina, and in the shortness of the ascending part of the premaxilla, so that a definite
part of the anterior nares is bounded by the superior maxilla. In the great width of the
orbits and of the skull in the interzygomatic region they also approach each other. On
the other hand the nasals are much longer in Ommatophoca than in Cystophora, and
the superior maxillae articulate with their outer border as far as the tip, and do not leave
the anterior part of this border free ; the palate plates of the palate bones are much
shorter in Ommatophoca than in Cystophora, and they differ from each other in the
number of lower incisors and in the shape of the crowns of the post-canine teeth.
Monachus, Fleming.2
Premaxilla articulating with side of nasal ; anterior nares oblique and a short distance
in front of the infraorbital foramina ; no postorbital process. Nasals not greatly elongated.
Inner wall of orbit defective ; zygomata expanded and the width greater than cranial
width. Hard palate slightly emarginate, the posterior border considerably behind
maxillary root of zygoma and some distance in front of the hamulars, which are rudi-
mentary, and well in front of the anterior border of the glenoid fossae ; the posterior
border of vomer just visible in palatal cleft, and arching forwards to join vomerine crest
about junction of superior maxilla and palate. Pterygoids everted and running almost
horizontally forwards, with large foramen between the upper border and the basis
cranii ; the interpterygoid width considerable. Tympanic bulla swollen, scarcely ridged,
apex truncated ; basi-occipital perforated mesially ; mastoid low ; par-occipital prominent ;
.2-2 1-1 5-5
occipital condyles confluent. Dentition — in. .y—?, c. ^ _}, p. c. g_g = 32.
1 The description of Ommatophoca and Monachus has been written from the crania in this Museum, and I am
indebted to Professor W. H. Flower, C.B., for permission to examine these and other crania of the Seals in the national
collection.
2 The generic name Monachus was suggested by Dr. Fleming in 1822. In a footnote to his Philosophy of Zoology,
vol. i. p. 187, he says, " Some Seals as Phoca monachus are said to have four incisors in each jaw. Such will probably be
constituted into a new genus under the title Monachus." In 1827, F. Cuvier used the name Pelagios {Mem. Mus. Hist.
Nat., t. xi. p. 196), and in 1829 (Diet. d. Sci. Nat., t. lix.) Pelagius, but Fleming's name has obtained general use for this
Mediterranean Seal.
REPORT ON THE SEALS. 67
Monachus monachus (Hermann). Monk Seal. Mediterranean Sea.
Phoca monachus, Herman]], Abliandl. d. k. Akad. d. Wiss. Berlin, vol. iv. p. 501, 1779.
Plioca albiventer, Boddaert, Elenchus Animal., p. 170, 1785.
Pelagios monachus, F. Cuvier, Mem. Mas. Hist. Nat., t. xi. p. 196, 182-1.
Monachus albiventer, Gray, Brit. Mus. Catal., p. 19, 1866.
Post-canines moderately large, very oblique, and set close together; two rooted in the
upper jaw, except the first, mostly three-cusped, with the central cusp large, the anterior
and posterior small, and with a prominent cingulum ; the last upper post-canine set
transversely. Mandible with angle and subcondyloid process not very strong and
apparently continuous with each other ; coronoid broad, not pointed, lower border of
body not incurved.
Cystophorin^e.
Anterior nares in a vertical or almost vertical plane, and the upper part of the opening
in the plane of the infraorbital foramina ; beak considerably prolonged in front of the
opening ; premaxilla not articulating with nasal, horizontal part thick and with a tubercle.
Greatest width of palate behind last molar, almost in line with posterior edge of maxillary
root of zygoma and the transverse part of palato-maxillary suture. Basi-occipital not
perforated mesially. Zygomata very bulging, the widest part of the arch in the middle
and wider than the cranium. Posterior end of malar almost reaching glenoid fossa.
Zygomatic root of maxilla not passing far back below malar. Inner wall of orbit defective.
Pterygoids almost vertical in direction.
r> *.-*.- -2-2 1-1 5 - 5 on
Dentition — in. , c. , p. c. = 30.
1-1 1-1 * 5-5
This subfamily contains the genera Cystophora and Macrorhinus.
Cystophora, Nilsson.
Ascending part of premaxilla short and not reaching the nasal, so that the upper half
or third of anterior nares is bounded by superior maxilla, the outline of which is concave.
Tympanic bulla very swollen, the inner two-thirds smooth, and separated by a strong
oblique ridge from the outer third which is roughened and continuous with the wall of
a short external auditory meatus, the aperture of which opens directly outwards. A
distinct depression in superior maxilla below infraorbital foramen. The orbital orifice
of the infraorbital foramen opens in the lower part of the slope formed by the anterior
part of the floor of the orbit, and below this foramen the bone passes almost vertically
downwards to the posterior edge of the maxillary root of the zygoma. Mastoid part of
temporal relatively prominent. Posterior border of vomer short and articulating with
vomerine crest of palate in the plane of the posterior nares. Crowns of post-canines
68 THE VOYAGE OF H.M.S. CHALLENGER.
plaited rather than lobed, last upper molar with two fangs. Adult males with inflatable
narial sac. First and fifth toes of hind foot longer than the rest, nails rudimentary or absent.
Cystophora cristata (Erxleben). Crested Seal.1 North Atlantic and Arctic Oceans.
Plwca cristata, Erxl., Syst. Eeg. Anim. p. 590, 1777.
Cydophora cristata, Gray, Zool. Voy. "Erebus" and "Terror," p. 4, 1844; and Brit. Mus.
Catal., p. 41, 1866.
Occipital crests strong, two sagittal crests formed apparently of the upper border of
the temporal ridges ; slight ridge along line of sagittal suture. Nasals elongated, not
attenuated, expanding into a somewhat lobate form at the tip ; outer border of nasal
articulating only with frontal and superior maxilla, but the most anterior part of this
border is free. Anterior nares capacious, the upper third being the widest part,
mesethmoids and maxillo-turbinals reaching the opening. Antorbitals strong, post-
orbitals indicated by a tubercle. Hard palate truncated at posterior border reaching
close to hamulars and a little in front of the anterior border of the glenoid fossa.
Zygomatic of temporal curving upwards to reach the tip of orbital process of malar.
A distinct groove between tympanic bulla and mastoid temporal ; slight par-occipital
processes. Mandible with a longitudinal, incurved, subcondyloid ridge separated by a
notch from the incurved angle of the bone ; lower border of body incurved ; coronoid
elongated.
Macrorhinus.
Macrorhine, F. Cuvier, Mem. Mus. Hist. Nat, xi. p. 200, 1824.
Premaxilla with only a horizontal part, lateral boundaries of anterior nares formed
exclusively of superior maxilhe, the outline of the nares concave. Tympanic bulla not
much swollen in adult male, but relatively flattened and roughened, its outer part being
prolonged into an expanded tympanic plate, which forms the wall of a remarkably
elongated external meatus, the aperture of which opens outwards. Depression in
superior maxilla below infraorbital foramen not very definite. The orbital orifice of the
infraorbital foramen opens below the slope formed by the anterior part of the floor of the
orbit, which slope is separated by a considerable interval from the posterior edge of the
zygomatic root of the maxilla. Mastoid temporal scarcely distinguishable as a process.
Posterior border of vomer articulating with vomerine crest of palate in front of truncated
border. Mandible with a distinct subcondyloid process, not incurved or greatly
elongated, and separated by distinct notch from angle ; lower border of body everted, and
with a wide arch between the two lateral halves ; coronoid not elongated. Adult males
with a proboscis. Humerus without a supracondyloid foramen.
1 A specimen of this Seal was caught at St. Andrews in 1872. R. Walker, Scottish Naturalist, November 1872.
REPORT ON THE SEALS. 00
Macrorhinus leoninus (Linnaeus). Elephant Seal. Southern and Antarctic Oceans.
Phoca leonina, Linn., Syst. Nat., ed. x. p. 37, 1758.
Phoca elephqntina, Molina, Saggio sul Stor. Nat. del Chili, p. 280, 1782.
Morunga elepltantina, Gray, Zool. Voy. "Erebus" and "Terror," p. 8, pis. ix., x., 1844.
Macrorhinus anguirostris, Gill, Proc. Essex Inst., vol. v. p. 13, 1866.
This animal has been described in so much detail in Part I of this Report that it is
not necessary for me to repeat its characters here.
Dr. Gill, Mr. Allen, and other American zoologists have regarded the Californian
Sea Elephant as distinct from the southern species, and have named it Macrorhinus
anguirostris. Dr. Gill's description1 was based on the examination of the skull of a female
from Lower California, and the name Macrorhinus anguirostris was conferred owing to
its narrowed and produced snout, as compared with that of a skull from the South Seas,
figured by Dr. Gray in the Zoology of the "Erebus" and "Terror," which was at one time
regarded as an adult female, but which is now known to be a male not full grown. If
this character of the snout be the only difference between them, and Mr. Allen has stated
that the Northern and Southern Sea Elephants differ very little in size, colour, and other
external features, it cannot have much if any value as a mark of specific difference, for
from my comparison of the male and female crania of the Southern Elephant Seal
(Part I.) it will be seen that the male is much broader than the female in the prenasal
region, owing to the greater size of the incisor and canine teeth.
The differences between the skulls of Cystophora cristata and Macrorhinus leoninus
are seen to most advantage in the region of the premaxillary bones and anterior nares,
in the shape of the tympanic bulla and the relative length of the external auditory
meatus, in the position of the orbital orifice of the infraorbital canal, the relative size
of the mastoid temporal, the place of articulation of the vomer with the palate bone, and
the configuration of the lower jaw. When taken collectively these differences are, I
think, sufficient to justify the separation of the genus Macrorhinus from Cystophora.
Trichechida
The family Trichechida? contains only a single genus amongst existing mammals,
although two fossil genera have been described, Alacthcrium and Trichechoclon. The
existing genus is usually called Trichechus, but the old Linnsean term Odobainus has
recently been revived for it by some zoologists, and Allen has consequently named the
family Odobsenidse.2
1 Proc. Essex Inst., vol. v. p. 13, 1866 ; and Proc. Cliieago Acad. Sci., vol. i. p. 33, 1866.
- I may refer to Mr. Allen's valuable History of the North American Pinnipeds for a full discussion of the question
of the generic term which should be given to the Walrus.
70 THK VOYAGE OF H.M.S. CHALLENGER.
Trichechus (Odobasnus), Linnaeus.
Horizontal part of premaxilla very short and thick, and with a strong tubercle ;
ascending part articulating with nasal. Superior maxilla swollen in front, and with
large alveoli for lodgment of canine tusks ; these alveoli are in the transverse plane of
the front of the face. Anterior nares relatively small, vertical in direction, and with
scarcely any premaxilla in front of the opening. Occipital crests distinct and meeting
mesially. No sagittal crest. Crests on frontal and temporal hones marking limit of
temporal muscle. A slight ridge along line of squamous suture. Interorbital and
interzygomatic region of frontal not very constricted. Hard palate truncated imme-
diately in front of strongly projecting hamulars and in line with anterior border of
glenoid fossae ; transverse part of palato-maxillary suture well behind the posterior edge
of maxillary root of zygoma ; vomer entirely concealed by hard palate, its posterior
border articulates with vomerine crest of palate considerably in front of the truncated
border. Zygomatic arch not bulging, its greatest width is at the zygomatic process of the
temporal, and the interzygomatic j>art of the skull is much below the greatest width of the
cranium ; zygomatic process of temporal short and not nearly reaching the orbital process
of the malar. Orbits relatively small ; infraorbital foramen large and opening below the
floor of the orbit. Tympanic bulla not swollen, but uniformly roughened on inferior surface
and continuous with the short thick wall of the external auditory meatus, the mouth of
which looks directly outwards. Foramen lacerum posterius relatively small. Par-
occipitals not detached processes. Basi-occipital not perforated mesially. Occipital
condyles converging anteriorly and sometimes continuous. Condyloid foramen imme-
diately in front of condyle.
Permanent dentition-
. 1-1
in. - - ,
0-0
1-1
c. -
1-1
• 3-3
m. - — -,
1-1
c. -
3-3
3-3
5 — 5
Milk dentition — in. - — , c. - — , p. c. - — = 34.
3 — 3 1-1 4-4
The last and penultimate upper molars and the last lower molar often rudimentary or
absent. Mandible with a massive, incurved, subcondyloid tubercle separated by a notch
from the feeble angle ; condyle with scarcely any neck, coronoid short and broad, sigmoid
notch very shallow, lower border of body everted and with a wide arch ; symphysis
ankylosed. Pineal body remarkably developed.
REPORT ON THE SEALS. 71
Trichechus (Odobwnus) rosmarus, Linnaeus. Morse, Walrus, or Sea Horse. North
Atlantic and Pacific Oceans.
Odobxnus rosmarus, Linn., Syst. Nat., ed. i., 1735.
Phoca rosmarus, Linn., Syst. Nat., ed. x. p. 38, 1758.
Trichechus rosmarus, Auctorum.
Odobsenus rosmarus, Steenstrup, Ofversigt k. Vetensk. Akad. FdrhandL, Bd. xvi. p. 441, 1859;
Zeitschr. f. gesaramt. Natunv., xv. p. 275, 18G0.
„ „ Malmgren, Ofversigt k. Vetensk.-Akad. Forhandl., 1863 (1864) (quoted by
Allen).
The above description of the skull of the genus Trichechus has been written from the
comparison of the crania of a number of specimens in the Anatomical Museum of the
University of Edinburgh, which are all believed to be from the North Atlantic Walrus,
so that they may be regarded as comprising both its specific and generic characters. On
the supposition entertained by the majority of naturalists that the Walruses of both the
North Atlantic and North Pacific are of the same species, they would, I doubt not, also
coincide with the last-named animal, but as I have not had the opportunity of examining
the skull of a specimen known to be from the North Pacific I cannot speak with absolute
certainty. Mr. J. A. Allen has indeed attempted to show that specific differences separate
the Walruses of these two oceans from each other, and, reviving an old name given by
Illiger, he has distinguished the North Pacific animal by the name of Odobmms (Triche-
chus) obesus. In thus subdividing the genus he is also supported by Mr. H. E. Elliott.
In making this division Mr. Allen attaches great importance to a difference in the
relative development of the frontal and occipital regions in the two animals. In the
Atlantic species, he says, the narrow facial breadth is in striking contrast with the great
occipital breadth, whereas in the Pacific species the two regions are more equally
developed. The interorbital constriction is both relatively and absolutely much narrower
in the Pacific animal ; the tusks are longer and thicker, generally more convergent and
less incurved in the Pacific, whilst in the Atlantic animal they are divergent and strongly
incurved. In the Pacific species the front profile is nearly vertical, and the anterior
edge of the nasals is very little posterior to the front border of the base of the tusk, and
the orbits are more anterior ; in the Atlantic animal the front profile is very oblique,
the muzzle is smaller and the nasals scarcely pass beyond a vertical line drawn from the
hinder border of the base of the tusk.
With regard to these characters I would point out that the increase in frontal breadth
of the North Pacific animal, which makes it approximately equal to the occipital breadth,
would necessarily be occasioned by an increase in thickness of the canine tusks, and
the consequently greater development of the superior maxillary bones for their lodg-
ment, and as it is stated that in the Pacific animal the tusks are thicker than in the
Atlantic specimen, it is possible that these differences, as described by Allen, are not
specific, but are simply due to certain specimens having thicker tusks than others. With
72 THE VOYAGE OF H.M.S. CHALLENGER.
regard to the length, thickness, and curvatures of the tusks, I find considerable
differences in the series of crania in the Anatomical Museum of the University. As a
general rule they diverge from each other so as to be two, three, or even four inches
further apart at the tip than at the root, but in one very fine specimen with long and
relatively thin tusks projecting 15 inches beyond the alveolus, the distance between their
tips was a trifle less than that between their roots ; it may be a question therefore if this
character is more than individual or perhaps sexual. As to the front profile and the
relation of the transverse plane of the anterior border of the nasals to the base of the tusk,
I find that in my specimens this plane sometimes corresponds with the posterior border,
at others it extends up to or indeed a little in front of the middle of the base of the tusk,
so that it is obviously a variable feature.
Mr. Allen also refers, though without attaching so much importance as with the other
characters, to a difference in the intermaxillaries in these animals. Usually, he says, in
the Pacific Walrus the interinaxillse extend posteriorly for two-thirds the length of the
nasal, whilst in the Atlantic animal these bones do not enter into the dorsum of the skull,
but end at the anterior border of the nasals. In those of my specimens, in which these
bones had not yet ankylosed with their neighbours, I noticed considerable variation ; in
four each premaxilla articulated with the anterior two-thirds of the outer border of the
nasal, in two each premaxilla reached the anterior border of the nasal and then seemed to
terminate, but an elongated sutural bone was intercalated on each side between the
superior maxilla and about the middle third of the outer edge of the nasal ; in one the
left premaxilla articulated with the anterior two-thirds, whilst the right bone only
reached the tip of the nasal, but beyond it was a sutural bone similar to that above
described. This intercalated bone obviously represents the detached upper end of the
premaxilla. It is obvious that a bone presenting such variations in arrangement in the
skull of the Walrus as does the premaxilla cannot have much importance attached to it
for purposes of classification.
As regards the external features of difference Mr. Allen states that the two animals
are similar in size and probably in general contour, though the facial outline is modified
by the differences in the skull already considered, but the mystacial bristles are shorter
in the Pacific than in the Atlantic Walrus.
On the whole I think it is doubtful whether these animals should be regarded as
specifically distinct ; I would rather consider them as varieties of one species.
Otariida
In no family of mammals, probably, have more diversities of opinion been expressed
by zoologists, both with respect to the number of species in the family and their arrange-
ment in genera and subfamilies, than in the Otariidse. These divergences are to be seen
REPORT ON THE SEALS. 73
Loth in the descriptions of different authors and in those of the same author at different
times. It is, therefore, a matter of much difficulty to construct a classification of the
Eared Seals which wdl prove satisfactory and conclusive, and it is doubtful whether
the specimens in our museums are yet sufficiently numerous to give definite data
of the variations in the skull engendered by age and sex, so that we may avoid
confounding; the sexual and asfe modifications with those that have a generic or
specific value.
Dr. Gray in his latest writings ' made in all eighteen species of Eared Seals, which
he arranged in nine genera. Professor Peters in his latest monograph 2 recognised only
thirteen species, which he classed in three genera, as follows : —
Otaria.
sp. jubata.
Eumetopias.
sp. stelleri.
gillespii (misspelt gilliespii).
cinerea.
hookeri.
Arctocephalus.
sp. pusillus.
falklandicus
brevipes.
elegans.
forsteri.
gazella.
philippii.
ursinus.
Mr. Allen in his History of the North American Pinnipeds divides the Otariidas into
the subfamdies Trichophocacese and Ouliphocaceae. In the Trichophocacese, which are
distinguished by a harsh pelage without under-fur, he places tbe genera Otaria, sp.
jubata; Phocarctos, sp. hookeri; Eumetopias, sp. stelleri; and Zalophus, spp. califomianus
and lobatus. In the Ouliphocacese, which are distinguished by a soft pelage with abundant
under-fur, are the genera Callorhinus, sp. ursinus; and Arctocephalus, spp. austral is
(falklandicus), antarcticus (pusillus), and forsteri. This division, which is essentially
based on differences in the character of the pelage, corresponds to the older and more
popular nomenclature of Hair-Seals or Sea Lions, and Fur-Seals or Sea Bears. Dr.
Burmeister arranges 3 the Hair-Seals into the genera Otaria, sp. jubata, Eumetopias, spp.
stelleri and califomianus, and Phocarctos, spp. cinereus and hookeri ; and the Fur-Seals
into the genera Arctophoca, spp. falclandica (australis), cinerea, and forsteri, and
Arctocephalus, spp. ursinus, philippii, gazella, and pusillus. Professor Flower again '
includes all the Eared Seals in the single genus Otaria, and he regards the different Sea
1 Ann. and Mag. Nat. Hist., ser. 4, vol. xiii; Hand List of Seals, 1874.
2 Monatsber. d. k. preuss. Akad. d. IViss. Berlin, August 9, 1877.
3 Die Seehunde der argentinischen Kiisten, Buenos Aires, 1883.
4 Catalogue of Vertebrated Animals in Museum of Royal College of Surgeons, pt. ii., 1884 ; and Article Mammalia,
Ency. Brit., 9th ed.
(ZOOL. CHALL. EXP. — PART LXVIII. 1888.) Yyy 10
74 THE VOYAGE OF H.M.S. CHALLENGER. .
Lions and Sea Bears as merely species of that genus. A similar view would also appear
to be entertained by Dr. St. George Mivart.1
The position which Mr. Allen, Dr. Burmeister, and to some extent Professor Peters,
have taken up, that a sharp line of demarcation separates the pelage of the Sea Lions
and Sea Bears, owing to the absence of an under-fur in the former and its presence in
the latter, is apparently not quite free from doubt or absolutely to be accepted. Professor
Peters, indeed, admits that both in young and aged Fur-Seals the fur is very sparse.
In the young Fur-Seal from Juan Fernandez described on p. 51 there was no differentia-
tion of the hair into over-hair and under-kair or fur. Dr. Murie states 2 that beneath
the hair of the Sea Lion, Otaria jubata, which is short, firm and thick in the pile, there
is a reddish underwool, very sparsely scattered and which sensibly diminishes with age.
Mr. J. W. Clark 3 also points out that in a specimen which he examined of the Grey Sea
Lion (Arctocephalus cinereus), from New South Wales, whilst the hair was short, stiff,
and bristly, a red and sparse under-fur was present. Dr. Gray, in the Supplement to
his Catalogue of Seals and Whales,4 dwells at some length on the variability displayed
in the length and abundance of the under-fur, which, indeed, he says may be present or
absent in accordance with the season at which the animals are observed. But with
reference to his remarks it should be stated that he does not always differentiate, with
sufficient precision, the Hair-Seals from the Fur- Seals. For he places the Sea Lion of
the North Pacific (Eumetopias stelleri) amongst the Fur-Seals, whilst it is without doubt
a Hair-Seal.5 If the presence in some Eared Seals of an under-fur and its absence in
others were absolute, then undoubtedly it would furnish a divisional character of much
value. It is, however, without question, that in the Fur-Seals the thick coat of under-
fur constitutes a most important structural feature-
There is also, I think, sufficient evidence to show that in the cranial construction
of the different species of Eared Seals differences do exist of such a kind as to support
the view which so many zoologists have entertained that they possess a generic value,
though whether these differences are so important as to justify the breaking up of the
group into six or more genera, is, I think, very doubtful. The great Sea Lion of the
southern hemisphere differs, however, in so many particulars from the Fur-Seals of the
same seas, that it may fairly be separated from them by a distinct generic name, and to
it, therefore, along with Peters and Allen, I restrict the name Otaria, although, in
adopting this separation, it must be admitted that the passage from the Sea Lion of
South America to the Fur-Seals is graded by such forms as Steller's Seal, the Californian
Sea Lion, and the Hair-Seal from the Auckland Islands.
1 Proc. Zool. Soc. Lond., May 19, 1885.
2 Proc. Zool. Soc. Lond., January 28, 1869.
3 Proc. Zool. Soc. Lond., March 18, 1884.
4 London, 1871, p. 9; also in Ann. and Mag. Nat. Hist, vol. iv. p. 264, 1869.
5 See Elliott's elaborate description of this Seal in his work on Alaska already cited.
REPORT ON THE SEALS. 75
On consideration of the whole question, therefore, I am not disposed to split up the
family Otariiclaj to the extent which has been done by Gray, Allen, and Burmeister, or
even by Peters in his earlier memoirs ; but to accept for the present at least the view
which Peters, unfortunately with too great brevity in the specification of the generic
characters, had adopted in his last monograph,1 and to arrange the species under the
generic terms Otaria, Eumetopias, Arctocephalus.
Otaria, Pdron.
Otaria, Peron, Voy. aux terres austr., ii. p. 37, 1816.
Professor Peters defines this genus as follows : — " Ears short (15-20 mm. long); hair
stiff and without under-fur. Bony palate elongated up to, or almost up to the hamular
pterygoids." In addition, I may state that the palate reaches almost as far back as
the transverse plane of the anterior borders of the glenoid fossae and is truncated ;
borders of palate are elevated so that its surface is concave ; in the male deeply so and
with the hamulars converging so as closely to approximate. Posterior nares contracted.
Vomer entirely concealed by palate, and not articulating with the floor of the nose
until it reaches the vomerine crest of the superior maxilla. Infraorbital foramen opens
in floor of orbit immediately above posterior border of maxillary root of zygoma. Pre-
maxilla articulates with outer border of nasal.
Dentition — post-canines .
F 5-5
Otaria jubata (Forster). Southern Sea Lion. South Atlantic and Pacific.
Phoca jubata, Forster, Descript. anim., p. 66, 1775.
,, „ Schreber, Die Saugthiere, iii. p. 300, pi. lxxxiiiB, 1778.
Otaria leonina, Gray, Brit. Mus. CataL, p. 59, 1866.
The skull has been described with so much detail in Part I., in the male, female,
and young, that it is unnecessary to repeat the characters here. Last upper molar
immediately behind posterior border of the zygomatic root of the maxilla. Mandible
with a very massive quadrequilateral subcondyloid process inflected strongly inwards ;
angle with tubercle distinct from subcondyloid process, lower border of body everted.
Muzzle broad in male. I may repeat that only one species has been referred in this
Report to the genus Otaria.2
1 Monatsber. d. k.preuss. Akad. d. Wiss. Berlin, August 9, 1877.
- It should be stated that Burmeister, so far back as 1868 [ZtiUchr. d. gcssamt. Naturwiis., Bd. xxxi. p. 294),
expressed the opinion that the Otaria godeffroyi of Peters is the same animal as the Otaria jubata of Forster, and that
the Otaria ullox of Tschudi and Peters is the female of Otaria jubata.
76 THE VOYAGE OF H.M.S. CHALLENGER.
Eumetopias, Gill.
Eumetopias, Gill, Proc. Essex Inst., vol. v., 1866.
This genus was established by Gill, and Steller's Sea Lion is by some zoologists the
only species which is included in it. Peters, however, has placed in this genus Steller's
Sea Lion, the Californian Sea Lion, the species Eumetopias cinereus from the Australian
Seas and the species Eumetopias hookeri from the Auckland Islands. The characters as
laid down by Peters are as follows : — " Ears longer than in Otaria, but with similar
hair; posterior border of the hard palate far removed from the hamular processes."
Eumetopias stelleri (Lesson). Steller's Seal or Sea Lion. North Pacific.
Otaria stelleri, Less., Diet. Class. Hist. Nat., t. xiii. p. 420, 1828.
Gray, Brit. Mus. Catal., p. 10, 1866.
Eumetopias stelleri, Allen and Bryant, Bull. Mus. Comp. Zool., vol. ii. p. 46, 1871.
Vertex sinuous or "^-shaped in outline from behind outwards.
PremaxUla articulates with about anterior third of outer border of nasal. Anterior
nares well in front of infraorbital foramina. Postorbitals very large and quadrilateral.
Inner wall of orbit very defective. Sagittal crest moderate in male, feeble in female.
Skull relatively broad at frontal constriction. Hard palate truncated, not reaching further
back than about the level of the middle of the zygomata, and well in front of the hamulars;
the borders of the palate scarcely elevated and its surface almost plane. Vomer with its
posterior border completely concealed and reaching the vomerine crest of the superior
maxillae. Hamulars strong and curved outwards. Tympanic not swollen or definitely
triangular, relatively small and roughened, and with one or two ridges projecting vertically
from it ; the bony wall of the external meatus is short ; mastoid massive. Post-canines
, the last upper is double rooted and is situated distinctly behind the maxillary root
of the zygoma and the transverse part of the palato-maxillary suture ; between the last
and the penultimate tooth is a considerable gap which gives the impression that a tooth
had at one time been developed in it, though Mr. Allen states that no evidence has as
yet been seen of the presence of a tooth in this gap. Mandible with angle not verv
prominent ; subcondyloid process massive, vertically elongated, but not projecting so
much inwards as in Otaria jtibata ; lower border of body neither inverted nor everted.
The skull originally described by Dr. Gray as Arctocephalus monteriensis is now regarded
as a specimen of Steller's Sea Lion. A most interesting account of the habits and
characters of this seal has been given by Mr. H. E. Elliott in his work on Alaska already
frequently referred to, in which he points out that in this species also the adult male is
about twice the weight and bulk of the adult female, and Mr. Allen states that the
REPORT ON THE SEALS.
77
average length of ten old male skulls is 375 mm., their breadth 221 mm. ; whilst the
mean length of two old female skulls is 296 mm., and their mean breadth 157 mm. In
the examination, therefore, of the adult skulls of the Sea Lions, both northern and
southern, differences in size are of no value as indicative of specific characters, but are
only of sexual importance.
Eumetopias californianus (Lesson). Californian Sea Lion.
Otaria cdliforniana, Lesson, Diet. Class d'Hist. Nat., xiii. p. 420, 1828.
„ GiJIespii, M'Bain, Proe. Roy. Phys. Soc. Edin., vol. i. p. 422, 1858.
Aretocephalus gilliespii, Gray, Brit. Mus. Catal., p. 55, 1866 {error e).
Zalophus gillespii, Gill, Proc. Essex Inst., vol. v. p. 13, 1866.
The late Dr. James M'Bain, E.N., of Edinburgh, was the first naturalist to describe
the skull of this animal 1 from a specimen obtained from the Gulf of California, which
was given to him by his friend Dr. Gillespie. I purchased this skull after Dr. M'Bain's
death for the Anatomical Museum of the University of Edinburgh. This animal is
generally recognised as distinct both from its southern neighbour, Otaria juhata, and
from the northern or Steller's Sea Lion, though whether a special genus Zalophus should
be established for it, or it should be regarded as only a distinct species of Otaria, or of
Eumetopias, is stdl a moot point amongst zoologists. Although Dr. M'Bain's description
embraces many of the characters of the cranium, and particularly the prominent sagittal
crest on the vertex of the skull, yet I may refer to several additional points and
measurements.
Table IX. — Skull of Eumetopias californianus.
mm.
Extreme condylo-premaxillary length,
292
From front of premaxUla to occipital crest,
268
From basion to optic foramen,
116
From optic foramen to premaxillary tubercle,
159
Extreme interzy somatic width,
168
Extreme width immediately behind external meatus,
150
Greatest width of palate, ....
47
Width between outer sides of base of upper canines, .
56
Width between outer sides of base of upper lateral incisors,
33
Width between outer sides of base of lower canines,
42
Length of palate to incisor teeth, .
128
Height from basion to middle of occipital crest,
102
Smallest interfrontal width in plane of upper surface,
25
Length of nasals, .....
51
Greatest width of anterior nares, .
31
Greatest width at postorbital processes,
68
Greatest length of mandible, . .
214
Greatest width at condyles of lower jaw,
152
Greatest height of sagittal crest, ....
30
1 Proc. Roy. Phys. Soc. Edin., vol. i. p. 422, 1858.
78 THE VOYAGE OF H.M.S. CHALLENGER.
This specimen is evidently an adult male. The suture between the premaxilla and
maxilla has disappeared, though there is an indication that the premaxilla had articulated
with about the anterior third of the outer border of the nasal. The basicranial synchon-
droses are obliterated. The muzzle is markedly more contracted than in Otaria jubata,
and the anterior nares are more oblique in direction than in Arctocephalus. In its
general form, however, the skull has many points of resemblance with Arctocephalus.
The mastoids are irregular and very prominent. The tympanic bulla is roughened and
a strong process projects vertically downwards from it, immediately external to the
carotid canal ; this canal opens into the anterior part of the foramen jugulare.
Palate is neither elongated nor truncated as in Otaria, but slightly emarginate and
converging behind last molar ; its posterior border is well in front of the slender hamular
pterygoids, and about opposite the middle of the zygomatic arch ; borders of palate
scarcely elevated, so that its surface is almost flat, and its widest part is opposite the
last post-canine. Posterior border of vomer is concealed by palate, but its superior
or sphenoidal articulation is in part seen in the emarginate border of the palate ;
vomer not articulating with floor of nose until it reaches vomerine crest of maxilla.
Infraorbital foramen opens into floor of orbit distinctly in front of posterior border of
zygomatic root of maxilla. Postorbitals triangular, and recurved at the apex. Sagittal
crest in the male very high and thin, reaching forwards to the postorbital processes.
The length-breadth index of this cranium, calculated on the interzygomatic width, is
57'5, and on the width behind the external meatus is 51 "0.
5 - 5
Dentition — post-canines ., relatively large, closely approximated, the most
o — o
posterior being below the zygomatic root of the maxilla and in line with or a little
in front of its posterior border. Mandible with a Quadrilateral subcondyloid process
much longer than broad, and inflected inwards ; angle marked by a sbght tubercle ;
lower border of body neither inverted nor everted. Muzzle narrow. The Phocarctos
elongatus of Dr. Gray is probably this species.
Eumetopias hookeri (Gray). The Auckland Island Hair-Seal.
Arctocephalus Hookeri, Gray, Zool. Voy. "Erebus" and "Terror," p. 4, pis. xiv., xv., 1844.
Otaria Hookeri, Peters, Monatsber. d. k. preuss. Akad. d.Wiss. Berlin, May 17, 1866, p. 269, 1867.
Mr. J. W. Clark has definitely established the presence of a large species of Eared
Hair-Seal on the Auckland Islands,1 and an examination of the crania has satisfied him
that they correspond exactly with the skulls brought home by Sir J. C. Ross's Antarctic
expedition, which Dr. Gray named Arctocephalus hookeri. The skull of this Seal is
1 Proc. Zool. Soc. Lond., November 18, 1873, with two characteristic figures of the skull, which is also figured in
Dr. Gray's Zoology of the Voyage of the " Erebus '' and " Terror," pi. xv.
REPORT ON THE SEALS. 79
distinguished by its great length in relation to the breadth both of the cranial box and
in the zygomatic region. Premaxilla articulating with about anterior half of outer border
of nasal. Tympanic with peg-like process from the back of its inferior surface ; mastoids
not very prominent. Palate almost truncated, ending behind about opposite the middle
of the zygomatic arch, and well in front of the hamular pterygoids, its surface hollowed
out in front, but flattened behind, and with its margins converging so that the posterior
nares are constricted. Post-canines g^ ; the last upper distinctly behind both the
maxillary root of the zygoma and the transverse part of the palato-maxillary suture ;
upper post-canines with one large cusp and either without a secondary cusp or with only
one, except in the last two, the crowns of which are bicuspidate or tricuspidate and the
fangs double rooted ; the lower post-canines have not unfrequently an anterior secondary
cusp and the last and penultimate teeth may have three cusps. Mandible elongated,
massive, and almost of the same vertical height in the whole length of the body. The
presence of an additional upper post-canine in this species as compared with Steller's Seal
and the Californian Sea Lion has induced some naturalists to place it in a separate
genus, as Phocarctos hookeri.1
I measured one of the type skulls in the British Museum, which was 268 mm.
in extreme condylo-premaxillary length and 120 mm. in greatest width immediately
behind the external meatus ; from the front of the cranial box to the posterior border
of the base of the postorbital process was 3 1 mm. , and from the same border to the pre-
maxillary tubercle was 139 mm. Another specimen from Campbell Island, New Zealand,
was 293 mm. long, 145 mm. wide in its greatest interzygomatic diameter, and 126 mm.
wide immediately behind the external meatus ; from the front of the cranial box to the
posterior border of the base of the postorbital process was 40 mm., and from the same
border to the premaxillary tubercle 144 mm. The length-breadth indices of these skulls
calculated on the width behind the external meatus were 44 "7 and 42 respectively, and
of the Campbell Island specimen calculated on the interzygomatic diameter was 49.
These skulls have a much lower index than any of the other crania of seals measured in
this Report.
Eumeto'pias cinereus (Peron). Grey Sea Lion of New Zealand and Australia.
Otaria cinerea, Peron, Voy. aux terres austr., ii. p. 54, 1816.
,, „ Peters, Monatsber. d. k. preuss. Akad. d. Wiss. Berlin, May 17, 1866, p. 272, 1867.
This Hair-Seal was first noticed by Peron, but as his account of the animal was too
brief to afford much distinct information of its characters, some naturalists, e.g., Mr. Allen,
have treated it as a mythical or undeterminable species. In a recent memoir, however,
1 See Peters, Monatsber. d. h. preuss, Akad. d. Wiss. Berlin, November 1, 1866, p. 671; and Allen, History of
North American Pinnipeds, p. 209.
80
THE VOYAGE OF H.M.S. CHALLENGER.
Mr. J. W. Clark ' gives a careful description of the skins and the more salient features of
the skull of several specimens of the Grey Sea Lion from the Seal Eocks near Port
Stephens, New South Wales, which animal he identifies with the Otaria cinerea ofPeron.
The Anatomical Museum of the University, and the Museum of Science and Art,
Edinburgh, have recently purchased from Mr. Edward Gerrard, junior, skeletons of a
Seal from Victoria, belonging to this species, and the following description is drawn up
from the examination of the skulls of an adult male and female, and of a young animal,
the dimensions of which are given in the accompanying table : —
Table X. — Skulls of Eumetopias cinereus.
Male.
Female.
Young.
mm.
mm.
mm.
Extreme condylo-premaxillary length, ....
301
245
205
From front of premaxilla to occipital crest,
295
225
From basion to optic foramen, ....
125
100
88
From optic foramen to premaxillary tubercle,
173
136
106
Extreme interzygomatic width, ....
179
137
113
Extreme width immediately behind external meatus,
175
123
100
Greatest width of palate, .....
38
32
27
"Width between outer side of base of upper canines,
62
44
35
Width between outer side of base of lower canines,
30
27
Length of palate to incisor teeth, ....
127
105
87
Height from basion to middle of occipital crest, .
104
79
72
Smallest interfrontal width in plane of upper surface,
18
15
36
Length of nasals, .....
60
47
34
Greatest width of anterior nares, .
43
29
26
Greatest length of mandible, ....
218
169
133
Greatest width at condyles of lower jaw, .
163
120
92
The occipital and sagittal crests are moderately developed in both the male and female,
but have not appeared in the young skull ; the sagittal crest scarcely reaches the con-
stricted part of frontal ; in the male a strong parietal tubercle like that seen in the adult
Otaria jubata is present. A marked character of the skull is its elongation in the adult
cranium in front of the cranial box, and this'is especially noticeable in the frontal con-
striction between the anterior wall of that box and the postorbital processes. At the
beginning of this constricted part the skull is pinched in laterally, and in front of this
constriction it widens somewhat before it reaches the postorbital processes.2 The nasals
1 Proc. Zool. Soc. Lond., March 18, 1884, p. 188.
2 In comparing with each other the skulls of the Seals too much importance must not be attached to differences in
the length and degree of the constriction immediately in front of the cranial box as indicative of specific distinction.
In the comparison of the young and adult skulls of Macrorhinus leoninus and Otaria jubata in Part I. of this Report,
it is shown that this constriction is both much shorter and less marked in the young than in the adult skull of the
same species. In an interesting paper on Cranial Variation in Mvstela pennanti, Erxl. (Proc. Zool. Soc. Lond., Feb. 16,
1886), Mr. Oldfield Thomas has noted how much the interorbital constriction in this animal also is increased in the
aged skull.
REPORT ON THE SEALS. 81
are more elongated than in Gillespie's Seal. The premaxilla articulates with a little more
than a third of the outer border of the nasal. The lateral borders of the hard palate are
almost parallel so that it is of almost uniform width throughout ; the dentary border is
elevated so that the anterior third of the surface is concave, but the posterior third is
flattened ; the hinder border is moderately emarginate, though in the adult male a mesial
cleft, due to imperfect ossification, separates the two palate bones for a short distance
posteriorly ; this border is about midway between the maxillary root of the zygoma and
the glenoid fossa, and well in front of the hamular pterygoids. The vomer has the usual
arrangement of this bone in the Eared Seals. The tympanies are roughened, and, except
a moderate ridge behind, with no special development of processes ; the mastoids are
very prominent, separated by a deep groove from the tympanies, and projecting almost
vertically downwards. The widest part of the zygomatic arch is at its glenoid end, from
which it rapidly diminishes from behind forwards ; in the male the arch is massive
in relation to the size of the skull. The post-canine dental formula is g— g ; these teeth
all possess a cingulum, and in addition to the large central cusp both a much smaller
anterior and posterior cusp, though in the last upper molar these accessory cusps have
almost disappeared. All the post-canines except the first in the lower jaw and the two
last in the upper jaw, are set somewhat obliquely in their sockets, but with distinct
diastemata in the adults ; the last upper is smaller than the rest and placed distinctly
behind the maxillary root of the zygoma. The mandible is massive in the male, with a
broad coronoid, a massive quadrangular subcondyloid process and slight angle ; the lower
border of the body is thickened and slightly inverted. As regards the pes it should be
stated that digits II., III., and IV. have strong black nails ; I. and V. only rudimentary
nails. The toes are almost equal in length, though I. and V. are a little shorter than the
intermediate toes. The toe-flap of digit I. projected 117 mm. beyond its rudimentary
nail.
The length-breadth index calculated on the interzygomatic width of the adult male
skull was 59, of the adult female 55, and of the young skull 55 ; calculated on the width
behind the external meatus the indices were 58, 50, and 49 respectively.
In the number of its post-canine teeth Eumetopias cinereus corresponds with
Eumetopias hookeri, but it differs from it in having the anterior and posterior cusps
much more distinctly marked and more general, and in so many of the teeth being set
obliquely ; further, it has not so great a constriction of the posterior nares and back
of the palate as is seen in the latter Seal. Mr. J. W. Clark says that it is distinguished
from the Otaria albicollis of Peron by the presence of the anterior and posterior cusps
in the post-canine teeth. As regards the species which has been named Otaria albicollis
it should be stated that Peters regards both it and an animal named Eumetopias lobatus
by Gray as identical with Eumetopias cinereus, and Allen is apparently of the same
(zool. cuall. exp. — part lxviii. — 1888.) Yyy 11
82 THE VOYAGE OF H.M.S. CHALLENGER.
opinion. In Dr. Gray's figure of Eumetopias lobatus1 only five post-canine teeth are
shown in the upper jaw, but in form, cuspidation, and oblique setting they correspond
closely with the teeth in my male skull. In an imperfect skull in the Museum of the
Eoyal College of Surgeons of England, marked Otaria lobata, there are only five upper
post-canines.
Arctocephalus, F. Cuvier.
Arctocephales, F. Cuvier, Mem. du Mus., xi. p. 205, 1824.
Arctocephalus, F. Cuvier, Diet. d. Sci. Nat., t. xxxix. p. 554, 1827.
Professor Peters gives the following definition of this genus : — " With longer ears.
o o o o
Below the contour hairs a thick under-fur, which in both quite young and old animals is
very sparing. Structure of skull and bony palate like Eumetopias." To these characters
maybe added, post-canines ftt-, the last upper being distinctly behind the maxillary root
of the zygoma. It is difficult to say how many species are to be referred to this genus.
That it contains several is however undoubted. The Fur-Seal of the Pribylov Islands is
quite distinct from the South American Fur-Seal, both of which again differ from that of
the Cape of Good Hope, and from the Fur-Seal of Australia and New Zealand. The
Fur-Seal of Kerguelen Island is apparently a distinct species, and so also, perhaps, is the
one from Juan Fernandez. It is possible also that there are other species.
Arctocephalus australis (Zimmermann). South American Fur-Seal.
Phoca australis, Zimmermann, Geogr. Gesckichte, iii., 1783.
,, faUdandica, Shaw, General Zoology, i. pt. 2, p. 256, 1S00.
Euotaria nigrescens, Gray, Zool. Voy. "Erebus" and "Terror," p. 12, 1875.
Otaria faUdandica, Abbot, Proc. Zool. Soc. Lond., 1868, p. 192.
Arctophoca falclandica, Burmeister, Die Seehunde der argentinischen Kiisten, Buenos Aires,
1883.
I have described, in the first part of this Eeport, the skull and skeleton of this
animal with considerable detail, from specimens shot in the Messier Channel. The
skull closely corresponds with certain crania in the British Museum from the Strait of
Magellan, labelled Arctocephalus nigrescens, and also with the specimens in the Museum
of the Royal College of Surgeons of England from Lobos Island, River Plate, and from
the Falkland Islands, labelled Otaria australis or Otaria falhlandica, so that the specific
terms falklandicus and nigrescens are obviously synonyms of australis. This Seal
frequents therefore the southern part of the South American continent on both the
1 Zoology of Voyage of " Erebus " and " Terror," vol. i., Mammalia, pi. xvii.
REPORT ON THE SEALS. 83
Atlantic and Pacific coasts, the Falkland Islands to the east, and perhaps Juan Fernandez
and Masafuera to the west. Its geographical distribution almost exactly corresponds with
that of Otaria jubata.
The distinguishing characters of the skull are as follows; the facial part narrow,
slender, and somewhat elongated, the nasal bones are almost in the same plane as the top
of the cranium, i.e., horizontal ; sagittal crest moderate ; tympanic with two or three
strong ridge or peg-like processes projecting vertically downwards ; mastoid massive,
separated from the tympanic by a broad and deep groove. The 6th post-canine not
much smaller than the others. Upper and lower post-canines with a cingulum, a large
cusp, and a small anterior cusp, the last two also with a small posterior cusp; mandible
arching slightly outwards from symphysis to angles.1
Arctocephalus gazella (Peters). Kerguelen Island Fur-Seal.
Otaria (Arctophoca) gazella, Peters, Monatsber. d. k. preuss. Akad. d. Wiss. Berlin, June 10,
1875, p. 393, 1876.
This animal has apparently a more slender configuration of skeleton than the South
American species. Nasal boues in the same plane as the top of cranium ; sagittal crest
absent ; tympanic almost flattened, and with feeble processes projecting from its posterior
part ; mastoid moderate and with broad shallow groove separating it from the tympanic.
The 6th post-canine much smaller than the others. Upper and lower post-canines with a
feeble cingulum, with one large cusp and no secondary cusps. Mandible as in preceding
species.2
1 A. Nehring has recently described (Archiv f. Naturgesch., 1887, Heft i. Taf. ii.) three crania from the River
Trainandahy, Rio Grande do Sul, south coast of Brazil, which he considers to be a new species and names it
Arctocephalus gracilis. His specimens were one male and two females, but they were all young. In the absence of
adult crania it would be hasty to pronounce them to belong to a new species. Burmeister has indeed described crania
of Arctocephalus australis from the north of the mouth of the Rio de la Plata, which is not far to the south of the Rio
Grande, so that it is not unlikely that the skulls described by Nehring are the young of Arctocephalus australis.
For the opportunity of reading Nehring's paper I am indebted to Mr. Oldfleld Thomas. In a more recent communica-
tion (Sitzungsb. d. Geselhch. naturf. Freunde zu Berlin, Dec. 20, 1887, p. 207), Nehring states that Professor Dr. Goldi
has intimated to him that a Fur-Seal has been taken at Ponta Negra, near Rio de Janeiro, which corresponded with the
Seal named by Burmeister Arctophoca falclandica, i.e., Arctocephalus australis.
2 In addition to the two carcases of young Fur-Seals and the two skeletons of the same procured at Fuller's
Harbour, Kerguelen Island, described on p. 36 as Arctocepltalus gazella, the Challenger collection contained the skeleton
of a young specimen killed at Betsy Cove, Kerguelen. This skeleton was overlooked until after Part I. of this Report
had been printed off. All the epiphyses of the long bones of the limbs and those of the vertebrae were unankylosed
and the cranial sutures were unossified, but the occipito-sphenoid synchondrosis was closed. The skull was immature,
so that the specific characters were not strongly marked, but there can be no doubt, I think, that the animal was a
young specimen of Arctocephalus gazella.
84 THE VOYAGE OF H.M.S. CHALLENGES.
Arctocephalus piisillus (Schreber). Fur-Seal of Cape of Good Hope and of Crozet
Islands.
Phoca pusilla, Schreber, Die Saugthiere, iii. p. 314, pi. lxxxv., 1778.
,, antarctica, Thunberg, Mem. Acad. St. Petersb., iii. p. 222, 1811.
Arctocephalus Delalandii, Gray, Brit. Mus. Catal., p. 52, 18G6.
Otaria pusilla, Peters, Monatsber. d. k. preuss. Akad. d. Wiss. Berlin, May 17, 1866, p. 271, 1867.
To this Seal the specific name Arctocephalus delalandii was applied by Dr. Gray,
who has given the following description of the skull in the British Museum. Hinder
aperture of palate narrow, with a rather acute, ovate anterior edge, surface of palate
concave ; facial part of skull rather short, forehead flattened from nasal bone to vertex ;
teeth large ; lower jaw rather short, strong. Further, he says that the hair is rigid,
the under-fur small in quantity, reddish-brown.
I have also examined this specimen. It is obviously an aged male, for the occipital
and sagittal crests are strong, a parietal tubercle projects immediately in front of the
occipital crest, the two nasals are ankylosed together, and the teeth are worn and
broken. The extreme length of the skull is 280 mm., its interzygomatic width is 177
mm., and its width behind the meatus 174 mm. It measures 51 mm. from the front
of the cranial box to the posterior border of the postorbital process, and 137 mm.
from the same border to the premaxillary tubercle. The mastoids are powerful and
project downwards and outwards ; tympanic roughened and with shallow ridges
projecting vertically. The post-canines though injured are obviously not so large as
in Eumctopias hookeri. Mandible with a faint angle and with a subcondyloid process
vertically elongated and not flattened from before backwards. The length-breadth
index calculated on the interzygomatic width is 63, and on the width behind the
meatus 62.
Several years ago I described the cranium of an Eared Seal in the Anatomical Museum
of the University of Edinburgh, which had been obtained from the Cape of Good Hope.1
I regarded it as a distinct species, and, owing to a mesial cleft in the palate between the
two palate bones, named it Arctocephalus schisthyperoes. The skull was 8'1 inches
(206 mm.) long, 5"1 inches (130 mm.) broad between the zygomatic arches, and
4'3 inches (109 mm.) immediately behind the external meatus. Although the teeth
were but little worn, yet the basicranial synchondroses were ossified, and the cranial
sutures had almost disappeared, so that I regarded the skull as an adult but not
aged.
Dr. Gray criticised my description of this skull2 and stated that it was evidentl)7 the
skull of a half-grown animal with the sutures apparent ; he believed the cleft palate to
1 Journ. of Anal, and Phys., November 1868, vol. iii.
2 Ann. and Mag. Nat, Hist, vol. iv., 1869, p. 264.
REPORT ON THE SEALS.
85
be an individual abnormality, and referred the specimen to Arctocephalus delalandii.
Mr. Allen has accepted Dr. Gray's view and has made Arctocephalus schisthyperdes a
synonym of the Seal which has been variously named Arctocephalus pusillus, Arcto-
cephalus antarcticus, and Arctocephalus delalandii. Professor Flower states that a skull
of an Otaria {Arctocephalus) pusilla in the Museum of the Royal College of Surgeons of
England1 is very like the specimen to which I gave the name of Arctocephalus schisthy-
perdes. Mr. J. W. Clark also refers it to the same species, and considers that the cleft
palate is an individual variation similar to what he has seen in a skull of Otaria
(Arctocephalus) ursina.2
I now agree with the view which has been expressed both by Dr. Gray and Mr.
Clark that the cleft condition of the palate is an individual variation, due, without doubt,
Fig. 2.— This figure represents the palate of the Seal which I named Arctocephalus schisthyperiks ; reproduced
from Journal of Anatomy and Physiology.
to imperfect ossification. But I cannot accept Dr. Gray's statement that the cranium
which I described was that of a half-grown animal, for although the sutures between the
facial bones are distinct, those of the cranial box have practically disappeared, so that I am
still of opinion that the skull is adult though not aged. Undoubtedly the skull is very
much smaller than that (apparently an old male) in the British Museum, to which Dr. Gray
originally gave the name of Arctocephalus delalandii, and which would now be called
1 Catalogue of Bones of Mammalia, pt. ii. p. 193, 1884.
2 Proc. Zool. Soc. Lond., March 18, 1884, p. 194. In Mr. Clark's explanation of figure 6, in which a view of the
palate is given, cinerea is obviously a misprint for ursina.
86 THE VOYAGE OF H.M.S. CHALLENGEK.
Arctocephalus pusillus, and if my specimen is to be referred to that species it is probably
to be regarded as that of a female, for we now know much more definitely than we
did a few years ago that the females of the Eared Seals are very much smaller than
the males.
Arctocephalus ursinus (Linnaeus). Fur-Seal of North Pacific.
Ursus marinus, Steller, Nov. coniru. Acad. Petropol., vol. ii. p. 331, pi. xv., 1751.
Phoea ursina, Linn., Syst. Nat., ed. x. p. 37, 1758.
Callorhinus ursinus, Gray, Proc. Zool. Soc. Lond., 1859, p. 359.
Otaria ursina, Peters, Monatsber. d. k. preuss. Akad. d. Wiss. Berlin, May 17, 1866, p. 273, 1867.
Facial part of skull in front of antorbital process short, relatively broad, and with the
nasal bones sloping downwards so as to give an aquiline character to the profile view
of the face. Anterior nares almost terminal and nearly vertical. This appearance is so
characteristic and peculiar as to have induced Dr. Gray to give it generic value, and to
form for this species the genus Callorhinus, with which arrangement Mr. Allen coincides.
Tympanic distinctly ridged, one ridge running antero-posteriorly ; mastoid massive.
Premaxilla articulating with about one-half of outer border of nasal. Hard palate some-
what emarginate, its posterior border well in front of hamular pterygoids, which are
distinct and curved outwards. Extreme condylo-premaxillary length of an adult male
242 mm., interzygomatic width 141 mm. ; width behind external meatus 129 mm.
From front of cranial box to posterior border of base of postorbital process 51 mm. ;
from same border to premaxillary tubercle 118 mm. The length-breadth index calculated
on the interzygomatic width is 58, and on the width behind the meatus 53. The 6th
post-canine is not much smaller than the others, with one large cusp and no secondary
cusps, usually single-fanged, and well behind the maxillary root of the zygoma. The
mandible has no definite angle, and the subcondyloid process is massive and strongly
inflected.
I may refer to Mr. Elliott's work on Alaska for the most complete account of the
appearance and habits of this Seal which has yet been published. Mr. Allen has stated
that in the character of the pelage this Seal differs in no marked way from the Fur-Seal of
the South Pacific, Arctocephalus australis, though the latter is much greyer than the
former ; but in Arctocephalus ursinus the toe-flaps of the pes are greatly developed,
their extension beyond the digits being nearly equal to the length of the rest of the foot,
140 mm., whilst, as I have stated, in Arctocephalus australis (p. 40) they did not pass
more than 110 mm. beyond the toes. Though the males of the two species are almost
equal in size the female of Arctocephalus australis is very much larger than that of
Arctocephalus ursinus.
REPORT ON THE SEALS. 87
Arctocephalus forsteri (Lesson). Fur-Seal of New Zealand and Australia.
Otaria forsteri, Less., Diet. Class. Hist. Nat., t. xiii. p. 421, 1828.
„ „ Clark, Proc. Zool. Soc. Lond., 1875, p. 675, pis. lxx.-lxxii.
Mr. J. W. Clark lias given a good description of this Seal1 from an examination of a
skin and several crania, supplemented by drawings of the animal made by Sir James
Hector. Amongst other characters he states that the hair is coarse in the young, and
black when wet, and with a dense under-fur of a yellow colour. In old animals the
hairs are tipped with white ; snout tapering, obliquely truncated, nostrils on sloping-
surface. Manus with digits I., II. , III. indicated by prolongations, whilst IV. and V.
are indicated merely by a wavy edge. Pes with hair extending as far as nails ; digits
II., III., IV. with strong black nails, in I. and V. nails rudimentary. Hinder edge of
palate rounded and some distance in front of hamulars, surface of palate flat behind.
Teeth thirty-six ; the post-canine formula being 531, small and conical, the first four
upper with an anterior cusp, the 5th tricuspid, the 6th simple ; lower molars all with
anterior cusp.
Mr. Clark is disposed to regard the Fur-Seal from St. Paul's and Amsterdam Island as
a variety of Arctocephalus forsteri, and it is not improbable that the specimen from one
of these islands, which Professor Peters named Arctocephalus elegans,2 is of the same
species.
Arctocephalus philippii (Peters).3 Fur-Seal of Juan Fernandez.
Otaria Philippii, Peters, Monatsber. d. k. preuss. Akad. d. Wiss. Berlin, May 17, 1866, p. 276,
pi. ii. figs, a, b, c, 1867.
This name has been given by Peters to the Fur-Seal of Juan Fernandez and Masa-
fuera. It is a question whether this animal is a distinct species or only Arctocephalus
australis. Mr. Allen holds the latter opinion as to its position, and he gives both
Arctocephalus philippii and Arctocephalus argentatus as synonyms of Arctocephalus
australis. The general form of the skull of both Arctocephalus philippii and Arcto-
cephalus argentatus, as figured by Peters,4 corresponds closely to that of the specimens
of Arctocephalus australis described in the first part of this Report, but in these crania
the rudimentary anterior cusp of the upper molars is more definite than is represented
in Peters' figures. The amount of development of the rudimentary anterior and posterior
cusps is, however, without doubt, variable in different crania of the same species of Seal
1 Proc. Zool. Soc. Lond., December 7, 1875.
2 Monatsber. d. k. preuss. Akad. d. Wiss. Berlin, May 18, 1876.
3 Idem, op. cit., May 17, 1866.
* Peters described and figured as Otaria argentata {Monatsber. d. k. preuss. Akad. d. Wiss. Berlin, November 9,
1871) a skull from Chili.
88 THE VOYAGE OF H.M.S. CHALLENGER.
both in this and in other genera. Only five upper post-canines are figured by Peters in
his specimen of Arctocephalus philippii, the 5th being in line with the posterior border
of the maxillary root of the zygoma ; iu Arctocephalus argentatus, however, six upper
post-canines are represented ; possibly the absence of a sixth upper post-canine in Peters'
specimen of Arctocephalus philippii may be an individual peculiarity. In the British
Museum is a skull marked Arctocephalus nigrescens from Juan Fernandez, and presented
by the Government of Chili, which in its general appearance resembles my specimens of
Arctocephalus australis, and it also has six upper post-canines. The under surface
of the tympanic is however smoother, and the lower border of the mandible runs almost
straight from symphysis to subcondyloid process. The young Fur-Seal described on
p. 52 of this Eeport was the pup of the Fur-Seal of Juan Fernandez, but its immature
condition prevents me from saying definitely whether it was Arctocephalus australis
or another species.
PART III.
BRAIN OF ELEPHANT SEAL AND OF WALRUS.
The Brain has been examined and described in only a few species of the Pinnipedia.
As might naturally be expected, the brain of the Common Harbour Seal, Phoca vitulina,
is the one which has most frequently attracted attention, and descriptions, often
undoubtedly very brief, but in many cases illustrated by figures, have been given by
Tiedemann, Vrolik, Daubenton, Cuvier, Leuret, Bellingeri, Owen, Broca, Krueg, Mivart,
and Theodor. Rosenthal has written a short description of the brain of Halichcerus
grypus. Mivart has made a few observations on some of the convolutions of the brain of
Cystophora cristata. Murie has written an elaborate description and figured the brain of
Otaria jubata, and Mivart has figured and described some of the convolutions of Otaria
(Eumetopias) gillespii. The only observation on the brain of the Walrus to which I can
find a reference is by Sir Richard Owen, who states, in the course of an account of the
dissection of a female Walrus which died in the Zoological Gardens, " the brain weighed
1 lb. 9 oz. avoirdupois ; its convolutions and structure were described," but no further
statement is made regarding them.
Numerous anatomists have, however, described in more or less detail the brains of
various Carnivora, more especially the Dogs, Cats, and Bears ; and through their researches
the plan of construction of the Carnivorous brain and the arrangement of the convolutions
have been worked out with considerable detail. The convolutions and sulci have also
received names, though not unfrecmently confusion has arisen both through different
parts being similarly named and through the same part being differently named by
investigators. The affinity between the Pinnipedia and the proper Carnivora is shown
by certain resemblances in brain-structure and arrangement, and as in the study of the
brain in the Seals and Walrus the anatomist finds it necessary to refer frecpiently to
the brain of the Dog, Cat, &c, it may be useful to give in this place references to the
principal sources of information on the brain of this order of mammals.
(zool. chall. exp. — paet Lxvin. — 1888.) Yyy 12
90 THE VOYAGE OF H.M.S. CHALLENGES.
LlTERATUKE OF THE BRAIN IN CARNIVORA AND PlNNIPEDIA.
Gall, F. J., and G. Spuezheim, Anatoniie et Physiologie du Systeine Nerveux, Atlas, pis. xxxiii., lxxvii.
Paris, 1810-1819.
Tiedemann, F., Icones cerebri simiarum et quorundam nianimalium rariorum. Heidelberga?, 1821. [He
figures the brain of Phoca vitulina in plates ii. and iii.]
Vrolik, W., Specimen anatoinico-zoologieuin de phocis, speciatim de Phoca vitulina. Trajecti ad Ehenum, 1822.
Daubenton, S. J. M., (Euvres completes de Bufl'on, t. xii. Paris, 1828. [He refers to the brain of the Seal.]
Rosenthal, F., Zur Anatomie der Seehunde, Nova Acta Cees. Leqp.-Carol. Acad., Bd. xv. part 2, 1831.
Bell, T., Carnivora in Todd's Cycl. of Anat. and Phys., vol. i., 1836. [He figures the brain of a Lion.]
Owen, Eichard, The Anatomy of the Cheetah, Traits. Zool. Soc. Lond., vol. i., 1833, p. 133. On the Anatomy
of the Walrus. Proc. Zool. Soc. Lond., vol. xxi., 1853, pp. 103-106, and Ann. and Mag. Nat. Hist.,
ser. 2, vol. xv., 1855, pp. 226-229. Comparative Anatomy of Vertebrates, vol. iii., 1868. [He figures
the brain of Phoca vitulina."]
Leuret, Fr., and P. Gratiolet, Anatomie comparee du systeme nerveux. Paris, 1839-1857. Atlas and Text.
[They figure the brain of Phoca vitulina.']
Cuvier, G., Lecons d'anatomie comparee, t. iii. art. v. Paris, 1845.
Bellinqeri, C. F., Anatomia di una Foca vituliua, Mem. reale Acad, di Torino, ser. 2, t. ix., 1848, pp. 651-664.
[He describes briefly the brain of Phoca vitulina.]
Dareste, Camille, Troisieme memoire sur les circonvolutions du cerveau chez les Mammiferes. Ann. d. Sci.
Ned. (Zool.), ser. 4, t. iii. 1855, pp. 65-111.
Pansch, Adolf, Ueber die typische Anordnung der Furchen und Windungen auf den Grosshirnhemisphaien
der Menschen und der Affen. Archiv f. Anthropologic, Bd. iii., 1868. Beitriige zur Morphologie des
Grosshirns der Saugethiere. Morphol. Jahrbuch, Bd. v., 1879. Ueber gleichwerthige Eegionen am
Grosshirn der Carnivoren und der Primaten. Centrdlbl. f. d. Med. Wiss., 1875, No. 38. Bemerkuugen
fiber die Faltungen der Grosshirns und ihre Beschreibung. Archiv f. Psycliiatrie, Bd. viii. Heft 2, 1877.
Flower, W. H., On the Anatomy of the Proteles, Proteles cristatus (Sparrmann). Proc. Zool. Soc. Lond.,
Nov. 11, 1869, p. 474. On the Anatomy of vElurus fulgens, Fr. Cuvier. Proc. Zool. Soc. Lond., 1870,
pp. 752-769. On the Bush-Dog, Icticyon venaticus. Proc. Zool. Soc. Load., 1880, pp. 70-76, pi. x.
Gervais, P., Memoire sur les formes cerebrales propres aux Carnivores vivants et fossiles. Nouv. Archives du
Mus. d'hist. Nat., t. vi., 1870.
Fritsch, G., and E. Hitzig, Ueber die elektrische Erregbarkeit des Grosshirns. Reichertu. du Bois Eeymond's
Archiv, Bd. xii., 1870, p. 300.
Lussana e Lemoigne, Fisiologie dei centri nervosi encefalici. Padova, 1871.
Garrod, A. H , Notes on the Anatomy of the Binturong (Artictis binturong). Proc. Zool. Soc. Lond., 1873,
p. 196. Notes on the Anatomy of Helictis subaurantiaca, Proc. Zool. Soc. Lond., 1879, pp. 305-307.
Also, in Collected Scientific Papers, 1881.
Murie, James, Anatomy of the Sea Lion (Otaria jubata). Trans. Zool. Soc. Lond., vol. viii, 1874. [He figures
the brain of this animal.]
Wilder, Burt G., The Outer Cerebral Fissures of the Mammalia, especially the Carnivora, in Papers, chiefly
anatomical, read at the meeting of the American Association, August 1873, Bulletin of the Cornell
University, 1874. Cerebral Variation in Domestic Dogs and its bearing upon scientific phrenology; also
in Papers chiefly anatomical, ut supra. The Brain of the Cat, read before the American Philosophical
Society, July 15, 1881.
Hitzig, E., Untersuehungen fiber das Gehirn. Berlin, 1S74.
Betz, , Anatomischer Nachweis zweir Gehimceutra. Centrdlbl. f. d. Med. Wiss., 1874, pp. 578, 595.
Fereier, David, The Functions of the Brain. London, ed. 1, 1876 ; ed. 2, 1886.
Benedikt, M., Der Eaubthiertypus am meuschlichen Gehirne. Centrdlbl. f. d. Med. Wiss., 1876, p. 930.
Meynert, Th., Die Wiudungen derconvexen Oberflache des Yorderhirnes bei Menschen, Allen und Eaubthieren.
Archiv f. Psycliiatrie, Bd. viL, 1877.
REPORT ON THE SEALS. 91
Broca, Paul, Anatomic comparee des circonvolutions cerebrales ; Le grand lobe limbique et la seissure lirabiquc
dans la serie des Mamrniferes. Revue d'Anihropologie, ser. 2, t. i., 1878. [He figures the brain of the
Seal]
"Watson, M., and A. H. Young, On the Anatomy of Hyaena crocuta (H. maculata). Proc Zool. Soc. Land.,
1879, pp. 79-107.
Krueg, J., Ueber die Furchen auf der Grosshirnrinde der zonoplacentalen Situgethiere. Zeitschr. f. toiss. Zool.,
Bd. xxxiii., 1880, pp. 595-672, pis. xxxiv.-xxxviii. [He figures the brain of Phoca vitulina, and copies
Murie's figures of Otaria jubataJ]
Miclucho, Maclay, Remarks about the Circonvolutions of the Cerebrum of Canis dingo. Proc. Linn. Soc.
N.S.W., 1881, vol. vi.
Schwalbe, G., Lehrbuch der Neurologie, Erlangen, 1881.
"Wilder, Burt G., and Simon H. Gage, Anatomical Technology as applied to the Domestic Cat. New York
and Chicago, 1882.
Langlet, J. N., The Structure of the Dog's Brain. Journal of Physiology, vol. iv. p. 248, no date.
Mivart, St. George, Notes on the Cerebral Convolutions of the Carnivora. Journ. Linn. Soc. Land. (Zool.),
vol. xix., 1884, pp. 1-24. [He figures the brains of Phoca vitulina and Otaria gillespii.~\
Familiant, Victoria, Beitrage zur Vergleichung der Hirnfurchen bei den Carnivoren und den Primaten.
Inaugural Dissertation, Bern, 1885.
Flesch, Max, Versuch zur Ermittelung der Homologie der Fissura Parieto-occipitalis bei der Carnivoren.
Festschrift fur Albert Kblliker, Leipzig, 1887.
Turner, W., The Pineal Gland (Epiphysis cerebri) in the Brain of the Walrus and Seals. Proc. Roy. Soc.
Edin., December 19, 1887, vol. xiv. and Journ. of Anat. and Phys., January 1S88, vol. xxii. p. 300.
Theodor, Fritz, Das Gehirn des Seehundes (Phoca vitulina). Bericht der naturf. Gesellsch. zu Freiburg,
I. B., Bd. iii., 1887.
BRAIN OF ELEPHANT SEAL (Pis. VIII. , IX.).
Weight and External Form of the Brain. — The brain of Macrorhinus leoninus
which I have examined was removed from the skull of the young female (d) killed at
Christmas Harbour, Kerguelen, on January 4, 1874, and at once placed in spirit for
preservation, without the pia mater having been stripped off. It reached me in good
condition, and weighed, after several years' immersion in spirit, 1 lb. If oz. avoirdupois
(17f oz.). As the brain loses considerably in weight from the action of spirit, the
normal weight of this organ is always greater during life than after being in spirit, so
that the fresh brain would have weighed in all probability several ounces more than is
expressed by the above figures. As the male brain, where the body of the animal
exceeds in size and weight that of the female, is heavier than the female brain, in all
probability the brain of the adult male Elephant Seal would be several ounces more than
that of this young female.
The principal dimensions of this brain were taken with callipers, and are stated in
millimetres in Table XI.
As the brain loses both weight and bulk, and to some extent shape, after pro-
longed immersion in spirit, it is necessary to correct, as far as possible, the dimensions
and form of a hardened brain. This may to some extent lie done by taking a cast of
the cranial cavity of the skull of the animal. I accordingly asked my assistant, Mr. H.
J. Stiles, M.B., to make a cast of the cranial cavity of the skull of the Elephant Seal from
92
THE VOYAGE OF H.M.S. CHALLENGER.
which the brain had been removed, and I append in Table XII. a few measurements
of this east, which, although they include the thickness of the dura mater, give most
probably a closer approximation to the size of the brain during life than from the
measurements of the organ itself.
Table XL — Brain of Elephant Seal.
Extreme length of cerebrum,
Greatest breadth of ,,
Greatest height of „
Antero-posterior length of cerebellum,
Greatest breadth of
Length of pons Varolii,
Breadth of ,, ,,
Length of medulla oblongata,
Greatest breadth of „
Length of olfactory bulb,
Breadth of ,, ,,
of optic nerve,
of optic commissure,
of 3rd nerve,
of sensory root of 5th nerve,
of motor root of ,, ,,
of portio dura or facial nerve,
of portio mollis or auditory nerve,
mm.
Ill
116
63
52
92
24
27
24
26
16
6
4
8
2
7
1-5
2
5
From these dimensions it will be seen that the cerebrum had in the spirit-preserved
specimen almost retained its original length, but had diminished greatly both in breadth
and height, so that the form of the cerebral hemispheres had become greatly modified.
As the cast represents the normal form of the brain the description of the general shape
of the cerebrum has been written from it.
Table XII. — Cast of Cranial Cavity of Elephant Seal.
mm.
Extreme length of cerebrum, ........
114
Greatest breadth of cerebrum, .......
149
Greatest height of cerebrum, .......
82
Length of olfactory bulb, ........
21
Breadth of olfactory bulb, ........
10
On a vertex view the cerebrum formed a triangle, the apex of which was in front and
the base behind ; the apex was somewhat truncated, and the base possessed the breadth
of 149 mm., so that the cerebrum was considerably broader than long, and the rounded
angles of the base fitted into the hollows of the squamous temporals. The anterior ends
REPORT ON THE SEALS. 93
of the olfactory bulbs appeared for a short distance in front of the anterior end of the
cerebrum. The two hemispheres were parallel, and formed the sides of the mesial
longitudinal fissure, but at the posterior end they diverged slightly from each other
so as to expose a small portion of the middle lobe of the cerebellum. The space between
the diverging hemispheres was occupied by a mesial plate of bone continuous with the
upper surface of the ossified tentorium.
The cerebellum projected behind the base of the hemispheres, and the surface of the
cerebellum which was exposed was the posterior or occipital, the general direction of
which curved from above downwards and backwards. The anterior or tentorial surface
again sloped downwards and forwards, and was completely concealed by the cerebral
hemispheres, except the small portion of the middle lobe above referred to. In my paper '
On the Anatomical relations of the surfaces of the Tentorium to the Cerebrum and Cere-
bellum, I pointed out that in the brains of the Carnivora the surface of the cerebellum
which is exposed behind the cerebrum is the occipital, or that which corresponds to the
inferior surface of the human cerebellum, and not the anterior or tentorial surface, which
is the superior surface of human anatomy. At the time when that paper was published I
had not seen the brain of the Seal in situ, but in the summer of the same year I had the
opportunity of seeing the brains both of a young Phoca grcenlandica and a Halichcerus
grypus in the crauial cavity. In the Greenland Seal the cerebellum was below the
hinder part of the cerebrum, and its occipital surface was almost vertical, though with a
slightly forward direction. In Halichcerus grypus the occipital surface of the cere-
bellum was posterior and almost vertical, the vermiform process being the most projecting
part ; the cerebellum was below the cerebrum, but, owing to a slight divergence of the
cerebral hemispheres posteriorly, a part of the vermiform process could be seen between
them when the brain was looked at from above. In the Elephant Seal the cerebellum
was apparently exposed to a greater extent than in the Greenland and Grey Seals.
The base of the brain was comparatively flattened, owing to the shallowness of the
middle cranial fossae. The olfactory bulbs were almost vertical in direction, in conformity
with that of the cribriform plate of the ethmoid bone. The olfactory peduncle was
21 mm. long. It was remarkably slender, more so even than in the human brain, and
was almost entirely concealed in the olfactory sulcus. It terminated posteriorly in a
slight elevation, situated in front of the inner end of the Sylvian fossa, and of the locus
perforatus anticus. This elevation, the trigonum or tuber olfactorium, was 16 mm. long
and 5 mm. broad, and was directed backwards and outwards into the Sylvian fissure. It
is possible that another root had passed inwards to the great longitudinal fissure, but it
was not clearly marked, for the surface of the brain was somewhat abraded at this spot.
The optic nerves, commissure, and tracts were all very distinct, and the last named
curved backwards on the outer aspect of the crura cerebri. The third nerves arose from
1 Proc. Roy. Soc. Edin., March 3, 1862, vol. iv. p. 549.
94 THE VOYAGE OF H.M.S. CHALLENGER.
the inner aspect of the crura cerebri. The fourth nerves had been torn away in the
removal of the brain. The hypophysis or pituitary body was situated behind the optic
commissure and between the third pair of nerves. It was about the size of a small hazel
nut and small on its surface, though a shallow depression on each side indicated a
division into an anterior and a posterior lobe. It was hollowed out internally into a
cavity continuous with the infundibulum. On raising the pituitary body the tuber
cinereum was seen surrounding the base of the infundibulum. The crura cerebri were
short and flattened on the ventral surface.
Convolutions and Sulci. — In entering on a description of the sulci and convolutions
of the brain, either of the Carnivora or of the suborder Pinnipedia, one of the difficulties
experienced by the anatomist is the selection of the terms to be employed. The literature
of the carnivorous brain is extensive, more especially in recent years ; and as many
authors have employed their own terms without much reference to the nomenclature
adopted by other writers, it is sometimes difficult to decide which name should be
selected in description. After some consideration I have thought it advisable not to
limit myself to the terminology of any single anatomist, but to select from the writings
of various authors such of the names as seemed to be most appropriate.
Each hemisphere of the cerebrum of the Elephant Seal was rich in convolutions and
intermediate sulci.
The Sylvian fissure, fissure of Sylvius (s). This was the largest sulcus, and com-
menced on the base of the brain in the Sylvian fossa, situated in the region of the
locus perforatus anticus. It passed almost transversely outwards to the side of the
hemisphere, and was then continued upwards and backwards for 32 mm. on the side of
the right hemisphere, but not so far on the left, and from it an offshoot ascended almost
vertically for 13 mm. The suprasylvian fissure sprang out of it, and seemed as if it were
an anterior branch of bifurcation.
The Crucial fissure (Leuret), fissura cruciata (c).1 This fissure was not visible in the
norma verticalis, for it was situated so far forward that the brain had to be looked at
from the front in order to see it, so that it corresponded with Leuret's description of its
position in the brain of the common Seal. In the Elephant Seal it extended at first
obliquely and then almost transversely outwards for 30 mm. from the mesial longi-
tudinal fissure. It formed a well-marked feature in this region of the brain, and a large
sigmoid gyrus (sgc) was bent around its outer end.
Bounded above and in front by the crucial fissure, and behind by the basal part of
the Sylvian fissure and fossa and the locus perforatus anticus, was a well-defined area on
the hemisphere, which rested on the sphenoid and frontal bones where they formed
the roof of the orbit. This supraorbital area2 obviously corresponded in position to
the orbital surface of the frontal lobe in the human brain, and like it was subdivided
1 Frontal fissure, Owen. 2 Supraorbital convolution, Leuret ; Orbital convolution, Langley.
RETORT ON THE SEALS. 95
into convolutions by certain fissures. The most obvious of these sulci was the olfactory
fissure (ol), in which the slender olfactory peduncle was lodged, and which was situated
parallel to the mesial longitudinal fissure. Between and parallel to the olfactory and
longitudinal fissures was the gyrus rectus (re), which extended from the locus perforatus
anticus forward to the prorean convolution at the anterior end of the hemisphere. The
rhinal fissure (rh) (Wilder),1 formed the outer boundary of the tuber olfactorium ; it was
shallow and was prolonged forwards into the olfactory fissure and backwards into the
Sylvian fissure. External to the tuber olfactorium and the olfactory peduncle was an
intraorbital fissure 2 (io) closely resembling what I have described in the brain of Man and
the Chimpanzee as the triradiate fissure? Its radiations by breaking up the supra-
orbital area contributed materially to the convoluted character of this part of the brain.
Between it and the olfactory sulcus was a convolution which I shall name internal
supraorbital (isc), and between it and the praesylvian fissure was a convolution which may
appropriately be named external supraorbital (esc).
Returning again to the crucial fissure, at the spot where it changed from the
oblique to the transverse, a short sulcus (fissura prsecruciata, pc, Krueg) proceeded
forwards and inwards from it, which along with the crucial and mesial longitudinal
fissures marked off a convolution 27 mm. long, and as this arrangement occurred in both
hemispheres a lozenge-shaped area was produced. This area was first noticed by St.
George Mivart in the brain of the Arctoid Carnivora, and was named by him Ursine
lozenge. In the Elephant Seal, as Mivart has also described in Phoca vitulina, this
lozenge could not properly be seen until the hemispheres were slightly separated from
each other (ur). It is of small size as compared with the corresponding area in the
brain of Ursus maritimus. The prorean convolution {pre) was 17 mm. long ; it was
bounded externally by the prorean fissure, and internally by the mesial longitudinal
fissure ; it was immediately above the gyrus rectus, whilst the ursine lozenge was placed
above and to its inner side.
The convolutions which lie around the Sylvian fissure were much more complex in
arrangement than in the brains of the Dogs, Cats, and Bears, which was due in the
Elephant Seal to their greater tortuosity and the more numerous secondary sulci.
Owing to the various modes of nomenclature which have been adopted by different
anatomists in describing the tiers of convolutions which surmount the Sylvian fissure,
the selection of the names to be applied to them in this description has been difficult.
Leuret, in those brains where the tiers were four in number, simply distinguished them
by numbers — I, II, III, IV — in their order from below upwards from the Sylvian to the
mesial longitudinal fissure. Broca also named them from below upwards the 1st, 2nd,
1 Ecto-rhinal, Owen.
- Intraorbital fissure of the carnivorous brain, Flower and Langley.
3 The convolutions of the Human Cerebrum topographically considered, Edinburgh, 1866. Notes more especially
on the Bridging Convolutions in the Brain of the Chimpanzee, Proc. Boy. Soc. Edin., February 19, 1866, vol. v. p. 578.
96
THE VOYAGE OF H.M.S. CHALLENGES.
3rd, and 4th parietal convolutions. Ferrier called them the 1st, 2nd, 3rd, and 4th
external convolutions, but he numbered them in the opposite direction from Leuret and
Broca, the first being next the longitudinal fissure whilst the fourth bounded the Sylvian
fissure. Owen preferred the following descriptive terms from above downwards — medial,
medilateral, supersylvian, and Sylvian folds or convolutions. Pansch named them from
above downwards — marginal, suprasylvian, outer Sylvian, inner Sylvian ; whdst Langley
called them from above downwards — superior, suprasylvian, ectosylvian, and Sylvian.
If numerical terms are employed, then I thiuk the plan pursued by Ferrier of number-
ing the convolutions from above downwards is to be preferred to that of Leuret and
Broca, as the order of arrangement is thus brought into conformity with the numbering
of the convolutions of the frontal, occipital, and temporo-sphenoidal lobes in the human
brain, where in each lobe the highest convolution is the first. If on the other hand
descriptive terms are used, then I prefer Owen's name of suprasylvian for the convolution
immediately above the Sylvian convolution, instead of outer Sylvian or ectosylvian as
employed by Pansch and Langley ; whilst the highest convolution may appropriately be
called sagittal or marginal, and the one immediately below it mediolateral. Moreover, I
shall call the fissure which separates the Sylvian from the suprasylvian convolution
the suprasylvian fissure ; that between the suprasylvian and mediolateral convolutions
the lateral fissure ; whilst that between the mediolateral and marginal convolutions is the
mediolateral or sagittal fissure. On both the numerical and descriptive methods the
following terms are synonymous in brains with four tiers of convolutions : —
1st external convolution
1st curved fissure
2nd external convolution
2nd curved fissure
3rd external convolution
3rd curved fissure
4th external convolution
Sagittal or Marginal convolution.
Mediolateral or Sagittal fissure.
Mediolateral convolution.
Lateral fissure.
Suprasylvian convolution.
Suprasylvian fissure.
Sylvian convolution.
Along with Flower and Ferrier I shall call the convolution which bounds the crucial
fissure in front, behind, and externally the sigmoid gyrus (sgc).
The Sylvian and suprasylvian convolutions were bounded in front and below by
the jprsesylvian fissure (ps), Owen,1 which passed forwards, upwards, and inwards to the
anterior part of the cerebrum, but did not reach the mesial longitudinal fissure. It
was separated from the triradiate fissure by the external supraorbital convolution,
and from the crucial fissure, above which its inner end was situated, by the sigmoid
convolution. Between the prassylvian and Sylvian fissures the anterior limbs of two
convolutions were situated, which were separated from each other by the suprasylvian
fissure. The more posterior and narrower of these two convolutions was the anterior
1 Supraorbital fissure, Flower and Langley.
REPORT ON THE SEALS. 97
limb of the Sylvian convolution (syc). The commencement of this limb was at first
concealed in the Sylvian fissure, but it became superficial as it passed upwards and
forwards ; it then wound tortuously above the apex of that fissure to become continuous
with the posterior limb of the Sylvian convolution, which was a broad convolution
on the side of the hemisphere behind the fissure and formed a part of its posterior
lip.
The Sylvian convolution was bounded above by the suprasylvian fissure (ss), which
arose out of the fissure of Sylvius just before that fissure passed backwards and upwards ;
in its course backwards the continuity of the suprasylvian fissure was broken by the
passage across it of a bridging convolution ; but it was prolonged downwards and forwards
behind and below the posterior limb of the convolution of the Sylvian fissure, where it
formed the fissura suprasylvia posterior (ssp), which did not extend into either the
Sylvian fissure or the fissura rhinalis posterior. The suprasylvian convolution (ssc)
formed the tier above the suprasylvian fissure ; its anterior limb, which lay next behind
the praesylvian fissure, passed almost directly upwards and forwards, and then turning
backwards became tortuous and was subdivided by short sulci ; but its posterior limb was
prolonged downwards and forwards below and behind the Sylvian convolution to join the
uncinate convolution, and to form with it the inner end of the posterior boundary of the
transverse portion of the fissure of Sylvius.
The fissura coronalis (co) (Owen), commenced at the outer end of the sigmoid
gyrus which it bounded externally ; it was continuous in one hemisphere with about
the middle of the praesylvian fissure, though in the other it was separated from it by
a short gyrus continuous with the suprasylvian convolution. It was separated by a
bridging convolution from the lateral fissure. It curved upwards and inwards, but
did not cmite reach the mesial longitudinal fissure, and it formed along with the
praesylvian fissure the anterior boundary of the suprasylvian convolution. The
fissura lateralis (I) bounded the suprasylvian convolution above, and was continued
backwards in a tortuous course and almost reached the posterior border of the
hemisphere.
Between the lateral fissure and the mesial longitudinal fissure two slender gently
wavy convolutions passed from before backwards. The most internal of these was the
sagittal convolution1 (sac), which formed the marginal convolution of the longitudinal
fissure. It was in part divided by longitudinal sulci into two secondary convolutions,
the more internal of which dipped in places into the longitudinal fissure so as to become
concealed within it. The sagittal convolution commenced as far forward as the posterior
limb of the sigmoid gyrus ; it also bounded the inner end of the coronal fissure, whilst
behind it reached the posterior border of the hemisphere and then inclined to the ten-
torial surface.
1 Medial fold of Oweu.
(zoou chall. exp. — part Lxvin. — 1888.) Yyy 13
98 THE VOYAGE OF H.M.S. CHALLENGER.
The convolution which was placed immediately to the outer side of the sagittal con-
volution was the mediolateral or 2nd external convolution (mlc). It extended forward to
the coronal fissure where it formed a tortuous fold — the coronal gyms ; on the vertex it
was narrow, but as it passed backwards it formed a broad tortuous convolution, subdivided
by sulci, which assisted the sagittal convolution in forming the posterior boundary of
the hemisphere. Between the mediolateral and sagittal convolutions was the medio-
lateral fissure (ml); it was not continuous with the coronal fissure iu either hemisphere,
and on the right side both it and the sagittal and suprasylvian convolutions were so
pushed inwards by the highly tortuous suprasylvian convolution, that it approached
close to the mesial longitudinal fissure.
The corpus callosum and the other mesial structures in the cerebrum were then
divided longitudinally, and the pons and cerebellum were removed by cutting through
the crura cerebri. The convolutions and sulci on the mesial and tentorial surfaces were
thus exposed, and the following arrangement was recognised. The corpus callosum (ccl) was
44 mm. long ; posteriorly it had a rounded free end or splenium, whilst anteriorly it bent
down to the base of the brain to form the genu ; it could be easily torn up into trans-
verse fasciculi of nerve fibres. A septum lucidum occupied the hollow of the genu
between it and the fornix. The splenial fissure (sp), Krueg, was well marked. It com-
menced immediately behind the lobus hippocampi and curved backwards, upwards, and
then forwards behind the splenium, from which it was separated by the gyrus hippocampi ;
it then ran forwards above the corpus callosum, but separated from it by the callosal
convolution, and was continuous at the anterior end of the hemisphere with the crucial
fissure. It was not interrupted in its course in either hemisphere by a superficial
bridging convolution. An offshoot of this fissure was prolonged upwards and forwards
to the sagittal border of the hemisphere 31 mm. behind the crucial fissure. Sixteen mm.
below the splenium the splenial fissure gave origin to a branch which I have named the
postero-horizontal fissure (ph) ; it ran horizontally backwards almost as far as the posterior
border of the hemisphere. The hippocampal fissure (h) was situated between the
hippocampal gyrus and the taenia hippocampi, and curved round the splenium to become
continuous with the callosal fissure, which separated the callosal convolution from the
corpus callosum.
These fissures marked out very distinctly an arched convolution, the great limbic
lobe of Broca,1 comparable with the gyrus fornicatus of human anatomy, which may
conveniently be divided into callosal and hippocampal convolutions, the latter of which
terminated in the uncinate gyrus or lobus hippocampi.- The lobus hippocampi (Ik) was the
inferior end of the gyrus fornicatus, and formed the inner part of the posterior lip of the
1 Strictly speaking, Broca's term " le grand lobe limbique " includes also tlie olfactory lobe, i.e., the olfactory bulb,
peduncle, tuber, and roots.
2 Lobus pyriformis or natiform protuberance of some authors.
REPORT ON THE SEALS. 99
Sylvian fossa; it was bounded externally by a short postrhinal fissure (pr), which was not
continuous with the splenial fissure, but was prolonged forwards into the S)dvian fissure,
and across that fissure into the rhinal fissure which formed the outer boundary of the
tuber olfactorium. A slender prolongation of this tuber passed backwards, but concealed
within the Sylvian fissure, to become continuous with the uncinate gyrus. The hippo-
campal gyrus {he) was prolonged from the uncinate gyrus backwards and upwards, and
was marked by shallow depressions due to the pressure of the small arteries which turned
round the gyrus to enter the choroid plexus situated in the transverse fissure of the
cerebrum. Opposite the splenium it was continued into the callosal gyrus by a slightly
constricted part or isthmus. The callosal gyrus (ee) was prolonged forwards, at first hori-
zontally, and then bending down in front of the genu it formed the genual part of the
callosal gyrus, and reached the end of the mesial longitudinal fissure on the base of the
brain in front of the optic commissure. The suprasplenial fissure of Krueg (sps) could
scarcely be said to exist in the right hemisphere, but in the left the convolution which
intervened between the splenial fissure and the margin of the hemisphere was partially
divided by a fissure running horizontally into an upper and a lower tier. This fissure was
the suprasplenial ; the convolution between it and the splenial fissure was the supra-
splenial convolution {sspc), whilst the convolution between it and the free edge of the
mesial longitudinal fissure was that aspect of the sagittal or marginal gyrus which was
directed to the mesial surface of the hemisphere. The suprasplenial fissure terminated
anteriorly on the dorsum of the brain behind the crucial fissure. The posts2)lenial
fissure (psp) of Krueg was not definitely marked, but the surface of the cerebrum,
which was situated below the postero-horizontal fissure, was divided by fissures into
four slender convolutions running parallel to each other ; below the lowest of these
was a fissure which opened into the splenial fissure, and then ran backwards and
outwards to the border of the hemisphere. Should this fissure represent the postsplenial
fissure, then the slender convolutions might be collectively regarded as representing
the splenial convolution.
I can make no definite statement as to the presence of the Island of Reil, unless
the concealed part of the anterior limb of the Sylvian fissure be regarded as repre-
senting it.
The Pineal body or Epiphysis cerebri, after the cerebral hemispheres were separated
from each other, was seen to project backwards immediately behind the corpus callosum.
It was 17 mm. long, 9 mm. in greatest breadth, and 6 mm. in greatest vertical diameter.
In shape it was like a three-sided pyramid with the apex forwards. The inferior surface
rested in its anterior half on the corpora quadrigemina, and in its posterior half on the
anterior part of the middle lobe of the cerebellum, whilst the two lateral surfaces were in
relation with the sides of the two cerebral hemispheres in the limited region in which it
lay between them. By its apex it projected forwards to the cleft between the two optic
100 THE VOYAGE OF H.M.S. CHALLENGER.
thalami. In Phoca vitulina the pineal body also projected behind the corpus callosum,
and resembled in shape and in its relations to corpora quadrigemina, cerebrum, and cere-
bellum the epiphysis cerebri in the Elephant Seal. It was 1G mm. long, 8 mm. in greatest
breadth, and 6 mm. in greatest vertical diameter.1 The length of the cerebrum in this
specimen was 78 mm. (3 inches). Dr. James Murie is, I think, the only anatomist who
has systematically described the brain of an Eared Seal, and he states that in Otaria
jubata the jjineal body is " relatively large," but he does not give its actual dimensions,
though, if I may judge of its size as represented in his fig. 44, it does not seem
to have been more than about 8 mm. long. It would appear, therefore, that in the
Seals this body is undoubtedly larger than in Mammals generally, though, as will be
shown later on, it is when largest in them only about one-half as big as in the Walrus,
and does not project so far back as to be visible between the two hemispheres of the
cerebrum.
Cerebellum. — This was of large size, and consisted of a middle and of two lateral lobes.
On the tentorial aspect of the cerebellum the middle lobe was greatly elevated above the
lateral lobes, and from its summit the surface sloped rapidly downwards and outwards to
the sides of the organ. At the superior border of the cerebellum, which corresponded to
the ossified tentorium, there was a slight notch opposite the termination of the middle
lobe. On the ventral surface the middle lobe formed the roof of the 4th ventricle and
was situated in a fossa between the two lateral lobes. The middle lobe was separated in
the greater part of its extent from the lateral lobes or hemispheres by a deep fissure on
each side. Each lateral lobe, much thicker when in apposition with the middle lobe
than at the borders of the hemisphere, was separated into a tentorial and an occipital
surface by a deep fissure, which corresponded to the great horizontal fissure of the human
cerebellum, but owing to the different plane occupied by the cerebellum in the Elephant
Seal, it may more appropriately be called the vertical transverse Jissure. The sur-
face of both the middle and lateral lobes was subdivided into numerous folia, but as
this surface was much broken up by fissures possessing considerable depth, and often
tortuous in direction, the folia were short, and did not have the broad plate-like character
one sees in the human cerebellum. These fissures were especially marked on the occipital
surface of the hemispheres, on which they ran from within outwards, but were not quite
symmetrical on the two sides.
Pons Varolii. — The pons had the usual form. Its mesial line on the ventral surface
was marked by a shallow groove for the basilar artery, and this surface consisted of the
superficial transverse fibres. It gave origin at the posterior part of its lateral and ventral
aspect to the two roots of the 5th nerve, the motor root being immediately internal to the
sensory. The sensory root was much thicker than the motor, and its fasciculi were
1 That the pineal gland in Phoca is larger than is usual in the Mammalia was recognised by Ehlers, Zeitschr.f. wiss.
Zool, Bd. xxx. p. 628, Supplement, 1878.
REPORT ON THE SEALS. 101
more compactly enclosed in a common envelope of connective tissue. The Gth nerve,
about equal in size to the same nerve in man, arose from the posterior border of the pons
in the groove between it and the anterior pyramid.
Medulla oblongata or Bulb. — The Bulb was sharply differentiated above by the groove
between it and the pons, but the demarcation between it and the spinal cord below was
not so clear, for the decussation of the pyramids was not very distinct in the ventral
longitudinal fissure. On each side of this fissure a definite anterior pyramid was seen,
much more distinct where it entered the pons than near the spinal cord. On each side of
this pyramid was a slight ovoid elevation which was continuous with the anterior pyramid
on its inner side, but was more clearly defined on its outer border by a shallow fissure.
Outside this elevation and in the interval between it and the pons the trapezium was
very distinct. The restiform body formed a definite elevation on the side of the medulla.
The dorsal surface was hollowed in the usual way into the 4th ventricle, which was
prolonged forwards to the dorsal surface of the pons. The 7th or facial nerve arose
immediately behind the sensory root of the 5th from the groove between the pons Varolii
and the trapezium. The 8th or auditory nerve arose immediately behind but lateral to
the 7th, from the outer part of the trapezium in close relation to the cerebellum ; as it
passed outwards it grooved the ventral surface of the hemisphere of the cerebellum. The
9 th or glossopharyngeal nerve arose immediately behind the auditory from the outer part
of the trapezium. The 10th or pneumogastric nerve arose by a number of distinct fasciculi,
some of which were situated mesial to the others ; they were placed behind the glosso-
pharyngeal and passed outwards in relation to the ventral surface of the hemisphere of
the cerebellum. The 11th or spinal accessory nerve was a cord of considerable magnitude;
its roots arose from the side of the medulla behind the pneumogastric and also from the
side of the cervical cord between the anterior and posterior nerve roots. The roots of the
12th or hypoglossal nerve came out of the medulla at the fissure which marked the outer
border of the ovoid elevation above referred to, so that this " elevation " can scarcely be
regarded as the homoloorie of the " olive " which in the human medulla lies to the outer
side of the roots of the hypoglossal nerve.
Arteries of the Brain. — Two vertebral arteries converged and joined on the ventral
surface of the medulla oblongata to form the basilar. From each vertebral two small
spinal arteries passed backwards in relation to the ventral surface of the spinal cord, and
a much larger branch, a postero-inferior cerebellar, was distributed to the posterior part
of the occipital surface of the cerebellum. The basilar artery ran forwards mesially, at
first in relation to the ventral surface of the medulla oblongata, and then to the corre-
sponding surface of the pons as far as its anterior border. When in line wdth the posterior
border of the pons the basilar gave off a pair of large branches, antero-inferior cerebellar,
which passed outwards to supply the more anterior part of the occipital surface of the
cerebellum. As it lay in the groove in the pons several small transverse branches arose
102 THE VOYAGE OF H.M.S. CHALLENGER.
from the basilar. The artery divided at the anterior border of the pons into three pairs
of branches, two of which passed to the tentorial surface of the cerebellum as the superior
cerebellar arteries, and the third pair passed to the tentorial surface of the cerebrum
behind the Sylvian fissure as the posterior cerebral arteries. At the inner end of the
Sylvian fossa was a short trunk, apparently the divided internal carotid artery, which was
joined with the posterior cerebral by a posterior communicating artery. From this short
trunk a middle cerebral or Sylvian artery ran outwards in the Sylvian fossa and fissure
to supply the cerebrum both in front of and behind the fissure ; another branch, an
anterior cerebral, ran forwards to the mesial longitudinal fissure, which it entered along
with its fellow of the opposite side, it ascended in front of the corpus callosum and then
ran backwards above it to supply the mesial face of the hemisphere. The two anterior
cerebrals were connected close to their origin by a large transverse anterior communi-
cating artery. A small branch, apparently a choroid artery for the supply of the choroid
plexus of the lateral ventricle, was seen at the inner end of the Sylvian fossa. In their
general arrangement these arteries at the base of the brain resembled the well-known
circle of Willis in the human brain.
BRAIN OF WALRUS (Pis. VIII., IX., X.).
Weight and External Form, of the Brain. — I have been fortunate to examine three
specimens of the brain of Trichechus rosmarus. The first was procured for me in 1865,
by my then pupil Mr. (now Dr.) Charles Moon of Dundee, from an animal (a) killed by
an officer of a whaling ship. I dissected it in the course of the following year, and
drawings were made in October 1866 by my then pupil Mr. (now Professor) Richard
Caton of Liverpool. The description of the brain and the drawings were at that time
reserved for future publication. Since then I have received two additional specimens,1
one from a young animal (b), the other from a larger specimen (c). The following
description is based on an examination of all three specimens, and the drawings have been
revised with the help of the two additional brains.
The brain of specimen a weighed after the removal of the membranes and hardening
in spirit 24 oz. 7 drachms avoirdupois ; that of b 13|- oz. ; that of c 26 oz. The two
cerebral hemispheres in c weighed 20^ oz.; the pons, medulla, and cerebellum 5f oz.
Brains a and c were therefore even after prolonged immersion in spirit heavier than the
brain of the specimen examined by Sir Richard Owen, which was probably weighed
immediately after removal.
The principal dimensions of the brains were taken with callipers, and are stated in
millimetres in Table XIII.
1 These were removed from the crania and brought to me by the late Mr. C. E. Smith and by Mr. Peffers.
REPORT ON THE SEALS.
103
Table XIII. — Brain of Walrus.
a.
h.
c.
mm.
mm.
mm.
Extreme length of cerebrum,
128
89
121
Greatest breadth of cerebrum,
140
109
142
Greatest height of cerebrum,
58
66
Antero-posterior length of cerebellum,
"eo
62
58
Greatest breadth of cerebellum,
94
87
112
Length of pons,
31
20
30
Breadth of pons,
42
30
38
Length of medulla oblongata,
19
24
Breadth of medulla oblongata,
23
29
A cast of the cranial cavity of an adult Walrus gave the following as the three great
dimensions of the cerebral hemispheres — length 136 mm., breadth 174 mm., height
105 mm. All these dimensions were considerably in excess of the largest of my three
brains, so that even when allowance is made for the thickness of the cerebral membranes
included in the cast, and for some loss of size from the action of spirit, it is obvious that
none of my spirit-preserved specimens represented the full adult magnitude of the organ.
Viewed from the vertex the cerebrum possessed the form of a broad triangle, the
apex of which was forward and truncated, whilst the base was directed backwards ;
the sides of the triangle were convex, and the junction of the sides and base was
rounded so that the greatest transverse breadth of the cerebrum was distinctly in front
of the base. About midway between the base and apex the side of each hemisphere was
deeply constricted in the region of the Sylvian fissure (PI. X. fig. 1). This con-
striction formed a definite feature in the configuration of the hemisphere ; it curved
upwards, inwards, and backwards, and corresponded to a crescent-shaped ridge of bone
on the inner aspect of the cranial wall.
The olfactory bulbs curved upwards in front of the anterior end of the cerebrum, and
were almost vertical in direction so as to be adapted to the cribriform plate of the
ethmoid. The mesial longitudinal fissure was occupied by the falx cerebri, and the
mesial surfaces of the hemispheres were parallel to each other and to the falx for the
greater part of their length. Posteriorly they diverged from each other and exposed a
portion of the middle lobe of the cerebellum, and the posterior end of the pineal body
(fig. 1, P). The angle of the divergence was occupied above by a thick mesial plate of
bone continuous with the upper surface of the ossified tentorium and below by the pineal
body. The tentorial surface of the lateral lobes of the cerebellum was under cover of
the hinder part of the cerebrum, but the occipital surface of the cerebellum was almost
vertical, and directed backwards and seen behind the cerebrum.
The base of the brain was comparatively flattened. The olfactory bulbs in the larger
104 THE VOYAGE OF H.M.S. CHALLENGER.
brains were 22 mm. long by 11 mm. broad. Each possessed a peduncle which in two
brains was 3 mm. broad but 6 mm. in the third. The peduncle was placed on the
olfactory sulcus, but was not concealed within it. This peduncle terminated behind in
a distinct tuber olfactorium (to), 21 mm. long by 6 mm. broad, which passed backwards
and outwards into the Sylvian fossa and joined the anterior end of the lobus hippocampi ;
from the olfactory tuber a band, which formed an inner root, passed inwards to the mesial
longitudinal fissure and the gyrus rectus. The optic tracts and nerves were from 2 to
3 mm. broad ; the nerves and commissural end of each tract were rounded cords, the
outer part of the tract formed a flattened band winding round the outer side of the crus
cerebri, and was traced to the posterior end of the optic thalamus ; the optic commissure
was smaller than in the human brain. Behind the commissure was a broad tuber cinereum,
from which the dilated infundibulum proceeded to the pituitary body. This body, the
hypophysis cerebri, was indented as if divided into two lateral and two median lobes, of
which the postero-median was much smaller than the antero-median and the lateral (PI. X.
fig 6). Corpora albicantia were not recognised. The crura cerebri were massive, diverged
from each other, and had between them the tuber cinereum and grey matter of the locus
perforatus posticus. The ventral surface of each crus was flattened and marked with
fasciculi, some of which ran in the long axis of the crus, whilst others formed on the
surface a raised bundle, which curved from within outwards. The third nerve was a little
larger than in man, and arose from the inner side of the crus. The fourth nerve was
similar in size and position to the human nerve.
Convolutions and Sulci. — The Sylvian fissure (s) commenced in the Sylvian fossa at
the locus perforatus anticus ; it passed at first almost transversely outwards, and then
mounted upwards and somewhat backwards in the notch on the side of the hemisphere
already referred to, and ended in two short branches of bifurcation. The Crucial fissure (c)
was not visible on the vertex, but was situated at the anterior end of the hemisphere
immediately above the olfactory bulb ; it was short and passed outwards and slightly
downwards. The sigmoid gyrus which bounded it was comparatively slender, and in
brain c, though not in a, was concealed at its outer end in the coronal fissure owing
to the overlapping of that fissure by the broad anterior end of the mediolateral
convolution. It is doubtful if either a prrecruciate fissure or ursine lozenge can be
said to exist.
The sujiraorbital area of the hemisphere was bounded in front and above
by the crucial fissure, and behind and below by the Sylvian fossa and com-
mencement of the Sylvian fissure and the locus perforatus anticus. In this area
the olfactory fissure was situated parallel to the longitudinal fissure and concealed
by the olfactory peduncle; a well-marked rhinal fissure (rh) extended backwards and
outwards from the olfactory fissure, and, bounding externally "the tuber olfactorium,
passed deeply into the Sylvian fissure. An intraorbital fissure (io), which, whflst repre-
REPORT ON THE SEALS. 105
senting the triradiate fissure, was sometimes not furcated, was also present. The gyrus
rectus (re) was definitely marked between the olfactory and mesial longitudinal fissures ;
between the olfactory fissure and the intraorbital fissure was a well-marked internal
supraorbital gyrus (isc), and between the intraorbital and prsesylvian fissures was a
relatively broad external supraorbital gyrus (esc).
The bridging convolutions were not so numerous in the brain of the Walrus as in the
Elephant Seal, and the arraugement of the convolutions in four successive tiers, with
intermediate sulci, above the Sylvian fissure, was more simple and more easily deter-
mined. But it should be stated that the convolutions in opposite hemispheres of the
same cerebrum were not perfectly symmetrical, and that one of the larger brains bad
more frequently bridging convolutions than the other. The convolutions, lying in rela-
tion to the Sylvian fissure, were bounded in front by the pr&sylvian fissure (ps), which,
commencing on the supraorbital area, ran outwards and then upwards on the side of
the hemisphere to become continuous on the vertex with the lateral or second curved
fissure. The external supraorbital gyrus separated it from the intraorbital fissure, whilst
between its upper end and the cruciate fissure the anterior part of the mediolateral
and the sigmoid convolutions were interposed. The Sylvian convolution (syc) immediately
bounded the Sylvian fissure ; its anterior limb consisted of a narrow, tortuous part
which was at its commencement so sunk into the Sylvian fissure as to be concealed
within it. As it ascended it emerged from the fissure, and formed the immediate
boundary of the apex of the fissure, round which it bent, and was continued behind
into a broad convolution situated on the surface of the hemisphere, which formed the
posterior Up of the Sylvian fissure and consequently the posterior limb of the Sylvian
convolution. The broad posterior limb was partially divided into two parallel gyri by a
fissure, which in one brain ran almost vertically, in another obliquely.
Between this Sylvian convolution and the mesial longitudinal fissure three distinct
convolutions intervened, which were separated from each other by fissures both on the
vertex and anteriorly, though the two uppermost blended with each other posteriorly.
The general direction of these convolutions was antero-posterior, but they dipped down-
wards towards the under surface of the hemisphere both in front of and behind the
Sylvian fissure. The convolution next above the Sylvian convolution was the supra-
sylvian convolution (ssc), which was strongly developed and tortuous both in front of and
behind the Sylvian convolution ; it showed a tendency, both in front of the Sylvian con-
volution and opposite the apex of the fissure of Sylvius, to subdivide into two secondary
gyri lying parallel to each other. The Sylvian and suprasylvian convolutions were
separated from each other by the suprasylvian fissure (ss), which was partially concealed
within the Sylvian fissure anteriorly, owing to the depression of the corresponding-
portion of the Sylvian convolution, but it was very distinct on the surface of the hemi-
sphere behind, and formed the fissura suprasylvia posterior (ssp).
(zool. chall. Exp.— part lxviii. — 1S88.) Yyy 1-t
106 THE VOYAGE OF H.M.S. CHALLENGER.
Bounding the mesial longitudinal fissure was the sagittal convolution (sac), which
commenced at the anterior end of the hemisphere at the posterior limb of the sigmoid
gyrus, and then passed back as the marginal convolution of the longitudinal fissure to
where the hemispheres diverged from each other, when it inclined outwards to reach the
tentorial surface of the hemisphere, though in one specimen it reappeared for a short
distance at the posterior border. Between the sagittal and suprasylvian convolutions
an intermediate mediolateral convolution (mlc) was placed, which broadened out in front,
ascended from the anterior border of the supraorbital area, and then passed backwards
to reach the posterior border of the hemisphere, down which it extended behind the
suprasylvian convolution. The coronal fissure was a short sulcus, not continuous with
either the prsesylvian fissure or the mediolateral fissure, from both of which it was
separated by short intermediate gyri. In brain c, where the outer end of the sigmoid
gyrus was overlapped by the mediolateral convolution, the coronal fissure was partially
concealed by it, and this broad anterior end of the convolution may be called the coronal
gyrus. The lateral fissure (I) ran at first upwards and backwards, and then curved
downwards to reach the tentorial border of the hemisphere ; it formed the boundary of
the suprasylvian convolution in front, above, and behind. The mediolateral convolution
was separated from the sagittal convolution by a definite mediolateral fissure (ml) running
antero-posteriorly, which almost reached the sigmoid gyrus, but was separated from the
coronal fissure by a narrow bridging convolution ; behind it reached the posterior border
and tentorial surface of the hemisphere. In the brain drawn in PL X. figs. 1, 3, this
fissure was not bridged across, but in the left hemisphere of one of the other specimens
a secondary gyrus passed across it about the middle of its length. The sagittal and
mediolateral convolutions were wider in front than behind, and formed a larger proportion
of the hemisphere anterior to the Sylvian fissure, whilst on the other hand the Sylvian
and suprasylvian convolutions were wider behind than in front, and formed much the
larger portion of the postsylvian part of the hemisphere.
The prorean convolution was short, and not beak-like as in the Dog, and was con-
cealed by the olfactory bulb.
The convolutions and sulci on the mesial and tentorial surfaces of the hemisphere
were examined after the pons and cerebellum had been removed, and the corpus callosum
mesially bisected. In the larger brains the corpus callosum was 50 mm. long, and was
distinctly differentiated from the grey surface of the convolution. One of the best
marked fissures on these surfaces of the hemisphere was the splenial fissure (sp.) of
Krueg. In its general direction it curved behind the splenium of the corpus callosum,
from which it was separated by the gyrus hippocampi. It was not quite uniform in its
disposition in the two larger brains.
In the one brain (a) (PL IX. fig. 3) it commenced well forwards on the tentorial
surface, and was separated from the postrhinal fissure by two narrow convolutions, which
REPORT ON THE SEALS. 107
connected the hippocampal gyrus with the Sylvian and suprasylvian convolutions ; the
ascending part of this fissure ran backwards behind the splenium, and then curved
upwards and forwards so as to get above it ; here it was interrupted by a bridging con-
volution, beyond which it was continued horizontally forwards above and parallel to the
corpus callosum, but separated from it by the callosal convolution ; it ended anteriorly
in two branches, both of which reached the margin of the hemisphere, the one ended a
little above the inner end of the crucial fissure, but the other was continuous with the
crucial fissure itself. Behind and below the end of the splenium the splenial fissure gave
off a postero-horizontal fissure (ph), which, running horizontally backwards, extended
almost to the posterior border of the hemisphere. Immediately in front of the
interrupting convolution an offshoot of the splenial fissure was prolonged upwards and
slightly backwards to the sagittal margin of the hemisphere.
In the other well-grown brain, the part of the splenial fissure in relation to the hippo-
campal gyrus was interrupted by two bridging convolutions in the left hemisphere, but
by only one in the right. The part above the callosal convolution was not interrupted by
a bridging convolution in the left hemisphere, though it was so in the right. In both
hemispheres this fissure terminated anteriorly by becoming continuous with the inner
end of the crucial fissure. In both hemispheres, also, a postero-horizontal fissure (ph)
extended backwards from the splenial fissure almost to the posterior border of the
hemisphere (PL X. fig. 5).
The hippocampal fissure (h) was situated above the hippocampal gyrus, between it
and the taenia hippocampi ; it curved round the splenium, and became continuous with
the callosal fissure ; at the bottom of this fissure, between it and the taenia hippocampi,
was the dentate gyrus.
The great arched convolution, gyrus fornicatus or the great limbic lobe of Broca,
was differentiated by the splenial, hippocampal, and callosal fissures, and consisted of
callosal and hippocampal convolutions with the uncinate gyrus or lobus hippocampi.
The lobus hippocampi (Hi) was the inferior end of the hippocampal convolution, and formed
the inner portion of the posterior lip of the Sylvian fossa ; it was demarcated on its outer
lateral side by the postrhinal fissure (pr) which was continued forwards into the Sylvian
fissure, to become through it continuous with the rhinal fissure ; the tuber olfactorium
was also continuous with the uncinate gyrus, but the intermediate band was small and
so deeply lodged in the Sylvian fissure as to be recognised with some difficulty. The
recurved part of the lobus hippocampi was continuous with the taenia hippocampi and
with the band of grey matter at the bottom of the hippocampal fissure which in human
anatomy is called fascia dentata or dentate gyrus. The hippocampal gyrus (he) was
prolonged from the uncinate gyrus backwards and upwards, and was marked by shallow
arterial depressions similar to those described in the Elephant Seal. The ccdlosal gyrus (cc)
at first passed horizontally forwards and then bent downwards in front of the genu of
108 THE VOYAGE OF H.M.S. CHALLENGER.
the corpus callosum, where it formed the genual j>art of the callosal convolution, to reach
the basal part of the mesial longitudinal fissure. The suprasplenial fissure (ssp) of
Krueg was as a rule elementary, but in the right hemisphere of brain c it was a distinct
fissure situated on the mesial surface of the hemisphere parallel to the splenial fissure,
and separated from it by a distinct gyrus, which may be called the suprasplenial gyrus.
It was continuous behind the splenium with the splenial fissure, whilst it terminated
anteriorly in a sulcus, which indented the sagittal gyrus above the crucial fissure.
Between the suprasplenial fissure and the free edge of the mesial longitudinal fissure
was that aspect of the sagittal gyrus which was directed to the mesial marginal
surface of the hemisphere. The postsplenial fissure (psp) of Krueg was situated
behind the ascending part of the splenial fissure, and ran backwards and upwards
nearly to the posterior border of the hemisphere below the postero-horizontal fissure ;
it was separated from the splenial fissure by the splenial convolution (sj)c), which is
consequently bounded in front by the splenial and behind by the postsplenial fissure.
I could not speak with any precision of the Island of Reil, unless the concealed part
of the anterior limb of the Sylvian convolution be regarded as representing it ; for the
lower end of this limb of the Sylvian convolution passed deeply into the fissure, and
was concealed by the anterior limb of the suprasylvian convolution, which for some
distance therefore formed the anterior lip of the fissure of Sylvius.
Interior ofi the Cerebrum. — A vertical transverse section through the right hemisphere,
immediately in front of the anterior pillar of the fornix, showed the fibres of the corpus
callosum extending outwards to become continuous with the white core of the hemisphere.
Immediately below the anterior mesial part of the corpus callosum the right half of the
septum lucidum formed a vertical lamina which was relatively thick. Laterally to the
septum lucidum was the lateral ventricle, the inner part of which was vertical, but the
outer part extended horizontally outwards below the corpus callosum, though it curved a
little downwards at its lateral limit. When the ventricle was opened into by slicing
away the corpus callosum the nucleus caudatus of the corpus striatum was seen to form
a large and well-defined pear-shaped body at the anterior part of the floor ; but the
ventricular chamber was not prolonged in front of the caudate nucleus. The greatest
transverse diameter of this nucleus was 27 mm., and its antero-posterior diameter was
30 mm.
The optic thalamus was behind and to the inner side of the nucleus caudatus, a
shallow groove in which the tsenia semi-circularis could be seen, being placed between
them. Its upper surface was covered by the fornix and choroid plexus, on removing
which structures this surface was seen to be 25 mm. in transverse and 35 mm. in antero-
posterior diameter.
The fornix was prolonged in a curve backwards, outwards, downwards, and forwards
into the descending horn of the ventricle as the tsenia hippocampi, and followed the
REPORT ON THE SEALS. 109
curve of the hippocampus major. Internal to it the choroid plexus was also prolonged
downwards, to become continuous through the great transverse fissure with the pia
mater covering the gyrus hippocampi. Where the cavity of the ventricle curved down-
wards and outwards into the horn an indication of a recess was seen in its posterior horn,
but it did not amount to a cornu, and there was no elevation which could be called a
hippocampus minor. The inner surface of the optic thalamus formed the lateral wall
of the third ventricle. The corpora quadrigemina were well marked, and the testes
overlapped the nates.
The Pineal body or Epiphysis cerebri was remarkable for its size. In specimen c it
measured 30 mm. in its long and 18 mm. in its greatest transverse diameter; in b it
measured 29 mm. by 13 mm. It was somewhat pyriform in shape, with the apex directed
forwards to the optic thalami, whilst the base, which was free, projected backwards so as
to be visible, when the brain was looked at from above, between the two cerebral hemi-
spheres, where they diverged from each other posteriorly. It possessed three surfaces —
one was inferior, and rested in almost its whole length on that surface of the middle lobe
of the cerebellum which was in relation with the tentorium, and this surface was some-
what depressed below the level of the corresponding surface of the hemispheres of the
cerebellum for its lodgment. The other two surfaces were lateral, and in relation to the
inner and jjosterior border of the cerebral hemispheres, between which the epiphysis was
placed. These surfaces were slightly concave in their anterior two-thirds, so as to be
adapted to the convex borders of the hemispheres ; but more posteriorly, where the pineal
body projected between the hemispheres, they were somewhat convex, and mounted up-
wards to form a ridge in the inter-hemispherical interval (PI. X. fig. 1). The pineal body
was separated by the tentorium from the cerebrum, and was closely tied down to the
cerebellum by the arachnoid and the pia mater ; so close indeed was this relation, that
in brain a of the Walrus, which I dissected as far back as 1865, I mistook the pineal
body for a special thickening of the pia mater covering the middle lobe of the cerebellum.
The apex of the epiphysis passed forwards in front of the cerebellum and superficial to the
corpora quadrigernina to the region of the optic thalami, but, owing to this part of the
brain being somewhat friable from imperfect preservation, I could not ascertain its exact
connections, though there can, I think, be little doubt that, as in other Mammals, it
was attached to the thalami by a pair of peduncles.
The very remarkable size of the epiphysis cerebri in the brain of the Walrus, and its
unusual development also in the brain of the Seals, are of especial interest in connection
with recent important observations on the connections and homology of the pineal body
Elders showed ' that in the Plagiostomata the epiphysis cerebri is lodged in a depression
in the cartilaginous cranium, whilst retaining its connection with the brain through its
1 Zcitschr.f. u-iss. Zool., Bel. xxx., Supplement, 1878.
110 THE VOYAGE OF H.M.S. CHALLENGER.
peduncle. Sir Richard Owen makes use1 of this and other facts connected with the
extension of the pineal body into or towards the cranium in these Fishes and in
Reptdes in support of the hypothesis that the conario-hypophysial tract represents the
passage of the gullet to the neural aspect of the body and the formation of a neural
mouth. But additional interest has quite recently attached to the pineal body by the
discovery, as the result of independent research, both by H. W. de Graaf 2 and W. Baldwin
Spencer 3 during the year 1 886 of a mesial pineal eye in the Lacertilia. By these naturalists
the mesial foramen in the parietal bone in this group of Reptiles has been seen to be occu-
pied by an eye, and Mr. Spencer has worked out in a number of species of Lizards the
structure of this eye and its connections, from which it would appear that the pineal eye
is connected by an elongated stalk or peduncle with the thalamencephalon. This peduncle
grows out of the optic thalami ; at first it passes upwards in the interval between the
cerebral hemispheres and the optic lobes, and then runs forwards on the dorsal aspect of
the cerebrum, to end in the mesial eye, situated in the parietal foramen.
In the Mammalia this apparatus has practically disappeared, and is represented only
by the aborted structure which we call the pineal body, though it should be stated that
in the Horse, as M. Chauveau has pointed out,4 it may occasionally assume larger dimen-
sions, and project backwards so as almost to touch the cerebellum. But in the Seals to
some extent, and in the Walrus in a more remarkable manner, the pineal body has retained
a greater magnitude than is customary in Mammals. The direction, however, which this
body takes in these Mammalia is different from that of the stalk of the pineal eye in the
Lizards. For in these Reptiles the direction of the peduncle is at first upwards and then
forwards, so as not to overlie either the optic lobes or the cerebellum, whereas in the
Walrus and Seals the direction of growth is always backwards. Two factors may operate
in the cranial cavity of the Walrus and Seals to induce the backward direction to which I
have referred, viz., the growth of the tense unyielding tentorium, and the backward
development of the hemispheres of the cerebrum. Through lying subjacent to the ten-
torium the growth of the elongated pineal body in the direction either of the parietal
bone or of the superior part of the occipital bone is effectually prevented, and the only
course which it can take is towards the cerebellar region of the occiput. Similarly the
posterior development of the cerebral hemispheres, which overlie both the optic lobes and
the cerebellum, would by the compression of the pineal body between the cerebrum and
cerebellum assist in giving it a backward direction. It is possible, also, that these same
factors may operate in producing the aborted condition of this body which one finds in
the Mammalia as compared with the Lizards. For the pressure exercised by the growth
1 Rep. British Assoc, York, 1881, p. 719 ; Aspects of the body in Vertebrates and Invertebrates, Lond., 1883.
2 Zool. Anzeiger, March 29, 1886.
3 Quart. Journ. Micr. Sri., October 1886.
4 Traite dAnatomie compared des animaux domestiques, 185", p. 650, fig. 177.
REPORT ON THE SEALS. Ill
both of the tentorium and cerebral hemispheres upon an elongated structure, like the
pineal stalk, would occasion atrophy both of the stalk and of the pineal eye, and a con-
secpient disappearance of the mesial parietal foramen in the mammalian skull. It is
conceivable, however, that the atrophy might begin distally by bone formation closing up
the parietal foramen, as a result of which both eye and stalk would disappear. But what-
ever cause has been in operation to lead to the disappearance of both pineal eye and stalk,
only the proximal end of the pineal organ, where it arises from the thalamencephalon, is
left in the Mammalia. It is, however, of interest to note that in at least one Mammal,
viz., the Walrus, this proximal part may retain such magnitude as to be visible between
the hinder ends of the cerebral hemispheres, so that it does not present so aborted or
residual a character as in other Mammals. But the special function, if any, which may
be discharged by the pineal body in this animal will be difficult to ascertain.
The Cerebellum was a massive organ and consisted of a middle lobe and of two
lateral lobes or hemispheres. The middle lobe on the tentorial surface was separated by
a furrow on each side from the corresponding lateral lobe, and the distance from this
furrow to the extreme lateral border of the hemisphere was 54 mm. At the superior
margin of the cerebellum the middle lobe was concealed in a cleft which separated the
two hemispheres from each other. Inferiorly where the middle lobe formed the roof of
the 4th ventricle it was depressed between the two lateral lobes of the cerebellum. Each
hemisphere was divided into a tentorial and an occipital part by a vertical transverse
fissure, and the surface of the tentorial aspect was split up into numerous broad plate-
like folia by intermediate fissures. The occipital aspect was similarly subdivided, but
there was also evidence of a division of this aspect into lobelets by broader fissures.
Thus about opposite the middle of the side of the medulla oblongata a broad and deep
fissure curved outwards and forwards, so as to divide this aspect of the cerebellum into
an anterior and a posterior lobelet. The folia which bounded the fissure dipped into it in
a whorl-like manner, so that the fissure may be distinguished as the vorticose fissure (v).
On raising the anterior lobelet the superficial transverse fibres of the pons could be traced
without any difficulty into the white core of the hemisphere.
The Pons Varolii was broader in the middle than either in front or behind. It had
the usual mesial groove for the basilar artery, and the superficial transverse fibres were
very distinct, and could readily be traced outwards into the hemispheres of the
cerebellum. The 5th nerve arose from the side of the pons by a large sensory and a
small motor root. The motor root was at first on the inner side of the sensory, and then
passed outwards in contact with its ventral surface to join the inferior maxillary division
of the ganglion. The sensory root was 1 3 mm. in its transverse diameter, and expanded
anteriorly into a flattened Gasserian ganglion which gave off the three divisions of the
nerve ; some fibres of this root entered the substance of the pons, but others passed
backwards between the facial and auditory nerves to the anterior and outer part of the
112 THE VOYAGE OF H.M.S. CHALLENGER.
medulla oblongata. The 6th nerve had been torn away in a and c, but in b was
situated in the groove between the pons and anterior pyramid. The 7th or facial nerve
arose from between the pons and medulla external to the anterior pyramid. The 8th
or auditory nerve arose from the lateral aspect of the medulla immediately behind the
pons.
The Medulla Oblongata or Bulb was much injured in specimen a, but in good order in
the two others. It possessed on each side of the mesial fissure a distinct anterior pyramid
which passed above into the substance of the pons. Immediately external to this pyramid
but not reaching the pons was an elongated oval swelling 14 mm. long and 4 mm. in
greatest transverse diameter; this swelling was bounded both on its inner and outer borders
by a shallow groove. The roots of a nerve, which were unfortunately torn across in c
close to the medulla, emerged from the groove at the inner border ; in b they were entire
and were the roots of the hypoglossal or 12th cranial. The swelling is to be regarded
as like the olive in the human medulla, where the hypoglossal nerve arises from the
groove on its inner side between it and the anterior pyramid. In Phoca vitulina a
swelling was also seen in the same region but not separated by a groove from the anterior
pyramid so that it is a part of that structure. External to the upper end of the anterior
pyramid was a body which apparently represented the trapezium, though it was not
marked with transversely arranged bundles of nerve fibres. Below this body the side
of the medulla swelled out into a restiform body, the surface of which was marked by
arciform fibres running from before backwards around the side of the medulla. Springing
out of the restiform body were some nerve roots, which in c were torn across close to the
medulla, but were entire in b, and were the origins of the 9th, 10th, and 11th cranial
nerves. Of these nerves the 11th or spinal accessory was large, and its spinal roots were
traced for a short distance along the side of the cord. The dorsal surface of the medulla
was hollowed out into the 4th ventricle, which was prolonged forwards on to the corre-
sponding surface of the pons.
Nothing is known of the development of the convolutions and fissures in the
cerebrum of the Walrus, and I have not met with any description of the order in which
the convolutions and fissures appear in the hemispheres of the Seals. Some years ago I
was presented by one of my pupils, Mr. T. G. Ker, with twin foetuses of Phoca grcen-
landica which he had extracted from the uterus of the mother, when acting as surgeon
on a ship engaged in the Seal fishing. The foetuses were preserved in rum, and after
they came into my possession I removed the brains. The length of the foetus from the
snout to the tip of the pes was 222 mm. The cerebrum was 20 mm. long and 22 mm.
wide. The cranial surface of each hemisphere was quite smooth, except that about the
junction of the anterior and middle thirds a very shallow furrow passed from the mesial
longitudinal fissure transversely outwards for 8 mm.; it seemed to be a slight tear on
the surface (PI. VIII. fig. 4). Low down on the outer side of the hemisphere a shallow
REPORT ON THE SEALS. 113
depression marked the position of the future fossa and fissure of Sylvius. With a pocket
lens one could also see the commencement of the differentiation of a Sylvian convolu-
tion about this fissure. On the tentorial surface of the hemisphere a shallower fissure
was also seen, which was probably the beginning of the splenial fissure.
I do not intend to enter into a discussion of the question whether the fissures on the
surface of the hemispheres are primarily due, either to unequal growth of the cortex
in different parts, or to unequal resistance offered to the growth of the cortex, or to
both causes acting in different parts of the same brain. I would, however, state that
in stripping off the pia mater from certain parts of the hemispheres of the brains
which I have dissected I have been struck with the tension and consequent pressure
exercised by the arteries on the surface of the cortex in the direction of their course.
This was well seen in the fissure of Sylvius occupied by the large middle cerebral
artery. Also in a less degree by the arteries which ran in the pia mater occupying the
great transverse fissure of the cerebrum, and which as they turned round the hippocampal
convolution undoubtedly indented its surface by their pressure.1 In these localities there-
fore there seems to be sufficient evidence to show that fissures may be produced and deepened
by the tension of the arteries, and doubtless the same cause operates also elsewhere.
Comparison of the Convolutions of the Seals and Walrus with
THOSE OF THE CaRNIVORA AND OF APES AND Man.
M. Leuret, in his well-known Anatomie comparee du Systeme Nerveux, both figures
and describes the cranial surface of the brain of a Seal, probably Phoca vitxdina. He
considers that the convolutions in this animal are analogous to those of the Ungulata,
especially the Pig, though without resembling them throughout, and in his arrangement
of the Mammalia, according to the grouping of their convolutions, he places the
Ungulata, Edentata, and Marsupialia between the Carnivora and the Seals. He recog-
nises only three convolutions in the hemisphere of the Seal — one internal, on the inner
surface, which is obviously the gyrus fornicatus or great limbic lobe of Broca ; one
external bounding the fissure of Sylvius and very irregular ; one stqierior extending
from before backwards on the top of the hemisphere and forming two tiers, with two
subdivisions in front and three behind, whilst he regards the supraorbital convolution as
only an offshoot of the two anterior subdivisions. Sir Richard Owen again has recog-
nised in the brain of Phoca a prefrontal lobe in front of the frontal crucial fissure ; an
orbital fold above the orbit ; Sylvian, supersylvian, medilateral, and medial folds or
convolutions arranged in tiers above the fissure of Sylvius ; it is obvious, however, from
1 Johannes Seitz has recently published an elaborate memoir (Ueber die Bedeutung der Hirnfurchung, Jahrbiicher
fiir Psychiatre, 1887) on the signification of the fissures in the hemispheres, in which he associates them with the places
of entrance and emergence of the blood and lymph vessels of the brain — that they are in fact nutrient fissures.
(ZOOL. CHALL. EXP. — PAKT LXVIII. — 1888.) Yyy 15
114 THE VOYAGE OF H.M.S. CHALLENGER.
his diagram of the outer surface of the hemisphere (op. cit., fig. 93, p. 118) that he does
not consider the interval of separation between the medial and medilateral folds to be
as definite as those between the other convolutions on the same surface. Krueg, who has
also studied the brain of Phoca vitulina, devotes his description to an account of the
fissures, and does not even name the convolutions. It would seem, however, both from
his description of the fissures and accompanying figures of the brain, that he only recog-
nises three tiers of convolutions on the outer surface of the hemisphere, whilst a well-
marked splenial fissure on the inner surface individualises the hippocampal and callosal
convolutions. Paul Broca's account of the brain of a Phoca is principally taken up with
a description of the great limbic lobe and its relation to the olfactory apparatus.
In the brain of Phoca vitulina, two specimens of which I have dissected, I found on
the outer surface of the hemisphere a distinct fissure of Sylvius, with its Sylvian
convolution, the anterior limb of which was narrower than the posterior, and at its com-
mencement concealed within the fissure of Sylvius. When this fissure was widely opened
out, prolongations of the Sylvian convolution were traced deeply into it, and occupied the
position of an insula. Above the Sylvian convolution were a suprasylvian fissure and
convolution, the latter of which showed at its summit a disposition to subdivide into
two parallel gyri for a short distance. This convolution was bounded above by a
lateral fissure, between which and the mesial longitudinal fissure were two slender con-
volutions running antero-posteriorly ; the lateral of these was apparently the mediolateral
convolution, whilst the medial one bounded the longitudinal fissure and was the sagittal
convolution ; as in Owen's figure, however, the fissure which separated the sagittal
(medial) from the mediolateral convolution was not continuous, but was bridged by short
annectent gyri. As this mediolateral fissure was imperfect and not prolonged far forward
in front, the coronal fissure was not continuous with it. In one instance the coronal was
prolonged backwards into the lateral fissure, but it might be separated from it by an
intermediate bridging convolution. In this region, therefore, the brain of Phoca vitulina
closely corresponded in the arrangement both of convolutions and fissures with the
Elephant Seal, though in the latter, from its greater size, the convolutions were bigger,
also I think more tortuous, and certainly with a greater number of bridging convolutions.
In the Walrus, again, the four tiers of convolutions were more definitely expressed
on the outer surface of the hemisphere, partly owing to the comparative absence of
bridging convolutions, and partly because the mediolateral fissure formed so definite a
plane of separation between the sagittal and mediolateral convolutions. In this animal
also the anterior limb of the Sylvian convolution was narrower, and sunken into the
fissure of Sylvius much more than either in Phoca or Macrorhinus, and from Dr. Murie's
description and plate of the brain of Otaria jubata (op. cit., fig. 40) it is obvious that
a corresponding depression occurred also in that of the Eared Seal. This narrowing and
depression were more marked than in Leuret's figures of the Brown Bear, Coati, and
REPOET ON THE SEALS. 115
Otter in pi. vi. of his Atlas. But I find that in the brain of an Otter (Lutra vulgaris),
of the Badger (Meles taxus), and Batel {Mellivora indica) in the University Museum a
similar sunken condition of this limb of the Sylvian convolution exists.
From the examination of these brains of Triehechus, Phoca, and Macrorhinus, I
am disposed to regard these animals as more or less approximating in the arrangement
of the convolutions of the outer face of the hemisphere to those Carnivora which
possess four tiers of convolutions in relation to the fissure of Sylvius. This arrangement
is found in the Dog, Jackal, Fox, and Wolf.1 From Dr. Murie's figures of the brain of
Otaria jvbata it would appear that in that animal, whilst the Sylvian and supra-
sylvian convolutions are quite definite, yet that the subdivision of the marginal convolu-
tion of the longitudinal fissure into mediosagittal and mediolateral convolutions is so
partial that the arrangement seems to be intermediate to that which one finds in
Triehechus and the Canidad on the one hand, and the Cats on the other. In regarding
this affinity in the general arrangement of the convolutions of the cranial surface of
the hemisphere in the Seals with those of the Canidae, it must be kept in mind that in
the Dogs the convolutions are less tortuous, and with fewer secondary fissures and
gyri than in the Pinnipedia.
The hemisphere of the cerebrum of Phoca vitulina possessed on the mesial and
tentorial surfaces a distinct gyrus fornicatus, or great limbic lobe, which was divided into
uncinate, hippocampal, and callosal convolutions, and was differentiated on its peripheral
side by the splenial fissure or the limbic fissure of Broca. This fissure was bridged in
its posterior part by a short retrolimbic gyrus, the pli de passage retrolimbique of
Broca. The splenial fissure had not always the same termination at its upper and
anterior end, for in the same brain I have seen it prolonged forwards into the crucial
fissure in one hemisphere, but in the other separated from it by a bridging convolution.
Both the suprasplenial convolution and fissure existed in the region above the corpus
callosum, though in one hemisphere the fissure was bridged by a short gyrus. Neither
the postsplenial fissure nor the splenial convolution was distinctly differentiated, and
the tentorial surface was subdivided into narrow convolutions. At its inferior end the
splenial fissure was continuous with the postrhinal fissure, and through it with the
transverse part of the fissure of Sylvius, across which it was prolonged into the rhinal
fissure, which defined the tuber olfactorium externally. The tuber was distinctly
prolonged into the uncinate gyrus across the bottom of the fissure of Sylvius. Imme-
diately to the outside of the connecting band between the tuber and uncinate gyrus
was the concealed portion of the anterior limb of the Sylvian convolution, which
apparently represented the Island of Reil. The supraorbital area possessed a gyrus
rectus, olfactory fissure, intraorbital fissure, internal and external supraorbital convolu-
tions. The olfactory peduncle was very slender, more so indeed than would be imagined
1 St* pi iv. iii Lenret ami Gratiolet's Atlas.
116 THE VOYAGE OF H.M.S. CHALLENGER.
from the figures published by Tiedemann and Leuret. The crucial fissure was at the
anterior end of the hemisphere, and about 14 mm. in its transverse diameter, and had
the usual relation to the sigmoid gyrus, with which the sagittal convolution was con-
tinuous. No prsecruciate fissure could be seen on the cranial surface of the cerebrum,
but, when the hemispheres were separated from each other, a short fissure was recognised
passing downwards from the crucial fissure, which apparently was the prascruciate fissure,
whilst the short convolution which it differentiated represented the ursine lozenge,
situated as Mivart has stated entirely on the mesial surface of the hemisphere. The
prorean convolution was continued into the gyrus rectus.1
The convolutions and sulci on the inner and tentorial surface of the hemisphere of
Macrorhinus corresponded in essential particulars with those of Phoca. Some differences
are, however, to be noted. Thus in Macrorhinus the splenial fissure was not continuous
with the postrhinal fissure, neither was it bridged across superficially by a retrolimbic
pli-de-passage, though there was a short gyrus projecting backwards from the hippo-
campal convolution which may represent it. In both hemispheres the splenial fissure
was continued into the crucial fissure ; the demarcation of the splenial from the sagittal
convolution by a continuous antero-posterior suprasplenial fissure was less marked in
Macrorhinus than in Phoca.
In the Walrus, also, the splenial and postrhinal fissures were not continuous with
each other. In one brain (a) (PI. IX. fig. 3) there was no retrolimbic bridging convolu-
tion, which was present however in both hemispheres of another specimen, and in one of
these hemispheres was represented by two convolutions. In two brains the splenial
fissure joined anteriorly the crucial fissure. The definition of the suprasplenial convolu-
tion and fissure varied in opposite hemispheres. Both brains possessed postero-horizontal
and postsplenial fissures and a splenial convolution. The olfactory peduncle and bulb
were larger than in the Phocidse.
In Otaria jubata, if I may judge from Dr. Murie's drawings of the brain of that
animal, the postrhinal and splenial fissures were not continuous with each other ; the
splenial fissure was bridged by a retrolimbic convolution ; the splenial fissure was not
prolonged directly into the crucial fissure ; the suprasplenial convolution and fissure were
not sharply differentiated ; the olfactor}^ apparatus was more like in size the same parts
in the Walrus than in the Phocidce.
Dr. St. George Mivart has recently introduced into the study of the brain in the
Carnivora and Pinnipedia the consideration of the area which he has named the Ursine
lozenge, and has pointed out that it constitutes a well-marked feature in the anterior part
of the dorsal surface of the cerebrum of Otaria gillespii. I have already stated that, in
both Phoca and Macrorhinus, but especially in the former, this area is rudimentary, and
concealed in the mesial fissure of the cerebrum. In the Walrus, again, the single con-
1 Theodor's essay on the brain of Phoca vitulina did not come into my hands until after this Report was in proof.
REPORT ON THE SEALS. 117
volution which represents this area was not definitely defined. Dr. Mivart attaches
much importance to the presence of the ursine lozenge in the Pinnipedia, as indicating
phylogenetic relations to the ursine group of the Carnivora.
I shall now compare the convolutions on the mesial and tentorial surfaces of the
hemisphere in the Pinnipedia with the corresponding surfaces in the brains of several of
the Canidse, and the brains which I have examined are those of the Dog, Jackal, and Fox.
In all these animals the postrhinal fissure joined the splenial fissure as in Phoca vitulina.
The splenial fissure on the tentorial surface was not bridged superficially by a retro-
limbic convolution. The lobus and the hippocampal and callosal divisions of the gyrus
fornicatus were definitely expressed. The splenial fissure was continued at its anterior
end into the crucial fissure, which was placed in the anterior third of the dorsum of the
hemisphere. In none of these brains was a suprasplenial convolution differentiated from
the sagittal convolution by a suprasplenial fissure, though in the Dog's brain an indica-
tion of such a fissure was present. The crucial fissure was bounded by the sigmoid
gyrus, which was continuous with the sagittal convolution. Immediately external to the
sigmoid gyrus was the coronal fissure, which was continued backwards into the medio-
lateral fissure, but not forwards into the prsesylvian fissure. There was neither prse-
cruciate fissure nor ursine lozenge. The olfactory peduncle was both relatively and
absolutely larger than in the Seals and Walrus notwithstanding the much smaller brain,
and the continuity of its large root with the lobus hippocampi was plainly marked across
the fissure of Sylvius.
In the Polar Bear (Ursus maritimus) the postrhinal fissure was deep, and passed
back towards the splenial fissure, but was separated from it by a slender retrolimbic
gyrus partially sunk in the fissure. The anterior end of the splenial fissure was not con-
tinuous with the crucial fissure, but bifurcated ; the posterior branch reached the dorsum
of the hemisphere as a sulcus in the ursine lozenge, the anterior passed horizontally
forwards in front of the knee-like bend of the callosal convolution. The suprasplenial was
not differentiated from the sagittal convolution by a suprasplenial fissure, although there
was an indication of such a fissure posteriorly. The tentorial surface possessed both a
postsplenial fissure and a splenial convolution. The ursine lozenge was large, being
34 mm. long by 42 mm. wide. It formed a large proportion of the anterior third of the
dorsum of the hemispheres, and was partially intersected by small sulci, one of which
was the posterior branch of bifurcation of the splenial fissure. The crucial fissure was
40 mm. long. The sigmoid gyrus which enclosed it was strongly developed, and. its
posterior bmb was continuous with the sagittal convolution. The coronal fissure was
behind and to the outer side of the posterior limb of the sigmoid gyrus, and was
prolonged backwards into the 1st curved fissure, but not forwards into the prsesylvian
fissure. The Polar Bear had three distinct convolutions above the Sylvian fissure. It
seemed at first as if they represented the Sylvian, suprasylvian, and marginal convolutions,
118 THE VOYAGE OF H.M.S. CHALLENGER.
and as the last named was partially divided by an antero-posterior fissure into two, it
looked as if it might represent both the sagittal and the mediolateral convolutions of the
Doo- and Walrus. On opening up the Sylvian fissure I found to my surprise that a
definite arched convolution was completely concealed within it. It was separated from
the convolution which bounded the Sylvian fissure by a deep fissure which was also
concealed. Its anterior limb, not quite so bulky as the posterior, was continued into
the supraorbital area immediately external to the rhinal fissure, and to the outer root of
the olfactory peduncle. Its posterior limb reached the postrhinal fissure and the lobus
hippocampi. I could not but think that we had here, more completely than either in
tbe Walrus or Seals, a sinking into the Sylvian fissure of the convolution which ought
to have bounded it, so that both the Sylvian convolution properly so called, and the
suprasylvian fissure, were concealed within it. If this be a proper explanation of the
arrangement, then the three convolutions on the cranial aspect would be sagittal, medio-
lateral, and suprasylvian ; whilst the two complete curved fissures between them would
be the mediolateral and lateral. The 1st curved fissure therefore into which the coronal
fissure is prolonged, would then as in the Dog be the mediolateral fissure. The olfactory
apparatus was large, and the external root formed a thick broad band of connection with
the lobus hippocampi, so that the Sylvian fossa was shallow.
In the Badger (Meles taxus) the postrhinal fissure was deep and prolonged towards the
splenial fissure, from which it was separated by a short retrolimbic gyrus ; anteriorly the
splenial fissure was continuous with the crucial fissure ; a short prsecruciate fissure marked
off a small ursine lozenge, consisting of a single convolution, and situated about the
junction of the anterior and middle third of the dorsum of the hemisphere. The supra-
splenial was not differentiated from the sagittal convolution. The crucial fissure was
18 mm. long, and bounded by a relatively large sigmoid gyrus, the posterior limb of
which was continuous with the sagittal convolution. Below and behind the sigmoid
gyrus was the coronal fissure, which was continued backwards into the 1st curved
fissure, but not forwards into the prsesylvian fissure. Only three convolutions sur-
mounted the Sylvian fissure, the anterior limb of the Sylvian convolution was partly
concealed in that fissure, the suprasylvian and marginal convolutions were distinct, and
the latter was not divided into a sagittal and a mediolateral convolution. The olfactory
apparatus was large.
In the Ratel (Mellivora indica) the postrhinal fissure was deep and separated from
the splenial fissure by a short and partially concealed retrolimbic gyrus. The callosal
convolution was relatively wide and closely resembled in its proportion the corresponding
convolution in the Otter as figured by Broca {pp. tit., fig. 1, p. 399). The splenial fissure
terminated a little in front of the middle of the dorsal surface of the hemisphere in
the crucial fissure ; a short prsecruciate fissure was also present, and between it and the
crucial fissure was a distinct ursine lozenge formed of a single convolution. The marginal
REPORT ON THE SEALS. 119
convolution was very narrow, and the suprasplenial convolution and fissure were absent,
though it is possible that this convolution was potentially present in the callosal con-
volution. The crucial fissure was 17 mm. long, and enclosed by a broad sigmoid gyrus
which was continuous by its posterior limb with the marginal gyrus. The coronal fissure
which bounded it was prolonged backwards into the 1st curved fissure, but not forwards
into the praesylvian fissure. Well-defined Sylvian and suprasylvian convolutions were
present, but only a slight indication of a division of the marginal convolution into
sagittal and mediolateral convolutions was visible. No arched convolution was con-
cealed within the Sylvian fissure. The olfactory apparatus was large.
My dissection of the inner and tentorial surface of the hemisphere of the Otter (Lutra
vulgaris) closely accords with Paul Broca's figures and description.1 In this animal the
crucial fissure was 14 mm. long; the sigmoid gyrus was relatively large; the coronal
fissure was not continuous with the prsesylvian fissure ; Sylvian, suprasylvian, and mar-
ginal convolutions were present ; the anterior limb of the Sylvian was almost entirely
concealed in the fissure, and there was evidence of separation of the marginal convolution
into sagittal and mediolateral by a short mediolateral fissure which was interrupted ; but
the coronal fissure should be regarded as prolonged into the fissure bounding the
upper aspect of the suprasylvian convolution, which may therefore be termed lateral.
In the Coati (Nasua rufa) the postrhinal was separated from the splenial fissure by a
short retrolimbic gyrus ; the splenial did not join the crucial fissure, but terminated
behind it in a sulcus in the sagittal convolution, which did not reach the margin of the
hemisphere. The marginal part of the sagittal convolution was relatively wider than
in the Otter and Eatel. The crucial fissure was distinct, but owing to an injury to this
part of the brain, I could not speak with certainty of the presence of a prsecrueiate
fissure leading forwards and inwards from the crucial fissure ; a small convolution in
front of the crucial fissure apparently represented the ursine lozenge, a convolution
which Mivart also considers to exist in the brain of this animal. Only three tiers of
convolutions were present.
In the Weasel (Mustela vulgaris) the postrhinal fissure was separated from the
splenial by a retrolimbic gyrus which was broad in relation to the size of the hemisphere.
The splenial fissure ended in the crucial fissure on the dorsum of the hemisphere.
No prascruciate fissure was visible on the dorsum, but on opening up the crucial fissure
a very short sulcus indented the convolution which formed the boundary of the crucial
fissure and marked off the anterior boundary of a minute ursine lozenge. In the Ferret
(Mustela furo) , however, a short but distinct praacruciate fissure differentiated the anterior
boundary of a minute ursine lozenge. The splenial fissure ended in the crucial fissure on
the dorsum of the hemisphere. The splenial was separated behind from the postrhinal
1 Figures of the ursine lozenge in the brains of Ursus maritimns and Mcllivora indica have been given by St. George
Mivart in his memoir already quoted, and its presence in the brains of the Otter, Badger, Coati, and other Arctoid
Carnivora is described by him.
120 THE VOYAGE OF H.M.S. CHALLENGER.
fissure by a short retrolimbic gyrus. In both the Weasel and Ferret the marginal part
of the sagittal convolution was much narrower than the callosal convolution. In both,
also, the olfactory apparatus was largely developed. In the Coati, Weasel, and Ferret,
the relations of the sigmoid gyrus to the coronal fissure, and of that fissure to the 1st
curved fissure, closely corresponded to the arrangement in the Badger and Eatel.
I have examined in the Felidse the tentorial and mesial surfaces of the hemisphere
in the brains of the common Cat {Fells domesticus) and the Tigef (Felis tigiis). In the
Cat one retrolimbic gyrus, and in the Tiger two, separated the splenial from the post-
rhinal fissure, and in the latter a third bridging convolution crossed the splenial fissure
immediately behind and above the splenium. In both, the crucial fissure was situated
in the anterior part of the dorsum of the hemisphere, and was not joined by the splenial
fissure, which in both animals reached the margin of the hemisphere behind the crucial
fissure. In neither animal was there an ursine lozenge. In the Tiger the convolutions
were more subdivided by secondary fissures than in the Cat, and on the tentorial surface
both a postsplenial fissure and a splenial convolution were present. Both animals had
a large olfactory apparatus connected by a strong tract with the uncinate convolution.
In the common Cat the coronal fissure was short and cut off by an intermediate
narrow gyrus from the prsesylvian fissure in front and the 1st curved fissure behind ; it
bounded the sigmoid gyrus externally. In the Tiger, in which the sigmoid gyrus was
large and tortuous, the coronal fissure formed its outer boundary, and though not pro-
longed forward into the prsesylvian fissure, it was continued backwards into the 1st curved
fissure. In both the Cat and Tiger the sagittal convolution was continuous with the
posterior limb of the sigmoid gyrus.
It is well known that in the Felidse the differentiation of the convolutions on the
cranial surface of the hemisphere into four tiers is not so precise as in the Canidas. The
convolution which bounds the Sylvian fissure is, in all probability, homologous in both
families. In the Tiger the suprasylvian convolution was differentiated in its whole length
from the Sylvian convolution by the suprasylvian fissure, and from the sigmoid gyrus
and sagittal convolution by the 1st curved fissure. There was no distinct mediolateral
convolution, but a convolution which might represent it was partially differentiated from
the sagittal convolution by an imperfect mediolateral fissure.. In the common Cat the
sagittal and the 2nd external convolution were distinctly differentiated from each other
by an intermediate fissure, but the Sylvian and suprasylvian convolutions were partially
blended together, especially in their posterior limbs.
In the series of brains examined the coronal fissure was seldom continued forward
into the prsesylvian fissure, but it was very frequently prolonged backwards into one of
the curved fissures on the cranial aspect of the hemisphere,1 though sometimes it was
1 In Leuret's figure of the brain of the Lion, the coronal fissure is continuous with the 1st curved fissure, but in
Victoria Familiant's figure of the brain of this animal these fissures are separated from each other, as in the common
Cat, by an intermediate bridging convolution.
. REPORT ON THE SEALS. 121
interrupted by a short bridging convolution. When prolonged into a fissure it joined
that which lay next to the marginal convolution or the 1st curved fissure. But this was
not necessarily morphologically the same in all these brains. Where four tiers of con-
volutions were differentiated, it was, of course, the mediolateral fissure, but when only
three tiers were differentiated, then it probably represented the lateral fissure, as in these
brains both the mediolateral fissure and convolution were eithei absent or only imperfectly
differentiated. The coronal fissure formed the outer boundary of the sigmoid gyrus. The
coronal gyrus wras the anterior part of the 2nd external convolution, which in those
brains that possessed four tiers of convolutions "was the mediolateral convolution ; but,
when only three tiers were present, it was most probably represented by the suprasylvian
convolution.
The crucial fissure varied materially in its position in the genera of the Carnivora
and Pinnipedia. In the Seals and Walrus it was so far forward as not to be seen on
the dorsum of the hemispheres, but only at the anterior end of the cerebrum. In the
Cat and Tiger it was visible in about the anterior fourth of the dorsum of the hemi-
spheres ; in the Dog, Weasel, Ferret, and Coati at about the junction of the middle and
anterior third ; in the Badger, Polar Bear, and Ratel it was even further back, so as to be
just in front of a line dividing the dorsum of the hemispheres into an anterior and a
posterior half. This variation in the position of the fissure necessarily affected that
of the sigmoid gyrus which bounded it in front, behind, and on the outer side, and in
those brains in which the fissure was elongated and far back, this gyrus formed a well-
marked convolution on the dorsum of each hemisphere. When the crucial fissure was
elongated both it and the sigmoid gyrus were continued downwards on the outer surface
of the hemisphere,1 and the direction of the coronal fissure, which formed the outer
boundary of the sigmoid gyrus, was from below obliquely upwards and backwards.
It will now be of interest to compare the convolutions of the cerebrum in the Carnivora
and Pinnipedia with those in Man and Apes, with the view of endeavouring to ascertain
if any correspondence in their arrangement exists, and to what extent, in these orders of
Mammals. The importance of instituting this comparison has already, indeed, presented
itself to several anatomists, and various attempts have been made to harmonize the
arrangement of the convolutions of the Carnivora with those of Man and Apes. The
desirabdity of arriving at some definite conclusion on this matter is owing both to the
interest of the subject from a purely morphological point of view, and to its physiological
value in connection with the numerous experiments which have of late years been made
for the determination of the functions of the cerebral cortex.
It wdl be obvious, if in the brains of these different orders one or two leading fissures
1 The anterior limb of the sigmoid gyrus is sometimes called gyrus prxcruciatus (pr&frontalis), the posterior limb
gyrus postcruciatus {postfrontaKs).
(ZOOL. CHALL. EXP. — PAET LXVIII. — 1888.) Yj'J' 16
122 THE VOYAGE OF H.M.S. CHALLENGER.
and convolutions can be identified as without question morphologically alike in their
development and relations, that a certain basis would be obtained from which it may be
possible to extend the comparison to other parts of the surface. A most important
investigation conducted in accordance with this method was published by Dr. Paul Broca,
on Le grand lobe linibique et la scissure limbique. In the course of this memoir he
reviewed the arrangement of the limbic lobe or convolution, and showed that it can be
identified throughout the mammalian series. It consists of a callosal and hippocampal
portion with a lobus hippocampi, and forms the boundary both of the corpus callosum
and the transverse fissure of the cerebrum. Moreover it is continuous with the roots of the
olfactory lobe, more especially through the lobus hippocampi, though the band of union
varies materially in thickness in the brains of different orders. In the proper Carnivora,
for example, the connecting band is large and very distinct, in the Pinnipedia it is less
marked, and in Man and Apes it is reduced to a fine thread.
In the Carnivora proper the rhinal fissure is distinct and continued across the fissure
of Sylvius into the postrhinal fissure, which again is prolonged towards the splenial fissure,
though frequently the exact continuity is interrupted by a superficial retrolimbic con-
volution. In Phoca the rhinal and postrhinal fissures resemble those in the Carnivora
proper, though relatively they are somewhat smaller. In Macrorhinus and Trichechus
the retrolimbic convolution is nearer the lobus hippocampi, so that the postrhinal
fissure is shorter. In Man and Apes, owing to the absolute and relative diminution in
size of the olfactory apparatus, the rhinal fissure is scarcely recognisable, and the
postrhinal fissure cannot be said to be continuous with it.
The limbic lobe is differentiated on its peripheral aspect by the fissure which has
been named the splenial fissure in the brains of the Carnivora and Pinnipedia described
in this Report. In Man and Apes the calloso-marginal fissure represents that part
of the splenial fissure placed peripherally to the callosal convolution, whilst the collateral
(occipito-temporal) fissure is apparently the representative of that part of the splenial
fissure which forms the peripheral boundary of the hippocampal convolution.
In the larger Carnivora and the Pinnipedia the supraorbital area possesses an olfactory
sulcus, a gyrus rectus, an intraorbital fissure, internal and external supraorbital con-
volutions ; though in the brains of the smaller Carnivora, especially when the olfactory
apparatus is relatively large, these fissures and convolutions are scarcely if at all
differentiated. In Man and Apes these parts are also seen, and the intraorbital fissure,
from so frequently trifurcating, was named by me the triradiate fissure.
The fissure of Sylvius forms a recognisable feature in the brains of the Carnivora, but
where it begins as the Sylvian fossa on the under surface of the brain, it is usually shallow,
owing to the thickness of the olfactory root which passes backwards to join the lobus
hippocampi. In the Pinnipedia, and still more in Man and Apes, owing to the diminished
size of this root, the Sylvian fossa is much deeper. In the Carnivora and Pinnipedia
REPORT ON THE SEALS. 123
tlie fissure of Sylvius passes upwards and with only a slight inclination backwards on the
cranial surface of the hemisphere, its length being regulated by the length of the Sylvian
convolution which bounds it. In Man and Apes, more especially the Anthropoids, it
is longer than in the Carnivora, and passes upwards and with a marked inclination back-
wards ; its backward direction being more decided in Man than in the highest Apes.
Up to this point there does not seem to be much difficulty in finding a morphological
correspondence between the fissures and convolutions in the brains of these orders of
Mammals, but beyond this stage many difficulties undoubtedly present themselves. In
the human brain, for example, the magnitude and direction of the convolutions of the
frontal lobe, the fissure of Rolando, the parieto-occipital fissure, the definite occipital lobe
lying behind that fissure, the calcarine fissure, the elongated convolutions of the temporo-
sphenoidal lobe, and the convolutions of the insula are all characteristic features, which
are repeated though in a less pronounced form in the brains of Apes except that in the
latter the distinctness of the occipital lobe is more accentuated. In the Carnivora and
Pinnipedia again the presence of three or four tiers of convolutions with their inter-
mediate fissures surmounting the fissure of Sylvius, the existence of a crucial fissure,
and also in many genera of a prsecruciate fissure and ursine lozenge, are noticeable
characteristics, and at first sight seem so divergent from the human arrangement as to be
apparently irreconcilable with it.
In the Human cerebrum four elongated convolutions running obliquely from above
downwards and forwards intervene between the fissure of Sylvius on the cranial surface
and the gyrus and lobus hippocampi on the tentorial surface ; viz., the superior, middle,
and inferior temporo-sphenoidal convolutions, and the occipito-temporal convolution. In
the Ape's brain the differentiation of the three temporo-sphenoidal convolutions is more or
less distinct in various species, but the occipito-temporal convolution is frecpuently not
differentiated from the inferior temporo-sphenoidal gyrus.
The apparently corresponding region in the brain of the domestic Cat is short and
stunted, but in the larger brain of the Tiger it is more elongated ; in the Dog's brain it
is a little longer than in the Cat ; in Phoca, Macrorhinus, and Trichechus it is also well
marked and the convolutions are tortuous. In these Carnivora and Pinnipedia three
convolutions lie behind the fissure of Sylvius on the cranial aspect of the hemisphere, for
they are almost vertical in direction and the most posterior forms the boundary of the
hemisphere at the junction of its cranial and tentorial surfaces. These convolutions are
the posterior limbs of the tiers of convolutions which surmount and arch above the fissure
of Sylvius. In the larger Carnivora and Pinnipedia a fourth convolution, varying in its
degree of differentiation, but not recognisable in the brains of the smaller Carnivora, is
situated on the tentorial surface peripherally to the hippocampal convolution, and
separated from it by the splenial (limbic) fissure, which fissure is usually bridged by the
retrolimbic convolution. In their relations to the Sylvian fissure on the one hand, and
124 THE VOYAGE OF H.M.S. CHALLENGER.
to the gyrus and lobus hippocampi on the other, these four convolutions in the Carnivora
and Pinnipedia might seem at first sight as if they approximated to the temporo-
sphenoidal and occipito-temporal convolutions in Man and Apes, though in Man they are
greatly elongated and approach the horizontal in their direction, in conformity with the
direction of the fissure of Sylvius. Moreover, they project in front of the uncus or
lobus hippocampi so as to form the tip of the temporo-sphenoidal lobe and the greater
part of the boundary of the Sylvian fossa, so that the lobus hippocampi with the short
postrhinal fissure is not visible at the base of the human brain, but is displaced inwards
on to the tentorial aspect. But further, in the brains of the Carnivora and Pinnipedia
the lobus hippocampi appears as a distinct protuberance on the base of the brain, and
itself forms the posterior boundary of the Sylvian fossa. These differences in the two
types of brain might seem to be accounted for simply by the great development and the
change in direction of the convolutions of the temporo-sphenoidal lobe in the brains of
Man and Apes, causing in them displacement of the lobus hippocampi to the inner
surface of the hemisphere, and its concealment, when the hemisphere is looked at from
the cranial aspect, by the greatly elongated temporo-sphenoidal convolutions.
But I am of opinion that this does not express the whole difference between these
brains in this region. In the description of the brain both of the Walrus and the Seals
I have indicated that the Island of Reil may find its representative in these animals in
the anterior limb of the Sylvian convolution, which is more or less concealed within the
fissure of Sylvius ; and in the brain of the Polar Bear I have shown that an entire
arched convolution is concealed within that fissure. If I am right in this indication,
then I believe that the Island of Reil, which in the brain of the Ape and still more in
that of Man is entirely concealed within the Sylvian fissure, is either the homologue of
the Sylvian convolution of the carnivorous brain, or that the Sylvian convolution in
the Carnivora potentially represents both that convolution and a rudimentary insula.
In the true Carnivora the Sylvian convolution was as a rule superficial and on the
cranial aspect, though in the Otter and Badger indications of the depression of its
anterior limb within the fissure were seen. In the Seals and Walrus the concealment
of this convolution was still more marked, so that the brains of these animals form
apparently in this particular a transition to those of Man and Apes, in which the
concealment of the Island is complete. On the supposition therefore that the Island
of Reil in Man and Apes is morphologically related to the Sylvian convolution of
the Carnivora, the superior temporo-sphenoidal convolution in the Human and Ape's
brain cannot be regarded as corresponding with the posterior limb of the Sylvian con-
volution, but with that of the convolution of the tier immediately above and behind the
Sylvian convolution, i.e., the 3rd external convolution of Ferrier or the suprasylvian
convolution of my description. The sinking of the Sylvian convolution into the fissure
may perhaps to some extent be associated with a diminution in magnitude of the
REPORT ON THE SEALS. 125
olfactory apparatus. When that is large the Sylvian fissure is shallow, but when the
olfactory peduncle and roots diminish in size, as in the Seal and Walrus, the fissure
deepens and the Sylvian convolution becomes partially concealed, untd in Apes and
Man, with a still greater diminution in the importance of the olfactory sense, the fissure
attains its maximum depth. In this connection, however, it must be remembered that
the Polar Bear, though with an arched convolution concealed within the Sylvian fissure,
yet possesses large olfactory nerve roots.
This view of the homology of the convolutions in this region enables one to harmonize
the results of physiological experiment with anatomical arrangement, and to remove a
difficulty which is experienced so long as the superior temporo-sphenoidal convolution
is regarded as corresponding with the posterior limb of the Sylvian convolution. Dr.
Ferrier, from his experiments, determined that the areas marked (14) in his figures
were the auditory centres. Thus when these areas in the superior temporo-sphenoidal
convolution were stimulated in Monkeys the opposite ear became pricked, the head and
eyes were turned to the opposite side and the pupds became widely dilated ; whilst
stimulation of areas (14) in the 3rd external convolution of the brain of the Dog and
Jackal also produced a pricking or retraction of the opposite ear, and stimulation of a
similar area in the Cat produced both pricking of the opposite ear and turning of the
head and eyes to the opposite side. Hence these areas in the carnivorous and Ape's
brain are regarded as physiologically the same ; though in the Ape the convolution
stimulated bounds the Sylvian fissure, whfist in the Carnivora it is separated from that
fissure by an intermediate convolution. On the theory that the Sylvian convolution
either becomes the Island of Eeil or blends with the insula and sinks into the fissure, the
3rd external convolution would then become the boundary of the fissure and its posterior
limb would be homologous with the superior temporo-sphenoidal convolution of the brain
of Man and Apes, whilst the suprasylvian or 3rd curved fissure would become lost in the
Sylvian fissure, and be represented by the sulcus insulae. This theory is somewhat
different from the conception of the relation of parts in this region entertained by
Ferrier, who suggests that the Sylvian convolution is in the Monkey's brain represented
within the lips of the Sylvian fissure, overlapping and concealing the Island of Reil.
Ferrier has also shown that electrical stimulation of the posterior limb of the Sylvian
convolution gives no definite reactions, and similarly stimulation of the Island of Reil
is not followed by movements except after increased irritation, when some move-
ments of the mouth and tongue occur, which he considers may be due to conduction
of the stimulus to the motor areas situated immediately anterior to the part irritated.
Ordinary stimulation in both instances therefore produces no definite results, showing
that neither of these convolutions responds to the electrical stimulus, and although the
experimental result is negative, it is certainly not adverse to the view that they are
homologous with each other. Confirmation of this theory is also furnished by the fact
126 THE VOYAGE OF H.M.S. CHALLENGER.
that stimulation of the area marked (16) on the lower end of the anterior limb of the
Sylvian convolution in Dr. Ferrier's figures of the brain of the Dog and Cat is occasionally
associated with movements of the lips, whilst similar movements are produced by irrita-
tion inside the fissure of Sylvius in the Monkey,1 doubtless due therefore to irritation of
the areas (9) and (10) which lie in proximity to the fissure.
To harmonize the arrangement of the convolutions of the frontal, parietal, and
occipital lobes of the human and Ape's brain with the tiers of convolutions which in the
Carnivora surmount the fissure of Sylvius, is undoubtedly a task of some difficulty.
Several anatomists have, however, attempted to do so. M. Broca, in his memoir already
quoted, has argued with great emphasis, that in the brain of the Primates the character
which dominates over all others in importance is the enormous development of the
frontal lobe, from whence results the backward position and the obbque direction of the
fissure of Rolando. The position which I took up many years ago2 that the fissure of
Rolando, or central fissure, should be regarded as forming the posterior limit of the frontal
lobe and the plane of demarcation between it and the parietal lobe, is now generally
accepted. It becomes therefore a matter of some moment to determine if possible the
fissure in the carnivorous brain which corresponds to the fissure of Rolando in Man and
Apes, the oblique and backward direction of which must be borne in mind.
Broca regarded the fissure of Rolando as represented in the Carnivora by the prae-
sylvian fissure, so that he practically confined the frontal lobe in these animals to the
region in front of and below that fissure, which has been named in this Report the supra-
orbital area. Schwalbe is apparently inclined to attach some weight to this view ; but
owing to the divergence in development of the Carnivora and Ungulata on the one hand,
and the Primates on the other, he does not consider it possible to make a strict comparison
between the convolutions and furrows of these orders of Mammals. I believe that the
limitation of the frontal lobe to the area in front of the prsesylvian fissure would be too
great a restriction of that lobe, which on developmental and other grounds may, I think,
be shown to extend further back in the hemisphere.
At the first glance there might seem to be a strong likeness between the crucial
fissure in the carnivorous brain and the fissure of Rolando. They are both directed more
or less vertically and transversely downwards on the cranial surface of the hemisphere, and
each is bounded in front and behind by a gyrus having a corresponding direction ; in the
Carnivora the gyri are the anterior and posterior limbs of the sigmoid gyrus ; in Man and
Apes they are the ascending frontal and parietal convolutions. These general resemblances
have led more than one anatomist to regard them as homologous. But in discussing the
homology of the crucial fissure it is important to attend to its relative period of ajjpearance
1 I am indebted to Dr. Ferrier for this information, which he wrote to me in reply to a request as to the area in the
Monkey's brain which corresponds to (16) in the brain of the Dog.
2 The Convolutions of the Human Cerebrum topographically considered, Edinburgh, 1866, p. 11. Notes more
especially on the Bridging Convolutions in the Brain of the Chimpanzee, Proc. Roy. Soc. Edin., 19th Feb. 1866, vol. v. p. 578.
REPORT ON THE SEALS. 127
on the surface of the cortex, and to its relations to the splenial fissure on the mesial
aspect of the hemisphere. There can, I think, be no doubt that the anterior and upper
part of the splenial fissure in the brains of the Carnivora and Pinnipedia corresponds
with the fissure which is known as calloso-marginal in Man and Apes. Both the
splenial and calloso-marginal fissures are separated from the corpus callosum by the
callosal convolution of the limbic lobe, and each runs in this part of its course about mid-
way between the corpus callosum and the free upper margin of the hemisphere. In the
Canidse, the Badger, Eatel, Weasel, Ferret, Elephant Seal, and Walrus the splenial
fissure was continuous with the crucial fissure, but in the Cat, Tiger, Coati, and Polar Bear
they were not continuous ; whilst in a Phoca vitulina the two fissures were continuous
in the one hemisphere, but not in the other. In those cases in which the fissures were not
continuous, the splenial ended in or near the margin of the mesial longitudinal fissure
in proximity to the crucial fissure and usually a little behind it.
In the human brain the calloso-marginal fissure turns round the genu of the corpus
callosum and then runs backwards about midway between the corpus callosum and the
margin of the great longitudinal fissure ; when a little in front of the splenium of the
corpus callosum it bends upwards to reach the margin of the hemisphere somewhat behind
the fissure of Bolando. Where it makes this bend a fissure is prolonged for a variable dis-
tance backwards from it, but does not reach the collateral fissure, for it is so interrupted
by convolutions in this region which are continuous with the precuneus or quadrilateral
lobule, that the callosal convolution loses immediately above the splenium its sharp line
of demarcation superiorly. Both in the human brain and that of the Ape the limbic
lobe, where the callosal and hippocampal convolutions approach each other, possesses a
less definite differentiation peripherally than is the case both in the Carnivora and in
Mammals generally, a condition which is apparently clue to the much greater develop-
ment of bridging convolutions at its splenial end. In the Walrus, for example, the
bridging convolution in this region (PI. IX. fig. 3) is a single narrow gyrus, whilst in
the human and Ape's brain they correspond to the broad base of the preecuneus. Not
unfrequently I have seen one or more short fissures arise from the calloso-marginal about
opposite the genu and indent transversely the superior frontal convolution at the anterior
end of the cerebrum, which was bent around each fissure like a short sigmoid gyrus.
In its direction and relation to the calloso-marginal fissure any one of these fissures
resembled the crucial fissure, but cannot be morphologically the same as the fissure of
Rolando, which is situated much further back on the side of the hemisphere, and which
has no definite relation with the calloso-marginal fissure. It is obvious that the crucial
fissure is not of primary importance, as it is not always present in gyrencephalous
Mammals, and in those Carnivora, such as the Dog and Cat, in which its development
has been examined, it has been shown by Pansch to appear subsequently to the splenial
fissure having assumed a certain depth, so that it has only a secondary value.
128 THE VOYAGE OF H.M.S. CHALLENGER.
We must therefore look elsewhere in the carnivorous brain for the homologue of the
fissure of Eolando.
Owen, from a comjiarative survey of the brain in a large number of gyrencephalous
Mammals, was led to regard the coronal fissure in the Caruivora as corresponding with
the fissure of Rolando in Man, which he also called the coronal fissure. Pansch and
Meynert took a similar view of the homology of the coronal fissure, and Pansch held that
the fissure of Rolando was the anterior part of the upper or first curved fissure and was
the second in order of position from before backwards of the three primary fissures which
occur on the cranial surface of the hemisphere of all Mammals with convoluted brains.
In the Primates they were placed radially above the Sylvian fissure, but in other
Mammals the first or most anterior was more vertical, whilst the second and third had
a sagittal direction. Pansch's determination of the homology was based on the relative
period of appearance and on the depth of the fissure in various Mammals, and guided by
these considerations he regarded the ascending frontal and parietal (or the central)
convolutions as having their morphological equivalents in the anterior parts of the 4th
and 3rd convolutions of Leuret, i.e., the sagittal and mediolateral convolutions of the Dog
or the 1st and 2nd convolutions of Ferrier.
Ferrier, who was at one time disposed to agree with the anatomists who looked upon
the fissure of Rolando as the homologue of the crucial fissure, now holds with those who
consider it to be represented by the coronal fissure, and he supports his present opinion
by the result of his experiments on the cerebral cortex in Monkeys and Carnivora.
From a comparison of these results it would seem that a number of the effects produced
are reconcdable with the view that parts of the brain in front of the fissure of Rolando
and of the coronal fissure in these animals are physiologically homologous. Thus the area
marked (12) by Ferrier, which in the Monkey includes the posterior half or two-thirds
of the superior and middle frontal convolutions, and in the Dog is situated on the anterior
limb of the sigmoid gyrus, when stimulated occasioned in both animals wide opening of
the eyes, ddatation of the pupils, and turning of the head and eyes to opposite sides ;
stimulation of the area (3), which in the Monkey lies in the upper end of the ascending
frontal convolution close to its sulcus, and in the Dog in the 1st external or sagittal con-
volution just behind the crucial sulcus, produced in both animals movements of the tail ;
stimulation of the area (4), situated in the Monkey in the upper end of the ascending frontal
and anterior margin of the adjacent part of the ascending parietal convolution, and in
the Dog in the back of the posterior limb of the sigmoid gyrus, produced corresponding
movements in the fore limbs of both animals ; stimulation of (5), situated in the Monkey
in front of (3) where the superior frontal joins the ascending frontal, and in the Dog in
the sigmoid gyrus about opposite the outer end of the crucial fissure, occasioned in both
animals an extension forward of the opposite fore limb. It will also be observed that the
area marked (12) in both animals was the most anterior region to respond to electrical
REPORT ON THE SEALS. 129
stimulus. These experiments all indicate a homology between both limbs of the sigmoid
gyrus in the Dog and the ascending, superior, and middle frontal convolutions in the
Monkey, which is incompatible with the view that the crucial fissure is the homologue of
the fissure of Eolando, but quite reconcdable with the theory that the coronal fissure
and fissure of Rolando are homologous ; for in the respective 1 trains both the coronal and
Eolando's fissures lie behind the areas stimulated, with the exception of a small part
of (4), which just touches the ascending parietal convolution.
On the other hand, it is more difficult to reconcile some other of Ferrier's experiments
with this conclusion as to the homology of the two fissures. For stimulation of area (1),
placed in the Monkey in the postero -parietal convolution just in front of the parieto-
occipital fissure, and in the Dog in the posterior limb of the sigmoid gyrus just behind the
crucial fissure, produced in both animals an advancement of the opposite hind limb as in
walking ; in the Monkey the area stimulated was distinctly behind the fissure of Rolando,
in the Dog well in front of the coronal fissure. Again, the areas (7) (8), which when
stimulated gave rise to movements of the zygomatic muscles and upper lip, he in the
Monkey in the ascending frontal convolution, and therefore anterior to the fissure of
Rolando ; but in the Dog the one is situated in the coronal part of the 2nd external
convolution, the other in the anterior composite convolution formed by the junction of
the anterior ends of the 2nd and 3rd external convolutions, and both therefore are behind
the coronal fissure.
If we regard, however, the whole evidence based on comparative anatomy, on the
depth and relative time of appearance of the fissures and on the results obtained by
stimulating the brain in front of the fissures, we may, I think, fairly assume the
fissure of Rolando to be homologous with the coronal fissure in the carnivorous brain.
The sigmoid gyrus with the adjoining part of the sagittal convolution, and in animals
which have an ursine lozenge that area also, would therefore represent the superior,
middle, and ascending frontal convolutions in the brain of Man and Apes.
But there are other fissures in the brains of these Mammals the homologies of which
it is desirable, if possible, to determine. A well-marked fissure in the carnivorous brain
is the praesylvian or supraorbital fissure. It is the most anterior of the three primary
fissures described by Pansch as appearing on the cranial surface of the brain of the fcetal
Dog, and it separates in this animal the anterior limbs of the four tiers of convolutions
from the supraorbital area and the prorean convolution. If we place side by side the
hemispheres of the Human and the Dog's brain we can see in the former two fissures, the
prsecentral fissure of Ecker and the ascending branch of the Sylvian fissure, one or other of
which would appear to represent the prassylvian fissure in the Dog. Meynert was of opinion
that the praesylvian fissure is homologous with the ascending branch of the Sylvian fissure.
Broca objected to this * on the ground that the ascending branch, whilst present in Man
1 Sur le cerveau du Gorille, Revue d 'Anthropologic, s£r. 2, t. i.
(ZOOL. CHALL. EXP. — PART LXVIII. — 1888.) - Yyy 17
130 THE VOYAGE OF H.M.S. CHALLENGER.
and the Anthropoid Apes, is absent in the non-anthropoid Apes ; and that it is not likely
that a character which is only found in the most highly developed brains of the Primates
should, when absent in the lower Apes, reappear in the Carnivora in which the frontal
lobe is only rudimentary. In connection with this matter it should be stated that the
area (8), which Ferrier associates with elevation of the upper lip so as to display the
canine teeth, is situated in the Monkey in the lower part of the ascending frontal con-
volution and in the Dog at the anterior composite end of the 2nd and 3rd external
convolutions ; whilst the area (9), which he associates with the opening of the mouth,
movements of the tongue, and not unfrequently barking or growhng, is situated in the
Monkey in the lowest part of the ascending frontal convolution, where the inferior
frontal convolution springs from it ; and in the Dog in the composite convolution
formed by the junction of the lower ends of the anterior limbs of the 3rd and 4th
external convolutions. Areas (8) and (9) are placed therefore in the Monkey
immediately behind the prsecentral fissure and in the Dog immediately behind the prse-
sylvian fissure.
The results obtained by experiment would seem therefore to harmonize with the
conclusion founded upon more purely anatomical data, and I think it probable that the
praesylvian and praecentral fissures are homologous.
A number of years ago I described in the brain both of the Chimpanzee and of Man '
a fissure within the parietal lobe which I named the intraparietal fissure.2 It is
situated in the first instance behind and parallel to the ascending parietal convolution,
and then runs almost horizontally backwards to separate the ascending and postero-
parietal convolutions from the supramarginal gyrus or convolution of the parietal
eminence, and it may be seen in the brain of a sixth month's human fcetus. Pansch
subsequently recognised the importance of this fissure and regarded it as the third
and most posterior of the three primary fissures on the cranial surface of the brain ; he
believed it to be homologous with the anterior part of the middle or second curved
fissure of the Dog — the lateral fissure of this Report.
In the Human brain the intraparietal fissure is separated from the Sylvian fissure by a
convolution which, under the name of supramarginal gyrus, or, as I have termed it, the
convolution of the parietal eminence,3 forms a single tortuous tier. In the Dog, again,
two tiers lie between the lateral and Sylvian fissures, viz., the suprasylvian and Sylvian
convolutions, separated from each other by the suprasylvian fissure. Pansch makes
no attempt to explain this difference, and, in the absence of such explanation, diffi-
culties at once suggest themselves as to accepting his view of the homology of the intra-
parietal fissure and the anterior part of the 2nd curved fissure. But, on the theory
1 See niyrnernoirs already quoted, pp. 95, 126.
2 Ecker and some other anatomists have misnamed it the interparietal fissure.
3 Relations of the Convolutions of the Human Cerebrum to the Outer Surface of the Skull and Head, Journ. of
Anat. and Phys., voL viii. p. 142.
REPORT ON THE SEALS. 131
which I have expounded on p. 124, e.s., that the Sylvian convolution of the Dog subsides
in Man and Apes into the Sylvian fissure, and that the suprasylvian and Sylvian fissures
become, as it were, thrown into one, then one can account for the presence of only a
single convolution on the lower aspect of the intraparietal fissure, which convolution
represents the anterior limb of the suprasylvian convolution of the Dog, and is the supra-
marginal convolution in the higher brains. Ferrier's experiments also to some extent
bear out this view, for his area (11), stimulation of which produces retraction of the
angle of the mouth, is situated in the Monkey partly in the lower end of the ascending
parietal, but more so on the adjoining supramarginal gyrus, and in the Dog in the
anterior limb of the suprasylvian convolution.
One of the most noticeable fissures of the cerebrum of Man and Apes is that which
under the name of parieto-occipital fissure separates the parietal from the occipital lobe.
In the human brain it is as a rule more strongly marked on the mesial than on the
cranial aspect, owing to the development on the cranial surface of strong bridging
convolutions which pass across its upper end. In the brains of Apes it is as well marked
on the one surface as on the other, though in the brain of both the Orang and Chimpanzee
superficial bridging convolutions sometimes obscure its upper end.1
Almost all WTiters have stated that this fissure is absent in the brains of the
Carnivora, so that in them the occipital lobe is not differentiated from the parietal part
of the brain. In a recent memoir, however, Max Flesch has described in the brain of the
Brown Bear (Ursas arctos) a short fissure as arising from the highest part of the fissure
which he calls middle-curved or suprasylvian, but which I have named in this Pieport 2nd
curved or lateral fissure, and as passing towards the mesial longitudinal fissure, near its
hinder end, though without reaching it. He represents it as arising by a stem about 3 mm.
long, and then as bifurcating into an anterior and a posterior part, of which the latter
is apparently the deeper. On the mesial surface of the hemisphere, however, there is no
fissure which could be regarded as parieto-occipital. He considers that in the brain of
the Bear the upper curved fissure is only partially present, as in the short coronal fissure
and one or two other short fissures near it. He associates the appearance of a parieto-
occipital fissure as in direct relation to the disappearance of the 1st curved fissure, also to
the metamorphosis of a part of this fissure into the fissure of Bolando (central fissure) and
to the disappearance (Riickbildung) of the crucial fissure. He obviously considers that
in the corresponding part of the brain of the Felines there is an indication of a parieto-
occipital fissure, but that this fissure is absent in all Carnivorous brains where the three
curved fissures are completely developed.
I have examined the brain of Ursus maritimus with the object of seeing if a
corresponding fissure existed in it. In the right hemisphere a shallow fissure situated
1 See my Notes more especially on the Bridging Convolutions in the Brain of the Chimpanzee, Proc. Roy. Soc. Edin.,
\u\. v. p. 578.
132 THE VOYAGE OF H.M.S. CHALLENGE?..
at a corresponding spot did for about 4 mm. indent the marginal convolution in the same
reo-ion as in Ursus arctos, and a somewhat longer one was present in the left hemisphere.
These fissures were so short and shallow that they gave me the impression of being quite
subordinate furrows. On the other hand, the parieto-occipital fissure in the brain of
Man and Apes is one of primary importance ; it appears in the human foetus at about
the fifth month, and is especially marked on the inner surface of the hemisphere ; whilst
neither in Ursus arctos nor Ursus maritimus was there any evidence of a fissure which
corresponded with the internal parieto-occipital fissure of the human brain.
Dr. Murie, in the course of his description of the brain of Otaria jubata, employs to a
large extent the terminology of human anatomy, and believes that he can recognise in
the brain of this Eared Seal the majority of the convolutions and fissures present in the
human brain. Amongst other fissures he describes, by the name of internal perpendicular,
the fissure which is more usually named parieto-occipital. He figures it as indenting
the marginal convolution towards the hinder end of the hemisphere, and as present both
on the mesial and cranial surfaces. In conformity with the method of nomenclature
which he has adopted he has named the convolution in front of the inner part of this
fissure the quadrilateral lobule of the parietal lobe, whilst that which lies behind it he
names the internal occipital lobule. In the brains both of the Elephant Seal and Walrus
the marginal convolution was indented in a position almost corresponding to that in
Otaria jubata, by a continuous fissure both on the cranial and mesial surfaces, the
length of which was, however, variable in the different brains, especially on the mesial
-surface. Partly owing to this variability, and partly because we have no information on
the development of this fissure in the Pinnipedia, I am not prepared to say that it is
homologous with the parieto-occipital fissure of the human brain.
The evidence obtained from experiments on the cerebral cortex has established the
important fact that stimulation of the occipital lobe in the brain of the Monkey produces no
definite reaction ; whilst stimulation of the angular gyrus, both in its anterior and posterior
limbs (13), affects the pupils and occasions movements of the eyes to the opposite side, so
that this convolution is a visual centre.1 In the Dog also the most posterior parts of the
1st and 2nd external convolutions do not respond to stimulus, whilst a portion of the
2nd external convolution in front of the most posterior part (13), when stimulated, gives
reactions similar to those obtained from the angular gyrus in the Monkey. There is
reason to think, therefore, that the most posterior parts of the 1st and 2nd external
convolutions of the Dog are potentially equivalent to the occipital lobe in the brain of
the Monkey, although they are not differentiated by a parieto-occipital fissure, whilst the
2nd external convolution immediately in front of the part which does not respond to
stimulus and the angular gyrus are homologous with each other physiologically. In all
probability these convolutions are also anatomically identical, for Gratiolet, who was the
1 See the researches of Ferrier and other experimentalists.
REPORT ON THE SEALS. 133
first to differentiate the angular gyrus (pli courbe) in the brain of Man and Apes,1 places it
behind the supramarginal gyrus, i.e., behind the tier of convolutions immediately above
the Sylvian fissure, and therefore in a position corresponding to what that part of the
2nd external convolution which gives a similar response to stimulus would assume were
this convolution in the Dog's brain pushed backwards by a great development of the
frontal lobe.
The general results arrived at in this comparison of the brains of these Mammals
are to some extent to be regarded as tentative and provisional. For, until the
development of the fissures and the development and structure of the convolutions have
been worked out with greater detail than up to this time has been done, it will not be
possible to speak with certainty on all the points which have to be considered in a
detailed comparison of the cortex of the cerebrum in the Carnivora with that of Man
and Apes. Further, it should be stated that in this, as in other organs of complex
constitution, it does not follow that all the parts which are seen in the more highly
developed brains are of necessity present, even in a rudimentary condition, in those
whose organisation is not so complicated. It must also be remembered that whilst the
brains of the Carnivora, and still more so those of the Pinnipedia, are highly convoluted,
those of such Apes as the Marmoset Monkey (Ha/pale jacchus) are smooth on the
surface, and, with the exception of the large surfaces separated by such fissures as the
Sylvian and hippocampal, have no definite subdivision into morphological areas which
are capable of being recognised by the naked eye. But both in the Marmoset Monkey
and in such other New World Apes as OEdijous,2 in which the convolutions are either
absent or rudimentary, the cerebral hemispheres are prolonged forwards to the front of
the olfactory bulbs and backwards above the cerebellum to an extent which is not seen
in the Carnivora. In this respect, therefore, these brains, though either without con-
volutions or having them only feebly developed, are more highly organised than is the
case in the Carnivora proper or in the Seals.
From the point of view of the hypothesis of evolution there would be no reason to
think that the smooth-brained lower Apes had originated out of the Carnivora, at least
after the cortex of the cerebrum in this latter order had begun to assume a convoluted
arrangement. If they had been derived from a carnivorous animal with a convoluted
brain, then in all likelihood the convoluted character of the cerebrum would not have
disappeared in the process of evolution. If the higher Apes have been derived by descent
from the lower Apes, then the hemispheres in the former with their complex arrangement
of fissures and convolutions have been evolved from a smooth-brained stock and not from
an animal with such an elaborate arrangement of convolutions as is possessed by either a
Dog or a Seal. Hence the acceptance of this hypothesis is not inconsistent with the
1 Memoire sur les plis CL-rcbraux de l'liomine et des primates, Paris, 1869. 2 See Gratiolet, op. cit.
134
THE VOYAGE OF H.M.S. CHALLENGEE.
fact that the convolutions of the brain in the Apes assume from the first their own
method of arrangement, and not necessarily that of the orders of Mammals with con-
voluted brains which are lower in the series. Beyond therefore a certain general
correspondence in the arrangement of those fundamental parts of the cortex which serve
a similar purpose in these various orders, one does not find it possible to determine the
presence of convolutions arranged in a precisely corresponding manner in the brains of
the Carnivora and Pinnipedia on the one hand, and of Man and Apes on the other. In
each of these orders the developmental process which gives rise to the disposition of
the fissures and convolutions is regulated by the vital and mechanical necessities of
the animals constituting the order, as well as by the conditions of hereditary descent.
Subject to the qualifications and reservations which have been just expressed, and with
the proviso that the homologies of the cortical areas of the cerebrum are in many instances
histological and physiological rather than morphological, the following summary of the
corresponding fissures and convolutions in the Dog and the Monkey has been drawn up
in a tabular form : —
Table XIV.
Dog.
Monkey.
Fissures.
Fissures.
Sylvian,
Hippocampal,
Splenial, .
Sylvian.
Hippocampal.
Collateral and calloso-inarginaL
Olfactory, .
Intraorbital,
Olfactory.
Triradiate.
Coronal,
Rolando's.
Prresylvian,
Prsecentral.
Anterior part of lateral,
Intraparietal.
Convolutions.
Convolutions.
Callosal, .
Callosal.
Hippocampal,
Lobus hippocampi,
Gyrus rectus,
Internal supraorbital,
External supraorbital,
Sylvian,
Hippocampal.
Uncinate or uncus.
Gyrus rectus.
Internal supraorbital.
External supraorbital.
Island of Reil iu whole or in part.
Posterior Hmb of suprasylvian,
Sigmoid gyrus, part of sagittal
composite convolutions,
;onvolut
ion, ant
Superior temporo-sphenoidal.
Ascending, superior, middle, and inferior frontal con-
volutions.
Anterior limb of suprasylvian,
Supramarginal or convolution of parietal eminence.
Part of 2nd external convolution posteriorly,
Angular gyrus.
Most posterior part of 1st and 2nd external con
volutions,
Occipital lobe.
PART IV.
VISCERA OF ELEPHANT SEAL.
The heart and some of the .abdominal viscera, and the male and female genitalia, of
specimens of Macrorhinus leoninus had been removed and preserved in spirit.
The heart was from the female killed at Christmas Harbour, and was as big as the
heart of a large bullock ; it showed a slight cleft where the two interventricular
grooves met at the apex, and each surface was almost equally divided between the two
ventricles.
A broad flattened thymus gland overlapped the ascending aorta and trunk of the
pulmonary artery. It measured 150 mm. in transverse and 154 mm. in antero-posterior
diameter. It was unequally divided into two lateral lobes, of which the left was about
twice the size of the right, and the left in its turn was almost completely subdivided
into two portions by intermediate connective tissue. Each lobe was subdivided into
numerous lobules, which had no appearance of having undergone fatty degeneration.
Two lymphatic glands about the size of walnuts were attached by areolar tissue to the
ventral surface of the thymus.
When the thymus was removed the ascending aorta was seen to emerge from under
cover of the pulmonary arterial trunk. Its transverse diameter externally about the
middle of its length was 66 mm., but immediately between the origin of the left
subclavian and the attachment of the ductus arteriosus the transverse diameter of the
arch was only 34 mm. A great contrast was presented between the dilated condition of
the ascending and transverse parts of the arch as compared with the descending part, for
immediately beyond the ductus arteriosus the transverse diameter of the aorta was only
29 mm. The rapid diminution in the calibre of the artery immediately beyond the
origins of the great vessels for the head, neck, and anterior limbs would without doubt
facilitate the flow of blood into these vessels.
The ascending aorta close to its origin gave rise to the pair of coronary arteries for
the supply of the heart's walls. From the middle of its ventral surface a thymic branch
nearly as large as the human radial entered the thymus and was distributed to its
substance. From the transverse part of the arch the wide but short innominate artery,
136 THE VOYAGE OF H.M.S. CHALLENGER.
the left common carotid, and left subclavian arose ; the innominate almost immediately
divided into the right subclavian and common carotid. From the right side of the base of
the innominate a branch as large as the human ulnar proceeded, which passed backwards
to the bifurcation of the trachea where it was cut across ; it was probably a bronchial
artery and in its course it supplied some large lymphatic glands placed at the side of the
trachea.
There was only one anterior vena cava, which received immediately in front of the
right bronchus a large azygos vein. The posterior vena cava was large where it opened
into the right auricle.
The uteri were bicornuated and non-gravid. In the largest uterus the corpus was 4^
inches (114 mm.) long, and each cornu was 1\ inches (190 mm.) long. The walls of the
uterus were tough and densely fibrous. In the cervix the wall was 20 mm. thick, in the
corpus 5 mm., in the cornu 4 mm. The mucous membrane was elevated into strongly
projecting parallel folds, which had a direction corresponding to the long axis of the
cavity ; these folds closely resembled the appearance which I have previously described
in the non-gravid uteri of Halichcei^us grypus and Cystophora cristata.1 The ovary was
about the size of a walnut, and was enveloped by a sac-like expansion of the peritoneum.
The testicles were 7 inches (178 mm.) long, 3 inches broad, and about lj inch thick.
A large and projecting epididymis ran along one border of each gland.
The kidneys were multilobulated, 6£ inches (159 mm.) long, 3 inches broad, and
2 inches thick. The lobules were about the same size as one finds them in the kidney
of Globiocephalus melas.
Nearly four feet of the small intestine had been preserved. It was firmly contracted
so that the lumen was closed, and the transverse diameter of the tube was only f-ths of an
inch.
The pyloric end of the stomach was preserved. It contained no food, but a quantity
of sand and fine gravel, the largest particle of which was about the size of a coffee bean.
After the Challenger had returned home there was forwarded to Sir Wyville Thomson,
from the Cape of Good Hope, a small box labelled " Seal's Ballast Bag." It contained
a dried and somewhat shrivelled membranous hollow organ, 11^ inches (292 mm.) long
by 5tj inches (140 mm.) in its greatest circumference. The cavity of this organ was to a
large extent occupied with smooth pebbles, flattened at the sides as if from mutual
attrition. I have not removed them from the cavity of the organ, as it would be
difficult to replace them, so that I cannot state the exact number, but there are certainly
upwards of twenty. They vary in size ; one of the largest is 1^ inch (38 mm.) in its
long diameter, and there are several of almost equal dimensions, but the smallest is
not much larger than a coffee bean. There is but little doubt that this so-called
" ballast bag" is the dried stomach of a Seal.
1 On the Placentaticm of Seals, Trans. Roy. Soc. Edin., 1875, vol. xxvii. p. 275.
REPORT ON THE SEALS. 137
That Seals do take stones into the stomach has been observed both by the seal
fishermen and by naturalists. Captain Henry Pain, of the S.S. " Scanderia," when writing
upon the habits of the Sea Lion, says1 that he has seen upwards of twenty-five pounds
weight of stones, some of which were the size of a goose's egg, in " a pouch " inside the
animal, obviously the stomach. He states that as these animals get thin they have the
power of throwing the stones up, a sufficient quantity only being retained to keep the Seals
from coming up too freely to the surface. Mr. Elliott relates2 that he has opened the
stomach in many specimens of Callorhinus ursinus, and that in old bulls he has seen stones
which weigh half a pound, and in one stomach he found about five pounds of large
pebbles : he also possesses the stomach of a Sea Lion in which more than ten pounds of
stones were present, some of which weighed two and three pounds. Mr. Robert Brown,
in his account of the Pinnipedia of the Greenland Seas,3 states that he has often seen
small stones or gravel in the stomach of the Walrus, and that this is a habit which
it possesses in common with PJwca barbata and even Beluga catodon. The intelfigent
keeper of the Seals in the Zoological Gardens, London, informs me that he is familiar
with this practice, and that he has seen the Sea Lion both swallow large pebbles and
subsequently disgorge them.
Various uses have been ascribed to this peculiar habit of the Seals. The prevailing
opinion amongst sailors is that the animals swallow the stones as ballast to enable them
to dive so as to catch fish, and that they can at will disgorge them. Mr. Elliott
considers that their use is, by grinding against each other, to destroy the numerous
Nematode worms with which the stomach is infested. Others again maintain that they
serve the same purpose as the stones in the gizzard of a fowl, and assist in the trituration
of the food. I am myself inclined to favour this view, for a Seal literally " bolts "
entire the fish which serve as its chief food, without any mastication, and the action of
the pebbles on the fish so swallowed would without doubt, through the movements of the
muscular wall of the stomach, most materially assist the gastric juice in the trituration and
chymification of the food.
1 Proc. Zool. Soc. Lond., 1872, p. 681.
2 Quoted in Allen's History of North American Pinnipeds, p. 353.
3 Proc. Zool. Soc. Lond., June 25, 1868, p. 430.
(ZOOL. CHALL. EXP. PAET LXVIII. 1888.) Yj7 18
138 THE VOYAGE OF H.M.S. CHALLENGER.
It had been my original intention to have dissected the carcases of the specimens of
Arctocephalus gazella and of Macrorhinus leoninus collected by H.M.S. Challenger,
and to have compared the anatomy of their muscular, vascular, and nervous systems, so
far as these were uninjured, with the corresponding parts in species of the genus Phoca
which I had from time to time collected and stored in the Anatomical Museum of the
University. As time went on I found that, from the pressure of many duties, I should
be unable to overtake this part of the work. I entrusted the specimens therefore to a
former pupil, Dr. William C. Strettell Miller, in whose skill as a dissector I had confidence,
to make the dissections and to draw up an account of his observations. This he has
now done so far as regards the muscular system and the nerves which supply the muscles,
and his description appears as an Appendix to this .Report. The time occupied in
dissecting, comparing, and describing the specimens was upwards of a year, and the
dissections were very carefully conducted. I should also say that the responsibility of
this part of the investigation rests with Dr. Miller, for beyond giving an occasional
superintendence over the dissections, and an opinion on some point when he was in doubt,
I have not interfered in the conduct of his work.
APPENDIX TO THE REPORT ON THE SEALS.
The Myology of the Pinnipedia. By Wm. C. Strettell Miller, M.D. Edin.
INTRODUCTION.
This memoir embodies the result of a series of dissections of Seals, commenced in
November 1886, in the anatomical department of the University of Edinburgh, at the
request of Professor Sir William Turner, to whom I am much indebted for his kindness
in placing the material for this research at my disposal.
The animals which I have dissected are as follows : — an adult Phoca vitulina, a half-
grown imperfect specimen of Phoca hispida, an imperfect specimen of a pup of Phoca
barbata, two partially damaged young specimens of Arctocephalus gazella brought by the
Challenger from Kerguelen Island, and a specimen of Macrorhinus. leoninus, also brought
by the Challenger from the same island, which, unfortunately, was so imperfectly pre-
served that the hind limbs were the only parts which could be dissected. In addition,
a half-grown specimen of Phoca vitulina was kindly given to me by Professor Cossar
Ewart for this investigation.
The plan which has been pursued in drawing up this description has been to take
the arrangement found in Phoca vitulina as the standard, and to describe fully the muscles
as seen in that animal, and then to point out wherein the other Seals agreed with
or differed from it ; but as Arctocephalus is far removed from Phoca vitulina in its
structure, its myology has had to be described in considerable detail. It must be
understood that when I use the term Phocinse in the course of my narrative I mean by
it Phoca vitulina, Phoca barbata, and Phoca hispida. Opportunity has frequently been
taken to compare the result of my own dissections with those of previous writers.
The muscles have been arranged in natural groups, and so far as possible the nervous
supply of each muscle has been stated.
The memoirs on the myology of the Pinnipedia which I have consulted in connection
with my dissections are as follows : —
Duvernoy, G. B., Sur les organes du mouvement du Phoque Commun. Mem. du Museum, torn, ix., 1822.
Vrolik, W., Specimen Anatornico-Zob'logicuni de Phocis, speciatiru de Phoca vitulina. Trajecti ad Rhenum, 1822.
140 THE VOYAGE OF H.M.S. CHALLENGER.
Rosenthal, F., Ueber die Sinnesorgane der Seehunde. Nova Acta Acad. Cmes. Leqp.-Carol., 1831.
Humphry G. M., On the Myology of Orycteropus capensis and Phoca communis. Journ. Anat. and
Physiol, May 1S6S.
Murie, J., Upon the Anatomy of the Pinnipedia. Trans. Zool. Soc. Lond., vols, vii., viii.
Lucae, J. C. G., Die Robbe und die Otter. Abhandl. Senckmberg. Naturf. GesellscK, 1873-75.
Cunningham, D. J., Report on some points in the Anatomy of the Thylacine (Thylacinus cynocephalus), Cuscus
(Phalangista maculata), and Phascogale (Phascogale calura), collected during the Voyage of H.M.S.
Challenger in the years 1873-1876; with an account of the Comparative Anatomy of the Intrinsic
Muscles and the Nerves of the Mammalian Pes. Zool. Chall. Exp., vol. v. part xvi.
Deemal Muscles.
The Panniculus carnosus in Phoca vitulina is an extensive sheet of subcutaneous muscular fibres
subjacent to pelt, blubber, and the deep fascia, which is dense and coarse in some regions, fine and
transparent in others. Dorsally, the panniculus almost covers the axial part of the animal. It
stretches from above the orbits to the root of the tail, ending as two finger-like prolongations
between the tail and the dorsal surface of the legs. Upon the ventral aspect, it begins at the
lower lip, covers all the neck, recedes from the presternum, exposing a small part of the sterno-
mastoid and cephalo-humeral muscles ; overlaps the pectoral,1 with the exception of a margin near
the sternum, and extends almost longitudinally, from the side of the thorax to the femur, which
it crosses midway between the trochanter and the external condyle, to join the finger-like pro-
longations. The belly, therefore, wants a subcutaneous muscular layer, and the abdominal part of
the pectoral and the external oblique lie next the investing fascia.
It is capable of division into five parts by the direction of the fibres : — a. The Platysma, on
the ventral side of the neck. b. The Lateral cervical, between the platysma ventrally and the
cervico-scapular dorsally. c. The Pectoral, on the sternal pectoral muscle, d. The Cervico-scapular
covers the back of the neck, and stretches from the orbit to the spine of the scapula, e. The
Dorso-abdominal extends from the scapular spine to the tail.
a. The Platysma is pale, and covers the muscles between the rami of the lower jaw, and
extends backwards to the junction of the presternum with the meso-sternum. The fibres are
longitudinal where they spring from the side of the presternum, but turn outwards at their anterior
ends ; some of these terminate over the sterno-mastoid, and partially overlap it near its insertion.
The remainder become more and more obliquely directed outwards ; at the anterior termination of
the presternum they become transverse, and over the thyroid cartilage also are transverse. The
most posterior fibres join the lateral cervical part ; all in front of these terminate after passing
over the ramus of the lower jaw, by mingling with the fibres of the lateral cervical, round the angle
of the mouth, and insert themselves into the skin round the lower lip.
b. The Lateral cervical is also pale, and passes forwards from the side of the neck. It begins
over the cephalo-humeral and sterno-mastoid muscles near their insertions. The fasciculi ascend
to the angle of the mouth, the lower lip, and the zygoma, where they end. The sides of this
longitudinal band are joined dorsally by the cervico-scapular part, and ventrally by the platysma.
c. The Pectoral is fan-shaped, and rests upon the presternal and sternal parts of the pectoral
muscle, which originate from the presternum and meso-sternum. The inner margin of the fan
1 Only one specimen had a pectoral panniculus.
REPORT ON THE SEALS. 141
lies next the sternum; the posterior border extends from about 3 inches to the outer side of
the xiphi-sternum to the forearm, and there joins the dorso-abdomiual part. The band of fibres,
representing the handle of the fan, stretches outwards over the inner surface of the forearm, and
blends with its deep fascia. A few fibres of the anterior border turn over the shoulder to join the
cervico-scapular part. In the large Phoca vitulina the pectoral part was wanting.
d. The Cervico-scapular begins 1J inch posterior to the spine of the scapula, covers the back of
the shoulder, the dorsum, and the posterior part of the side of the neck, and reaches as far forwards
as the orbits. Laterally it joins the lateral cervical part, and in the ventral thoracic region the
pectoral part over the great humeral tuberosity. It arises about 2 inches posterior to the scapular
spine, from the aponeurotic band covering the spines of the vertebra;. This aponeurosis runs to a
point anterior to the scapula, from whence it takes origin from the ligamentnm nucha;, as far
forwards as the occipital bone, and then from a fine fibrous slip continuous from this forwards
between the parietal eminences to opposite the orbit.
The posterior fibres are transverse, and sweep over the deltoid and the trapezius. A little
further forwards they turn round the humerus to the ventral thoracic region to unite with the
pectoral part over the shoulder joint. From here the fibres begin to turn upwards, and curl round
the posterior lateral region of the neck, stopping upon the cephalo-humeral muscle, so that part of
it is uncovered by the panniculus. The middle fibres have the same direction, and join the
lateral cervical part. The bulk of the anterior fasciculi are directed forwards and outwards, and
terminate upon the vertex of the cranium, midway between the lambdoidal and coronal sutures,
near the posterior termination of the zygoma. The fibres, arising from the fibrous slip, extending
from midway between the lambdoidal and the coronal sutures to near the root of the nose,
have a peculiar distribution.1 The most posterior fasciculus stretches onwards and outwards, to
end above the middle of the orbit ; the next fasciculus stretches outwards and ends to the inner
side of the last. In this way is formed a muscular slip, which, with its fellow of the opposite
side, forms a V with a leg resting above each orbit. A little anterior to the root of the zygoma
a muscular slip about half an inch broad ascends, and a few of its fibres join the two or three which
cross the line midway between the lambdoidal and coronal sutures. The fibres separated by this
arrangement are connected by fasciae stretching between these groups. This part is with difficulty
removed from the cephalo-humeral and anterior part of the trapezius, which lie beneath. In the
large Phoca vitulina the cervico-scapular part was fused with the dorso-abdominal and not defined
as in the smaller specimen. The fibres were very coarse and stronger than the dorso-abdominal.
They had gaps between them, and the fibres were closely adherent to the fascia above.
c. The Dorso-abdominal part. The extreme hindmost fibres lie in the hollow between the tibia
and the sides of the caudal vertebra; behind the great trochanter of the femur. Anteriorly, the
fibres extend nearly as far forwards as the spine of the scapula. The intermediate ones embrace
the back, the flank, and the side wall of the thorax, and reach as far forwards laterally as the outer
edge of the abdominal pectoral. From the plane of the xiphi-sternum to the flipper the fibres
intermingle with the pectoral part. Posterior to the elbow joint and the forearm it passes from
the side of the thorax over the outer surface of the ulna. It arises from the spines of the vertebra;
by a broad aponeurotic band, which is continuous posteriorly with the deep fascia over the tail.
This fibrous origin is narrow at the hinder extremity, expands over the middle of the back, and,
1 They represent the occipito-frontalis.
142 THE VOYAGE OF H.M.S. CHALLENGER.
as already shown, tapers off to a point between the scapulas. The fibres between the tail and the
tibia are directed obliquely upwards and outwards, and remain so for about l£ inch ; the anterior
I inch is continuous with the hindmost fibres of the lateral abdominal part of the pectoral. Those
in front of the portion joining the pectoral upon the leg sweep more obliquely forwards and
outwards over the great trochanter of the femur, slightly cover its shaft, and overlap for about half
an inch the outer margin of the abdominal pectoral at the level of the knee joint. The fibres
lying between the knee and the level of the xiphi-sternum become more oblique in their direction
forwards and outwards the nearer they get to the axillary fold. Their lateral ventral terminations
cover the lateral portion of the abdominal pectoral for half an inch. The fibres anterior to the plane
of the xiphi-sternum are very obliquely directed upwards and outwards ; some of them end by
mingling with the pectoral panniculus, others branch out along the dorsum of the forearm, and
join its deep fascia ; a few are lost in the fascia over the triceps and the external condyle ; and
lastly, those between the spines of the vertebras and the external condyle of the humerus fade
away among the almost transverse fibres of the cervico-scapular part, posterior to the spine of the
scapula. It is closely adherent to the latissimus dorsi until it reaches the elbow joint and the
posterior border of the ulna ; here it leaves the latissimus, and passes over the olecranon and the
outer surface of the forearm to join the panniculus descending from the neck. At this position
there is a quantity of fat, no doubt to facilitate the movement of the elbow. In the large Phoca
vitulina a small fasciculus joined the abdominal pectoral muscle near the axilla. It was abund-
antly supplied with nerves and vessels, coming through the digitations of the external oblique,
between the latissimus dorsi and the lateral part of the abdominal pectoral.
The fibres of the cervico-scapular, dorso-abdominal, and pectoral parts are of the ordinary red
colour, and the two latter are of uniform strength. The cervico-scapular fibres are coarse, and inter-
mixed with fibrous tissue. Every part is closely connected by fibrous strands with the cutaneous
structures above and the muscles beneath. No part is directly inserted into bone. The cervico-
scapular and the dorso-abdominal are connected to the deep fascia, which is bound to the outer
side of the tendon of the pectoral muscle, and so indirectly to the humerus. In the large Phoca
vitulina, the panniculus terminated abruptly over the dorsum of the scapula, and did not run
gradually into the deep fascia on the dorsum of the forearm. All the dermal muscles were com-
posed of red muscle-fibres, and the platysma and lateral cervical had no fascial line dividing them,
but were intimately blended.
In the specimens of Arctoccphalus gazclla the panniculus was destroyed.
Myology of the Fore-Limb.
The fore-limb of the Phocinas and of Arctocephalus gazella has inserted into its bones the
superficial muscles of the back which are arranged in two layers. The First Layer consists of the
Cephalo-humeral, Trapezius, and the Latissimus dorsi.
Before describing this layer, let us glance at the human trapezius. It has a cephalic and
a vertebral origin, and a twofold insertion ; part of the insertion going to the clavicle and the
remainder to the scapular spine. The latter may be regarded as in two parts, because the lower-
most fibres form a tendon which glides upon the smooth surface at the vertebral end of the spine,
REPORT ON THE SEALS. 143
and is fixed to a tubercle at the outer edge. What is found in the Seals is a complete division
of the muscular sheet resembling the human trapezius into three muscidar masses — (1) the
cephalo-humeral, which represents the clavicular fibres of human anatomy ; (2) the trapezius
anterior part, the fibres fixed to the spine; and (3) the trapezius posterior part, the fibres
forming the tendon attached to the tubercle.
The Ccphcdo-humcral as in the Carnivora generally forms a bulky mass. It is triangular, the
base rests upon the ligamentum nuchas, and the apex upon the shoulder, and is under cover of
the cervico-scapular part of the panniculus. It arises from the occipital ridge, from the fascial
slip anterior to the ligamentum nuchas, and from the anterior half of the ligamentum nuchas.
The fibres trend oblicmely backwards and outwards, and cover the side of the neck ; at the
shoulder they converge and are inserted by a short tendon into the upper end and anterior edge
of the great humeral tuber above the pectoral insertion, with which the tendon blends; and into
the transverse ligament, stretching between the two tubera over the biceps. The fibres are a little
coarser than those of the trapezius ; there is a cellular interval between the cephalo-humeral and
the anterior part of the trapezius, which is not distinct, but continuous. Above the shoulder the
atlanto-humeral muscle appears between the anterior part of the trapezius and this muscle, before
it reaches the hmnerus. From the shreds of this muscle traced in Phoca barbata and in Phoca
hispida it appears to be disposed as in Phoca vitulina.
In Arctocephalus gazclla the lacerated condition of the muscle in both specimens compels me
to pass over the origin. The muscle is larger and better developed than in the other specimens ;
it extends from the head to about 1 inch posterior to the anterior angle of the scapula, and
partially overlaps the anterior part of the trapezius ; above and behind the shoulder it forms a
broad muscidar band of which the anterior two-thirds blends with the sterno-mastoid, whereas
in the Phocinas it only touches the trapezius, and is a small bundle of muscle near its insertion.
It is inserted into the humerus between the insertion of the deltoid posteriorly, and the insertion
of the sterno-mastoid anteriorly.
Professor Humphry in his description does not use the names cephalo-humeral and trapezius,
anterior and posterior parts, but includes the whole mass under the name trapezius.
In the Phocinas and Arctocephalus the cephalo-humeral pulls the humerus forwards and rotates
it inwards ; this action is much greater in the latter, for the insertion is lower down upon the
shaft, and it also abducts. In the Phocinas it is supplied by the spinal accessory nerve. In
Arctocephalus the nerve was destroyed ; both receive branches from the cervical nerves.
The Trapezius (proper) is in two parts, an anterior and a posterior.
The anterior part with its fellow of the opposite side forms a trapezium. It is opposite the
vertebral border of the scapula, and arises from the posterior half of the ligamentum nuchas,
from the spines of the first six dorsal vertebras, and from the supra-spinous ligaments. The fibres
of this muscle are distributed in a threefold manner. The most anterior fibres pass anterior to
the scapidar spine at its axillary termination, and are partially concealed by the atlanto-humeral
muscle before being inserted into the anterior surface of the great humeral tuber, to the inner side
of this muscle; a few fibres do not reach the bone but blend with the atlanto-humeral. The
posterior fibres attach themselves to the vertebral end of the spine. The ones intervening between
the most anterior fibres and the posterior must be studied in two layers, a superficial, and a deep.
The superficial layer passes over the scapular spine and terminates upon the surface of the deltoid.
144 THE VOYAGE OF H.M.S. CHALLENGER.
The deep layer attaches itself to the whole extent of the scapular spine, to its posterior lip, and
posterior border ; some fasciculi immediately below the spine are confluent with the deltoid. In
the larger Phoca vitulina it arises from the 1st dorsal vertebra to the 5th. This part was destroyed
both in Phoca barbata and in P/wca hispida.
In Arctocephalus gazella the insertion was traceable, and for the sake of clearness it is advan-
tageous to divide this attachment into an outer and an inner half. The outer half consists, as in the
intervening fibres of Phoca vitulina, of a superficial and a deep set of fibres. The superficial ones
of the outer half blend with the deltoid posterior to the scapular spine. The deeper fibres of this
half are inserted into the outer half of the spine, into its posterior lip, and posterior border, to
within one-eighth of an inch from the axillary termination. The fibres of the inner half cross the
spine of the scapula to blend with the deltoid.
As in the Earless and Eared Seals, owing to absence of a clavicle the spine of the scapula does
not possess a well-formed acromion process but ends abruptly about 1 inch from the axillary
border, a modification in the attachment of the fibres is occasioned. In the Phocina? the most
anterior fibres correspond with those in human anatomy attached to the acromion, and cross to the
humerus over the gap formed by the stunted spine ; in Arctocephalus the anterior part of the
trapezius is smaller and does not pass to the humerus. In both, some of the fibres of the anterior
part blend with the deltoid and must at times work in unison with it. The posterior part of the
trapezius is at right angles to the deltoid, and the anterior part nearly in a plane with the deltoid
fibres. This part will draw the shoulder forwards and slightly backwards with rotation upon the
ribs ; also pull the scapula towards the vertebral column.
In the Phocina? and Arctocephalus it is supplied by the spinal accessory and spinal and cervical
dorsal nerves.
The posterior part of the trapezius is an elongated triangular slip situated alongside of the
vertebral column. It arises from the 6th, 7th, 8th, 9th, and 10th dorsal spines, from the
supra-spinous ligaments, and from the lumbar aponeurosis as far back as the 12th dorsal trans-
verse process. It ascends over the posterior angle of the scapula, passes beneath the anterior part
of the trapezius, and is inserted by a tendon into the extreme vertebral termination of the scapular
spine, and by expansion of this tendon on either side into the scapula, between the vertebral end of
the spine and the vertebral border, and into the spine on the outer side. In the large Phoca vitulina
it arises from the 8th to the 14th dorsal vertebra?, the posterior half of the origin being tendinous.
There is some difference in the origins of this part hi Phoca barbata and Phoca hispida. In the
former it arises from the 10th, 11th, 12th, and 13th dorsal spines, and in the latter from the 9th,
10th, 11th, and 12th dorsal spines.
This part in Arctocephalus gazella is riband-like. It arises from the spines 'of the 8th, 9th, 10th,
and 11th dorsal vertebra?, and, in addition to the parts described in Phoca vitulina, has insertions
into the dorsal surface of the deltoid by its fibres blending with it. In all the specimens it drags
the scapula backwards, tilting the glenoid forwards at the same time.
In the Phocina? and Arctocephalus it is supplied by the spinal accessory and dorsal spinal nerves.
The Latissimus dorsi is rectangular ; it covers the back, the lateral aspect of the abdomen, and
the thorax. It is hidden by the dorso-abdominal panniculus. It arises from the spines of the
5th, 6th, 7th, 8th, and 9th dorsal vertebra? by muscular fibres, which touch those on the opposite
side of the spine ; from the lumbar aponeurosis as far back as the 5th lumbar vertebra ; and from
REPORT ON THE SEALS. 145
the outer surfaces of the five lower ribs below the insertion of the lateral division of the erector
spina;. These five origins interdigitate with the external oblique. The hindmost fibres curve
sharply forwards, the rest are almost longitudinal, the anterior ones most so. It passes over the
posterior angle of the scapula and divides into two parts ; the outer is inserted into the posterior
border of the abdominal part of the pectoral muscle ; the inner blends with the tendon on the
dorsal surface of the teres major, and is inserted with it into the inner border of the humerus, the
tendon being next the bone and the fleshy part of the teres above. The origin in Phoca larbata is
from the spines of the 5th dorsal vertebra; to the 3rd lumbar, otherwise the description is the same ;
and in Phoca hispida the origin is from the 4th dorsal spine, and the spines of all the remaining dorsal
vertebra;, from the lumbar aponeurosis as far back as the 4th lumbar spine, and for the rest as in
Phoca vitulina.
In Arctocephalus gazella it arises from the lumbar aponeurosis opposite the 3rd lumbar spine as
far forwards as the 12th dorsal vertebra, from the spines of the 12th dorsal vertebra to the spines
of the 7th dorsal by muscular fibres, and from the outer middle surfaces of the 9 th to the 15 th
ribs. The posterior fibres do not curve sharply forwards but ascend obliquely forwards and out-
wards, the anterior, as in Phoca vitulina, are almost transverse. The anterior border passes outwards,
touching the posterior angle of the scapula above the serratus magnus, and beneath the dorsi-
epitrochlear division of the triceps. At the middle of the posterior border of the latter muscle it
divides into two parts. The inner part has the same insertion as in Phoca vitulina, and the outer
part blends with the thoracico-abdominal part of the pectoral muscle opposite the lateral aspect of
the 5th rib. The inner part brings the fore-limb backwards and turns it inwards. The outer acts
on the pectoral muscle.
In the Phocina; and Arctocephalus it is supplied by dorsal and lateral cutaneous spinal and
lumbar nerves, and by the subscapular nerve.
The latissimus separates into two parts about midway between its origin and insertion, forming
an inner and outer division. In Arctocephalus the outer division of the latissimus goes to the
thoracico-abdominal part, in the Phocina; to the abdominal. There are no slips coming from the
posterior angle of the scapula.
The Second Laver of the superficial muscles of the back connected with the fore-limb consists
in the Phocina; of the levator anguli scapula;, three rhomboidei, and the atlanto-humeral ; but in
Arctocephalus gazella the muscles are levator anguli scapula;, two rhomboidei, and atlanto-scapular.
The Levator anguli scapulas in the Phocina; lies below the cephalo-humeral and the trapezius.
It is an elongated slip and arises by an aponeurotic band from the ventral surface of the transverse
process of the atlas. At its origin it is situated dorsally to the atlanto-humeral, coming in contact
with the anterior angle of the scapula and overlapping it, to be inserted into the scapula between the
spine and the vertebral border, and into the vertebral border between the spine and the anterior
angle. The ventral surface at the origin is posterior at the insertion, giving the muscle half a
turn.
This muscle in Arctocephalus gazella is an elongated triangle, and lies under cover of the
rhomboideus capitis over the back of the scapula. The origin was destroyed. It is inserted as in
the Phocina;, but falls short of the anterior end of the vertebral border of the scapula by 1 inch.
It moves forwards the anterior angle of the scapula and rotates it upon the back.
(zool. cuall. exp. — part Lxvin. — 1888.) Yyy 19
146 THE VOYAGE OF H.M.S. CHALLENGER.
In the Phocina? and Arctoccphalus the levator is supplied by the spinal accessory anterior to the
scapular spine and by the cervical nerves.
Rhomboidci. — There are three rhomboidei in the Phocinte, and these are named — a. PJiom-
boideus capitis ; b. Ehomboideus cervicis ; c. Pdiomboideus dorsi.
The Ehomboideus capitis is a long narrow band, lying below the cephalo-humeral and the
trapezius. It arises from the superior posterior angle of the parietal bone, to the inner side of the
origin of the temporal muscle, and from the margin of the adjacent occipital bone. Opposite the
spine of the scapula at the vertebral border, it passes beneath the rhomboideus cervicis, and is
inserted into the ventral side of the cartilaginous plate of the scapula, near its posterior angle,
between the insertions of the serratus. Professor Humphry has not described two separate
muscles coming from the neck and head, but one, and to this the name rhomboideus minor is given.
In the Phocina? it is supplied by filaments from the cervical nerves.
The Ehomboideus cervicis arises from the forward fascial prolongation of the ligamentum nucha?
opposite the occipital bone, and from the ligamentum nuchee. Until the fibres reach the middle of
this ligament, the muscle is a slender band, then it becomes broader, and the fasciculi are obliquely
directed to the base of the scapula. It is inserted into the vertebral border of the scapula
posterior to the spine, and into the vertebral border of the cartilaginous plate. Some of the
hindmost fibres run into those of the serratus magnus at its insertion. In the large PJwca
vitulina the origin is as far back as the 2nd dorsal vertebra. This muscle is not specially noted
by Humphry, but named rhomboideus minor with the last muscle. In Phoca vitulina it is supplied
by the 4th cervical, and in Phoca barbata and Phoca hispida from the 5th cervical.
The Ehomboideus dorsi is a small triangular muscle lying between the scapula and the serratus
magnus. It arises from the first four dorsal spines, and from the supraspinous ligaments. The
fibres go towards the posterior angle of the scapula on its ventral surface. It is inserted into the
axillary border of the cartilaginous plate, and to a very small extent into the axillary border of
the scapula. In the large Phoca vitulina the origin is from the 2nd dorsal vertebra to the 4th.
There is a slight difference in Phoca barbata, it arises from the 3rd, 4th, and 5th dorsal vertebrae.
To this muscle Professor Humphry has given the name rhomboideus major. In Phoca vitulina and
Phoca hispida it is supplied by a lateral nerve from the 1st intercostal space ; in Phoca barbata
by nerves from the 3rd and 4th, and 4th and 5th, intercostal spaces ; in the large Phoca vitulina by
a large dorsal branch passing between the 2nd and 3rd ribs.
In Arctoccphalus gazella, instead of three distinct muscles, there are only two, but these have
three insertions. They are the rhomboideus capitis (et scapularis) and the rhomboideus dorsi.
Bhomboidcus capitis (ct scapularis). — As the attachments of this muscle are vastly different from
the corresponding muscle in the Phocime, I have added " et scapularis " to emphasise the peculiarity.
The origin was mutilated. The fibres proceed backwards and slightly outwards, and cover half
the dorsal surface of the scapida anterior to the spine. It is inserted into the inner half of the
scapular spine, into the posterior lip, and into the scapula between the spine and the vertebral
border. Some fibres unite with those of the atlanto-scapular just anterior to the spine.
The Ehomboideus dorsi is of a rhomboid shape. It arises from the spine of the 7th cervical,
and then from the same spines as in Phoca vitulina. There is no division at the origin, but
as the fibres approach the vertebral border of the scapula they collect into two parts. The
anterior part has the same insertion as the rhomboideus cervicis of Phoca vitidina, but it
REPORT ON THE SEALS. 147
extends anterior to the spine for 1 inch along the vertebral border of the scapula. The posterior
part is inserted behind the anterior part into the vertebral border of the cartilaginous plate for
a slight distance ; then into the ventral side of this plate, reaching nearly to the posterior angle.
It closely resembles the insertion of the rhomboideus capitis of the Phocinaa, but is outside the
insertion of the serratus and not inside it on the ventral side of the plate.
In Otaria the muscle is regarded as the rhomboideus major and minor, but from its attachment
is considered more likely to be the minor. It is supplied by the dorsal and lateral intercostal nerves.
The rhomboideus dorsi approximates the posterior angle to the spinal column. All the others pull
forwards the anterior angle of the scapula, the rhomboideus capitis et scapularis having the greatest
efficiency in this direction.
The Cartifar/inous Plate of the Scapula. — This plate is found in many animals, and is of the
greatest magnitude in younger ones ; when present it gives origin and insertion to muscular fibres.
As an instance of the variety in shape, take the sheep as an illustration ; in it the plate extends
along the entire vertebral border of the scapula, and is of equal depth throughout : it only adds
another inch or so to the transverse length of the bone. In the Seals it is like a small triangle
with the apex beginning a little anterior to the spine and the base in a line with the axillary
border (PI. IV. fig. 1). When removed from the scapula the vertebral border is semilunar, but
when attached is almost straight and closely approaches the shape of the human vertebral border.
It enlarges the scapular surface, and one must remember its presence when examining a
macerated bone, otherwise the surface for muscular attachment may be undervalued.
The Ligamcntum nuchm. — Owing to the shortness of the neck in the Seal, especially in the
Phocina?, to the support given to the head by the water in which they spend most of their lives,
to the relative lightness of the cranium, and to their not requiring much up and down move-
ment of the head in search of food, this ligament is not the well-formed elastic band found in
many animals, which relieves the muscles of the neck, but a thin septum in a line with the
cervical spines. Its anterior termination is a thin fascial prolongation and ends upon the vertex
of the cranium ; in the cervical region it is much better marked, and altogether is not unlike this
ligament in man.
The Atlanto-Jmmcral arises from the aponeurotic band, which gives origin to the levator anguli
scapulas, and from the transverse process of the atlas anterior to this band. It goes towards the
shoulder under cover of the cephalo-humeral ; over the shoulder it emerges from beneath this
muscle, and is inserted into the anterior surface of the great tuberosity of the humerus in its upper
two-thirds. Professor Humphry does not give this muscle in his description, and it does not exist
in the Arctocephalus.
The representative of the atlanto-humeral in ArctoccpJialus gazclla is the atlanto-scapvlar. It
ai'iscs from the ventral surface of the transverse process of the atlas, by a tendon common to it
and the levator scapula?, and from the transverse process of the 2nd cervical vertebra. The fibres
pass backwards over the outer half of the dorsum of the scapula ; a few of the posterior internal
ones join the rhomboideus capitis. It is inserted into the outer half of the scapular spine along
its anterior lip. A small fasciculus joins the fibres of the deltoid outside the spine of the scapula
over the back of the shoulder.
In Otaria jubata and the Walrus the levator muscles are alike, and the insertions are the
same as that of the atlanto-scapular in Arctocephalus. From this I gather that there is no levator
148 THE VOYAGE OF H.M.S. CHALLENGER.
anguli scapulae in Otaria and Trichechus resembling in insertion the muscle of human anatomy. In
all the Phocinre and the two Arctoccphali I find there is one with a very similar insertion to that
found in man, so there is reason for giving the name atlanto-scapular to the muscle in Arctoccphalu*
corresponding to Dr. Murie's levator anguli scapula?.
In his paper on the Trichechus he points out that there is also a muscle which may be the
representative of the so-called levator-claviculfe. The atlanto-humeral has the same action as the
cephalo-humeral. The atlanto-scapular pulls the scapula forwards and rotates it.
Up to the present there have been differences in the muscles of Phoca barhata and Phoca
hispida, and special descriptions of various points have been required ; but now we come to a stage
in this myological study where all these agree, with only an occasional difference. It must there-
fore be remembered that the description of Phoca vitulina is also the description of Phoca barhata
and Phoca hispida, and only when a deviation occurs from the one selected as the standard animal
will their names be cited.
The Ventral Thoracic Eegion contains the pectoral muscle. The pectoralis minor and sub-
clavius are absent.
TJic Pectoral Muscle. — In consequence of the importance of this muscle both in swimming
and in moving on land, I have very carefully examined it. It has received numerous names.
Vrolik and Humphry call it the pectoralis major, Lucae the pectoralis, and Murie in the Otaria the
pectoral muscles, whilst in the Trichechus he divides it into three — (a) a fleshy pectoralis major, (b) a
second, (c) a third layer. It is situated in the pectoral region at its insertion, but the origin is more
extensive, for it covers the neck, chest, abdomen, and leg. The panniculus partially conceals it.
This most extensive muscle is divisible into three parts — (a) the presternal, (b) the sternal, and (c)
the abdominal. The presternal and sternal form one triangle, the abdominal another. The
presternal part arises from the fascia over the trachea 1 inch anterior to the presternum, and from
the side of it. It is separated, close to the junction of the presternum with the meso-sternum, by a
very faint cellular line, seen best on the under surface of the muscle. The fibres pass towards the
shoulder. The sternal part arises from the whole length of one side of the meso-sternum, and from
the cartilages of the eleven true ribs, and by an antero-posterior slip from the xiphi-sternum. A
cellular interval separates it from the third part. The abdominal part must be studied as three groups
of fibres. The first group arises posterior to the xiphi-sternum from 3£ inches of the linea alba ; the
second group, from the fascia over the external oblique muscle, by several finger-like prolongations,
which are shortest and most obliquely directed outwards near the middle line. Between these the
fibres of the external oblique are seen ascending to the ribs. The third group arises from the fascia
on the back of the leg. These hindmost fibres rest on the back of the leg, are continuous with the
hindmost fibres of the panniculus, turn round the leg, sweep over the femur, touch the outer side of
the patella, and course antero-posteriorly with the rest of the abdominal fasciculi, which are obliquely
turned outwards. All meet at the axillary border of the sternal part and disappear beneath it.
The three parts — presternal, sternal, and abdominal — converge on nearing their attachment to the
humerus. They are inserted in the following manner : — The presternal part blends with the sternal,
and the anterior third of this combination is inserted into the inner margin of the deltoid tuberosity of
the humerus, with the exception of a small part at the upper end. The posterior two-thirds join the
posterior layer of the deep fascia of the forearm, reaching near to the lower end of the ulna on the
REPORT ON THE SEALS. 149
posterior border of the flipper, and the anterior border of the radius about its middle.1 The abdominal
part, after disappearing beneath the sternal, is joined by the outer division of the latisshnus dorsi,
and then joins the portion of the sternal part which is inserted into the humerus. The humeral
portion is muscular on the under surface in the upper third and tendinous in the lower two-thirds.
In Arctoccpludus gazella there are also three parts having almost similar names. The pre-
sternal part is most anterior, and consists of a narrow muscular band. It arises from the lateral
anterior termination of the presternum by a small tendon. It is half under cover of the sternal
part, and courses almost transversely outwards to the shoulder over the sterno-cleido-mastoid,
which descends behind its inner half to gain the fascial slip occupying the position of the absent
clavicle. The outer half is between this muscle, which is now ventral to it, and the sternal part.
Over the shoulder it is lost among the fibres of the stemo -mastoid above and the sternal part be-
hind.2 The sternal part lies posterior to the presternal, and anterior to the thoracico-abdominal,
partly covering it. It is a fleshy mass of transverse fibres of considerable depth and breadth. It
arises from the lateral half of the presternum and meso-sternum ; and from the cartilages of the
four anterior ribs. It blends over the presternum and meso-sternum with the same part of the
opposite side. The fibres reach the anterior border of the humerus in a sheet of the same breadth
as at the origin. This wide bundle is inserted in its anterior half after blending with the thoracico-
abdominal part into the inner lip of the deltoid ridge of the humerus. The anterior three-
fourths of this insertion pass beneath a slip of the sterno-cleido-mastoid, which is adherent to the
pectoral, and end by dividing into two ; the outer slip becoming confluent with the origin of the
inner part of the brachialis anticus, and the inner by ending similarly upon the pectoral. The
posterior half is inserted obliquely into the deep fascia of the forearm, from the anterior side of
the bend of the forearm to the middle of the posterior border of the ulna. It is also attached
to the cartilaginous bar over the outer side of the forearm near the elbow-joint. The thoracico-
abdominal part is a large strong triangular sheet with the base in the mesial plane. It arises by
two divisions ; the ventral from the linea alba 1 inch behind the xiphi-sternum, and from the
outer half of the xiphi-sternum. This soon blends with the dorsal, which arises from the cartilages
of the 2nd to the 11th ribs ; from the 8th to the 11th ribs, opposite to the origin of the
ventral division, it only springs from the cartilages ; but anterior to the 8th rib it also has
origin from the side of the meso-sternum, and blends with its fellow as far forwards as the 4th
rib. Anterior to this, it does not blend over the meso-sternum ; for the sternal part intervenes
between the origins of the thoracico-abdominal parts of the opposite sides. The fibres pass to-
wards the humerus, partly under cover of the sternal part ; the posterior fibres ascend, the anterior
are transverse. At the level of the 5th rib laterally, the outer division of the latissimus dorsi
blends with it. After gaming these fibres along its outer edge, it blends with the sternal part and
is inserted with it. The under surface of the insertion is tendinous.
As a guide to the descriptions of this muscle in Phoea xitidina, Arctocepludus gazella, Of",'--
and Trichechus, a statement of the names used for the various divisions may be useful, as anatomists
differ much in their nomenclature.
In Phoea vitulina there are three parts — (1) the Presternal, (2) the Sternal, (3) the Abdominal.
1 The deep fascia over the anterior surface of the forearm divides into two layers — the anterior one gives attachment
to the panniculus, as has already been stated ; the posterior is for the pectoral muscle.
2 The fascial slip representing the clavicle is attached to the presternum internally, and passes outwards beneath
the inner half of the presternal part of the pectoral, to end by joining the sterno-cleido-mastoid going to the humerus.
150 THE VOYAGE OF H.M.S. CHALLENGER.
Professor Humphry mentions two divisions in Phoca vitulina — (1) the first or pectoral proper
includes the presternal and sternal ; (2) the second is the same as the abdominal.
In Arctoccplialus gazella there are also three — (1) the Presternal, (2) the Sternal, (3) the
Thoracico-abdominal. The last is so named because it covers the thorax and abdomen, and is not
wholly abdominal as in Phoca vitulina.
Dr. Murie in the Otaria gives three divisions — (1) a first division (which embraces the presternal
and sternal parts in Phoca vitulina and in Arctoccplialus), (2) a second (representing the abdominal
part in Phoca vitulina and the thoracico-abdominal in Arctoccplialus), (3) a third (not found in
Arctocephalus and Phoca vitulina, and called the sterno-scapular).
The same author in the Trichcchus gives three divisions — (1) a thick fleshy pectoralis major
(equivalent to the presternal and sternal parts in Phoca vitulina and Arctoccplialus), (2) a second
layer or pectoralis minor (which is the abdominal part in Phoca vitulina and the thoracico-abdominal
in Arctoccplialus), (3) a third layer (called the sterno-scapular by Dr. Murie in Otaria).
The muscle fibres are not arranged alike in the Phocinas and Arctoccplialus, but form muscular
layers of very different shapes. The presternal and sternal parts in comparing their form can be
combined, and this gives two masses for examination. The presternal and sternal parts in the Phocinae
form a large triangular layer, with the anterior and middle fibres transverse, and the posterior
obliquely directed forwards ; the base of the triangle springs from the whole of the presternum,
meso-sternum, and xiphi-sternum. In Arctoccplialus and in Otaria the same mass consists only of
transverse fibres, and stretches from the presternum and meso-sternum directly outwards to the
flipper. Judging by the drawing of the Trichcchus, the configuration of the same division is more
like that of the Phocinae, for it approaches the triangular shape, and the posterior fibres are not directly
transverse as in Arctoccplialus and Otaria, but obliquely directed forwards as in the Phocinae. The
abdominal part in the Phocinas approaches the triangular shape, and consists of an inner or mesial
belt of fibres directed forwards and outwards, and an outer or lateral belt running along the lateral
abdominal wall and a number of intermediate muscular bars or fingers filling in the triangle.
All these fibres go to the axilla. In Arctoccplialus the thoracico-abdominal part is a large badly
formed triangle nearly like that of Otaria and Trichcchus.
Humphry describes the second division (i.e., abdominal part) of the pectoral muscle in Phoca
vitulina as arising " from the linea alba, the pubes, and also from the margin of the ilium, covering
the fibres of the external oblique which were seen running transversely between the iliac and pubic
portions." This being a most interesting point in the anatomy of the pectoral muscle, I made a
series of dissections to ascertain the exact condition, and in the large Phoca had special opportunity
of investigating this among many other points, and wish to emphasise what was ascertained. A
group of fibres did come from the linea alba, also one from the back of the leg, and an intermediate
number of digitations from the fascia on the external oblique muscle whose hindmost ends did
not pass a line drawn from 3 inches behind the xiphi-sternum to the inner side of the patella,
so none reached the pubes. In this animal the digitations of some of the intermediate group of
fibres of the abdominal part reached the outer side of the rectus sheath.
As no other writer describes a presternal part, but includes it with the sternal, I give my
reason for so doing. In the Phocinas some were fresh specimens, and in these there was a slight
separation of the fibres at the junction of the presternum and meso-sternum ; but in Arctoccplialus,
the specimens being at least ten years old and preserved in brine, which had hardened the flesh and
REPORT ON THE SEALS. 151
cellular tissue, I could not be positive as to a natural division. However, as the presternal fibres
in the Phocinte and Arctocephcdus are the only ones of the pectoral muscle which are not inserted
into the humerus and forearm, but end among the fibres of the sternal part in the Phocinte, and
of the sterno-cleido-mastoid and sternal part in Arctocephalus, it shows how closely these prajsternal
parts are allied.
In Otaria the second division (i.e., the thoracico-abdominal) is inserted partly by fascia, which
joins the aponeurotic fascia of the forearm. In Arctocephalus there is no fascia going to that of the
forearm.
In Trichechus the second division, deep layer or pectoralis minor (i.e., the thoracico-abdominal
part), is inserted directly into the whole length of the shaft of the humerus, so there is no blending
of the second part with the first as in Arctocephalus and Otaria.
Neither in the Phocinse nor in Arctocephalus have I made out the third and smallest division
(sterno-scapidar) described in Otaria and Trichechus, and Professors Vrolik, Humphry, and Lucae
do not mention such a muscle in their researches on Phoca.
Dr. Murie inclines to the view that the two most superficial layers in Otaria are a divided
pectoralis major, and not the minor ; the third layer he classes as the sterno-scapular. In de-
scribing his Trichechus, he uses the names first layer or pectoralis major, second layer or
pectoralis minor, and third layer. Although the name pectoralis minor is used by Murie, I do
not think this muscle really exists in the Phocinse and other Seals, for as the insertion acts as a
guide in determining the identity of a muscle, and as there is a well-marked lesser tuberosity in the
humeri of the Phocinse, and a better representative of it in the Arctoccphcdi, still no fibres of the
pectoral muscle find their way to it, but all pass over to the deltoid ridge. If the pectoralis minor
did exist, the insertion' would be into the lesser tuberosity of the humerus, because the coracoid is
inside the shoulder-joint in the seals.
The Lateral Thoracic Region contains the Serraius macjnus, which covers the trunk and the
neck. This muscle arises by five muscular slips from the ventral transverse processes of the five
posterior cervical vertebras behind the scalenus anticus, from the outer and posterior surfaces of the
nine anterior ribs at the junction of the bones with their cartilages. The five lowest interdigitate
with the external oblique ; the slips from the first and second ribs are not divisible near their origin.
From the insertion into the base of the scapula it is seen that the muscle is fixed in three
ways. The five cervical slips are inserted into the vertebral border of the scapula, between the
anterior angle and the cartilaginous plate, to 1 inch posterior to its anterior end. The four
anterior thoracic digitations crossing from the trunk are inserted obliquely across the ventral
surface of the cartilaginous plate between the cervical part ending 1 inch posterior to the anterior
end of the cartilaginous plate, and the osseous posterior angle of the scapula. The five posterior
ascending from the trunk are inserted into the vertebral border of the cartilaginous plate, and
slightly into the ventral surface of it, extending as far forward as 1 inch posterior to the spine.
These five digitations are attached in the opposite order of origin ; the 5th goes into the posterior
angle, the 6th is placed anterior to it on the scapula, and the 9th is the highest. In the large
Phoca vitulina there were ten digitations from the trunk.
In Arctocephcdus gazella it arises by sixteen digitations, almost as in Phoca vitulina. The fibres
course to the base of the scapula. The six posterior digitations from the trunk form a strong
152 THE VOYAGE OF H.M.S. CHALLENGER.
bundle near the posterior angle of the scapula ; this is partly tendinous on its outer side, and is
inserted into the vertebral border of the cartilaginous plate at the posterior angle, posterior to the
insertion of the rhomboideus dorsi, which sends a few fibres to it. The five cervical slips and the
five anterior thoracic are inserted into the inner lip of the vertebral border, from 1 inch posterior
to the anterior angle of the scapula to the posterior angle, and into the ventral side of the cartila-
ginous plate, with the exception of a- small piece which is for the insertion of the rhomboideus dorsi,
and joins with the insertion into the posterior angle. The cervical digitations are distinct, while
those from the anterior thoracic region are not quite so, but touch each other from their origin to
their insertion. The posterior slip interdigitates with the latissimus dorsi, the five anterior to this
one with the external oblique.
The origins of the serratus in the Phocinas and in Arctocephalus differ in the number of
the digitations from the trunk; in the latter there are two more coming from the 10th and 11th
ribs. In the Phocinas it only interdigitates with the external oblique, whereas in Arctocephcdus it
interdigitates with it and the latissimus dorsi.
By the greater size of the cartilaginous plate in the Phocinas a change in the two insertions is
brought about. The plate in Arctocephalus is only a narrow bar, but in the Phocinas it is very wide.
In the Phocinas the cervical slips fix themselves upon the anterior vertebral border of the bone
and anterior end of the cartilaginous bar ; the anterior thoracic slips follow the junction of the bar
with the bone from the termination of the cervical slips to the posterior angle, and the posterior
thoracic begin where the last ended, and follow the vertebral border of the plate to its anterior end.
By this arrangement a circle of fibres surrounds the cartilage, and a clear space is left in the centre.
The rhomboidei cervicis, capitis, and dorsi are fixed to it in this order from before backwards.
In Arctocephalus the cervical and the anterior thoracic are attached to the scapula and ventral
surface of this plate as far as the posterior angle, leaving uncovered a small part of the vertebral
border of the plate near the posterior angle for the rhomboideus dorsi. The posterior thoracic slips
go to the posterior angle. In Otaria it arises from ten ribs, and in Trichechus from eight.
The digitations of the cervical serratus are in a plane with the digitations of the levator anguli
scapulas, and are so combined in many Mammals that one muscle is the result. The slips are not so
closely approximated in the Phocinas and Arctocephalus as to prevent a natural division. Professor
Humphry states that it forms a continuous sheet with the levator, and Dr. Murie says in Otaria " that
the serratus digitations were tolerably fused together, so that they formed but one continuous sheet."
In Otaria he makes special reference to its " two upper nuchal slips which are inserted quite on the
dorsal surface of the scapula," and in the Trichechus explains that " the highest, as in Otaria, is
more or less separate, and is inserted into the dorsum of the scapula between the angle and spine
on the vertebral border." When the levator anguli scapulas is not well developed and is absorbed
by the serratus, and the fusion has not been absolute, then the serratus must be scrutinised
closely to discover what has become of it. The difference in its anterior part in Otaria and
Trichechus by the outer slips going to the dorsum of the scapula, and the want of perfect fusion of
the slips, makes it doubtful as to the highest being serratus. The atlanto-scapular in Arctocephalus
is the levator anguli scapulas of Dr. Murie in the Otaria and Trichechus. To justify this observa-
tion, the myological researches of the various investigators on the Phocinas in which there is no
atlanto-scapular must be quoted. Professor Humphry describes a levator anguli scapulas inserted
into the base of the scapula. Professor Lucae in the same animal gives, in one of his plates, a
REPORT ON THE SEALS. 153
levator with a similar insertion, and in my dissections of the Phocinae and Arctoccpludus I find in all
a levator which corresponds to the nuchal slips called serratus in Otaria and Trichechus, so that I
regard the nuchal slips as the levator in the Otaria and Trichechus.
It pulls the scapula away from the spinal column, the posterior fibres rotate it outwards, and
the anterior fibres from the neck must pull the shoulder forwards.
The Muscles of the Shoulder. — In the Phocinaj are found the deltoid, subscapulars, sub-
scapulo-capsularis, supraspinatus, infraspinatus, teres minor, and teres major.
In Arctoccpludus the teres minor and subscapulo-capsularis are absent, but this animal possesses
in addition an episubscapularis.
The Deltoid is placed upon the infraspinatus behind the scapular spine. It is in the
form of a quadrant, and arises from the entire posterior border of the spine of the scapula,
above the spinal origin of the infraspinatus ; from the dense fascia between the outer termina-
tion of the spine and the shoulder-joint ; slightly from the dorsal surface of the capsule of the
shoulder-joint ; from the scapula internal to the vertebral end of the spine and internal to the
origin of the infraspinatus ; from the dorsum of the cartilaginous plate to a small extent ; and
from a narrow surface between the infraspinatus and the dorsi-epitrochlear muscles. The fibres
course towards the humerus, overlap part of the triceps, and cross the upper half of its dorsal
surface ; from the middle to the axillary end of the spine it receives some of the fibres of the
anterior part of the trapezius, then passes over the shoulder-joint where the atlanto-humeral partly
joins it along its anterior border. It is inserted into the lower half of the outer edge of the great
humeral tuberosity (deltoid ridge).
In Arctoccpludus gazclla it lies posterior to the scapular spine, and is almost rectangular. It
arises from the whole extent of the posterior lip of the spine, from the posterior border of it, from
the capsule of the shoulder-joint dorsally, from the scapula by tendinous fibres between the spine
and the vertebral border, from the vertebral border by muscular fibres, from the dorsum of the
cartilaginous plate alongside of the vertebral border, and from the adjacent sides of both to the
posterior angle. All the fibres incline to the outer surface of the humerus, a few to the outer end
of the spine blend with a small group of the atlanto-scapular. It is inserted into the outer rim of
the deltoid ridge, and into the dorsal part of the capsule of the shoulder-joint. The under two-
thirds of the insertion is tendinous. From the corner of the muscle joining the lowest part of the
deltoid ridge, a tendinous slip goes to the fibro-cellular bar lying upon the anterior border of the
radius.
A glance at the scapulre of the Phochue and Arctoccphalus impresses one with their dissimilarity
of mould. The spine, which is the boundary line between the supra- and infraspinous fossae, is
situated at the junction of the anterior third and the posterior two-thirds of the dorsum in the
Phocinaj. In Arctoccpludus it is at the junction of the anterior two-thirds and the posterior
third (PI. VII. fig. 2). The osteological differences between these two bones show clearly the
variety in form, function, and development of the soft structures which are attached to them, and
point to the likelihood of some muscles being present in the one and not in the other, which is the
case. The form and position of the origin of the deltoid in the Phocinse and Arctoccpluilus are
markedly unlike. In the former it approaches the shape of a gun, with the stock at an acute angle
to the barrel, in the latter it is like an old-fashioned scythe.' The barrel of the gun and the handle
(ZOOL. CHALL. EXP. — PART LXVIII. 18S8.) Yyy 20
154 THE VOYAGE OF H.M.S. CHALLENGER.
of the scythe are the spinal origins. The stock of the gun is placed at the vertebral border of the
scapula, and the butt rests upon the dorsi-epitrochlear muscle posteriorly, which is midway between
the spine and posterior angle ; the outer angle of the butt is between the infraspinatus and dorsi-
epitrochlear muscles, and the infraspinatus only fills the upper half of the infraspinous fossa. In
Arctoccphalus the blade of the scythe goes along the vertebral border to the posterior angle, and is not
much longer than the vertebral portion in the Phocinre, because the spine is low down on the
scapula. In Otaria, two layers are described on the left side, one on the right. In Trichechus,
the first part is the same as in Otaria, but wants the slip to the supinator longus. In the Phocinae
it is supplied by the circumflex nerve, and by a twig from the suprascapular. In Arctoccphalus
by the circumflex.
As there are no clavicular and extremely few acromial fibres, there is no covering from the
deltoid for the shoulder-joint anteriorly, and its action as an elevator of the fore-limb is nil. This
want in the deltoid is atoned for by the fixation of some of the fibres of the anterior part of the
trapezius to the deltoid and the humerus in the Phocinae only. Lucae and Murie regard it as
an external rotator, Murie adding also that it draws the humerus backwards. There must, how-
ever, be some slight elevating power through the anterior part of the trapezius.
The Subscajmlaris is a triangular muscle, and arises from the ventral surface of the scapula, with
the exception of a small part near the neck, from the vertebral border to 1 inch posterior to the
anterior angle ; between which and that part of the vertebral border opposite the vertebral end of
the spine, it lies to the outer side of the insertion of the serratus magnus ; from here it follows the
junction of the cartilaginous plate with the bone, taking origin from both along their line of
junction to the posterior angle ; and from the posterior angle for 1 inch along the axillary border.
Anterior to the neck its fibres are united with those of the supraspinatus, and the origin from the
posterior angle is tendinous. It converges and the under surface becomes fibrous ventral to the
glenoid cavity, and the fibres from the posterior angle go almost transversely to the humerus. As
the axillary border of the scapula is arched, the posterior half of the muscle does not lie upon the
venter of the bone, but is next the teres major. It is inserted into the capsule of the shoulder-
joint and into the lesser tuberosity of the humerus.
In Arctoccphalus it arises from the concave surface of the venter of the scapula ; from the
cartilaginous rim close to its junction with the bone ; from the posterior costa, with the exception
of half an inch at the posterior angle, which gives origin to the teres major. On the surface next
the bone there are three grooves corresponding to three ridges ; from these latter there are no
tendinous slips going into the substance of the muscle. The ventral surface has three deep furrows
planted upon it ; the first lies between the first and second ridges, the second between the second
and third, and the third a little above the axillary border, whilst in its substance there are several
tendinous slips. At the posterior angle the origin is tendinous, and this also gives origin to the
teres major. The fibres converge towards the shoulder-joint ; the anterior are in a line with the
anterior border, and blend with the episubscapularis ; the posterior, coming from the angle, are
blended with the teres major ; after leaving the angle they run parallel with its anterior border.
Between the teres major and the scapula it lies upon the long head of the triceps. It is im& i
into the venter of the capsule of the shoulder-joint, into the inner side of the lesser tuberosity, and
into the humerus below the tuber for one-fourth of an inch, where it is behind the anterior insertion
of the episubscapularis. In the Phocinae it is supplied by three scapular nerves, along its upper
REPORT ON THE SEALS. 155
border by a twig from the suprascapular, and below by another from the circumflex. In Arcto-
cephalus it is supplied by the subscapular nerve and at the insertion by the circumflex. It rotates
the limb inwards, but this powerful action is checked by the prominent lesser tuberosity coming
against the glenoid.
The Subscapnilo-capsularis 1 is a small muscular slip under cover of the subscapularis, and is
found in the Phocina? and not in Arctocephalus. It arises from the anterior part of the axillary
border of the scapula near the glenoid, and from the ventral surface of the scapula, and it is inserted
into the ventral aspect of the capsule of the joint, and into the humerus below the lesser tuberosity.
In Phoca barbata, in addition, it takes origin from the dorsal surface of the axillary border.
No notice is taken by the authors frequently quoted of this small muscle in Seals. According
to various anatomists it goes either into the capsular ligament or the humerus, but in the dissection
of the Phocinae it was seen going to both. It is supplied by the circumflex nerve.
The Epnsubscapndaris is found in Arctocephalus and not in the Phocinaa. It is a cylindrical
muscle overhanging the anterior border of the scapula, the subscapularis, and the supraspinatus
muscles. It arises from the inner half of the arched anterior border of the scapula, by a tendinous
band on the ventral side, and by muscular fibres on the dorsal edge, also by muscular fibres
from the outer half of this border, from the anterior surface of the neck to the glenoid cavity ; and
where the neck is covered by the capsule from the anterior surface of it. It makes a bed for itself
on the subjacent anterior portions of the supraspinatus and subscapularis, blending with both.
In the substance of the muscle, about the middle of the anterior arched border, there is a flat strong
tendon, and on both sides of this, throughout its whole extent, many of its fibres are attached. The
tendon is inserted into the upper anterior surface of the lesser tuberosity of the humerus ; and the
fibres coming from the outer half of the arched border of the scapula which do not ascend to this
internal tendon, go transversely outwards to the superior posterior of the lesser tuberosity, and
into the superior surface of the capsule of the shoulder. At the insertion this tendon and these
fibres are continuous, forming a hook over the lesser tuber ; lastly, it is inserted by a flat bundle of
fibres from the part which overlies the ventral anterior surface of the subscapularis, and after crossing
its insertion goes into the small part of the lower inner side of the lesser tuberosity, below the
insertion of the subscapularis, and into the inner border of the humerus above the teres major
insertion. Prom Murie's accounts of the Otaria and Trichcchus, I conclude that he inclines
to the episubscapularis being a derivative from the supraspinatus and subscapularis, but from
evidence gathered from the Phocinas, I consider it as formed from the subscapularis. It is supplied
by the suprascapular nerve. It tilts forwards and outwards the lower end of the humerus. It
is principally for forcing the fore-Hmb forwards through the water, and is in place of clavicular
deltoid fibres ; it also turns the limb inwards, thus preparing it for the backward stroke.
The Supraspincdus arises from the supraspinous fossa to the outer side of the insertion of the
levator anguli scapulae, from the anterior border of the scapular spine, and from the capsule of
the shoulder-joint. Anterior to the neck of the scapula it is fused with the subscapularis. At
the vertebral side the muscle is a thin sheet, but anterior to the neck it is thick and fleshy. It is
inserted into the outer surface of the lesser tuberosity, into the superior surface of the Ugament
stretching between the greater and lesser tubers, and into the upper end of the great tuberosity.
1 Wenzel Gruber, Ahhandl. am der menschl. und vergleich. Anat., 1854 ; also Macalister, Muscular Anomalies,
Dublin, 1872.
156 THE VOYAGE OF H.M.S. CHALLENGER.
In Arctocephalus it is a triangular muscle and arises from the supraspinous fossa, to within
half an inch from the glenoid cavity. Overhanging it upon the anterior border is the episubscapu-
laris, separated by a deep furrow. In considering this origin the configuration of the scapula must
be grasped. The supraspinous fossa is divided into two by a well-marked ridge, or diminutive
spine, anterior to which the suprascapular muscle is thick, whilst posterior it is thin ; at the outer
third of the ridge there is a trench between the fibres arising anterior and posterior to it, but no
division in the fibres internal to this. From the ridge and the partial trench, it is seen that this
muscle is a double one and consists of two parts, an anterior lying in front of the ridge, and
a posterior behind it. The anterior part goes to the great tuberosity of the humerus, and is
inserted into the capsule over the superior surface of the joint, into the pit on the anterior and
upper surface of the upward prolongation of the great tuber, into its upper anterior half and into
the posterior surface ; a fasciculus crosses from the great tuber to the tip of the lesser tuber, and
is inserted into the outer side of it, forming a narrow muscular bridge over the transverse ligament
and the biceps ; it joins the fibres of the pectoral below the great tuberosity. The posterior part
lying posterior to the ridge and above the spine is inserted into the pit or impression on the outer
side of the great tuberosity, above the pit for the infraspinatus, and into the capsule of the joint
superiorly. In Otaria and Tricheehus it has a single insertion.
Upon the great humeral tuberosity of the young Arctocephalus there are three depressions
for tendons, comparable in this respect with the human great tuberosity. In the human subject
these are for the supraspinatus, infraspinatus, and teres minor, but in the adult specimens the
two lower are fused and the upper and lower extremities of this combined depression are deeply
pitted, showing that the fibres going to either end act somewhat independently. As pointed out,
the spines of the scapulas in the Phocinee and Arctocephcdi are in very different positions upon their
respective bones. The accessory spine or ridge of bone in Arctocephalus bears the same relation
to its scapula as the only spine in the Phocinre. The origin of the posterior part of the supra-
spinatus in the former, disregarding the spines, is from the same site as the infraspinatus in the
latter, and both are inserted into the same part of the major tuberosity. The actions of the
posterior part are those of an infraspinatus, so it may be regarded as a transposed muscle ; and
the infraspinatus in Arctocephalus is functionally a large teres minor. If the infraspinatus were
placed above the spine in the Phocinas, and the spine changed to a lower latitude, then there
would be almost the same arrangement of these muscles in both. In the Phocinae it is supplied
by the suprascapular nerve from the 6th cervical ; in Arctoccphcdus by the suprascapular.
In the Phocinte it carries the fore-limb forwards, and in Arctoccplmlus the anterior part
raises the fore-limb with the episubscapularis and turns it slightly inwards. The posterior part
with the insertion, like the human infraspinatus, is a feeble elevator of the limb, but a powerful
rotator outwards, bringing the fore-limb backwards to the side.
The Infraspinatus lies beneath the deltoid and is similar to it in form. It arises from the
posterior border of the spine of the scapula ; from the scapula between the spine, and the origins of
the triceps posteriorly, and the deltoid internally. It goes towards the shoulder ; a little beyond the
spine its fibres blend with the tendon of insertion of the supraspinatus. Over the dorsum of the
neck of the scapula, it is between the supraspinatus and the teres minor. It is inserted into the
outer side of the great tuberosity of the humerus ; and into the capsule of the shoulder-joint,
lower than the supraspinatus.
REPORT ON THE SEALS. 157
In Arctocephahis it is under cover of the deltoid, and is triangular in form. It arises from the
dorsum of the scapula, anterior to the tendon of origin of the long head of the triceps, and the
dorsi-epitrochlear muscles ; from the posterior border of the scapular spine, beneath the origin of
the deltoid ; from the dorsal surface of the capsule surrounding the neck of the scapula. It
crosses the dorsal surface of the shoulder-joint as a round tendon, is closely adherent to the
capsule, and is inserted into a pit at the junction of the posterior bolder of the great tuberosity
with the head of the humerus, below the pit for the posterior part of the supraspinatus. In
Otaria it penetrates the capsular ligament, and strengthens it. In Trichechus it overlaps its large
fossa. In the Phocinre it is supplied by the suprascapular nerves from the 6th cervical. In
Arctoccplmlus by the circumflex. In the Phocinre and Arctocepkalus it rotates the fore-limb
backwards.
The Teres minor is a scanty muscular band which arises from a narrow line anterior to the
long head of the triceps, and posterior to the infraspinatus. It is inserted into the capsule of
the shoidder-joint ; and into the anterior side of the great tuberosity of the humerus, below the
infraspinatus. In Arctocephahis it is not found. In Otaria the fibres are lost upon the capsular
head of the triceps, and in Trichechus it is rather indistinct, if present.
In Arctocephalus the infraspinatus has the same action as the teres minor in the Phocinse, and
I infer that the infraspinatus in Arctocephalus does the same work as the teres minor, and that the
posterior part of the supraspinatus in the latter is functionally the same as the infraspinatus
of the Phocinse. In the Phocinre it is supplied by the circumflex, and is a feeble rotator outwards.
The Teres major is a triangular muscle, lying on the posterior angle and dorsal surface of the
scapula. The latissimus dorsi covers a portion of it. It arises from the scapula posterior to the
origins of the dorsi-epitrochlear and the long head of the triceps, to one inch from the glenoid
cavity ; and from the dorsum of the cartilaginous plate. About the middle of the posterior
surface it becomes muscular, and the lower margin of the tendinous surface of the inner half blends
with the inner tendon of the latissimus dorsi. It is inserted into the inner border of the humerus
below the subscapulo-capsularis for half an inch ; and slightly into the great bicipital hollow.
In Arctocephalus it is rectangular and arises from the ventral surface at the posterior angle of the
scapula, from the lower surface of the tendinous area which gives origin to the subscapularis
anteriorly, from the posterior costa, and from the posterior angle by muscular fibres ; a few fibres
come from the posterior angle of the cartilaginous rim, and from the serratus magnus. The
latissimus joins it along its posterior border ; anteriorly it blends with the subscapularis outside of
its origin for an inch. It is inserted into the middle third of the inner border of the shaft of the
humerus, and slightly into the great bicipital groove, below the episubscapularis. In Otaria and
Trichechus it is inserted from the middle of the shaft upwards to the internal condyloid ridge
with the dorsi-epitrochlear and first head of the triceps.
With the assistance of the human scapula a better reading of those of the Phocinre and the
Arctoccphali is obtained. In man there is an axillary border which has an adjacent surface on
the dorsum of the bone, and this is cut off from the infraspinous fossa by a ridge running from
the glenoid to the inferior angle. The adjacent surface is divided into two, the upper half for the
teres minor, the lower for the teres major. This ridge is present in the Phocinre and the
Arctoccphali, but is modified ; in the former the glenoid third on the dorsum is well marked, over the
remainder is faint, but on close examination can be seen and felt; and it ends at the vertebral border
158 THE VOYAGE OF H.M.S. CHALLENGER.
i me inch anterior to the posterior angle. In the Arctoccphali the ridge extends from the glenoid to
the posterior angle, and is distinct ; but the surface between it and the axillary border is limited.
In the Phocinse and Arctoccphali the long head of the triceps and the dorsi-epitrochlear arise from
the ridge ; from the large surface in the Phocinse posterior to the ridge only the teres major springs ;
as the corresponding surface for the same in Arctoccphalus is limited, it has a linear origin one inch in
length at the posterior angle. The insertion of the teres major is enormous, for it takes up about
one-third of the length of the shaft. Professor Humphry and I do not find it going to the tuber
major like Professor Lucae. The difference between Arctoccphalus and Otaria is in the insertion ;
in the latter it goes to the internal condyloid ridge with the dorsi-epitrochlear and first head of the
triceps. In the Trichcchus it is similar to Arctocepludus. The mode of junction of the teres ninjoi
and latissimus dorsi is interesting ; over the posterior border of the teres tendon the inner part of
the latissimus dorsi expands upon and is interwoven with the fascia over the dorsal surface of the
teres, and the two thus associated go on to the humerus, the tendon of the latissimus lying next
the bone. In the Phocinse it is supplied by the subscapular, in Arctoccphalus by the circumflex.
In all the specimens it draws the humerus downwards, inwards, and backwards.
The Anterior Brachial Eegion in the Phocinse and Arctoccphalus is composed of the biceps
and brachialis anticus. The coraco-brachialis is wanting.
The Biceps is a short muscular band lying on the internal surface of the humerus. It arises
from within the capsule of the shoulder-joint by a strong short tendon from the rudimentary
coracoid or beak above the glenoid cavity ; and passes out of the capsule below the transverse
ligament stretching from the greater to the two lesser tuberosities. It descends between the two
tubers, and fills the bed of the wide bicipital groove ; after dipping between the brachialis anticus
and the pronator radii teres, it forms a tendon which is inserted into the radial tuberosity on the
posterior border of the shaft. The external side becomes anterior at its attachment.
In Arctoccphalus, besides the one tendinous head, it arises from the edge of the glenoid cavity
on both sides of the beak, and is closely united with the under surface of the anterior part of the
capsule above the head of the humerus. The tendon emerges from the capsule between the greater
and lesser tuberosities, and passes below the tendon of the first part of the supraspinatus, and
under the transverse ligament as in Phoca. It then continues down the humerus as a flat muscular
band, partly tendinous on its under surface, filling the bed of the enormous bicipital groove. It
dips between the pronator radii teres and the inner part of the brachialis anticus, to reach the
bicipital tuberosity of the radius into which it is inserted. The external side is also anterior at the
attachment into the radius.
The coracoid is separate in young Seals from the glenoid, and in them is found to form a con-
siderable part of the glenoid cavity. If in the adult the coracoid were taken from the ovoid-shaped
glenoid it would make a decided break in its shape. When the cartilage is attached to the glenoid
it also covers a part of the coracoid. The glenoid and coracoid were inside the capsular ligament of
the joint in the Seals dissected, and it is the coracoid that principally gives origin to the biceps.
In the Phocinse it is supplied on the anterior lower surface by a small branch of the median
nerve, and on the posterior surface by the external cutaneous. In Arctoccphalus it is supplied by
the musculocutaneous nerve. In all the specimens it flexes the forearm, and when the manus is
in pronation it will turn it outwards.
REPORT ON THE SEALS. 159
The Brachialis anticus is situated on the outer surface of the humerus, and arises from its
external surface behind the deltoid ridge, and by a bundle of fibres from the lower end of this ridge.
It sweeps over the anterior border of the humerus ; the posterior two-thirds of the muscle near its
insertion goes between the humerus and the tendon of the biceps, and is inserted into the anterior
border of the ulna below the coronoid process ; the bundle of fibres from below the deltoid ridge
ends by forming the anterior third of the belly of this muscle, and is finally inserted into the bicipital
tuberosity of the radius, outside the tendon of the biceps.
In Arctocephalus gazella it is in two parts. The inner part arises from the deltoid ridge between
its two lips, having the tendons of insertion of the pectoral and stemo-cleido-mastoid on the inner
lip, and the cephalo-humeral and deltoid on the outer. It extends as high as the epiphysial
line of the great tuber, and down to the junction of the outer lip with the inner. The outer part
arises from the external surface of the humerus ; and from the capsule of the shoulder-joint, which
is behind the outer lip of the deltoid ridge, and in front of the external border of the shaft. The
fibres of the inner part springing highest from the deltoid ridge of the humerus remain anterior
to the insertion; the lowest are posterior. It is riband-shaped, with its anterior edge in the
same plane as the anterior border of the radius ; it passes between the biceps and the outer part
to be inserted by a tendon which splits in two, behind the tendon of the biceps. The outer
division of the tendon is attached to the capsule of the joint over the inner surface of the head of
the radius, and into the tubercle .of the radius on the outer side of the tendon of the biceps. The
inner division of the tendon of the inner part goes behind the tendon of the biceps, and is inserted
into the anterior border of the ulna below the lesser sigmoid cavity opposite the radial tuberosity.
The outer part, with the exception of a small triangular portion at the upper end of the origin, is
covered by the supinator longus ; and the deltoid fills in this triangle. In front of the elbow-joint
it crosses from the external surface of the humerus to the internal surface of the radius, and hi
doing this twists, so that the anterior fibres from the shaft are external and the posterior internal.
It is inserted by muscular fibres into the inner surface of the capsule, over the head of the radius,
higher up than the inner part ; and by a tendon into the ulna outside the tendon of the inner part.
The few fibres taking origin from the lower end of the deltoid ridge in the Phooinae, the
anterior surface of it in Arctocephalus, and to its inner side in Trichcchus, may be considered as the
equivalent of the fibres from the surface internal to the deltoid ridge in man.
The deltoid impression of the human bone is only for the deltoid muscle. In the Seals it is
an eminence, and acts like an additional surface, making compensation for the smallness of the
humerus. In these animals it is a downward continuation of the great tuberosity, and is planted
upon a thick vertical wall of bone in the Phocinse and a thin translucent one in the Arctocephali,
The inner edge of the surface in both is flush with the inner side of the vertical plate and has a
straight edge ; but on the outer side it overhangs the outer surface of the shaft, and has a slight
projection near the middle of its surface in the Phocinas, and in the Arctocepludi a gradual expansion
from the middle to its inferior extremity (PI. VII. fig. 3). Roughly the eminence in the Phocinse is
rectangular, and in the Arctocepludi triangular with the base downwards. The surface in the Phocina.-
gives origin to a few fibres of the brachialis anticus and insertion for -the supra- and infraspinati.
teres minor, cephalo-humeral, atlanto-humeral, trapezius (anterior part), pectoral, and deltoid. This
surface in Arctocepliali gives origin to the brachialis (inner part), and insertion to the supraspinatus
(anterior and posterior parts), infraspinatus, pectoral, cephalo-humeial, sterno-cleido-mastoid, and
160 THE VOYAGE OF H.M.S. CHALLENGER.
deltoid. These muscles for rotating the humerus have more leverage by being removed from the
shaft by a vertical plate.
The brachialis anticus muscle in the Phocinse is single, and divides into two for insertion into
the radius and ulna. Lucae gives the division for the ulna as in combination with the biceps, but
Humphry and I find it quite apart. In Otaria as in Arctoccphalus it has two heads of origin,
but in the former the outer head joins the inner head on the outer side of the elbow, whereas
in Arctoccphalus the head from the surface of the deltoid eminence divides into two, one going to
each bone of the forearm, and the outer has also two divisions for both bones. In the Phocinfe
it is supplied by the musculo-cutaneous and musculo-spiral. In Arctoccphahis it is supplied by
the musculo-cutaneous. It is a flexor of the forearm on the upper, and, like the biceps, will
rotate the forearm outwards when the manus is prone.
The Posterior Brachial Region in the Phocina? and Arctocephali consists of the triceps, which
has four heads — (a) the dorsi-epitrochlear, (b) the long head, (c) external licad, and (cl) internal head.
There is no subanconeus.
The Triceps, first head, or the dorsi-epitrochlear,1 is a thin muscle partially covered by the deltoid,
and arises from the dorsum of the scapula by a broad sheet-like tendon, extending from the vertebral
border of the cartilaginous plate to a spot posterior to and in a line with the middle of the scapular
spine. This tendon is continuous with the tendon of origin of the long head of the triceps, and it is
placed between the origin of the infraspinatus anteriorly and the teres major posteriorly. Above
the olecranon of the ulna it collects into a small muscular band, which runs over the border of the
olecranon, near its junction with the posterior border on the internal surface of the ulna. After
receiving a few fibres from the long head it is inserted, or rather moored by its lower edge, to the
junction of the olecranon with the posterior border of the ulna, and to one inch of the posterior
border below this junction. The band passes to the flexor minimi digiti, blends with it, and
terminates at the junction of the middle and upper third of this muscle. In Phoca barbata the
insertion overlapped both sides of the olecranon.
In Arctoccphalus gazella it is of a triangular shape, and arises from the dorsal rim of the inferior
costa of the scapula, by a sheet-like tendon which is one inch long, and extends transversely from the
posterior angle to the middle of the dorsal rim of the posterior costa of the scapula. It is placed
between the infraspinatus anteriorly, and the teres major and subscapularis posteriorly. It becomes
cylindrical over the olecranon, slightly overrides both sides of it, and there receives a few fibres from
the external head on its outer side, and is inserted into the olecranon from the middle tubercle to
the posterior, and into the posterior and outer upper third of the flexor carpi ulnaris.
The difference in this muscle in Phoca vitulina and Ardocephalus according to my dissections is
well marked. In the former the origin is far removed from the axillary border by the extensive
surface for the origin of the teres major, which in the latter is adjacent to this border. The insertion
in Phoca blends with the flexor minimi digiti, and in Arctoccphalus with the flexor carpi ulnaris.
The statement by Professor Humphry that the muscle reaches the paddle finds no support
from Lucae, and in none of the specimens did I see this ; perhaps the flexor minimi digiti was
included with it in his description. Lucae gives its insertion into the fascia of the front arm, &c,
1 This is Humphry's first division ; Lucae {op. cit, pi. ix. fig. 1) calls it the " triceps pars longa," and in his text
" portio longa tricipitis."
REPORT ON THE SEALS. 161
but I made it out as blending with the flexor minimi digiti. In Otaria it arises from the inferior
angle, but in Arctocephalus from the axillary border ; the fibres in the former run to the forearm,
but in the latter they blend with the flexor carpi ulnaris. Trichechus agrees with Arctocephalus
in the origin, but the insertion differs, for it ends in the antebrachial fascia of the forearm, and
is inserted into the olecranon by fascia only.
The second, or long head of the triceps,1 is a triangular muscle, and arises between the origin of
the teres minor anteriorly and the teres major posteriorly. The extent of its origin is from a spot
immediately posterior to and in a line with the middle of the scapular spine to the glenoid cavity ;
the portion arising from the scapula is thin and chiefly tendinous ; the remainder which arises from
the neck is muscular, and covers the under surface of the neck as well as the lower half of the back.
Tt also has origin from the capsule surrounding the neck. The lower half of this muscle lying next
the external head is tendinous ; it lies upon the internal and external heads, and is inserted into
the olecranon below the middle head, into both of its sides and into the tendinous surface on the
posterior of the external head.
In Ardoceplmlus gazclla it is also triangular, and arises from the outer half of the dorsal rim of
the posterior costa of the scapula, between the fibres of the subscapularis, which springs from the
posterior costa, and posterior to the infraspinatus, which overlaps it. It stretches transversely
from above the middle of the posterior costa to the under surface of the capsule of the shoulder-
joint, from which it also has origin. The ventral surface is tendinous for the play of the teres
major, while the dorsal is only tendinous near the olecranon above its insertion. As in Phoca it lies
on the external and internal heads, when viewed from the inner aspect of the hmb ; and it is inserted
into the anterior internal half of the border of the olecranon on the inner surface, which is opposite
the anterior and middle tubercles of the outer surface ; some of the fibres run amongst those of
the internal head over the quadrilateral surface of the olecranon on the internal side, and it is
tendinous on the outer surface near the olecranon. In Otaria and Tricheelius it joins the common
cubital insertion of the triceps.
The third or external head 2 arises from the capsule surrounding the head of the humerus, which
is continuous with the posterior surface of this bone, and from the same position on the outer
surface to midway between the anterior and outer borders, from the hollow between the head and
the shaft, and very slightly from the humerus below this. It overlies the internal head, and is
closely connected to the tendinous lower half of the long head which covers it : it then joins the
anterior tendinous side of the long head, and is inserted by a small fasciculus into the outer side of
the tip of the olecranon between the internal heads.
In Arctocephalus gazclla it arises from the capsule of the shoulder-joint at the lower posterior
surface of the glenoid cavity; from the capsule between this and the neck of the humerus; from the
neck to the middle of the outer surface ; from the capsule above the neck, and also from the
external border of the shaft in its upper half. It has an opening, near its origin from the capsule
on the external border, for the circum3ex vessels. It divides above the olecranon into two parts ;
the inner is inserted into the tendinous portion of the long head above the olecranon and into the
inner side of the olecranon, opposite to the anterior and middle tubercles to the outer side of the
long head, and a slip from it joins the dorsi-epitrochlear ; the outer is inserted into the superior
1 Tbis is Humphry's second division ; Lucae's middle head ; Murie's first division.
2 This is Humphry's third division; Murie's (Trichechus and Otaria) second division.
(ZOOL. CHALL. EXP. — PART LXVIII. — 1888.) Yyy 21
162 THE VOYAGE OF H.M.S. CHALLENGER.
surface of the olecranon between the inner and outer borders, as far back as the middle tubercle
(PL VII. fig. 4, in Arctocephalus australis); some fibres blend with the other head over the olecranon.
The second division is nearly alike in Otaria and Trichcchus, and its origin differs from that in
Arctocephalus, for in the former two it arises from the back of the humerus, whereas in the latter it
only comes from the external border.
The fourth or internal head 1 lies on the back of the humerus under cover of the external
head. It arises from the posterior surface of the shaft of the humerus beneath the origin of the
middle head, and from the posterior ligament of the elbow-joint. It is of a triangular form, but
the base is next the elbow-joint, and the apex below the inner side of the head of the bone. It is
inserted into the posterior ligament of the elbow-joint ; into the sides and tip of the olecranon ; and
into the quadrilateral surface behind the sigmoid cavity.
In Arctocephalus gazella it arises from the posterior surface of the shaft of the humerus, from
the capside of the shoulder-joint, from the ligament as in Phoca, and is inserted into the .quadri-
lateral surface behind the sigmoid cavity of the ulna. In Otaria and Trichcchus its disposition is
almost the same. It is the extensor of the forearm.
In ArctoccpJialus the musculo-spiral supplies the dorsi-epitrochlear, long, internal, and external
heads, the latter also has a twig from the circumflex nerve.
The Flexor or Inner Surface. — In the Phocinse the following are the muscles — anconeus
internus, palmaris longus, flexor communis digitorum, flexor carpi radialis, pronator radii teres,
flexor carpi ulnaris, abductor minimi digiti longus.
In Arctocephalus, instead of one palmar muscle there are three, and the abductor minimi digiti
is not found. Neither in the Phocinse nor Arctocephalus is there a pronator quadratus.
The Anconeus internus, called supinator quadratus by Lucae, is nearly double the size of the
anconeus externus. It arises from the back of the internal condyle below the supracondyloid
foramen, and is inserted into the inner side of the olecranon below the long head of the triceps.
In Arctocephalus gazella it arises, as in Phoca, from the posterior part of the internal condyle
above the palmaris longus, and behind the pronator radii teres. It crosses from the internal condyle
to the inner lip of the olecranon, and is inserted into it opposite the anterior and middle tubercles
of the outer surface. The long head of the triceps and the deep palmar have a bed for it in
their substance. It is present in Otaria and Trichcchus. It is a short extensor of the elbow-joint,
and also steadies it. It is supplied by the ulnar nerve.
The Palmaris longus in the Phocinse is in two parts : — a. The first part arises from the
posterior half of the hollow on the internal surface of the ulna, where the olecranon and the
posterior border of this bone meet. It is situated at the origin below the anconeus internus, and
higher up the shaft than the flexor carpi ulnaris. At the junction of the upper and middle thirds
of the ulna it divides into two slips, one being anterior, the other posterior. The latter soon splits
into two fine tendons, which are inserted into the deep fascia over the wrist. The anterior slip is
also tendinous, and disappears beneath the palmar fascia ; upon the under surface of this it widens
and emerges on the opposite side as two fascial slips, which descend to the heads of the 2nd and
3rd metacarpal bones. Here they are attached to the sheaths of the corresponding tendons, and
also to the tendons of the flexor subliruis for the 2nd and 3rd digits. It is with difficulty
1 This is Humphry's fourth division ; also Lucae's, and in his plate is the anconeus quartus.
REPORT ON THE SEALS. 1G3
removed from the palmar fascia, to which it is closely adherent. In Phoca barbata the origin was
partly destroyed, but the fragment appeared the same ; the insertion is similar, b. The second
part is a narrow tendinous slip, and arises from the internal condyle between the flexor carpi
radialis and the flexor communis digitorum (first head), and is inserted into the deep fascia over the
ulnar side of the wrist.
Humphry does not separate this muscle into its divisions, but our descriptions are very much
the same. Lucae's insertion is not anything like what Professor Humphry and I make it to be.
In Aretoecphalns there are three palmar muscles — the Palmaris longus; superficialis, and
profundus.
a. The Palmaris longus, the palmaris longus primus of Murie, has a fascial and a bony origin.
It arises from the internal surface of the internal condyle by a tendinous slip, also by a muscular
slip which is blended for a short distance with the flexor carpi radialis, and from the convex side
of a fascial band for 1 inch, and blends with the superficial palmar for half an inch on the inner
side. This slender muscle ends in a fine tendon at the junction of the upper third with the lower
two-thirds of the radius, and at the lower end of the radius the tendon dips beneath the broad
tendon of the deep palmar, upon the under surface of which it expands on both sides, forming a
triangle with the apex as the continuation of the tendon ; over the base of the 1st metacarpal it
is moored to the under surface of the tendon of the deep palmar muscle, and ends over the 1st
metacarpal.
b. The Palmaris superficialis, the palmaris longus secundus of Murie, is a slender broad layer of
muscular fibres lying upon the deep palmar. It arises from the convex side of a fascial band, with
the exception of that portion on its anterior side which gives origin to the palmaris longus. This
band stretches from the internal condyle over the deep palmar to which it is closely bound, and
ends upon the anterior edge of the dorsi-epitrochlear. Below the band on the posterior side its
fibres are blended with the dorsi-epitrochlear for half an inch, and on the anterior with the palmaris
longus for the same distance. It crosses the deep palmar muscle ; and is inserted into the skin
opposite the ulnar border of the base of the 5th metacarpal ; the bulk of the tendon lies along
the ulnar border of the 5th metacarpal, and is attached to the head of this bone, and to the
ulnar side of the whole of the 1st phalanx.
c. The Palmaris profundus is the palmaris longus tertius of Murie, and has an extensive origin.
It arises from the internal surface of the olecranon ; from the inner edge of the quadrilateral surface
behind the sigmoid cavity ; from the capside of the elbow-joint ; from the internal concave surface
of the ulna in its upper half, posterior to the ridge running down the shaft from the sigmoid
cavity ; and from the upper half of the posterior border of the idna by an aponeurosis. At the
level of the junction of the olecranon and the posterior border of the ulna on the surface of the
muscle is a slender aponeurosis which covers its whole breadth, and as it descends upon the muscle
towards the lower end of the shaft it becomes thick and strong. The muscle-fibres terminate abruptly
on its posterior surface, parallel with the posterior border of the lower half of the ulna. This
tendon is also common to the flexor carpi ulnaris. By its posterior border below, it is attached to
the pisiform bone, and to the anterior border of the flexor carpi ulnaris ; it gives a tendinous slip
to the radial side of the base of the 5th metacarpal, and another strong broad one to the base of the
1st phalanx of the 5th digit ; from the tendon to the 1st phalanx a slip joins the transverse liga-
ment, i.e., the metacarpophalangeal ; the rest of the broad tendon crosses the wrist obliquely from
164 THE VOYAGE OF H.M.S. CHALLENGER.
the ulnar side of the forearm to the radial side of the carpus, and descends as an aponeurotic band
of very considerable strength, covering the palmar aspect of the whole of the 1st metacarpal bone,
and the radial halves of the 1st and 2nd phalanges of the pollex to the phalangeal cartilage.
It passes over the tendons of the thumb, and is inserted into the radial side of the carpus, the 1st
metacarpal, and the 1st and 2nd phalanges of the pollex ; into the skin over the radial side of
the 2nd phalanx of the pollex ; into the phalangeal cartilage attached to the distal end of the 2nd
phalanx of this digit, and it is firmly fixed to the cartilaginous bar, running from the elbow down
the radial border of the radius, on the anterior side of the manus to the phalangeal distal cartilage
of the 1st digit. Over the middle of the radial side of the 1st metacarpal, the deep palmar tendon
gives a strong slip which goes down to the head of the metacarpal and the base of the 1st phalanx.
The insertion of the longus primus in Otaria and the palmaris longus in Arctoccphalus are
alike, so is the insertion of the longus secundus in the former, and the palmaris superficialis in the
latter, only the tendon of the superficialis goes down to the 1st phalanx of the 5th digit instead of
stopping at the distal end of the 5th metacarpal. The longus tertius in Otaria and the palmaris pro-
fundus in Arctoccphalus are not so close ; the latter is much more complicated, its origin being more
extensive and the insertion being very different. In Otaria it ends along the radial side of the 1st
digit. All are supplied by the median and ulnar nerves.
The Flexor communis digitorum is a combination representing and built up from — (a) the flexor
sublimis digitorum ; (6) the flexor profundus digitorum ; (c) the flexor longus pollicis.
It arises by three heads, a. The 1st head ' may be regarded as the flexor sublimis digitorum,
for it arises from the internal condyle below the palmaris longus (part two), and from the internal
lateral ligament, b. The 2nd head 2 resembles the flexor profundus digitorum, for it arises from
the inner surface of the ulna posterior to the internal lateral liagment down to 1 inch from
the wrist, c. The 3rd head3 corresponds to the flexor longus -pollicis as it arises from the posterior
half of the middle third of the radius below the pronator radii teres, and from the interosseous
membrane. The first and third heads are conjoined at the lower third of the forearm, and form
one belly ; the second head forms another, and these two bellies unite at the wrist, and thus a broad
tendon of considerable strength is formed, which immediately divides into an anterior and posterior
set of tendons ; in the anterior set there are three slender ones, in the posterior five stronger. The
anterior set is superficial and disposed like the flexor sublimis digitorum ; the posterior is deep,
and the first or outermost tendon, like the flexor longus pollicis, goes to the pollex, the remaining
four are distributed like the flexor profundus digitorum. Of the superficial set, or flexor sublimis
digitorum, the three tendons descend upon the surfaces of the deep tendons for the 2nd,
3rd, and 4th digits, opposite the middle of the metacarpal bones ; they enter the sheaths, and
over the bases of the 1st phalanges divide to give passage to the deep tendons, and then are
attached to the sides and heads of the 1st phalanges of the 2nd, 3rd, and 4th digits. Of the
deep set (five), the first tendon or flexor longus pollicis runs along the inner side of the pollex,
inside the sheath, and is attached to the head of the last phalanx. From this tendon a smaller one
springs, and unites with the sheath of the pollex at the head of the 1st metacarpal bone ; it
may be regarded as the only lumbrical. The remaining four tendons or flexor profundus digitorum
1 Humphry's flexor sublimis digitorum = Lucae's flexor communis digitorum (strongest head).
2 Humphry's flexor profundus digitorum = Lueae's flexor communis digitorum (second head).
3 Humphry's flexor profundus digitorum = Lucae's flexor communis digitorum (third head).
REPORT ON THE SEALS. 165
go to the heads of the terminal phalanges of the four fingers ; over the base of the 1st phalanges
they run through the slits in the superficial tendons. Humphry gives the flexor sublimis
digitorum as a distinct muscle with three tendons, but Lucae and I find that the three heads unite
to form a common mass, and out of this superficially the sublime tendons come, but cannot be dissected
out of this mass from their insertions to their origin.
In Aretocephalus it is also formed by three muscles — («) the Flexor sublimis digitorum, (b) the Flexor
profundus digitorum, and (e) the Flexor longus pollicis. The 1st head or Flexor sublimis digitorum is a
short band, which arises from the internal condyle of the humerus, and from the internal lateral ligament.
It is inserted into the anterior half of the flexor profundus digitorum, about the middle of the shaft
of the ulna. It is tendinous on its surface, b. The 2nd head or Flexor profundus digitorum is
covered by the sublimis in its upper half. It arises from the internal surface of the upper third of
the ulna, anterior to the ridge, from the whole breadth of the ulna as far as the lower third of this
bone, and from the internal lateral ligament which is continuous from the joint down the shaft. It
terminates in a strong tendon which widens over the carpus, c. The 3rd head or Flexor longus
pollicis arises from the whole of the inner surface of the shaft of the radius to its lower third, from
the capsule of the joint, from the interosseous membrane, and from the anterior border of the ulna
in its upper two-thirds. It descends over the carpus as a strong tendon, and joins the flexor
profundus digitorum. The single tendon thus formed soon divides into five slips, the 2nd, 3rd,
and 4th being double tendons which are anterior and posterior. The 1st or radial tendon
divides into two slips. The outer runs down the middle of the 1st metacarpal bone
and the 1st and 2nd phalanges, and is inserted into the head of the 2nd phalanx of the
pollex. The inner descends along the ulnar side of the 1st metacarpal, and is inserted into the
ulnar side of the base of the 1st phalanx of the pollex. These two pollical tendons come chiefly
from the muscle, having origin similar to the flexor longus pollicis. The 2nd, 3rd, and 4th
tendons, after splitting into anterior and posterior slips, descend over the middle of their metacarpal
bones, and over the proximal ends of the 1st phalanges the superficial slips are split opposite the
metacarpo-phalangeal articulations for the deep tendons ; and descend to the bases of the
2nd phalanges, into which they are inserted. They almost cover the 1st phalanges on their inner
surfaces, and are adherent to them. The 2nd, 3rd, and 4th deep tendons pass through
the openings in the superficial tendons, and then become anterior to the short flexors, and
are inserted into the heads of the terminal phalanges for the 2nd. 3rd, and 4th digits. The 5th
tendon is single and is inserted into the head of the terminal phalanx of the 5th digit.
The deep tendons terminate by dividing into three slips, a central strong and two lateral fine ones ;
the central slip terminates upon the terminal phalanx, the lateral pass from the sides of the central
and end on the sides of the same phalanx. The lumbricals are absent in Aretocephalus.
In Trichechus the origin of the flexor sublimis digitorum is the same as in Aretocephalus, and it
joins the head of the flexor profundus digitorum. The flexor profundus digitorum and the flexor longus
pollicis are nearly alike, except that in Aretocephalus the flexor longus pollicis has an additional origin
from the ulna. As in Aretocephalus, the flexor digitorum and flexor longus pollicis combine in the
palm for the flexor tendons. From an examination of fig. 4 (Murie) picturing the inner aspect of
the fore-limb, I find that the distribution of the tendons might be said to correspond. In Murie's
paper there is a perforating tendon to the 5th digit, not represented in the plate. As in the
Phocinas only, it has a lumbrical muscle to the pollex.
166 THE VOYAGE OF H.M.S. CHALLENGER.
In Otaria the deep and superficial flexors are very different from those of all the other species,
since they do not unite in the palm. Tha flexor sublimis digitorum has two heads which join, and it
divides into double slips for the 2nd, 3rd, and 4th digits, and a single for the 5th. The flexor profundus
digitorum comes from the ulna and radius and divides into two tendons for the pollex, forming its short
and long flexors, two short and long flexors for the index, and a single one for the 3rd digit ending
like a short flexor. From this short account it is obvious that the names flexor sublimis digitorum
and flexor profundus digitorum do not bring out the functions of these tendons, because both act as
short and deep flexors. In the Phocinse it is supplied by the median and ulnar nerves, in Arcto-
cephalus by the median. It has the usual actions.
The Palmar fascia in the Phocina?. is a quadrangular piece of fascia attached on its outer
border to the outer border of the lower end of the radius, and to the scapholunar bone. It extends
to the posterior border of the radius, where the superior angle of this side gives attachment on the
under surface to the anterior slip of part one of the palmaris longus.
The Flexor carpi radialis arises from the internal condyle between the pronator teres and the
first head of the flexor communis digitorum. It passes over the carpus, and then the tendon divides
into three. The outermost or anterior is inserted into the ulnar side of the base of the 1st
metacarpal bone ; the middle into the ulnar side of the base of the 2nd metacarpal bone ; and the
innermost or posterior into the base of the radial side of the 3rd metacarpal.
In ArctocepJudus it arises as in the Phocina?, but is placed between the origins of the pronator
radii teres and palmaris longus ; it is connected with the former for about half an inch. It is
cylindrical ; about the middle of the forearm it ends in a long, slender tendon which divides into
three very short slips. These are inserted into the ligament between the trapezium and trapezoid
bones ; into the ulnar side of the proximal end of the 1st metacarpal, beneath the first flexor brevis
muscle ; and into the radial side of the proximal end of the 2nd metacarpal.
In Arctocephalus the origin differs slightly from that in the Phocina? in its relations, and the insertion
of the 1st tendon in the former is beneath the flexor brevis into the radial side of the 2nd metacarpal
instead of the ulnar side. The 3rd tendon ends over the ligament between the scaphoid and the trape-
zoid bones, and is not long enough to reach the radial side of the 3rd metacarpal as in the Phocina?.
The origins in Otaria and Trichcchus are the same as in Arctocephedus ; the insertion in Otaria
is single, being only to the 1st metacarpal. In all, the insertion into the 1st metacarpal is constant,
and is the largest and strongest when more than one tendon is present. This gives increased
steadiness to the wrist-joint in flexion, which is necessary, because the pollex is not opposable
but bound up with the other digits by the integument, and is finger-like in actions. In all the
specimens it is supplied by the median nerve. It has the usual actions.
The Pronator radii teres arises from the internal condyle of the humerus, below the supra-
condyloid foramen, and is inserted into the inner surface of the radius, 1 inch from the lower
extremity of the shaft, by a quadrilateral bundle of fibres ; and into a very small extent of the
inner side of the anterior border, above the supinator longus.
In Arctocephalus it arises from the condyle as in the Phocina?, but is partly covered by the
flexor carpi radialis, and is united with it for half an inch ; and from the internal lateral ligament.
It descends as a flat muscle along the internal surface of the radius ; and is inserted into the
anterior border of the middle of the shaft for about one inch. In Otaria and Trickcchus it has
a slightly different arrangement from that seen in Arctocephalus.
REPORT ON THE SEALS. 167
It receives no fibres from the inner side of the coronoid process, for the upper end of the
internal condyle is at a greater distance from the coronoid and of greater length than in the
human subject. The bones of the forearm being naturally placed midway between pronation and
supination, a rotation of a quarter of a circle makes the hand prone. The mode of insertion in the
Phocinse and Arctocepludus is very different ; in the latter it is as is in human anatomy, but in the
former it is almost circular, and extends halfway across the inner surface of the radius near the
lower extremity. This strengthens the statement as to the absence of a pronator quadratus, for the
surface where it should be is partially occupied by the insertion of this muscle. In all the speci-
mens it is supplied by the median nerve, and has the usual action.
The Pronator quadratus. — No evidence was got of its presence in the Phocinse and Arctocepludus,
although Humphry states that he found it but small, and Lucae inconsiderable. There is none in
the Otaria, but in the Walrus it is fairly developed. If I had only dissected young specimens I
should be sceptical as to its absence in the Phocinse, but in an adult Seal no fibres were seen where
the above writers have described it. The dissection in this specimen was clone by turning aside the
structures without removing any tissue.
The Flexor carpi ulnaris arises from the inner surface of the ulna where the posterior border joins
the olecranon, and from the posterior border of the bone in its upper three-fourths, and is inserted
into the pisiform bone, from which the tendon passes on expanding and attaching itself to the bases
of the 3rd, 4th, and 5th metacarpal bones by joining the deep fascia adherent to them.
In Arctoce'plmlus it arises from the inner side of the 3rd or posterior tubercle of the olecranon,
to the slightest extent from the commencement of the posterior border of the ulna below the
tubercle, and from the remainder of the upper half of the posterior border by an aponeurosis
common to it and the deep palmar muscle. It lies posterior to the deep palmar, with which it is
united so closely that no division is seen in the bellies of the combined muscles. It is covered
anteriorly by the tendon common to the deep palmar, and posteriorly by the aponeurosis of origin
from the posterior border of the ulna. About an inch from the lower end of the ulna it forms a
strong tendon, which is inserted into the pisiform bone as in the Phocinse, and into the strong
fascia over the base of the 5th metacarpal. In Otaria and Triclwchus it is very like the corre-
sponding muscle in Arctoccplwhts.
All are agreed as to the pisiform being the chief point of insertion, but there are variations in the
ending of the tendon. From human anatomy we learn that there is close relationship between the an-
nular ligament and the flexor carpi ulnaris, and that the tendon terminates at the base of the 5th meta-
carpal, and this is nearly the arrangement I found in the Earless Seals ; in the Eared forms I am not
certain of the connection with the annular ligament. As the outer three digits in the Phocinre give
insertion to the flexor carpi radialis, and the bases of the three inner have this muscle fixed to them,
the whole series of metacarpal bases have each a flexor of the forearm acting upon them. Owing
to the posterior border of the ulna being arched with the concavity on the same side, and this muscle
passing in a direct line from the two extremities, the manus and bones of the forearm form two sides
of a triangle, with the pisiform as the apex. Hence this muscle must draw the hand to the inner
side, and also turn it a little to the outer side, as well as flex the manus. In all specimens it is
supplied by the ulnar nerve. It has the customary actions, and is in addition a powerful supinator
of the manus.
The Abductor minimi digiti longus. — Humphry names it flexor minimi digiti ; Lucae, abductor
168 THE VOYAGE OF H.M.S. CHALLENGER.
digiti V. It is not present in Otaria and Trichcchus. It arises from the inner surface of the ulna
where the olecranon and the posterior border meet, and passes down the posterior border of the
ulna on the flexor carpi ulnaris ; at the wrist it divides into two tendons — one is inserted into the
skin over the palmar surface of the 5th metacarpal bone, the other is bound to the sheath and
the deep fascia along the ulnar side of the 5th metacarpal, and ends opposite the ulnar side of the
head of the 1st phalanx into which it is inserted. It is both an abductor and flexor of the manus,
the flexing action commencing after the abduction is complete. It is supplied by the ulnar nerve.
The Manus. — The inner or palmar region consists of three groups of muscles. The First
Group is composed of two adductors, found only in Arctocephalus : —
The Adductor of the 2nd digit, named superficial interosseous in Otaria and Trichcchus, arises
from the base of the 3rd metacarpal between the two heads of the flexor brevis of the same digit.
It is inserted into the proximal extremity of the ulnar side of the 1st phalanx of the 2nd digit,
and is superficial to the flexores breves which it crosses.
The Adductor minimi digiti is the adductor minimi digiti in Otaria and is absent in Trichcchus.
It arises from the middle of the ulnar side of the 4th metacarpal and from the deep fascia
between the 4th and 5th metacarpals ; after passing downwards and backwards it is inserted into
the outer half of the shaft and head of the 5th metacarpal and base of the 1st phalanx.
The Second Group contains the flexores breves, which are the deep interossei in Otaria and
Trichcchus. In the Phocinre the first arises from the ulnar side of the metacarpal of the pollex ;
and is inserted into the ulnar side of the 1st phalanx of the 1st digit. In Arctocephalus it is disposed
as in the Phocinae. The 2nd interrosseus in the Phocina? is double and arises from the radial and
ulnar sides of the 2nd metacarpal. The radial head is inserted into the radial side of the base of
the 1st phalanx of the 2nd digit ; the ulnar head into the ulnar side of the same digit. It has the
same relations in Arctocephalus. The 3rd and 4th muscles in the Phocinse and Arctocephalus resemble
the last named. The 5th in the Phocina? is single and arises from the radial side of the 5th digit,
and is inserted into the same side of the base of the 1st phalanx of the same digit. In Arctocephalus
I did not observe any muscle for the 5th digit. In Otaria these muscles are in pairs for all the
digits but the 1st, which has only one. In Trichcchus the 1st and 5th digits have only one each,
the other digits two.
The Third Group embraces the following : —
The Abductor brevis jwllicis is the M. flexor pollicis of Lucae, and is wanting in Otaria and
Trichcchus. In the Phocinse it arises from the lower side of the process on the outer side of the
scapholunar bone, and from the lower border of this bone to the outer side of the tendon of the flexor
carpi radialis ; and is inserted into the front of the radial side of the base of the 1st phalanx of
the thumb. There is a sesamoid bone beneath its tendon.
In Arctocephalus it arises from the trapezium, from the upper and internal half of the 1st
metacarpal, from the radial side of the base of the 2nd metacarpal, and from the carpo-metacarpal
ligament. The greater portion is inserted into the ulnar side of the distal end of the 1st metacarpal,
and the remainder into the distal extremity of the radial side of the 2nd metacarpal. Humphry
does not mention this muscle, but Lucae describes it.
The Abductor minimi digiti is the flexor brevis minimi digiti in Otaria and is absent in
Trichcchus ; in the Phocinse it arises from the pisiform bone, and from the tendon of the flexor carpi
REPORT ON THE SEALS. 169
ulnaria ; and is inserted into the inner side of the base of the 1st phalanx. In Aretocephahis it
arises from the tendon of the flexor carpi ulnaris, from the pisiform bone, and from the tendon
of the palmaris profundus. It is well formed and is inserted into the ulnar side of the head of the
5th metacarpal ; and into the same side of the entire length of the 1st phalanx. Lucae names it
the M. flexor brevis digiti V. In Otaria Murie designates it the flexor brevis minimi digiti, and
in describing Trichcchus says " there is no trace whatever of the flexor orevis minimi digiti so well
developed in Otaria, PJwca fcetida, and Plwca vitulina." The origin and insertion coincide with those
of the corresponding human muscle, and the function is the same, being chiefly an abductor and
a flexor. The human muscle sometimes has accessory heads from the flexor carpi ulnaris and the
palmaris longus, and since a part of the origin in the Phocinse is from the former, and in Arcto-
ccphalus from the palmaris profundus and the flexor carpi ulnaris, there is much to support the name
abductor.
In Triclicchus there is an opponens pollicis, and also a palmaris brevis.
The Outer or Extensor Surface of the Forearm. — In PJwca vitulina, Plwca barbata, and
Plwca hispida the following muscles lie in this region: — Anconeus externus, supinator longus,
extensor carpi radialis, extensor communis digitorum, tensor fascise, extensor carpi ulnaris, supinator
brevis, extensor ossis metacarpi pollicis, extensor primi internodii pollicis.
In Arctoccphcdus, besides the above muscles, there is an extensor proprius pollicis, and out of the
extensor communis is formed the extensor minimi digiti. The tensor fasciae is absent. All the
specimens want the dorsal interossei and the pronator quadratus.
The Anconeus externus is a narrow slip and arises from the back of the external condyle of
the humerus, a little above the condyle ; it is inserted into the outer side of the tip of the
olecranon beside the inner head of the triceps, and into the upper half of the outer side of the
quadrilateral surface behind the sigmoid cavity.
In Arctoccphcdus it arises from the posterior part of the supracondyloid ridge, and also, as in the
Phocinse, from the back of the external condyle ; and is inserted into the olecranon on the external
lip, between the 1st and 2nd tubercles. In Arctocepludus it did not run into the triceps so
intimately as in the Phocinas. It is an extensor and lateral supporter of the elbow-joint. It is
supplied by the musculo-spiral nerve.
The Supinator longus is the most anterior of the extensors. It arises from the upper two-
thirds of the external border of the humerus, above the muscrdo-spiral groove on the outer border.
Below the head of the humerus it is covered by the external head of the triceps. After it crosses
over the external surface of the shaft of the humerus, it lies along the anterior border of the
radius, and is inserted into its anterior border, half an inch from the wrist, above the groove for
the muscle of the pollex.
In Arctoccphcdus it arises from the external border as in Plwca, above, where the musculo-
spiral nerve turns round the supracondyloid ridge, lying to the outer side of the extensor carpi
radialis, from the neck of the humerus, and from the capsule of the shoulder-joint beneath the
external head of the triceps. The extent of origin from the neck is from the external border to
the outer border of the greater tuberosity. The fibres descend between the outer head (part one)
of the brachialis anticus, and the extensor carpi radialis along the anterior border of the radius.
At the middle of the anterior border it forms a round tendon, which is inserted into the external
(ZOOL. CHALL. EXP. PART LXVIII. — 1888.) Yyy 22
170 THE VOYAGE OF H.M.S. CHALLENGER.
surface of the radius, to the outer side of the extensors of the pollex, at the junction of the epiphysis
with the shaft. In Trichechus it is single, and arises from the deltoid ridge and shaft of the
humerus. In Otaria it is double headed ; the external head comes from the upper end
of the external condyloid ridge and the internal from the deltoid ridge and joins the external
head.
Vrolik, under Section 23, describes the Extensor ossis metacarpi pollicis as the supinator longus.
Humphry describes it as inserted into the projecting margin of the radius and Lucae the same ; Dr.
Murie in Otaria into the outer side of the styloid process, and in Trichechus to the styloid process.
It appears to me that these authors have not sufficiently defined the exact place of attachment to
the radius. In the Phocinfe and Arctoccphalus there is only one groove for the tendons to the
thumb. In the former it is placed obliquely downwards and inwards across the anterior border of
the radius ; in the latter it runs down on the outer side of the anterior border ; one-quarter of the
groove in both is on the epiphysis. In the Phocinfe the supinator is inserted into the anterior
border of the radius above the groove, and the same in Arctoccphalus. It flexes the forearm, and
supinates it when prone. It is supplied by the musculo-spiral nerve.
The Extensor carpi radialis arises from the external supracondyloid ridge, lies to the outer side
of the supinator longus, and passes along the anterior border of the radius ; above the wrist it passes
below the tendon of the extensor ossis metacarpi pollicis, and then through the second division of the
annular ligament ; above the carpus it divides into three tendons. The outermost is inserted into the
base of the radial side of the 1st metacarpal bone, the middle into the dorsal surface of the radial
side of the scapholunar, and the innermost into the radial side of the base of the 2nd metacarpal bone.
In Phoca barbata there is a variation ; half an inch above the extensor ossis metacarpi
pollicis it divides into two tendons. The outer is inserted into the base of the 2nd metacarpal
bone. The inner gives off from its outer side a slip, which goes to the dorsum of the trapezium,
and then is inserted into the outer side of the 2nd metacarpal bone.
In Arctoccphalus it lies to the outer side of the supinator longus above the elbow. It arises
from the external condyle as in the Phocinfe, below where the musculo-spiral nerve turns round the
external border of the humerus, slightly from the anterior surface of the capsule of the joint over
the head of the radius, and from the external condyle. In the forearm it has the same relations
as in the Phocinfe. Before passing beneath the extensors of the pollex, it divides into two tendons of
equal size. The anterior or outer tendon is inserted into the "ulnar side of the base of the 1st
metacarpal. The posterior or inner into the upper third of the radial side of the base of the
2nd metacarpal. Vrolik, under Section 24, says of the extensor carpi radialis longus and
brevis that both rise from the upper part of the outer margin of the humerus. The brevis is inserted
into the lower part of the radius, and the longus into the outer face of the os navicularis. From
the insertions the short one is the supinator and the long the extensor carpi radialis.
Humphry and Lucae describe two metacarpal insertions. Otaria has the longus and brevis
as a common mass with two tendons. The insertions are similar to those in Arctoccphalus. In
Trichechus they are so united that they cannot be distinguished. The insertion is by a single tendon
attached equally to the 1st and 2nd metacarpals. In Phoca larbata the muscle forms two tendons
about the middle of the radius, in the other PhocinEe it is divided equally in the region of the carpus
into three, and in Arctoccphalus into two as in Phoca barbata. There is therefore an attempt to form
two muscles in Phoca barbata and Arctoccphalus, but as the origins in all the Phocinfe are from the
REPORT ON THE SEALS. 171
supracondyloid ridge, and in Arctoccphalus and Otaria from it and the external condyle, it is
difficult to comprehend whether it is a single muscle with a divided tendon, or the longus and
brevis united. The action of the extensor carpi radialis longus and brevis in human anatomy is
to extend the wrist, but after this is done the longus can flex the arm. In the Seals both actions
can be performed, and thus the function of this muscle is that of the longus and brevis.
The lower end of the radius in the Phocinse and Arctocephalus has a characteristic difference.
The scapholunar bone in the former has a very large radial tubercle, in Arctoccphalus a small one,
and in the latter the lower end of the radius articulates almost entirely with the scapholunar. In
the Phocinre the large tubercle seems to be formed at the expense of the scapholunar, for this bone
only articulates with half of the lower end of the radius : hence the outer lower half of the radius
is non-articular, the inner being the articular surface. It is supplied by the musculo-spiral
nerve.
The Extensor communis digitorum is a double muscle consisting of two separate origins ; these
are named primus and secundus. a. The extensor communis digitorum primus is named by Vrolik
M. digitorum extensores ; by Humphry, extensor communis digitorum ; by Lucae, mus. extensor
cpiatuor digit. ; and by Murie, extensor. It cerises from the supracondyloid ridge, below the extensor
carpi radialis. At the middle of the arm it forms a flat tendon, which passes through the third
division of the annular ligament. Above the bases of the metacarpal bones the tendon expands
and breaks into four tendinous slips, which pass down between the metacarpal bones to the radial
sides of the four fingers. At the middle of the 1st phalanges the tendons begin to expand
towards the ulnar sides of these bones, and at the heads of the 2nd phalanges the tendons cover
the entire dorsum. They proceed to the bases of the 3rd phalanges, where they are inserted.
The tendons adhere closely to the posterior ligaments of the joints of the digits, b. The extensor
communis digitorum secundus. Vrolik appears to call it the extensores digitorum communes breves,
Humphry the extensor secundus digitorum, Lucae the mus. abductor quatuor digitorum. It arises
from the supracondyloid ridge below the primus, and from the external condyle. It slightly over-
laps the primus ; and a little below the middle of the forearm divides into four tendons, which
pass through the fourth division of the annular ligament posterior to the primus. Two of the
tendons pass outwards beneath the tendons of the primus, and run down the ulnar sides of the
2nd and 3rd metacarpal bones. The 3rd runs down the ulnar side of the 4th metacarpal
bone, and the 4th divides into two, one going to each side of the 5th metacarpal. The tendons
of the 2nd, 3rd, and 4th metacarpals are inserted into the heads of the ulnar sides of these bones,
and into the dorsal surfaces of the proximal ends of the 1st phalanges, and also into their ulnar
sides. The tendon of the 5th metacarpal splits into two as before stated ; the anterior one is
inserted into the radial side of the head of the 5th metacarpal, and into the dorsum and ulnar
side of the proximal end of the 1st phalanx ; the posterior into the dorsum and head of the 5th
metacarpal.
In Arctoccphalus it arises from the external supracondyloid ridge, and from the extensive
lateral ligament beneath the muscle. It passes to the interosseous space and divides into two
slips, which cross the extensor pollicis proprius. From these two slips are formed the outer slip
or extensor communis digitorum, and the inner or extensor minimi digiti. The extensor communis
digitorum divides into four tendons. The 1st descends along the ulnar side of the 2nd metacarpal
bone ; the 2nd along the dorsum of the 3rd ; the 3rd descends upon the radial side of the 4th ;
172 THE VOYAGE OF H.M.S. CHALLENGER.
and the 4th runs down the radial side of the 5th metacarpal. The 1st, 2nd, and 3rd portions
expand over the heads of the metacarpals, and continue expanding until they reach the heads
of the 1st phalanges, to which they are strongly adherent. Then they pass over the posterior
phalangeal joints as fine aponeurotic sheets, which go to the terminal phalanges, and are bound
throughout their length to the bases of the phalanges and to the posterior ligaments over which
they pass. They are inserted into the dorsal surface of the heads of the 2nd, and the bases of
the 3rd phalanges of the 2nd, 3rd, and 4th digits to their radial sides. As has already been
shown, the extensor minimi digiti is derived from the origin of the extensor communis digitorum.
It divides upon the carpus into three tendons, the highest, the middle, and the lowest. The
highest is the shortest, and is inserted into the radial side of the base of the 5th metacarpal.
The middle is intermediate in size, and is inserted into the radial side of the middle of the 5th
metacarpal, between the bases of the 4th and 5th. The lowest is the largest, and goes over the
dorsum of the 5th metacarpal base as a narrow slip ; and is inserted into the head of the 1 st
phalanx, and descends from this to the terminal phalanx as a thin fibrous sheet.
Vrolik in Section 25 points out that " close by the origin of the extensor digitorum a muscle
peculiar to the Seal takes its rise," no doubt the secundus of the communis digitorum. Humphry
and Lucae found two extensors, one above the other. In Otaria the common extensors arise by a
common origin, but divide into three groups. The anterior, outer, or extensor communis digitorum
divides into three tendons for the 2nd, 3rd, and 4th digits. The middle or extensor medii digiti
has two tendons, one for the 3rd, and one for the 5th digit. The third, innermost, or extensor
minimi digiti, has tendons for the 5th metacarpal and for the 4th digit. In Trichechus according
to Murie the extensor communis digitorum has three tendons for the 2nd, 3rd, and 4th digits,
which go into the distal ends of the 1st phalanges. The extensor medii digiti is also found, but
adheres very closely to the extensor communis and ends in the interspace of the 4th and 5th
digits in four or five short tendons. The extensor minimi digiti divides into two sbps for the
5 th metacarpal.
In all the Seals and the Walrus the extensors of the 2nd to the 5th digits of the manus are of
the same type, each having an extensor mass subdivided into two sets of muscles. The Phocinae
have the two sets superimposed and they are described as primus and secundus. In Arctocephalus
the two consist of the extensor communis digitorum and the extensor minimi digiti as in man.
The Otaria and Trichechus have three muscles out of the two sets. The extensor communis being
one set, the extensor medii digiti and minimi the other. In the Common Seal, Murie states that
Duvernoy has noted that the index receives a tendon as well as the median digit, and to this
muscle in question he applies the term " extensor propre de l'index," but Murie finds in Otaria and
Trichechus no special indicator. The 1st digit in the Phocina? gives attachments to two extensor
tendons, and these are sbps from the primus and secundus division of the common extensor. Neither
of these can be called special indicators for they belong to the common group going to the other
digits. The index in these animals is a collective unit in the manus, and its action is very
much like that of the other digits, all being surrounded by the integument, as is seen by examining
the combined actions of the extensors. In the Phocinae the tendons of the extensor communis
primus are placed along the radial sides of the phalanges, and when extending the digits will also
adduct them, whereas the extensor communis secundus being on the ulnar side will extend and
abduct. In Arctocephalus the extensor communis simply extends while the extensor minimi digiti
EEPORT ON THE SEALS. 173
extends and abducts ; it therefore acts like the extensor communis digiti secundi in the Phocinae,
It may be as well to note that the Phocinae have a tendon from the extensor primus to
the 5th digit, as is seen in Arctocephalus, but not in Otaria and Trichcchus. The actions are as
usual.
The Tensor of the posterior annular ligament, named by Lucae tensor ligamenti carpi dorsalis
communis, springs from the extensor communis digitorum secundus, near the external condyle, and
consists chiefly of a narrow tendinous band. It is inserted into the middle of the upper edge of
the outer or posterior annular ligament. In Phoca hispida and Phoca barbata it could not be made
out owing to the condition of the specimens, but is most likely present in them. On referring to the
accounts of Otaria and the Walrus it will be seen that in them the extensor minimi digiti and extensor
medii digiti form one set of the extensors, and as the tensor comes out of the same set, I consider it
to be the representative of the extensor medii digiti of the Sea Lion and Walrus.
The Extensor carpi ulnaris in the Phocinae arises from the external condyle, from the outer
surface of the ulna beside the articular facet for the radius ; and from the ligament of the joint.
It passes through the fifth division of the annular ligament ; and is inserted into the middle of the
ulnar side of the 5th metacarpal bone.
In Arctocephalus the anconeus externus partly covers its origin. It arises from the external surface
of the olecranon, between the anterior and middle tubercles (PL VII. fig. 4) ; and from the outer edge
of the quadrilateral surface behind the great sigmoid cavity of the ulna. It descends to the carpus
on the inner side of the externus, and, after crossing the extensor proprius pollicis, is inserted into
the base and head of the ulnar side of the 5th metacarpal, but chiefly into the head of the first
phalanx on the ulnar side. In Otaria and Trichcchus it is inserted only into the 5th metacarpal. It
extends the manus, powerfully abducts it, stretching out the digits, besides aiding extension of the
forearm. It is supplied by the posterior interosseous nerve.
The Supinator brevis arises from the external condyle of the humerus, and from the ligament of
the elbow-joint on the external surface. It is inserted into the upper two-thirds of the anterior
border of the radial shaft, into the anterior third of the inner surface down to the pronator teres
insertion, and into the anterior two-thirds of the outer surface to the same insertion.
In Arctocephalus this muscle is hidden on the outer aspect of the radius by the downward
expansion of the extensive lateral ligament. It arises from the posterior and external surface of
the external condyle below the external lateral ligament, from the posterior aspect of the capsular
ligament covering the posterior surface of the condyle, from the capsule over the outer side of the head
of the radius, also from the outer half of the capsule covering the anterior aspect of the head of the
radius, and slightly from the front of the capsule covering the anterior side of the external condyle.
The fibres from the back of the condyle are mostly tendinous, and all are parallel to the radial
shaft. It is inserted into the anterior half of the neck on its outer surface, into the external surface
of the shaft of the radius, as far down as the lowest fibres of insertion of the pronator radii
teres, that is about the middle of the bone ; into the neck of the shaft of the anterior border to
the insertion of the same muscle, and slightly into the inner border below the neck.
In Trichcchus, as in Arctocephalus and the Phocinae, it has only one head of origin, but in
Otaria it has two heads, the additional one from the ulna below the coronoid process. It supinates
the forearm and steadies the joint. It is supplied by the posterior interosseous nerve.
In Arctocephalus the Extensor proprius pollicis covers a considerable part of the flexor carpi
174 THE VOYAGE OF H.M.S. CHALLENGER.
ulnaris. It arises from the outer surface of the olecranon between the middle and posterior tubercles,
from the ulna posterior to the ridge running down the shaft from the middle tubercle, and from
the external surface of the ulna, where the ridge ends, to 1 inch from the epiphysial line. It has
also fibres of origin from the fascial expansion of the external lateral ligament, which passes
beneath it to attach itself to the lower end of the ridge of the ulna. It forms a flat strong tendon,
and crosses obliquely forwards and outwards to the interval between the radius and ulna. At the
level of the epiphysial line it enters the annular ligament, and passes over the posterior inferior
corner of the lower end of the radius. It reaches the carpus and goes between the 1st and 2nd
metacarpal bones ; at the middle of the 1st metacarpal it begins to expand to the outer or anterior
side, the tendon crossing the head of the metacarpal. Above the wrist it is crossed by the extensor
carpi ulnaris, and over the wrist by the common extensors. Upon the base of the 2nd meta-
carpal it crosses the posterior tendon of the extensor radialis, and then descends upon the dorsum
of the head of the 1st metacarpal, over which it joins the outer side of the extensor prirni
internodii pollicis. It is inserted into the base of the 1st phalanx of the pollex, and sends a fine
aponeurosis to the terminal one. The posterior capsule of the joint of the 1st and 2nd phalanges
of the pollex is chiefly formed by it.
This muscle is wanting in the Phocinse, and comes under the heading " extensor pollicis et
indicis " in Dr. Murie's papers on the Pinnipedia. In Otaria it arises almost from the same locality
of the ulna, and is inserted into precisely the same part of the pollex as in Arctoccphalus. The surface
of the ulna which gives origin to this muscle in Arctocephalus, Otaria, and Trichcchus, is occupied
by the extensor primi internodii pollicis in the Phocinaa. It is supplied by the posterior inter-
osseous nerve.
The Extensor ossis metacarpi pollicis is the abductor pollicis of Lucae. It arises from the
external flat surface of the olecranon, beginning above the tip of the olecranon, and extending along
its border to where the posterior border of the ulna begins; from the anterior upper half of the
outer surface of the ulna; slightly from the external lateral ligament; from the outer surface of the
interosseous ligament ; and by a few fibres from the middle of the posterior border of the radius.
After receiving the radial fibres it forms a flat tendon which descends obliquely downwards and
forwards over the outer surface of the radius, crossing the extensor carpi radialis tendon above the
wrist. It enters the first division of the annular ligament and runs in the groove on the anterior
border of the radius above the styloid process, then over the palmar surface of the process or
tubercle of the scapholunar bone; and is inserted into the radial or anterior side of the proximal
end of the 1st metacarpal bone.
In ArctoccpJialus it arises from the external surface of the middle of the shaft of the ulna,
anterior to the ridge which descends from the middle tubercle, below the origin of the extensor
primi internodii pollicis, which is not covered by the attachment of the fascial expansion of the
external lateral ligament, from the posterior surface of the interosseous membrane, and by a small
surface from the external side of the posterior border of the radius in front of the ulnar origin.
It passes obliquely downwards and forwards, over the external surface of the radius, under cover
of the expansion of the external lateral ligament. Crossing the tendons of the extensor carpi radialis,
it enters the first division of the annular ligament on the anterior border and external surface of
the lower end of the radius, and runs along the side of the carpus to the outer side of the extensor
primi internodii pollicis; and is inserted into the radial side of the base of the 1st metacarpal.
EEPOET ON THE SEALS. 175
In Otaria "it takes origin from the outer surface of the olecranon, and from the ulna to as far
as about the middle of the latter, and is inserted into the prominent and anterior or outer corner of
the metacarpal of the pollex."
In Trichechus it is combined with the extensores primi and secundi internodii pollicis. This
muscle in the Phocinae covers that part of the ulna which, in Arctocephalus, gives origin to the
extensor ossis metacarpi pollicis and the extensor primi internodii pollicis, in Otaria gives origin
to the same muscles, and in Trichechus to the extensor ossis metacarpi, extensor primi internodii,
and extensor secundi internodii pollicis. It is supplied by the posterior interosseous nerve.
The Extensor primi internodii pollicis arises from the posterior third of the outer surface of the
olecranon; from the posterior border of the ulna in its upper half; and very slightly from the
shaft where the extensor ossis metacarpi begins to cross the radius. One inch above the wrist it
forms a tendon, which passes beneath those of the extensor communis digitorum secundus, and
enters the third division of the annular ligament with and below the extensor communis primus.
It is, inserted into the base of the 1st phalanx of the thumb on the anterior or radial side; and
into the head of the metacarpal.
In Arctocephalus the extensor digitorum and the extensor carpi ulnaris must be turned up
before the origin of this muscle is seen. It arises from the external surface of the olecranon
between the anterior and middle tubercles; from the edge of the quadrilateral surface in front
of the anterior tubercle; from the sigmoid cavity of the ulna; from the concave outer sur-
face of the ulna, as far as the origin of the extensor ossis metacarpi pollicis ; from the external
surface of the ligament of the capsule of the elbow-joint ; from the interosseous ligament ; and from
the upper half of the posterior quarter of the breadth of the shaft of the radius. Its tendon
crosses the radius, and goes through the same division as the extensor ossis metacarpi pollicis.
Opposite the head of the 1st metacarpal it is bound to the palmar fascia, and is inserted into the
radial side of the base of the 1st phalanx of the pollex, being joined to the extensor proprius
pollicis.
Lucae gives the insertion into the base of the 1st metacarpal, but Humphry gives a different
insertion into the back of the 1st phalanx of the pollex, which is the phalanx for the insertion
of the extensor primi internodii pollicis.
In Otaria this muscle is wanting In Trichechus it is inseparably united with the extensor
ossis metacarpi pollicis. That part of the ulna upon which this muscle is implanted in the Phocinae
gives origin in Arctocephalus, Otaria, and Trichechus to the extensor proprius pollicis. It is
supplied by the ulnar nerve.
According to those works on the Pinnipedia that I have had an opportunity of reading, the
outer surface of the ulna in the Phocinae and Arctocephalus gives origin only to muscles for the
pollex. The surface to which the fibres are attached, generally speaking, is the upper two-thirds of
the shaft. In Arctocephalus the outer surface of the ulna is divided into two by a ridge commencing
from the middle tubercle on the outer side of the olecranon, and becoming indistinct at the junction of
the upper two-thirds and the lower third of the shaft (PI. VII. fig. 4). In the Phocinae there is a slight
ridge dividing the external surface of the olecranon into two, which comes close to the posterior border
of the ulna f of an inch below the junction of the posterior border with the olecranon, runs down the
shaft close to it, and ends at the middle of the shaft. These ridges separate the muscle fibres,
which clothe the external surface of the shaft, into two groups of the same functional importance
176 THE VOYAGE OF H.M.S. CHALLENGER.
for the pollex. The posterior group gives origin to one muscle in all, and this is the extensor
proprius pollicis in Arctocephalus, the extensor pollicis et indicis in Otaria and Trkhechus ; but
as these three are exactly alike in their mode of insertion, practically they are the same, and all
mi"ht be as correctly named extensor proprius pollicis. In the Phocinse the muscle is called the
extensor primi internodii pollicis, but only provisionally, for its origin is exactly as in all the others,
but its insertion is half that of the proprius and half that of the extensor primi internodii, for
a part of the insertion crosses the base of the 1st phalanx of the pollex, and the other half
is continued down the radial side like the extensor primi internodii ; thus it has a compound
action.
The anterior group in the Phocinse and Otaria is similar, forming the extensor ossis meta-
carpi only ; in Arctocephalus two muscles come from this group owing to its division into an upper
and a lower half. The upper half is the extensor primi internodii pollicis, the lower the extensor
ossis metacarpi pollicis. In Trichechus there are three muscular elements — the extensor ossis
metacarpi pollicis, the extensores primi et secundi internodii, all combined at their origin, and indis-
tinguishable from the common mass of fibres. This affords evidence of a distinct portion of a bony
surface being reserved for a muscle discharging a special function with respect to a certain digit.
The area of origin in all these animals remaining constant and not increasing in size, whether there
is a single tendon or more, is probably novel. It leads to the conception that there must be
a method in the formation of distinct muscles out of common masses of fibres, though it may not
always be traceable. These muscles have the usual actions.
Myology of the Hind Limb.
The Ilio-Femoral Region includes the psoas and iliac muscles, with a varying set of muscles
in connection with the former. The following shows the various accessory muscles found in each
specimen. The meaning of the names is explained further on : —
The Psoas magnus or primus is present in the large Phoca vitulina ; in the small Phoca
vitulina with an ilio-femoralis et lumbalis anterior; in Phoca barbata with an ilio-femoralis et
lumbalis anterior; in Phoca hispida with an ilio-femoralis anterior; in Macrorhinus; and in
Arctocepluilus.
The Psoas minor or sccundus is found in all the above specimens and in the large Plwca
vitulina, on the right side, with a lumbo-femoralis posterior.
The Psoas tcrtius is common to all but Macrorhinus, and it has an ilio-femoralis posterior.
The Eiacus is found in all the specimens.
Before entering upon the details of the psoas, it is well to understand upon what grounds
the names are given, as well as to point out what peculiarities each muscle possesses, and the
similarities and dissimilarities in each dissection. As much interest in the anatomy of the Seals
centres around this group of muscles the localisation of the fibres is of importance. Upon the lumbar
vertebras ventrally there are two longitudinal fleshy masses, each including a psoas magnus and parvus.
As one of these muscles is attached in all the specimens, though with some modifications in detail,
to the pectineal eminence, I regard it as the equivalent of the psoas parvus of human anatomy, called
the secundus or minor in the text. The other muscle, lying to the outer side of the minor, has not
REPORT ON THE SEALS. 177
the usual femoral attachment of the psoas maguus, but its position in the trunk and the want
of the trochanter minor in the Phocince and Maerorhinus, indicate that the niagnus must wander
to some other point, and it has settled on the posterior ventral spine of the ilium. The names
maguus and minor do not suit the magnitudes of these muscles in the Earless Seals, for the minor is
by far the larger, and the major along side of it is small, but it is convenient to keep the names
used in human anatomy. The tertius is a small muscle not to my knowledge previously described
in the Seals, and called the iliacus by Dr. Murie in Otaria. It is located upon the junction of the
lumbar with the sacral vertebras under cover of the other two muscles, and is directed obliquely over
the pelvic brim to the femur. It exists in all the specimens excepting Maerorhinus. The want
of a psoas magnus to the femur is compensated for by fibres springing from the posterior ventral
spine of the ilium, or from the pectineal eminence with, in some instances, a prolongation of fibres
from the psoas magnus into this group, or a direct offshoot from the psoas minor. The name ilio-
femorahs anterior is given when the fibres are only derived from the posterior ventral spine, and
ilio-femoralis posterior when from the pectineal eminence. In two specimens (small Phoca vitulina
and Phoca barbata) an addition is required to the name ilio-fernoralis anterior. The psoas magnus
in them gives fibres to blend with the ilio-femoralis anterior and then the name ilio-femoralis et
lumbalis anterior may be adopted. Lastly, in the large Phoca vitulina, the psoas secundus gives a
group of fibres directly to the femur, and this is called the lumbo-femoralis posterior.
The Psoas magnus is called the psoas major and ilio-lumbalis by Lucae ; in Phoca vitulina it
arises from the sides of the ventral surface of the 3rd, 4th, and 5th lumbar vertebras to the inner
side of the psoas tertius, which it covers and crosses, to be inserted only into the posterior ventral
spine of the ilium. It has no ilio-femoralis anterior.
In the small Phoca vitulina most of the fibres are inserted into the posterior ventral spine
of the ilium. It has an extension of its fibres forming an ilio-femoralis et lumbalis anterior. In
Humphry's account of this muscle he points out that some of the fibres pass to the inner side of
the thigh ; this is the group which Lucae figures as the femoralis major and has not described.
In Phoca barbata this muscle has the same insertion and distribution as in the last species.
In Plwca hispida it is inserted into the anterior half of the posterior ventral spine of the ilium,
and slightly into the adjacent inner half of the ventral border of the outwardly directed wing of the
ilium. There is a great difference between this muscle and the corresponding one in the small Phoca
vitulina, for not one of the fibres proceeds beyond the spine. It has, therefore, no lumbalis fibres
from the psoas magnus, but it has an ilio-femoralis anterior.
In Maerorhinus leoninus it is inserted into the posterior ventral spine of the ilium, which is fused
with the pectineal eminence.
In Arctoccphalus (jazella this is much the largest of the group. In the dorsal region it arises
by a series of muscular slips, from the posterior halves of the last four dorsal vertebra;, from their
intervertebral discs, and from the ventral surfaces of the ribs and the ligaments of the rib joints.
In the lumbar region it arises from the whole of the ventral surfaces of the 1st, 2nd, 3rd, and 4th
lumbar vertebra;, and from their intervertebral discs and transverse processes. It courses directly
backwards, above the psoas minor, to the outer side of the tertius, becomes narrower, and is
inserted into the posterior ventral spine of the ilium. There is no ilio-femoralis et lumbalis anterior,
or ilio-femoralis anterior, in this animal nor in Maerorhinus leoninus.
The Ilio-femoralis et lumbedis anterior, in the small Plwca vitulina, is the ilio-psoas and ilio-
(zool. chall. exp. — part lxviii. — 1888.) Yyy 23
178 THE VOYAGE OF H.M.S. CHALLENGER.
fenioralis of the text of Lucae, and the ilio-femoralis major and minor of his drawing. It is in part
formed by a continuation onwards of some of the fibres of the psoas magnus, and by fibres spring-
ing from the posterior ventral spine of the ilium. A number of the superficial fibres of the psoas
magnus pass over the spine, and run with a fresh set of fibres which arise from the posterior
ventral spine. The fibres from these two sources proceed backwards and outwards, lying to the
outer side of the psoas tertius, and are inserted into the lower outer third of the inner border of the
femur.
In Phoca barbata it arises from the posterior half of the posterior ventral spine of the iliuni,
from the side of the sacrum, from the sacro-iliac ligament, and from the ventral surface of the
ilium along the anterior or upper border to the spine. A few fibres of the psoas magnus run into
it, and after receiving them it courses along the ©uter side of the psoas tertius, and unites with it
near the supracondyloid ridge on the inner border of the femur.
The Ilio-femoralis anterior in Phoca hispida arises from the posterior half of the posterior
ventral spine of the ilium, from the ventral surface of the sacro-iliac ligament, from the sacrum,
and from the ventral surface of the ilium posterior to the spine. It passes to the lower end of the
inner border of the femur, and is- inserted into the supracondyloid ridge in front of the psoas tertius,
and- is partly blended with it. Lucae had obviously recognised this muscle, and though he names it
in his plate ix., he does not describe it in the text.
The Psoas minor or sccundus in the larger Phoca vitulina, as in the Earless Seals, is the largest
muscle of this group. It arises from the ventral surfaces of the 14th and 15th ribs and their
rib-joints, from the sides of the 14th and 15th vertebrce, from the ventral surfaces of the bodies of
these vertebrae, and from the ventral surfaces of the transverse processes of all the lumbar vertebrae,
and is inserted into the pectineal eminence. A similar insertion has been recognised by the
authorities already named in the specimens they describe.
In the small Phoca vitulina, Phoca harbata, Phoca hispid a, in Macrorhinus, and in Arctoccphalus
it is much smaller than the psoas major, and is a thin fusiform band, which arises by short tendons
from the posterior aspect of the rounded tips of the transverse processes of the 2nd, 3rd, and 4th
lumbar vertebras. It crosses inwards ventrally to the psoas magnus and tertius, and is inserted into
the pectineal eminence.
In the large Phoca vitulina the lumbo-femoralis posterior is found on the right side only.
After dividing the tertius and turning the two ends aside, a muscular band is exposed, which is the
direct continuation of the psoas secundus* This is a flat riband-shaped band of fibres from the
psoas secundus passing over the insertion of this muscle into the pectineal eminence, and turning
outwards upon the psoas tertius to the lower end of the femur on the inner border into which it is
inserted. The secundus of the left side had no such distribution, and all its fibres ended in the
pectineal eminence.
The Psoas tertius in the large Phoca vitulina is the most inferior, and passes beneath the psoas
magnus over the pelvic brim to the lower inner border of the femur. It arises under co\er of the
magnus from the junction of the ventral surfaces and sides of the two last lumbar vertebras. It
lies upon the lumbo-femoralis on the right side and the iliacus on the left, and is inserted into the
termination of the femoral ridge on the inner border of the femur at the lower end.
In the small Phoca vitulina it is a band of muscular fibres 1 inch broad, stretching from the
lowest lumbar vertebra to the femur. It arises from the ventral surface of the hinder half of the
REPORT ON THE SEALS. 179
/
last lumbar vertebra, from the anterior half of the 1st sacral vertebra, from their intervertebral
discs, and from the anterior sacro-iliac ligament. It turns over the ilium between the anterior
and inferior spines and the pectineal eminence, going beneath the tendon of the psoas minor which
passes to the pectineal eminence. Hence it goes downwards to the lower end of the inner
border of the femur, and is inserted into the supracondyloid ridge, occupying the lower third of the
internal border of the femur above the condyle and below the ilio-femoralis et lumbalis anterior
found in this specimen.
In Phoca hispida it arises from the ventral surface of the side of the body of the last lumbar
vertebra, and from the intervertebral plates above and below {i.e., anterior and posterior to this
vertebra). It descends from the vertebral column over the iliac synchondrosis below the tendon of
the psoas minor or secundus, and is inserted into the supracondyloid ridge as the last.
In Phoca barbata it arises from the last lumbar vertebra, but only from the lower or posterior
part of its ventral surface, from the ventral surface 1st vertebra of the sacrum, from the inter-
vertebral disc anterior to the last lumbar, and from the disc between the last lumbar and the
sacrum. The course is the same as in the small specimen of PJwca vikdina, and it is inserted as the
other muscles.
In Arctoceplmlus gazclla it is situated to the inner side of the psoas minor, and arises from the
sides and ventral surfaces of the lower border of the second last lumbar vertebra, from the upper
half of the same part of the last lumbar, and from the intervertebral disc between it and the
2nd lumbar, and from the root of the transverse process of the last lumbar. From the sides of
the vertebral column it descends, partly hidden by the large psoas minor. At the level of the
insertion of the psoas major it passes beneath the tendon of the minor, and after crossing the
capsule of the hip-joint turns round the inner surface of the femur. It is inserted into the inferior
large surface of the small trochanter of the femur, behind the insertion of the iliacus and in front of
the insertion of the pectineus.
In Macrorhinus leoninus there is no psoas tertius coming from the vertebral column, but there
is an Plio-femoralis posterior. It arises only from the entire length of the outer side of the pecti-
neal eminence, reaching as far forwards as the tendon of insertion of the psoas magnus, which it
slightly overlaps. It is inserted into the supracondyloid ridge on the inner border of the femur,
just as are the ilio-femoralis anterior, the ilio-femoralis et lumbalis anterior, and the lumbo-femoralis
posterior found in the various animals.
The Iliacus, as a separate muscle, was found in three specimens. From the distortion of the
bony parts and the small size of the ventral surface of the ilium one might easily be led to suppose
that there was no iliacus. This idea would seem not unwarranted, seeing that the femur has no
trochanter minor in the Phocinre and Macrorhinus (PI. IV. fig. 4). But when one turns to a
large specimen of a Phoca, good reason may be found for the identification of this muscle.
In the large Phoca vikdina it arises from the ventral surface of the ilium between the inser-
tions of the psoas magnus and minor. This surface is equivalent to that portion of the ventral
surface of the human ilium, which is immediately above the ilio pectineal eminence. It passes
beneath the psoas secundus tendon on both sides, and the lumbo-femorahs on the right. The
former is embedded in it on both sides, and the latter also on the right. The pectineal eminence
being very prominent, and at its anterior end perpendicular to the ventral surface of the ilium, the
fibres which arise from this surface are at right angles to those from the ventral surface of the
180 THE VOYAGE OF H.M.S. CHALLENGER.
ilium, so that it appears like two muscles. According to these two directions of the fasciculi, it is
inserted in two ways ; the fibres from the ilium into the anterior part of the inner border of the femur,
the pectineal fibres into the femur posterior to the others. In Phoca the iliacus does not exist
according to Duvernoy ; Murie gives a description of it, and Humphry states that it was represented
by a few fibres.
In Macrorhinits leoninus it arises from the outer half of the ventral surface of the ilium, and
the inner half of the ventral border of the wing. It lies below the origin of the rectus toe th
outer side of the insertion of the psoas magnus. It passes backwards and inwards, and joins the
middle third of the outer side of the ilio-femoralis posterior. In all the specimens this muscle,
when present, takes origin between the insertions of the psoas magnus and minor, but in Maeror-
hinus the magnus is posterior to the iliacus. Its name is due to the fact that it joins the ilio-
femoralis posterior, and comes from the ventral surface of the ilium.
In Arctocephalus gazella it is a slender fasciculus, and arises from the ventral surface of the ilium
between the ventral posterior spine and the pectineal eminence, and from the anterior or ventral
sacro-iliac ligament. The psoas tertius lies upon it, and is adherent to its fibres, from origin to
insertion. The course is the same as that of the psoas tertius, but the fibres of the iliacus are fixed to
the capsule of the hip. It is inserted into the superior small surface of the small trochanter of
the femur above the psoas tertius (PI. VII. fig 7). In Murie's description of the iliacus, the
lumbar and iliac fibres are made a common muscle. I made the dissection by cutting away the
psoas tertius (Murie's iliacus) near its insertion and drawing it backwards, and found the iliacus
fibres upon the ventral surface of the iliaum, having a different point of insertion into the lesser
trochanter.
The muscles of the ilio-femoral region situated on the posterior and anterior spines of the
ilium, and around the pectineal eminence along the inner side of the femur to the inner condyle,
can only be differentiated by following closely the fibres from origin to insertion. The bulk
of the muscles are not divided by well-defined fibrous septa as in many other Mammals, and the
smaller muscles especially are in consequence difficult to isolate. The points of origin must be
sought for and considered beforehand, otherwise many artificial muscle bundles would be formed and
confusion result from divisions made. This has been most carefully attended to with this group,
as they are so closely applied that no definite result could have been otherwise obtained. In the
small specimens, the smallness of the space available for work, the fineness of the fibres,
and the presence of a quantity of fat added considerably to the difficulty of the problem.
However, sufficient evidence was obtained to put beyond doubt that a psoas primus, secundus,
and tertius were present in all but Macrorhinus leoninus, which lacks the tertius, and that in
all three there is a distinct iliacus. In the Phocinae this muscle is supplied by the anterior crural
nerve.
The interpretation of the muscles called " ilio-femoralis et lumbalis anterior " springing from the
psoas magnus and the posterior ventral spine, " ilio-femoralis anterior " from the posterior ventral
spine, "ilio-femoralis posterior" from the pectineal eminence, and " lumbo-femoralis " from the
psoas minor presents no intricacy. All these are inserted into the inner border of the femur with
trifling variations as to extent and locality. Some are directly connected with the psoaa, or are in
close proximity to their insertions. From this we see that all these various origins maintain a
connection with the psose, and that they are the representatives of it in the thigh.
REPORT ON THE SEALS. 181
The variations in shape and alterations in size of the ventral surface of the ilium have no doubt
been the cause of the want of exactness and the difficulty in describing the iliacus. In Arctocephalus,
which is a near relation of Otaria, the ventral surface of the ilium is well marked ; in my specimen
it was fully an inch long and half an inch broad. In the large Phoca vitulina it was also well
formed, in a large Phoca grcenlandica it was half an inch long by half an inch broad. In the
small Phoca vitulina three-quarters of an inch long by quarter of an inch broad. In Phoca hispida
barely half an inch long by two lines broad, and in a specimen of Phoca vitulina in which the pubic
bar and the ilium were not fused it was only a border. In Macrorhinus the ventral surface was in-
creased by the broadening of the ventral border of the wing of the ilium, and the ventral surface and
broad ventral border were not recognisable as such, for the one was continued into the other. Meckel,
Humphry, and Murie agree as to there being an iliacus arising from the ventral surface of the ilium
in Phoca vitulina. Murie believes the iliacus is present in Otaria and Trichechus, but describes
it as coming from the spinal column as well as from the ventral surface of the ilium, and calls
it a serni-dividecl iliacus. Instead of naming this an iliacus, I have named the spinal fibres
psoas tertius, and the iliac fibres as the iliacus. In Macrorhinus only is there an iliacus without
a psoas tertius. In the Phocinas and in Arctocephalus the psoas tertius lies upon the iliacus, and
in the small specimens is intimately fused with it.
The Ventral Femoral Eegion in the Phocinas, Macrorhinus Iconinus, and Arctocephalus gazella,
is composed of the tensor fascias femoris, sartorius, rectus femoris, vastus externus, and crureus.
The vastus internus and subcrureus are wanting.
The Tensor fascim femoris in Phoca vitidina arises from the fascia lumbo-dorsalis in its inner
half, from the erector spinas in its outer half. It forms a band, which descends between the
anterior ventral spine and the posterior ventral spine of the ilium, just touching both. After
crossing the iliac crest it sweeps backwards, forwards, and inwards. Above the external condyle
of the femur it forms a tendon which is inserted into the deep strong fascia over the head of the
tibia and fibula. The tendon extends from the middle of the head of the tibia to the middle of the
head of the fibula, on the outer side, and into the lowest three-fourths of the edge of the patella.
In Phoca hispida the origin is similar. It is inserted into the outer edge of the patella, into the
outer edge of the ligamentum patellae, and into the head of the fibula to the dorsal side of the
ligamentum patellse. In Phoca barbata it arises from the fascia over the erector spinas only, and is
inserted as in Phoca hispida.
In Macrorhinus it arises from the lumbo-dorsalis fascia, one inch and a half above the iliac
crest, and from the erector spinas. It descends over the anterior half of the crest of the ilium,
joins the dorsal border of the tendon of the sartorius above the patella, descends along its
outer edge to the tibia, and is inserted into the fascia over the head of the tibia, dorsally to the
sartorius.
In Arctoccpludus gazella it arises from the fascia lumbo-dorsalis, three-quarters of an inch from
the spinal column opposite the middle of the space between the 3rd and 4th lumbar vertebrae to
opposite the spine of the 5th. The muscular fibres commenoe at the edge of the erector spinas.
It passes backwards and forwards to the knee, crossing between the ventral anterior and posterior
spines of the ilium, and is inserted by muscular fibres into the dorsal half of the patella, ending in
the ventral border of the ligamentum patellae. Lucae considers it as a muscle cover, otherwise his
1S2 THE VOYAGE OF H.M.S. CHALLENGER.
view is not different from the above. Murie does not notice it in the text or the drawings
of Otaria and Trichechus. From the position of this muscle, reaching from the spinal column
over the "luteal muscle to the outer side of the knee, it must act as a flexor of the thigh and a
rotator outwards of the thigh and leg.
The Sartorius in Phoca vitulina is an elongated slip. It arises from the ventral anterior spine
of the ilium, descends backwards, forwards, and slightly inwards, lying to the inner side of the
tensor fasciae femoris. It is inserted into the upper edge of the patella. In Phoca hispicla it arises
also from a small part of the lower lip of the iliac crest, and is inserted as in Phoca vitulina, but
the fibres are also attached to the inner edge of the patella, into the ventral edge of the ligamentum
patella?, a few fibres descending along the inner side of the ligament to the head of the tibia. In
Phoca barbata it has the same relations as in Phoca vitulina.
In Macrorhinus Iconinus it arises from the lower edge of the ventral anterior spine of the ilium,
which is a continuation of the outer lip of its crest, and from half an inch of the outer lip of the
ilium. Its course is as in Phoca vitulina, but it is adherent to the rectus above the patella, and
is joined on the dorsal edge by the tensor fascia? femoris. The tendon is broad, and is inserted into
the outer edge, and into the outer two-thirds of the superior edge of the patella. The tendon
unites with that of the rectus and goes with it to the tibia.
In Arctoccphalus gazella there was a second sartorius, much smaller than the proper one and
posterior to it. The proper or anterior arises from the ventral anterior spine of the ilium, and
from the ventral border behind the ventral anterior spine for a slight distance ; it passes downwards
and outwards, and is inserted into the inner edge of the patella, sending a fascial expansion to the
fascia over the internal condyle to the head of the tibia. The posterior muscle arises from the
middle of the venter of the ilium by a slender tendinous slip, descends to the knee, and is inserted
over the internal border of the fascial expansion of the anterior one. Lucae only recognises part of
the external oblique as its analogue. It appears in Murie's drawings of Otaria and Trichcchus but
not in the descriptions. It flexes the thigh, and may slightly adduct and evert it.
A very noticeable fact is the uncertainty as to the precise insertion of this muscle, its tendency to
variation, and the fixity of the origin. Some may be in favour of regarding the tensor vagina?
femoris and the sartorius in the Thocina? as offshoots from the external oblique, but the dissections
appear to me to indicate that they are distinct from it. I look upon the external oblique as a
progressive muscle, increasing in size and carried backwards in adaptation to the crawling movement
of the Seals along the ground. In Arctoccphalus where there is no crawling the external oblique
follows the ilium and pubes. In the Phocina? it is supplied by the anterior crural nerve.
The Pectus femoris in Phoca vitulina is a single-headed muscle, and arises from the ventral
fourth of the anterior border of the outer surface of the ilium, from the outer surface between
the anterior ventral spine and the origin of the gluteus minimus on the outer side, which is about
the ventral half of the outer surface, and from below the capsule of the hip-joint, anterior to the
middle of the acetabulum. Tims it springs from the outer surface of the ilium, from a surface
bounded anteriorly by the anterior border of the ilium, posteriorly by the capsule of the hip-joint,
where it is attached to the ilium on the dorsal side by the origin of the gluteus minimus, and on
the ventral side by the ventral posterior spine of the ilium. The fibres form a rectangular band,
which is partly behind the sartorius and the tensor fascia? femoris. After passing downwards,
forwards, and slightly outwards, it is inserted into the upper edge of the patella, and is united with
REPORT ON THE SEALS. 183
the fibres of the vastus externus near the patella. In Plwca hispida it arises from the outer surface
of the ilium ventral to the ridge between the ventral anterior spine and the middle of the anterior
rim of the acetabulum. It does not arise from the ventral border, for the representative of the
iliacus comes from the ventral posterior spine. It is inserted into the upper edge of the patella
and is also joined above its insertion by a few fibres of the external vastus. In Phoca barbata it
arises from the fossa of the posterior third of the outer surface of the ilium ; and from the capsule
of the hip-joint, but not from the anterior border of the ventral surface. It is inserted into the
upper edge of the patella, and is also united to the fibres of the vastus externus near the insertion.
In Macrorhinus Iconinus it arises from the external surface of the ilium, to the ventral side of
the ridge running from the outer side of the ventral anterior spine to the middle of the
acetabulum. Where the ridge is covered by the capsule it takes origin from it, and arises also
from the outer half of the ventral surface of the ilium dorsal to the origin of the iliacus. Above
the patella it forms a broad flat tendon, which is joined on its dorsal side above the patella by
fibres of the external rectus, and is inserted into the upper edge of the patella.
In Arctocephalus gazella it arises from the external surface of the ilium ; and from the capsule
covering the front of the acetabulum. If a straight line be drawn from the ventral anterior
spine to the middle of the anterior rim of the acetabulum, a triangular surface is mapped out,
which is bounded ventrally by the ventral posterior spine, posteriorly by the rim of the
acetabulum, on the inner side by the outer edge of the brim of the pelvis ; and dorsally by the
line from the ventral posterior spine to the acetabulum. Within this space is the origin of the
muscle. It is inserted into the upper edge of the patella, and is joined on its outer side near the
insertion by fibres from the vastus externus. Lucae has it as united to the cruralis, but I found it
blended with the vastus externus. It is similar hi Otaria and Tricheehus.
As it is the reflected tendon in human anatomy that acts mostly upon the thigh, and this is
the origin in the Seals corresponding to it, it is probably only a flexor of the thigh. It is
relatively a much larger muscle than in human anatomy and will compensate for the vastus internus.
In the Phocinaj it is supplied by the anterior crural nerve.
The Vastus externus in the Phocinffi and Macrorhinus Iconinus is best seen in the last named
animal, and extends along the outer border of the femur to the epiphysial line of the external
condyle ; i.e., to the supracondyloid ridge, and is similar in all. It is placed upon the crureus
and covers a considerable part of it ; it arises from the capsule surrounding the neck of the femur,
from the shaft between the inner termination of the great trochanter on the neck, from the whole
length of the anterior edge of the great trochanter, slightly from the shaft below this, and from
the external border of the femur in its upper half. It is blended with the crureus and is
inserted into the upper and outer half of the patella, and into the capsule of the knee-joint, on a
level with the middle of the outer edge of the patella. No fibres descend further down the capsule.
In Arctocephalus gazella it is a rectangular muscle, lying to the outer side of the rectus femoris,
and partially overlapping the crureus. It arises from the front surface of the femur below the
great trochanter; from the front of the neck of the same; from the capsule of the hip-joint, and
from the external border of the shaft, almost reaching the external condyle. It is inserted into the
outer upper half of the patella, and blends with the rectus femoris. Lucae describes a combined
crureus and vastus externus. In Otaria it is attached to the whole anterior surface of the femur, as
also in Tricheehus.
\\
184 THE VOYAGE OF H.M.S. CHALLENGER.
In Macrorhinus, and in all the femora of the Phocinae, there is an oblique ridge running from
the top of the great trochantei on the front surface to the external condyle, which curves out-
wards in Macrorhinus to below the middle of the head of the femur before going to the external
condyle (PI. IV. fig. 4). In Arctocephalus the ridge is not well marked, but above the line on the
femur for the capsular ligament of the knee in the middle of the shaft, a ridge from the top of the
great trochanter runs down the upper third of the shaft in a line with this point (PI. VII., fig. 7).
This ridge marks off the vastus externus, even although the fibres of the crureus pass in below it
for a distance. In the Phocinae the vastus externus is supplied by the anterior crural nerve.
The Crureus. — In all the specimens there is a layer of muscular fibres without a natural division
covering the front of the femur, which may be the combined vastus internus and crureus, but
certainly is not the vastus internus only. For in Arctocephalus there is a large internal surface on
the femur, and if it were covered by muscular fibres going to the patella, that would afford
sufficient proof that it was the vastus internus. The fibres on the front surface, however, run from
the ventral side of the internal condyle to the corresponding surface of the neck, and do not cover
any part of the extensive internal surface of the shaft, therefore the group of fibres is more entitled
to the name of crureus in Arctocephalus and still more so in the Phocinas where there is no internal
surface.
In the Phocinaj it arises from the anterior surface of the femur, from the intertrochanteric
line. It extends down the shaft to the part which is covered by the capsular ligament of the knee-
joint, and is inserted into the upper edge of the patella beneath the rectus, into the outer upper
half of it (the muscular fibres descend no further), into the inner side of the ligamenturn patellae,
into the head of the tibia on the ventral side, and into the capsule of the knee-joint.
In Macrorhinus Iconinus it is similar, with the exception of its insertion into the inner side of
the patella. The muscular fibres stop at the lower inner edge of it, and are inserted into the inner
side of the head of the tibia by a tendon.
In Arctocephalus gazella it arises from the front surface of the femur, with the exception of the
surface occupied by the vastus externus, and is inserted into the capsule of the knee-joint, and into
the upper edge of the patella. In Otaria and Tricheehus it is combined inseparably with the
vastus externus.
Lucae describes the vastus externus and crureus as a combined muscle, thus recognising the
presence of both ; and I found them in all the dissections, including that of Arctocephalus. Although
the vastus externus is not perfectly separate, the direction of the fibres is an aid to its recognition
as a distinct muscle. In Macrorhinus the separation is further assisted by a tendinous surface upon
the posterior portion of the externus, and here the distinction of it from the crureus was easier. In
Phoca it was not so distinct, and less so in Arctocephalus; but in all it merited a special
description. Dr. Murie describes in Otaria and Tricheehus the internus and externus, and is
doubtful about the crureus. As already explained, the vastus internus would go over the internal
surface if it were present as it is in Arctocephalus; when this surface gives no origin to the
internus, then the crureus must be crowded out by the two lateral muscles encroaching upon its
surface of origin, for in the Phocinae, where there is no internal surface, there is the same collection
of fibres with the same insertion as in Arctocephalus, so I conclude that in Arctoccplmlus there is a
crureus and vastus externus, whereas Otaria and Tricheehus have a vastus externus and internus.
The extensor of the leg, as the name implies, will extend it and flex the thigh upon the pelvis. The
REPORT ON THE SEALS. 185
flexion of the leg only takes place in the Phocinas when on land, but this muscle will be a powerful
depressor of the hinder extremity in the water. In the Phocime it is supplied by a branch of the
deep anterior crural nerve.
The Internal Femoral Region of the Phocina? and Macrorhinus Iconinus contains the pectineus,
obturator externus, adductor longus, and adductor brevis.
In Arctoccphalus gazclla there are in addition the pectineo-superficialis vel femoralis, and adductor
magnus.
The Pectineus in the Phocinse is triangular and of small size. It arises from the ventral surface
of the pubic bar behind the ilio-pectineal eminence ; and slightly from the inner side of the bar. The
psoas tertius crosses it and the obturator is above it. It passes across the head of the femur to the
inner side of the posterior surface of the shaft ; and is inserted into the capsule, and into the upper
third of the inner surface of the back of the femur, extending across the femur one-half its breadth.
In Macrorhinus leoninus it arises from the posterior third of the ilio-pectineal eminence outside
the pelvic brim, from the anterior quarter of the pelvic brim, and slightly from its outer surface,
reaching as far as the capsule of the hip-joint. It passes downwards over the capsule, and is
inserted into the upper third, and the inner half of the hinder surface of the femur. If a line be
drawn from the middle of the inner border of the femur to meet the upper third of one drawn
through the centre of the long axis over the back of the shaft, the attachment is into the triangular
space below the neck inside the lines indicated.
In Arctoccpludus gazclla it is called "adductor brevis primus" by Murie. It arises from one-
sixth of the ventral surface of the pubic bar, this part bemg ventral to the acetabulum and posterior to
the pectineal eminence. It is covered at the origin by the pectineo-superficialis vel femoralis, and is
inserted, after crossing the capsule of the hip-joint, into the back of the femur behind the small
trochanter.
The Pectineo-superficialis vel femoralis is the same as the pectineus in Otaria and
Trichcchus. Upon the surface of the origin of the pectineus there is a band of fibres forming
a distinct muscle, which goes to the lower end of the inner side of the shaft of the femur.
It arises from the same sixth as the latter, but only from the crest of the os pubis, which
is to the inner side of the pectineal origin. It is inserted above the internal condyle of the
femur, at the junction of the front surface with the inner. This muscle may be looked upon
as an accessory psoas, and may be named pectineo-femoralis, for the insertion is similar to the
ilio-femoralis posterior in Macrorhinus. The pectineo-superficiabs is regarded as the pectineus in
Otaria and Trichcchus. It is an adductor. The superficiabs can flex the hip in addition. In
the Phocinae it is supplied by the obturator nerve.
The Adductors are wanting in the Phocinae, but seem to be worked into the fibres of the very
large obturator externus.
The Adductor longus in Macrorhinus Iconinus arises from the posterior outer half of the pubic-
bar, from the outer surface of the ascending ramus of the pubis, with the exception of a margin
near the symphysis, extending to the junction of this ramus with the descending ramus of tin-
ischium, which is in a line with the middle of the acetabulum, and very slightly from the
obturator membrane next the bone. It is inserted into the supracondyloid ridge below the psoas
tertius, and above the epiphysial line on the inner border of the femur.
(ZOOL. CHALL. EXP. PAKT LXVIII. — 1888.) Yyy 2-1
]Q6 THE VOYAGE OF H.M.S. CHALLENGER.
In a large specimen of Ardoccphalus gazella this muscle could probably be divided into two.
It arises from the outer hindward ventral half of the pubic bar, behind the obturator foramen, from
all the surface between the origins of the obturator externus, and the gracilis and adductor niagnus,
and from the hindward quarter of the ischial bar to the origin of the quadratus femoris. It is
inserted obliquely across the back surface of the femur, from the lower end of the great trochanter
to the middle of the inner surface of the shaft of the femur, ending at the junction of the
anterior surface with the inner. The adductor longus primus and secundus of Otaria form the
adductor longus of Ardoccphalus. In Trichcchus the adductor longus has only one head. It
adducts the thigh in Macrorhinus, but in Ardoccphalus besides adducting, the fibres upon the
posterior surface rotate it outwards, those on the inner surface inwards.
From the lower end of the great trochanter in Ardoccphalus, crossing the back of the femur, and
terminating at the middle of the junction of the posterior with the inner surface, is a ridge of bone
resembling the linea aspera of human anatomy and giving attachment throughout its entire length
to the adductor longus. This is a faint ridge in Macrorhinus, but it lies midway between the
upper and lower ends of the great trochanter, and terminates in the middle of the back of the shaft
at the junction of the upper third and lower two-thirds of the femur, the pectineus touching it.
In Ehoca grcenlandica there is a similar ridge to the last, but it ends at the middle of the inner
border of the femur ; the pectineus also lies above and on it. Humphry describes the adductors in
a general way. The name as he uses it is not altogether unsuitable, for the muscles have this action.
His adductor magnus I take to be the semimembranosus. Of his other two adductors, one is the
pectineus and the other probably the ilio-fenioralis.
The Adductor brevis in Macrorhinus leoninus. If the pubic bar be divided into fourths, the
adductor brevis arises from the second fourth behind the pectineal muscle, near the ilio-pectineal
eminence outside the brim of the pelvis, and slightly from the outer surface dorsal to this. It passes
upwards and outwards, and is inserted into the middle of the posterior surface of the femur, outside
the insertion of the pectineus. The exact spot is found by drawing a line from the lower end of the
great trochanter, across the back of the femur, when the middle of this line is the insertion surface.
In Arctocephalus gazella it arises from the pubic bar outside the brim, and from the ventral
half of it behind the origin of the pectineus. It lies between the adductor longus and the pectineal
muscles. It is inserted obliquely across the posterior surface of the femur, from the lower end of
the great trochanter to the insertion of the pectineus behind the small trochanter, and higher
than the longus which is upon the linea aspera. Murie describes this muscle as a primus and
secundus, but I think the primus is the pectineus and the secundus the above-mentioned muscle.
What he gives as the pectineus is inserted into the internal condyle of the femur, and the primus
below the neck and trochanteric fossa, which is the usual insertion of the pectineus. It rotates
outwards and flexes the thigh.
The Adductor magnus is called primus and secundus by Murie and is only found in Ardoccphcdus
gazella. It arises from the outer rim of the innominate bone, dorsal to the symphysis, extending
to the commencement of the dorsal border of the ischial bar. It passes forwards and outwards
from the pelvis to the knee, and is inserted into the lower half of the shaft of the femur at the
junction of the inner with the front surface, across the internal condyle, into the tibia immediately
behind it, and into the capsule of the knee-joint on the inner side. It adducts both the femur and
the tibia. It is a single muscle in Trichcchus.
REPOET ON THE SEALS. 187
The Obturator externus in the Phocinae covers the outer surface of the obturator membrane.
It arises from the outer surface of this membrane, from the outer surface of the pubic bar to half an
inch posterior to the front of the obturator foramen, from the outer surface of the ischial bar,
from the outer surface of the ischial tuber, and from the anterior half of the rami of the ischium
and pubes, posterior to the obturator foramen. The fibres pass upwards and forwards below the
capsule of the' hip-joint in four slips, which are closely attached but easily distinguished. The
ventral or first slip comes from the pubic bar, the dorsal or fourth from the ischial bar, and the
other two from the surface of the large obturator membrane. It is inserted into the obturator pit,
and into the outer half of the posterior or dorsal border of the great trochanter to the external
border of the femur. The slip from the ischial bar may be looked upon as the quadratus femoris,
but it is indistinguishably blended with the obturator externus. This conclusion is based upon
the continuation upon the great trochanter of the insertion of the large obturator.
In Macrorhinus leoninus it is very different from the former three muscles in its origin. It
arises in two parts. The dorsal part (or quadratus femoris) from the posterior half of the ischial
bar to where it turns down, from the outer surface of part of the ischial tuber, slightly from
the obturator membrane next the bar, and from the ischial bar posterior to the obturator
foramen. It is partially blended with the anterior part, and the part along the dorsal border,
forming a strong broad tendon, which gives off a tendinous slip from its ventral side. This slip
joins the adductor brevis, and is inserted along with it. The larger remaining part of the muscle
goes along the under surface of the neck of the femur, and is inserted into the whole of the dorsal
or posterior border of the great trochanter. The anterior part (or obturator externus proprius) arises
from the outer surface of the ischial bar from opposite the middle of the obturator foramen to
behind the acetabulum, from the same extent of the pubic bar, but only from its dorsal half, from
the obturator membrane lying next the bony origins, and from the posterior half of the concave
surface behind the acetabulum. It crosses the joint-capsule, and is inserted by a strong tendon
into the upper half of the posterior surface of the great trochanter. The insertion of this part is
like that of the obturator externus, while the dorsal is similar to the quadratus femoris.
In Arctoeephalus gazella it arises from the entire outer surface of the obturator membrane ;
slightly from the capsule of the hip-joint ; and from the inner half of the pubic and ischial bars
surrounding the foramen. It passes forwards and upwards, and is inserted into the digital fossa
on the back of the great trochanter by a strong tendon. In Otaria it is inserted into the lesser
trochanter. This muscle acts as a powerful rotator of the upper end of the femur. It .rolls the
thigh backwards and inwards to the side of the pelvis. Humphry describes the obturatores as
large, and says the quadratus femoris is not a distinct muscle. I believe it is indistinguishably
blended with the externus in the Phocinae.
In the Phocinte the digital pit is well marked, in Arctoeephalus it is like a groove, and in
Macrorhinus there is none. In the Phocinre the obturator externus covers all the ischial and pubic
bars and the obturator membrane to a little behind the foramen ovale. There is a slight attempt
at division into four slips, the slip over the pubic bar resembling the origin of the adductor longus
in the other Seals, and the ischial origin the quadratus femoris. In Macrorhinus the surface of
bone and membrane corresponding to the surface covered by the fibres in the Phocinaj is shared by
the obturator externus, quadratus -femoris, adductor longus and brevis. In Arctoeephalus the same
divisions exist but are not so simple. The muscle around and over the obturator membrane in
188 THE VOYAGE OF H.M.S. CHALLENGER.
Macrorhinus has divided into three masses, the longus occupying the pubic bar and adjacent
membrane, the quadratus femoris, the ischial and adjacent membrane, and the obturator externus
the front of the obturator foramen and the pubic and ischial bars on either side. The adductor
brevis is isolated and is almost upon the pelvic brim, and there is a large space in the centre of the
obturator foramen with no fibres. In Arctocephalus the obturator externus covers all the membrane
and bone surrounding the foramen. The brevis is on the pubic bar and the longus is behind the
brevis and runs round the posterior aspect of the obturator externus to the ischial bar, while the
quadratus femoris is anterior to its termination. The function of this area of bone posterior
to the acetabulum in the Seals is to give attachment to fibres which will rotate the femur outwards,
adduct, and flex at the hip-joint. In Arctocephalus there are separate muscles for these various
movements, in addition there is an adductor magnus, and each is separately inserted into the femur.
In Macrorhinus there is no magnus, the brevis is insignificant in comparison with the same in Arcto-
cephalus, and on the femur it is receding to the obturator extensor insertion. The longus is confined
to the internal border of the femur, whereas in Arctocephalus it crosses obliquely the back of the
femoral shaft. The quadratus femoris though separate at its origin in Macrorhinus is combined
with the externus at the insertion. In the Phoeinre all the fibres of the externus go to the digital
fossa near it. Though the origins in Macrorhinus are nearly like those in Arctocephalus the insertions
are not, but slightly resemble those of the Phocinre as regards the quadratus and obturator externus,
and those of Arctocephalus as regards the brevis, and are like neither in the longus. The movements
of the thigh in the Phocinas are the most imperfect, and this combined mass is sufficient for them.
In Macrorhinus a higher stage is reached as indicated by the separation into muscle bundles, and
in Arctocephalus there is sufficient differentiation of the muscular mass to enable the animal to walk
as well as swim.
The Gluteal Region of the Phocinae and Macrorhinus contains the gluteus maximus, medius,
minimus, pyriformis, obturator internus, and genielli.
In Arctocephalus in addition to these there is the quadratus femoris.
The Gluteus maximus in Phoca vitulina is the most superficial muscle of the gluteal region, and
is triangular in form. The base rests upon the vertebral spines and the apex upon the femur.
The dorsal head arises from the crest of the ilium between the two lips, extending from the
ventral anterior spine to the dorsal posterior spine; between the dorsal posterior spine and the
spine of the last lumbar vertebra it takes origin from the fascia covering the erector spina1, ami
also from the spines of the last lumbar, all the sacral, and the 1st caudal vertebra, by the fascia
which is an extension backwards of the lumbar aponeurosis, from the tendinous expansion over the
back of the sacrum, and from the dorsal sacro-iliac ligament. The ventral head is a narrow
riband-shaped fasciculus about an inch broad. It arises beneath the great division opposite the
level of the 3rd sacral vertebra, from the side of the dorsal sacro-iliac ligament. The fibres of
the dorsal division pass from their origin to the great trochanter and the external border of the
femur. Those coming from the crest of the iliiun go backwards to the lower and outer part of the
anterior border of the great trochanter. The portion lying between the dorsal posterior spine and
the three sacral vertebrae passes almost horizontally outwards, and the remainder between the
3rd sacral and the 2nd caudal go forwards and outwards to join the femur. The dorsal part is
inserted into the outer third of the anterior border of the great trochanter, goes obliquely
REPORT ON THE SEALS. 189
across the great trochanter to the middle of its posterior border, and then passes out
alone; the outer half of its posterior border, down the outer border of the shaft of the
femur, to the lower end of the external supracondyloid ridge. The ventral part is inserted
into the under surface of the dorsal part near the outer border of the femur, a few fibres
"•amino- the femur. In Phoca hi&pida the gluteal muscle was in a very bad condition. It
arises from the 4th sacral and 1st to 4th caudal vertebras, and the insertion is as in Phoca
barbata. In Phoca barbata it is smaller than in Phoca vitulina, and has three heads. The
anterior head arises from the aponeurosis over the erector spinas, by a band of muscular fibres
springing midway between the dorsal posterior spine of the ilium and the 1st sacral vertebra, and
from the dorsal sacro-iliac ligament. The second or posterior head arises from the posterior con-
tinuation of the same aponeurosis, which is attached to the spine of the 4th sacral vertebra, and
the 1st, 2nd, and 3rd caudal vertebras. The third or ventral head consists of a series of fibres
springing from the side of the dorsal sacro-iliac ligament, opposite the level of the 4th sacral and
1st caudal vertebras, beneath the posterior part. There is a space between the first and second heads
and the erector spinas, and the dorsal sacro-iliac ligament is uncovered by the gluteal muscles over
the sacrum. The fibres from the first head pass back and out, those of the posterior part pass out
and forwards over the great trochanter, and form one muscle by the anterior head joining the
posterior. The three heads are inserted into the femur by the ventral or third head joining the
under surface of the second, and sending a few fibres directly to the femur ; thus two heads are
left, the anterior and posterior, which are disposed like the dorsal head in Phoca vitulina.
In Macrorhinus leoninus there are three parts. The anterior part arises from the dorsal surface
of the dorsal sacro-iliac ligament, and from the inferior lip of the crest of the ilium, and after
joining the ventral part is inserted with it into the posterior half of the great trochanter. The
ventral part lies beneath the posterior part, and arises from the ventral surface of the dorsal sacro-
iliac ligament, and from the anterior surface of the 2nd and 3rd sacral vertebras. It goes to the
femur, and is joined, near the great trochanter on the anterior border, by the anterior part. The
posterior part arises from the fascia over the 1st to the 3rd sacral spines and the 1st caudal, and
from the dorsal surface of the dorsal sacro-iliac ligament. It passes transversely outwards to the
femur, and is inserted into the outer border of the femur below the great trochanter and into the
external condyle.
In Arctoccphcdus gazclla there are two heads. The anterior arises from the fascia attached to
the spines of the 1st, 2nd, and 3rd sacral vertebras, goes outwards to the femur, and is inserted
into the lower three-fourths of the posterior border of the great trochanter. The posterior arises
from the fascia attached to the spines of the 1st, 2nd, and 3rd caudal vertebras, and is partly
overlapped by the anterior head. The fibres pass transversely outwards, and are inserted into the
external border of the femur, into the capsule of the knee-joint, and into the head of the fibula.
Humphry and Lucae do not refer to the ventral part. In Otaria it has two parts, but there is
only one in Trichcchus. In the Phocinas and Macrorhinus the iliac part rotates the femur inwards,
tilts the lower end outwards, while the posterior part will rotate the femur outwards and flex the
thigh. In Arctoccphalus there are no iliac fibres, and consequently no rotation inwards and
forwards.
The Gluteus medius in the Phocinas is situated below the maximus, and arises from the lower
lip of the crest of the ilium, from the external surface of the ilium immediately below the lower
190 THE VOYAGE OF H.M.S. CHALLENGER.
lip, from the sides of the laminae of the 1st, 2nd, and 3rd sacral vertebrae below the dorsal sacro-
iliac ligament, and from the ventral surface of the ligament below the 1st and 3rd sacral vertebra?.
The anterior fibres pass backwards and outwards, the middle outwards, and the posterior forwards
and upwards, just like those of the gluteus maximus, but on a smaller scale. It is inserted into the
great trochanter of the femur. The anterior fibres are fixed to the middle of the front border of
the great trochanter; from here the fibres are attached obhquely across the trochanter to the
junction of the upper surface with the posterior border, where they fix themselves to the upper half
of the posterior border of the great trochanter.
In Macrorhinus leoninus it arises from below the inferior lip of the ilium, and behind the ridge
which extends from the ventral anterior spine to the middle of the acetabulum. It is inserted into
the outer surface of the great trochanter above the tubercle on the superior side of the posterior
border and the middle of the anterior border.
In A?xtocephalus gazella it arises from the inferior lip of the crest of the ilium, and from the
lumbar aponeurosis. The fibres pass backwards, and are inserted into the great trochanter from
midway between the anterior border to the posterior inferior corner of it. In the Phocinae and
Macrorhinus it tilts the lower end of the femur outwards, rotates the trochanter inwards, and pulls
the femur forwards. In Arctocephalus it draws the head of the bone inwards and forwards.
The Gluteus minimus in Phoca vitidina and Plwca hispida is beneath the medius, and arises from
the outer surface of the ilium behind the ridges passing from the ventral anterior spine to the
middle of the front of the acetabulum; this surface of origin is concave. It forms a narrow
muscular rectangle which can be divided into three slips. It is inserted into the upper inner half
of the front border of the great trochanter, and slightly into the surface of the trochanter adjoining.
In Phoca barbata the origin is similar, but the insertion is into the outer half as well as into the
inner half of the front border of the great trochanter.
In Macrorhinus leoninus it arises from the concave surface of the ilium behind the medius and
dorsal to the ridge. It is inserted into the anterior border of the great trochanter in its upper half.
In Arctocephalus gazella it arises from the external surface of the ilium dorsad to the feebly
marked ridge (already mentioned) and from the venter of the dorsal sacro-iliac ligament ; and is
inserted into the upper half of the front border of the great trochanter above the insertion of the
gluteus medius. In all the action is to rotate the femur inwards and forwards.
The Piriformis in the Phocinae cannot be recognised apart from the gluteus medius until the
dorsal sacro-iliac ligament is cut and turned aside. It arises from the ventral surface of the dorsal
sacro-iliac ligament posterior to the 1st sacral foramen, and from the sides of the ventral surfaces
of the 1st, 2nd, and 3rd sacral vertebrae. The fibres converge and are inserted, into the upper third
of the back of the great trochanter of the femur.
In Arctoccphcdus it closely resembles the same part in Phoca, but arises from the 2nd, 3rd, and
4th sacral vertebra;.
The Gemellus superior in Phoca vitidina and in Arctocephalus lies anterior to the tendon of the
obturator internus. It arises from the dorsal surface of the ischial liar posterior to the acetabulum.
The Gemellus inferior in Phoca vitulina and in Arctocephalus lies posterior to the tendon of the
obturator internus. It arises from the internal surface of the ischium below the tuber extending
to the origin of the obturator internus interiorly, and the obturator internus tendon anteriorly. For
the insertions of the gemelli in Phoca, see the obturator internus.
REPORT ON THE SEALS. 191
In Arctocephalus they are inserted into the great trochanter to the outer side of the obturator
interims.
The Obturator internus in Phoca vitulina arises from the internal surface of the obturator mem-
brane, and from the rim of bone around it, and forms a tendon which goes over the dorsal surface
of the ischial bar in its groove. The two gemelli meet over, surround, and conceal the tendon of
the obturator internus ; and all three are inserted together into the posterior border of the great
trochanter.
In Arctocepilialus, after scraping away the gemelli and isolating the tendon, the muscle is found
to arise from the obturator membrane, &c, as in Phoca ; and is inserted to the inner side of the
insertion of the gemelli into the posterior upper end of the great trochanter of the femur.
The Quadratics femoris is only found in Arctocephalus. It is triangular, and arises from the
dorsal half of the ischial bar, posterior to the gemellus inferior and anterior to the origin of the
semimembranosus. It passes forwards, outwards, and downwards, and is inserted by a tendon into
the lower half of the posterior border of the great trochanter.
These three rotate the femur outwards, bringing the thigh near the pelvis.
In Macrorhinus the dorsal part of the obturator externus represents the quadratus. In the
Phocinaj it is fused with the obturator externus and unrecognisable as the quadratus.
The Muscles fkom the Pelvis to the Leg. — In the Phocinne, Macrorhinus leoninus, and
Arctocephalus gazella the gracilis, semimembranosus (in Phoca vitulina it has an anterior and
posterior part), semitendinosus (with two heads), and biceps (which has a long head, the biceps,
and a short head named the sacro-peroneus) are present.
The Gracilis, also called symphysis tibialis in Lucae's plate, is a flat triangular muscle in the
Phocinre, stretching from the symphysis to the tibia. It arises from the symphysis pubis, and
radiates outwards to the lower leg, the superficial fibres only arising not from the bone but from
the linea alba. The posterior third of the latter is continuous with the fasciculi of the opposite side
over the symphysis. The anterior two-thirds is anterior to the pubic arch, the muscle of the right
side is beneath and overlapped by that of the left. It is inserted into the posterior two-thirds of
the ventral surface of the tibia, and many of the fibres end in a tendon near the shaft. The tendon of
insertion is combined with that of the semimembranosus and semitendinosus. In the substance of
the gracilis, near the ventral border of the tibia and parallel with it, is a long narrow tendon running
at right angles to its fibres. This is an indistinct white streak close to its anterior border, which
gradually widens and strengthens to a strong broadish tendon at its posterior border; at the bend
of the anterior surface of the astragalus it expands, forming, with the prolongations backwards of
the combined tendons of insertion of the semimembranosus and seniitendinosus, and gracilis, the
plantar fascia.
In Macrorhinus leoninus it arises from the symphysis pubis by the fibres of its deep surface,
from the linea alba in front of the pubis by the intermediate fibres, and by the superficial fibres
from the ligament stretching between the pubic bones. The fibres opposite the pubes are blended
with those of the opposite side. It is inserted into the posterior half of the tibia.
In Arctocephalus gazella it arises from the symphysis pubis, and from the ligament between the
pubic bones. The superficial fibres are continuous with those of the opposite side, and none of the
fibres reach further forward than the symphysis. It is inserted into the middle third of the ventral
192 THE VOYAGE OF H.M.S. CHALLENGER.
surface of the tibia, extending to the external border. It draws the leg inwards, and in Arctoccphcdus
will turn the leg inwards when progressing on land. In Otaria a few fibres overlap the external
oblique, but in Trichcchus it has no fibres covering this muscle, and so is similar to Arctocephalus.
In the Phocinae it is supplied by the obturator nerve.
The Semimembranosus is named Muse, pubo-tibialis by Lucae ; in PJioca vitulina it lies above the
gracilis and is partially hidden by it. It is in two parts, the anterior and posterior. The anterior
arises from the outer surface of the innominate bone posterior to the foramen, in front of the origin
of the posterior part, and above the origin of the gracilis. It is inserted into the posterior two-
thirds of the front of the tibia, its tendon combining with the gracilis. The posterior part arises
from the outer edge of the innominate, between the tuber ischii, from which it also has fibres of
origin, and the origin of the anterior part from the pubic bone. It is inserted into the tibia,
anterior to the seniiinembranosus (anterior part), and extends forwards to one-fifth from the upper
extremity of the tibia. In Phoca hispida the anterior part arises from the outer surface of the
innominate bone, from the body of the pubis upwards to where the pubis and ischium fuse, and
from the edge of the bone between the origin of the gracilis to the semimembranosus (posterior
part). It is inserted into the posterior half of the ventral surface of the tibia, in conjunction with
the semitendinosus and hinder three-fourths of the gracilis. The posterior part is placed above
the gracilis. It arises from the outer dorsal half of the pubic bone, between the tuber ischii and
the origin of the semimembranosus anterior part, slightly from the base of the tuber posteriorly, and
from the edge of the bone between these two points. It goes outwards and forwards, and is
inserted into the ventral surface of the tibia in its upper half, the fibres almost reaching the head
of the shaft. In Phoca barbata both parts are similar to the last, except that the origin does not
go up to the tuber ischii, and the insertion is smaller, not reaching so far forwards on the tibia,
but falling short of the head by a quarter of the length of this bone.
In Macrorhinus leoninus it is almost the same as is the posterior part in Phocinse. It arises
from the outer surface of the innominate bone, posterior to the origin of the adductor longus and
that part of the obturator externus which represents the quadratus femoris, extending backwards a
little more than halfway between the symphysis and the ischial tuber. It is inserted into the
anterior half of the ventral surface of the tibia, reaching almost to the head of the bone.
In Arctocephalus yazella it is closely allied to the posterior part in Phoca vitidina, Phoca hispida,
and Phoca barbata. It arises from the posterior third of the outer edge of the pubic bar, which is
behind the rudimentary tuber ischii, from the third of the posterior border of the innominate bone,
which is the continuation of the sitting bone downwards, and slightly from the surface of the
innominate adjoining the marginal origin. It is inserted into the front of the tibia in front of
the semitendinosus, the fibres terminating a quarter of the length of the tibia from the head of the
shaft, i.e., it is inserted into the second fourth of the tibia from the head. Lucae describes it as one
part in Phoca. In Otaria, what Murie names semitendinosus I call semimembranosus ; this also
applies to Trichcchus.
In all the specimens the posterior part is present ; in Macrorhinus and Arctoccphcdus the
anterior part is wanting. The insertion of the posterior part in all is nearer the tibial head, the
insertion in Macrorhinus almost touching it. The portion of bone giving origin to the anterior part
in the Phocinaa is utilised in Arctoccphcdus for the adductor magnus, and in the latter it has
wandered up the limb and divides its attachment between the femur and tibia. This is a case of
REPORT ON THE SEALS. 193
the migration of a muscle, the anterior part of the semimembranosus in the Phocinas having become
an adductor muscle in Arctoccphalus by changing to a more anterior po.sition in the hind limb.
The former adducts the thigh and leg and rotates the limb, and the latter adducts the leg and also
rotates it. In the Phocinas it is supplied by the obturator nerve.
The Scmitendinosus, which is named Muse, coccygo-tibialis by Lucae and the semimembranosus
by Murie, has two heads of origin. The dorsal head arises from the posterior half of the spine of
the last sacral vertebra, from the spine of the 1st caudal vertebra, and is continuous anteriorly with
the gluteus maximus. The ventral head arises from the side and ventral surface of the dorsal
sacro-iliac ligament, and from the transverse processes and bodies of the 4th and 5th sacral, and the
1st and 2nd caudal vertebra. The dorsal head is inserted upon the dorsal surface of the ventral
head, half an inch from the caudal vertebrae. The ventral, thus strengthened, passes to the posterior
two-thirds of the front of the tibia, into which it is inserted. In Phoca hispida the dorsal part was
not seen, and the ventral part was also in a bad state of preservation. It appeared to arise from
the 2nd, 3rd, 4th, and 5th caudal vertebrae, and to be inserted into the posterior two-thirds of the
front of the tibia above the semimembranosus. In Phoca barbata the dorsal head, or gluteal slip,
which is continuous with the hindmost fibres of the gluteus maximus, arises from the 2nd and 3rd
caudal vertebras ; the ventral head arises from the 2nd, 3rd, 4th, and 5th caudal vertebras. The
insertion is the same as in Phoca hispida.
In Macrorhinus leoninus there are two heads of origin. The dorsal head arises from the spine
of the 2nd caudal vertebra, under cover of the gluteus maximus, and from the dorsal sacro-iliac
ligament, as in the others. The fibres are obliquely directed backwards and outwards. It is
inserted into the posterior third of the second head near the commencement of the tendon, which is
midway between the origin and insertion. The ventral head arises from the sides of the 1st, 2nd.
3rd, and 4th caudal vertebrae near the anterior surface of their bodies. Midway between its origin
and insertion it forms a very strong broad tendon, which is inserted into the posterior half of the
ventral surface of the tibia ; the outer half covers the gracilis.
In Arctoccphalus gazclla it arises from the transverse processes of the 2nd, 3rd, 4th, 5th, and 6th
caudal vertebras, and from the sides of the bodies of these vertebras between the transverse processes
and their tubercles. It is inserted into the lower half of the ventral surface of the tibia. In Otaria
and Trichcchus it is inserted into the posterior half of the tibia like Arctoccphalus. The action is the
same as in the last muscle. Humphry, Lucae, and Murie do not refer to the double mode of origin.
In the Phocinas it is supplied by the obturator nerve.
The Biceps, or ischio-tibialis of Lucae, consists of two distinct muscles, and these are named the
Biceps or long head, and the Sacro-peroneus or short head (sacro-fibularis of Lucae). In Phoca vitulina
the long head is fan-shaped, and arises from the ischial tuberosity, and from the dorsal and outer
surface of it by a pointed fasciculus. It spreads out or radiates towards the fibula. Over the
peronei it extends from the head of the fibula almost to the malleoli. Here it is bound to the deep
fascia covering the peroneal muscles, and terminates by joining the strong fascia over the outer
muscles of the leg, which fascia is bound to the outer and ventral surface of the tibia. The portion
at the lower end of the fibula joins the sacro-peroneus and is inserted with it. The short head or
sacro-peroneus is riband-like, and arises from the under surface of the dorsal sacro-iliac ligament,
from the sides of the 2d and 3rd sacral vertebras. It goes obliquely backward and outward to the
lower outer third of the fibula. It is inserted, after joining the former, into the posterior quarter
(zool. chall. exp. — paet i.xviii. — 1888.) Yyy 25
194 THE VOYAGE OF H.M.S. CHALLENGER.
of the dorsal border of the fibula along with the tendon of the long head. In Phoca hispida the
long head is the same as in Phoca vitulina, and the short head or sacro-peroneus arises from the
4th sacral and 1st caudal vertebrae. In Phoca barbata the biceps (long head) is the same as in
Phoca vitulina, and the sacro-peroneus arises from the 2nd and 3rd sacral vertebrae.
In Macrorhinus leoninus the long head of the biceps arises from the dorsal sacro-iliac ligament
opposite the 1st caudal vertebra by a small slip, which blends posteriorly with the ventral part of
the semitendinosus, and joins the origin from the tuber ischii 1 inch behind it ; otherwise it is the
same as in Phoca vitulina. The sacro-peroneus or short head arises from the ventral and lateral
surfaces of the 2nd and 3rd sacral and 1st caudal vertebra;, and from the ventral and lateral
surfaces of the dorsal sacro-iliac ligament. It is inserted as in Phoca vitulina.
In Arctocephcdus gazella the long head of the biceps consists of three parts ; all three arise from
the sides of the sacral vertebras. The fibres are transverse and go to the outer anterior surface of
the tibia. The anterior part is slightly overlapped by the middle, but the fibres of the middle and
posterior parts touch each other. Over the back of the fibula these three form a tendon which
turns round the limb to the ventral border. This tendon forms also the deep fascia over the muscles
of the leg, and is attached to the tibia, but it does not appear to go to the fibula. The sacro-
peroneus or short head arises from the anterior surface of the 4th sacral and 1st caudal vertebrae,
and is inserted by a small tendon into the dorsal border of the fibula, over the dorsal malleolus. In
all the two heads bend the knee, roll the legs outwards, and adduct them. In Otaria and Trichechus
the long head is in two parts. In the Phocinae both the long and the short heads are supplied by
the small sciatic.
The Leg. — The Outer Tibiofibular Eegion in all the specimens has a tibialis anticus,
extensor proprius hallucis, and extensor longus digitorum.
The Tibialis anticus in the Phocinae and Macrorhinus is an elongated triangle with the base
at the knee-joint. It is partly under cover of the extensor communis digitorum, and arises from
the outer surface of the tibia in its anterior two-thirds, with the exception of a small triangular
surface at the upper dorsal part of the head of the shaft, from the ligamentiun patellar, from
almost the whole of the anterior two-thirds of the interosseous membrane, and by a small
fasciculus from the outer surface of the fibula posterior to the fusion of the bones. Almost at the
posterior third of the tibia it forms a strong tendon, which goes through the groove on the outer
side of the posterior extremity of the tibia, beneath the annular ligament, and divides into two
tendons of equal size. It is inserted into the proximal end of the metatarsal bone of the hallux
on its tibial and outer surface, and into the ventral tibial surface of the internal cuneiform.
In Arctoccphalus gazella it arises from the head and from the outer surface of the tibia in
its anterior four-fifths. Near the annular ligament it forms a tendon, which passes beneath it,
ventral to the extensor proprius hallucis, and crosses the tarsus, then expands and is inserted into
the proximal tibial surface of the 1st metatarsal on its outer side. In Otaria and Triclicchus, besides
the insertion, there is also as in Arctocepludus a tendon to the entocuneiform bone. In the Phocinae
and Macrorhinus it flexes the ankle, depresses the pes, and turns it outwards. In Arctoccphalus,
besides having these actions, it will in walking raise the foot on to the outer edge. Lucae gives only
one tendon of insertion, and that to the metatarsal. In the Phocinae it is supplied by the musculo-
cutaneous nerve (dorsal division).
EEPOET ON THE SEALS. 195
The Extensor proprius hallucis in the Phocinae and Macrorhinus lies along the dorsal side of the
extensor communis digitorum, under cover of the peroneus longus. It arises from the whole of the
ventral surface of the fibula, from a very slight margin of the outer surface of the interosseous mem-
brane next the shaft, and from the ventral border of the shaft from the termination of the outer
border to the junction of the middle and posterior thirds. It crosses from the dorsal side of the
extensor communis to its ventral side, runs beneath the annular ligament between the tibialis anticus
and the extensor communis digitorum, and goes over the tendon of the tibialis anticus to the
ventral surface of the tarsus, then it ascends gradually to the outer surface of the distal end of the
1st metatarsal. It is inserted into the distal tibial outer surface of the same, into the proximal end
of the 1st phalanx, and into the capsule of the joint between. In the PhoeinEe it is supplied by
the musculo-cutaneous nerve (ventral division).
In Arctocephalus gazella it is almost as large as the tibialis anticus in the same animal. It arises
from the anterior two-thirds of the ventral surface of the fibula, from the anterior half of the dorsal
border of the tibia, from a small triangular surface of the tibia posterior to its head, and between
its short outer border and its dorsal border, from the fusion of the tibio-fibular articulation beneath
the origin of the extensor longus digitorum, and from the interosseous membrane. It courses back-
wards between the tibialis anticus on its ventral side and the extensor longus digitorum on its
dorsal ; beneath the annular ligament it forms a tendon, which passes over the tibio-fibular joint
and the tarsus, and runs along the dorsal side of the 1st metatarsal. It is inserted, after the
expansion of its tendon, into the proximal end of the outer surface of the 1st phalanx. In Otaria
it arises from the fibula and interosseous membrane, and to the proximal end of the proximal
phalanx of the hallux. In the PhocinEe and Macrorhinus it extends the digit, and then flexes the
ankle, and depresses and abducts the pes. In Arctoceplwlus it only extends and flexes.
The Extensor communis or longus digitorum, named by Lucae the extensor quatuor digitorum, in
the Phocina? and Macrorhinus is an elongated triangle situated between the tibialis anticus and the
peroneus longus. The latter partially overlaps it on the dorsal side, and it partially overlaps the
tibiabs anticus, and crosses the extensor hallucis. It arises from the triangular surface of the tibia
posterior to the superior tuberosity, marked off inferiorly by a faint ridge, i.e., the short outer border
which extends from the middle of the outer surface of the superior tuberosity backwards and
upwards to the dorsal border of the tibia, from the tibia anterior to the fusion of the bones of the
leg, from the capsule of the joint, and from the outer surface of the tibia and fibula where they
fuse posterior to the origin of the peroneus longus. It forms a strong tendon, which passes back-
wards, crosses over the extensor hallucis, and goes beneath the annular ligament dorsal to the
extensor hallucis at the posterior tibio-fibular articulation. Having traversed this, it crosses the
ankle-joint and enters the groove on the middle of the outer side of the tarsus, which terminates
over the proximal end of the 3rd metatarsal bone. Here it expands and divides into four tendons.
The first or ventral passes obhquely over the ventral distal end of the 3rd metatarsal and expands
upon the upper proximal end of the 1st phalanx of the 2nd digit. The second tendon passes back
over the middle of the outer surface of the 3rd metatarsal, and expands upon it. The third crosses
obbquely over the middle of the dorsal side of the 3rd metatarsal, goes backwards upon the ventral
side of the 4th metatarsal, and expands upon the proximal end of the inferior side of the 1st
phalanx. The fourth crosses obliquely backwards and upwards from the proximal dorsal end of the
3rd metatarsal, over the middle of the outer surface of the 4th metatarsal, and reaches the ventral
196 THE VOYAGE OF H.M.S. CHALLENGEE.
distal end of the 5th metatarsal. These four tendons go backwards over the metatarso-phalangeal
joints, expand upon the outer surfaces of the 1st phalanges of the four digits, and are inserted into
the distal ends of the 1st phalanges of these digits, into the outer surfaces of the ligaments between
the phalanges, and into the proximal ends of the 2nd phalanges.
In ArctocepJialus gazella it is a long narrow muscle, and arises from the external condyle of the
femur, by a small fasciculus posterior to the insertion of the gluteus maximus, which passes back-
wards over the capsule of the knee-joint, gaining fibres from it, from the outer sides of the
head of the tibia and fibula adjoining the fused tibio-fibular articulation, also from the anterior
half of the outer border of the fibula by a fine but strong aponeurosis ventral to and touching the
tendon of origin of the peroneus brevis from the same border. Anterior to the malleolus it is a
strong tendon, which passes beneath the annular ligament, traverses the shallow groove on the
outer surface of the fibula, and the groove on the astragalus, lies on the outer surface of the tarsus
between the cuboid and cuneiform bones, and expands and divides into four slips over the bones of
the 3rd and 4th metatarsals. The first or ventral runs along the dorsal side of the 2nd metatarsal ;
the second along the middle of the 3rd metatarsal, the third along the ventral side of the 4th meta-
tarsal. The fourth crosses the outer surface of the middle of the 4th metatarsal, and runs down the
ventral side of the 5th metatarsal. At the metatarso-phalangeal articulation, the tendons begin
gradually to widen ; over the distal ends of the proximal phalanges they completely cover their
outer surfaces; and after passing over the joints and sharing in the formation of the posterior
ligament they are inserted into the proximal ends of the 2nd phalanges of the 2nd, 3rd, 4th, and
5th digits ; from their attachments, fine aponeurotic sheets are prolonged onwards, and end
imnoticeably over the phalanges. In the Phochife I did not find any origin from the femur, as
described by Lucae. In Otaria and Trichcchus there is no tibial origin. In the Phocinse, Macror-
hinus, and ArctocepJialus it extends the digits, and flexes the ankle. In the Phoenue it is supplied
by the musculo-cutaneous nerve (ventral division).
The Fibular Eegion in all the specimens has the same muscles. The peronei longus, quinti
digiti, and brevis.
The Peroneus longus, the peroneus primus of Lucae, in the Phocinre and Macrorhinus is a longi-
tudinal band of fibres. It arises from the small pit on the external condyle of the femur above
the depression for the origin of the popliteus, from which it is separated by the intervention of the
capsule of the knee-joint, which is attached to the femur between these two origins, and above it is
the termination of the insertion of the gluteus maximus; also from the external surface of the capsule
of the knee, from the outer surface of the fused tibio-fibular articulation, slightly from the adjacent
surfaces of both bones and from the fascia above the muscle. It courses backwards over the
tibio-fibular ankylosis, and lies in the hollow between the bones of the leg. About the junction
of the middle with the posterior third of the fibula it ends in a strong tendon, which leaves the
interosseous space and crosses obliquely backwards and upwards to gain the inferior groove on
the outer surface of the fibula, then it follows the bed of this groove, which runs to the dorsal
malleolus beneath the annular ligament. It passes over the tendons of the peronei brevis and
quinti digiti which lie on the calcaneo-astragaloid articulation, and, entering the dorsal groove on
the posterior outer corner of the os calcis, runs obliquely backwards, inwards, and downwards in the
groove upon the dorsi-plantar surface of the cuboid bone. It goes beneath the ligament stretching
REPORT ON THE SEALS. 197
from the plantar surface before the peroneal groove of the cuboid to the base of the metatarsal
of the 5th digit, and finally enters the groove on the plantar surface of the cuneiform bones. It is
inserted into the base of the proximal extremity, on the plantar and dorsal side of the metatarsal of
the hallux, the ligaments of the foot forming a channel for it by bridging over the grooves. The
high origin of this muscle from the femur will give support to the knee, and make up for the
absence of the external lateral ligament.
In Arctocephalus gazella it is situated on the outer side of the leg, and arises from the external con-
dyle of the femur, out of the same tendon which gives origin to the popliteus and plantaris muscles ;
and from the tibia and fibula at the tibio-fibular ankylosis, as in Phoca. It courses backwards
between the extensor communis digitorum and the dorsal malleolus, turns outwards over the inner
border of the fibula and gains the dorsal surface, enters the outer groove of this surface, runs over the
tendons of the peronei brevis and quinti digiti, traverses the inner groove on the dorsal surface of
the os calcis, and turns down into the groove of the cuboid bone, descending obliquely forwards over
the cuneiform bones to the proximal extremities of the metatarsals. It is inserted into the dorsal
proximal extremity of the 1st metatarsal bone. In Otaria and Trichechus it has origin from the
femur, and is inserted into the head of the 1st metatarsal, and joins the fascia to that of the 4th.
In the Phocinse it extends the ankle and turns the dorsal border of the foot outwards ; in
Arctoc&phalus only it will raise the heel in walking. Humphry and Lucae describe no tibial fibres.
In the Phocinae it is supplied by the musculo-cutaneous nerve.
The Peroneus quinti digiti in the Phocixue and Macrorhinus is a flat band-like muscle which
arises from the outer surface of the peroneus brevis, upon which it is planted, and from the outer
surface of the head of the fibula, dorsal to the peroneus longus. It passes backwards to the
posterior end of the fibula, and is closely adherent to the peroneus brevis, from which it is with
difficulty separated. About the lower third of the outer side of the fibula it forms a small tendon,
which passes through the annular ligament behind the malleolus in front of the tendon of the
peroneus brevis. It goes in this order over the inferior groove of the os calcis, and is inserted into
the outer and dorsal surfaces of the distal end of the 5th metatarsal, and the proximal end of the
1st phalanx of the 5th digit.
In Arctocephalus gazella it lies on the peroneus brevis, and arises from the head of the fibula
below the soleus, and from the anterior quarter of the dorsal surface of the fibula. It is adherent to
the peroneus brevis, passes backwards in the inner groove on the dorsal surface of the fibula upon
the tendon of the brevis, then it enters the groove of the os calcis which is on its outer surface,
and proceeds backwards upon the dorsal side of the 5th metatarsal bone to be inserted into the
proximal dorsal extremity of the 1st phalanx of the 5th digit, expanding before reaching it. In
the Phocinse it is supplied by the external popliteal nerve.
The Peroneus brevis, the peroneus secundus of Lucae, in the Phocinag and Macrorhinus
leoninus is the largest of the group, and arises from the outer surface of the head of the
fibula ; and from the anterior three-quarters of the outer surface of this bone, the fibres arising
from the anterior half being dorsal to the outer border, and the remaining fourth of the muscle
dorsal to the ventral border. Near the malleolus it forms a strong tendon, which goes with the
peroneus quinti digiti but to its upper side through the annular ligament, and in this order enters
the inferior groove on the os calcis, and is inserted into the dorsal surface and distal end of the 5th
metatarsal.
198 THE VOYAGE OF H.M.S. CHALLENGER.
In Arctocephalus gazella it is beneath the peroneus quinti digiti, and arises from the dorsum of
the head of the fibida below the peroneus longus, from the whole extent of its outer border, and
from the posterior three-fourths of its dorsal surface. It has the same course as the peroneus
quinti digiti, and is inserted into the dorsal proximal surface of the 5th metatarsal bone, but
before gaining the bone it broadens considerably. In Otaria it is very much the same as in Arcto-
cephalus. In Trichechus the tendon does not expand so much as in Otaria, In the Phoeinae it is
supplied by the external popliteal nerve.
In all the specimens the peroneus tertius and the peroneus quartus are absent.
In Trichechus are found the peroneus tertius and the peroneus quart i digiti, which latter is
diminutive. The peroneus longus is an extensor of the ankle, the peroneus brevis and the peroneus
quinti digiti are flexors and abductors of the foot, and the brevis and quinti digiti expand the toes.
The Inner Tibiofibular Eegion consists of a superficial and a deep group of muscles.
The Superficial Group in Phoca vitulina, Phoca hispida, Phoca barbata, and Macrorhinus
leonins is formed by the gastrocnemius and plantaris. In Arctocephalus gazdla, besides the other
muscles, there is the soleus.
The Gastrocnemius in the Phocinae is a two-headed muscle, and the inner head is more than
double the size of the outer. The inner head arises from the back of the femur above the internal
condyle, reaching up the shaft to the junction of the internal border with the supracondyloid ridge,
from the internal surface of the same condyle above the fossa for the internal lateral ligament, from
the internal lateral hgament extending to the junction of the anterior third with the posterior two-
thirds of the tibia, from the anterior third of the ventral border of the shaft ventral to the lateral
hgament, and from the capsular ligament of the knee-joint. The outer head arises from the outer
surface of the external condyle in common with the plantaris muscle, slightly from the outer half
of the surface of the femur above the same condyle, and by a few fibres from the back of the head
of the fibula. The two heads unite opposite the junction of the middle two-thirds with the
posterior third of the tibia, and form a tendon which widens near the os calcis, and is inserted into
the anterior aspect of the tuberosity of the os calcis.
In Macrorhinus leoninus the inner head arises as in Phoca vitulina, but covers more of the back
of the femur, also from the front surface of the internal condyle up to the patellar facet of the
femur. The outer head does not arise from the femur, but from the inner dorsal surface of the
head of the fibula. The fibres of the inner head join those of the outer head at the anterior third
of the tibia, and form a strong tendon, which is inserted as in Phoca vitulina.
In Arctocephahis gazella it is a single-headed muscle, and arises from the inner surface of the
internal condyle of the femur below the fossa for the internal lateral hgament, from the internal
lateral hgament, from the internal border of the tibia in its upper third, and from the capsule of
the knee-joint. It crosses the leg from the dorsal to the ventral side; one inch from the os calcis
it forms a tendon, which widens and is inserted into the os calcis to the outer side of the groove for
the plantaris tendon.
Humphry and Lucae give no connection with the fibula ; the bony attachments are the same in
Otaria and Trichechus. In the Phocinae and Macrorhinus it will powerfully extend the foot when
swimming ; in Arctoccplialus it also extends the foot in the water, and raises the heel in walking.
In the Phocinaa it is supplied by the great sciatic nerve.
REPORT ON THE SEALS.
199
The Solcus is found in Arctoccphalus gazclla, but not in the Phocinae and Macrorhinus. It is a
flattened elongated muscle, lying on the peronei brevis and quinti digit! Near the head of the
fibula it is a fine sheet, at the middle triangular, the apex being the origin, the base the free edge ;
over the posterior fifth it is a fleshy bundle. It arises from the dorsal surface of the head of the
fibula by a thin tendon, from the whole length of the inner border of the shaft by a fine aponeurosis,
and by muscular fibres from the inner surface of its posterior fifth, ve.itral to this border and dorsal
to the interosseous membrane. The fibres pass backwards, and are inserted into the proximal
surface of the tuberosity of the os calcis beneath the attachment of the gastrocnemius extending
further back on the dorsal side of the bone, and on nearing the lower or posterior border of the
fibula the inner surface becomes tendinous. It has the same action as the gastrocnemius. Murie
gives an origin from the outer condyle of the femur which I did not observe.
The Plantaris in the Phocinae lies below the gastrocnemius. It arises, as already mentioned, from
the femur with the outer head of the gastrocnemius, and descends along the ventral side of the
flexor longus hallucis, at the lower third of the leg it crosses to the dorsal side of the above muscle,
and enters the plantar surface between the gastrocnemius and the flexor longus hallucis, below the
backward prolongation of the tendons of the gracilis, semimembranosus, and semitendinosus which
form the plantar fascia. Beneath this it widens, and is moored to the dorsal side of the larger
combined tendon of the flexor longus hallucis and the flexor longus digitorum. Before reaching
this tendon, the accessorius is inserted into its dorsal side (fig. II., p. 201). It sends one slip, behind
its union with the combined tendon, to the distal end of the inner surface of the 5th metatarsal bone.
In Macrorhinus leoninus it arises alone from the same
part of the femur as the outer head of the gastrocnemius
and the plantaris in PJwca vitulina. It blends with the
insertion of the gluteus maximus on its outer side at the
origin. At the os calcis it enters the pes, as in Phoca vitu-
lina, and joins the dorsal side of the conjoined plantar
tendon of the flexor longus digitormn and the flexor longus
hallucis (fig. IV., p. 202).
In Arctocephalus gazella it is one-third the size of the
gastrocnemius, and arises, in common with the popliteus,
from the external border of the femur to the point of its
tendon from the external condyle. It courses backwards,
lying upon the soleus, partially covering the gastrocnemius,
and is situated on the dorsal side of the leg. Near the ankle
it forms a round tendon, wlndi occupies the groove on the os
calcis to the ventral side of the gastrocnemius. One inch
posterior to the distal end of the os calcis it widens and
divides into an anterior and posterior slip. The anterior
joins the plantar fascia (fig. I.). The posterior divides into
four slips, which are the superficial perforated tendons for
the 2nd, 3rd, and 4th digits. In the Phocinae it is supplied by the great sciatic nerve.
The Deep Group in all the specimens is alike. The muscles are the popliteus, flexor longus
hallucis, flexor longus digitormn, with its accessorius and lumbricales, and the tibialis posticus.
T.P.
jinnnJ
PLF.
Fig. I. — The plantar fascia of Arctocephalus.
Sm, semimembranosus ; St, semitendinosus ;
PI. , plantaris ; T.P., tibialis posticus ;
Pl.F., plantar fascia.
200 THE VOYAGE OF H.M.S. CHALLENGER.
The Popliteus in the Phocina? is a triangular muscle with a round tendon. It arises from within
the capsule of the knee-joint, from a shallow fossa situated below the termination of the external
supracondyloid ridge on the lateral surface of the external condyle. The tendon of origin turns
round to the back of the external condyle throughout its posterior surface. It crosses the back of
the knee-joint obliquely from without inwards, and is inserted into the upper third of the ventral
border of the tibia, into the inner part of the ventral tuberosity, into the whole extent of the dorsal
side of the internal lateral ligament, and into the inner surface of the tibia anterior to the feebly
marked oblique line posterior to this ligament.
In Macrorhinus leoninus the tendon arises below a slight depression on the lateral side of the
external condyle ; otherwise as in Phoca vitidina. It is inserted into the well-marked triangular
surface anterior to the oblique line of the tibia ; otherwise as in Phoca.
In Arctocephalus gazella it is larger than in the Phocinse. It arises from the external surface
of the external condyle by a strong round tendon, which forms part of the capsule of the knee-
joint, and by the same origin as the plantaris from the femur. The latter origin at once becomes
muscular and covers the round tendon. The two heads blend over the back of the knee-joint,
and cross between the outer condyle of the femur and the head of the fibula. It is inserted,
into the anterior two-thirds of the inner surface of the tibia dorsal to the internal lateral ligament
and ventral to the popliteal line. The internal lateral ligament only extends backwards to the
middle of the shaft. It bends the knee and rolls the leg inwards.
There is in all the specimens a groove upon the external condyle of the femur for the tendon of
the popliteus. The oblique line runs from the junction of the external and internal tuberosities on
the inner surface of the tibia, backwards and downwards to join the ventral border of the tibia.
This is very different from human anatomy, where it runs from the fibular facet of the tibia to
the internal border. The oblique line in Macrorhinus is more like what is seen in man. In the
Phocinae the muscle is supplied by the great sciatic nerve.
The Flexor longus hallucis is the flexor digitorum of Humphry; in Phoca vitidina it is an elon-
gated fusiform mass of fibre, and is the largest of the deep flexors of the back of the leg. It arises
from the inner surface of the fibula, going backwards to its posterior extremity, from the inner
surface of the head, and from the interosseous membrane. It just overhangs the ventral border of
the fibula, and does not encroach far upon the interosseous space. The flexor longus digitorum
touches its border and the tibialis posticus lies to its ventral side. Anterior to the inner surface of
the ankle-joint it forms a tendon which is broad, flat, and strong ; this runs in a groove on the
backward projection of the astragalus through a fascial tunnel formed by the annular ligament. In
Phoca hispida and in Phoca barbata the origins and insertions are similar to those in Phoca vitidina.,
but the development is much more perfect in the two former than in the latter, the bellies being
much larger and more fusiform. It can with safety be said that the bellies were enormous for the
size of these two animals.
In Macrm-hinus it is like that in Phoca vitidina, but in addition there was a dense fascia over
its anterior surface. The belly was the same as in Phoca vitidina, but only of moderate size.
In Arctocephalus gazella it arises from the inner surface of the head of the fibula, from the inner
surface of the anterior fourth of the shaft, and by an aponeurosis from the tibia, which gradually
passes from its dorsal border to the short inner border on the posterior two-thirds of the shaft.
Near the ankle it forms a tendon, which runs beneath the annular ligament in the groove on the
REPORT ON THE SEALS.
201
inner surface of the posterior extremity of the tibia, dorsal to the tibial groove, and at the posterior
border of the os caleis blends with the flexor longus digitorum. The insertion comes after the
insertion of the flexor longus digitorum in the Phocince and Macrorhinns.
In Arctoccphalus the inner surface of the tibia differs from other Seals ; for upon the posterior
third of the shaft is a border intermediate between the ventral and dorsal borders, and it is to
it that the origin of the flexor longus hallucis changes from the dorsal border. In Otaria and
TrMicchus it arises from the fibula only. In the Phocinse it is supplied by the great sciatic nerve.
The Flexor longus digitorum is the flexor quatuor digitorum of Lucae. In the Phocinre it is a
triangular muscle, and arises from the triangular surface of the fibula to the dorsal side of the
interosseous membrane behind the tibio-fibular fusion, and from the popliteal line on the inner
JZH PI
FLB
FLD.
SP
SP
J7ZB
ELD
Fig. III. — Tendons of the foot of Arctocephahis.
FLD, Flexor longus digitorum ; FLH, Flexor
longus hallucis.
SP n
Fig. II.— Tendons of the foot in the Phocina;. FLD, Tendon of
Flexor longus digitorum ; FLU, Tendon of Flexor longus
hallucis ; PI, Tendon of the Plantaris ; Ac, Insertion of the
Accessorius ; Gr, Tendinous slip from the plantar fascia, which
is formed by the Gracilis, Semimembranosus, and Semiten-
dinosus ; S, Tendinous slips to the flexor tendon sheaths ; SP,
Flexor sublimis digitorum (perforates) ; D, Flexor profundus
digitorum (perforans) ; M, Teudinous slips to the metatarsal
bones.
surface of the tibia behind the insertion of the popliteus to the middle of the shaft. The tibialis
posticus lies to its dorsal side, and the ventral border of the flexor longus hallucis behind. It forms
a tendon which crosses to the dorsal border of the tibialis posticus, and enters the dorsal furrow in
the large groove above the internal malleolus beneath its division of the annular ligament.
In Macrorhinns leoninus it is different, for there is a large popliteal line, and it arises from the
whole of it. The ventral tuberosity of the tibia forms more of the. internal surface than in the
(zool. chall. zxp. — paet lxviii. — 1888.) Yyy 26
202
THE VOYAGE OF H.M.S. CHALLENGER.
other Seals, for, while in the others the popliteal line forms part of the ventral border of the shaft,
and the anterior end of the popliteal line terminates in the Phocinse upon the inner dorsal side of
the ventral tuberosity, in this case it ends upon the dorsal tuberosity.
In Arctoeephalus gazella it arises from the posterior two-thirds of the ventral border of the
fibula, from almost the same extent of the dorsal border of the tibia, from the inner surface of the
FLD.
FLH
JDT.
Flo. IV. — Tendons of the foot of Macrorhiims. FLD., Tendon of Flexor longus digitorum ; FLH., Tendon of
Flexor longus hallucis; PI., Tendon of the Plantaris ; Or., Tendinous slip from the plantar fascia, which is
formed by the Gracilis, Semimembranosus, and Semitendinosus ; S, Tendinous slips to the flexor tendon
sheaths ; ST., Flexor sublimis digitorum (perforatus) ; DT., Flexor profundus digitorum (perforans).
shaft of the tibia between the intermediate and the dorsal borders to 1 inch from the ankle, and
from the posterior two-thirds of the interosseous membrane. The muscle ends in a strong tendon,
which traverses the groove of the astragalus and joins the tendon of the flexor longus hallucis. It
arises in Otaria and Trichcchus from the lowermost two-thirds of the shaft of the fibula and the
REPORT ON THE SEALS. 203
lowermost third of the tibia and the interosseous membrane. In the Phocinoe it is supplied by a
branch of the great sciatic nerve.
The union and insertion of the tendons of the flexor longus hallucis, and the flexor longus
digitorum in the Phocinre is as follows : — The tendon of the flexor longus hallucis is the largest
in the sole ; on its tibial side, 1 inch posterior to the os calcis, the tendon of the flexor longus
digitorum unites with it. Upon its fibular aspect the plantaris tendon expands, and is fused
with it by its outer surface. A slip from the plantar fascia of the gracilis, &c, blends with it
along the dorsal edge ; opposite the proximal extremities of the metatarsals this union of tendons
and plantar fascia divides. The part which corresponds to the flexor longus digitorum gives off two
slips. The ventral slip descends to the distal end of the terminal phalanx of the hallux, and is
inserted there. The dorsal slip goes to the proximal dorsal side of the 1st phalanx of the hallux,
into which it is inserted.
The rest of the tendons, which roughly are those of the flexor longus hallucis, break up in a
more complex manner. For the 2nd digit two tendons spring out of the common one. The anterior
is the superficial or perforated tendon, which gives off an anterior slip to blend with the sheath.
The anterior superficial tendon splits over the proximal end of the 1st phalanx, and is inserted into
the proximal end of the 2nd phalanx. The posterior or deep tendon gives off a posterior slip, which
is inserted into the distal end of the metatarsal ; then the deep tendon passes through the slit in the
superficial tendon, and is inserted into the distal end of the terminal phalanx of the 2nd digit. For
the 3rd digit there are three slips coming off separately. Two come off anterior and posterior to
each other. The third is a large, strong slip, springing from the main tendon between the flexor
tendons for the 3rd and 4th digits. This large slip is attached to the distal end of the 3rd meta-
tarsal. The anterior slip divides into an anterior and posterior part. The anterior is inserted into
the sheath, the other is the superficial tendon or posterior part, which is inserted like the other
superficial tendons. The posterior part from the main tendon passes through the same slit as the last,
and is inserted as in the former group. For the 4th digit, they are the same as for the 2nd digit,
with a slight difference. There is no posterior slip from the deep tendon, and the anterior slip to
the sheath is formed by the direct continuation of the plantaris tendon, which only fuses on its
under surface with the great tendon. Those for the 5th digit have the same attachments. The
difference in this group, as compared with the 2nd digit, is in the formation of the superficial or
perforated tendon, which is formed from the plantar fascia of the gracilis, &c, and only joins the
great tendon on its dorsal edge. This superficial slip gives off a small one to the distal end of the
metatarsal of the 5th digit.
In Macrorhinus leoninus the combined tendon on the plantar surface divides into four slips.
The ventral or internal one soon breaks into three. The ventral and middle slips of these three
are for the hallux, and have the same course and insertion as in Phoca vitulina. The dorsal of
these three is for the 2nd digit and forms an anterior and posterior tendon, which are the same
as those for the 2nd digit of Phoca vitulina, without the posterior slip for the distal end of the
metatarsal. In the 3rd digit they are similar to those in Phoca vitulina, without a slip from the com-
bined tendon. In the 4th digit they are also similar to those in Phoca vitulina, but the plantaris
muscle forms a greater part of the tendons. In the 5th digit the tendon is chiefly formed by the
plantaris tendon and by the plantar fascia of the gracilis, &c. It comes off in one slip and divides
into two, which have the same insertion as in Phoca vitulina, but there is no slit in the superficial
204 THE VOYAGE OF H.M.S. CHALLENGER.
tendon. All the slits of the other superficial tendons are lateral, and not antero-posterior as in the
other Seals.
In Arctocephalus gazclla the insertion of the combined flexor tendons out of the union of the
flexor longus digitorum and the flexor longus hallucis forms a rectangular band, which divides at
the base of the 2nd metatarsal bone into two broad tendons, the ventral and the dorsal portions.
The ventral portion also divides into two, forming the flexor longus hallucis tendon and the first
or inner flexor longus digitorum tendon. The ventral or flexor longus hallucis slip runs backwards
to the terminal phalanx of the hallux, and after expanding is inserted chiefly into its proximal
plantar surface, and into the whole of the plantar surface of this phalanx, by the prolongation of
the tendon of insertion over the surface of the terminal bone. The first or inner flexor longus
digitorum tendon, formed out of the dorsal part of the ventral division of the main tendon, is
described with the following deep tendons. The outer or dorsal main portion breaks into three
slips, that for the 5th digit coining off higher than the other two. The four long flexor tendons thus
formed go backwards along the plantar sides of the 2nd, 3rd, 4th, and 5th digits to the distal
phalanges ; opposite the bases of the 1st phalanges they pass through the aponeurotic tunnels in
the short flexors formed from the plantaris, becoming anterior to them, and are inserted into the
phalanges as the tendon of the flexor longus hallucis.
The action of these combined muscles in the Phocina? and Macrorhinus is to bring the pes
to the middle line and to bend the digits. In Arctocepludus they will raise the heel in walking,
otherwise they are the same.
There is an important difference in the relation of the flexor longus digitorum and flexor
longus hallucis in the Phocinffi, Macrorlvinns, and Arctocephalus. In the first two the flexor
longus hallucis is to the dorsal side of the flexor longus digitorum from origin to insertion, but in
the last the flexor longus hallucis is superficial to the flexor longus digitorum and crosses anterior
to the ankle to the ventral side of the pes; and the flexor longus digitorum in the pes lies dorsal
to it — the reverse of what is found in the Phocinas and Macrorhinus.
While recognising the intermingling of the tendons of the flexors, I find it impossible to work
out how far the tendons of the flexor hallucis and flexor longus digitorum cross each other to assist
in forming the flexors of the digits, and therefore I have described only what is easily made out.
Humphry writes " in the case of the pollex the superficial tendon did not divide as in the
other toes." In my dissections of the Phocinas I find that the slip out of the combined tendon
comes off singly, and very soon divides into two long slips, one being the flexor longus hallucis and
the other the flexor brevis hallucis. The same author also explains that " the tendons of each muscle
(flexor longus hallucis and flexor longus digitorum) contributed some fibres to each of the tendons
(with the exception presently to be mentioned), but the deep tendons were derived mainly from
the flexor longus pollicis, the flexor digitorum being distributed chiefly to the superficial tendons.
The superficial tendon of the 4th digit was in one foot, and that of the 5th hi both, derived from
the plantaris." The flexor longus digitorum in the Phocinse is not crossed by the flexor longus hallucis
as in human anatomy, but they pass each other along their contiguous edges without crossing. The
two tendons for the most part keep their own side in the pes. The flexor longus digitorum gives
off the tendons for the hallux and 1st digit, the flexor longus hallucis for the 3rd, 4th, and 5th
digits. I find that the plantaris forms the anterior slip for the sheath of the 4th digit and not
the superficial tendon, and in the 5th digit the tendons were principally formed by the flexor
REPORT ON THE SEALS. 205
longus hallucis, the plantaris giving a shp apart from the usual tendons to the distal end of the 5th
metatarsal.
The Plantar fascia in the Phocinte is formed out of the tendons of the gracilis, semimembranosus,
and semitendinosus, which are prolonged into the foot, while the tendon of the plantaris muscle is
interposed between it and the combined tendon of the flexor, and does not form a plantar fascia,
but strengthens the common tendon, and forms part of the flexors of the digits. In the
foot three layers are got from this modification, the first by the gracilis, &c, the second by the
plantaris, and the third by the flexor longus hallucis and flexor longus digitorum.
The Lumbricalcs in the Phocinfe and Macrorhinus may be represented by the anterior tendons
from the combined tendon going into the sheath of the digits.
The Lumbrical muscles in Arctocephalus gazclla are five in number. The first lies between the
long flexor tendons for the 1st and 2nd digits, coming out of the ventral main division. It arises
from the adjacent sides of these tendons and forms a small tendinous slip, which is inserted into
the distal dorsal side of the 1st metatarsal. The second arises from the surface and ventral side
of the long flexor tendon for the 3rd digit, and is inserted by muscular fibres into the tunnel in the
superficial flexor tendon formed out of the plantaris muscle. The third arises from the surface of
the deep tendon for the 4th digit, and ends upon it near the distal end of the 4th metatarsal bone,
like the last. The fourth arises from the dorsal side of the deep tendon for the 4th digit, passes
beneath the deep tendon for the 5th digit, and is inserted by a small tendon into the ventral side of
the distal end of the 5th metatarsal. The fifth comes from the tendon of a different muscle. The
superficial tendon for the 5th digit from the plantaris gives origin upon its surface to a lumbrical
muscle, which ends on the same tendon lower down. From the description of these slender
fusiform muscular slips it will be seen that there are five, four from the combined tendons of the
flexor longus digitorum and the flexor longus hallucis, and the fifth from the plantaris tendon. In
Otaria there are six, the sixth is derived from the outermost tendon of the flexor longus digitorum,
but there are no other differences.
The Accessorius is the M. caro-quadrata of' Lucae. In Phoca vitulina, Phoca hispida, and
Phoca barbata it is a triangular muscle, with its base directed outwards, and arises from the dorsal
surface and posteiior end of the os calcis to the inner side of the groove for the long peroneal
tendon. The fibres pass inwards and obliquely backwards over the dorsal border of the hindward
corner of this bone, forming a fine tendon which is inserted into the outer side of the tendon
of the plantaris, before this muscle reaches the combined tendon. In Macrorhinus leoninus it
was wanting, but most probably had decayed. In Trichechus it is absent, and was not noticed in
Otaria.
The Tibialis posticus in the Phocinse and in Macrorhinus leoninus is triangular, and lies to the
outer side of the flexor longus digitorum. It arises from the inner side of the interosseous mem-
brane, from the anterior two-thirds of the inner surface of the tibia, from the anterior third of the
ventral edge of the fibula near the interosseous membrane, and from the inner side of the dorsal
tuberosity of the tibia beneath the place of fusion of the tibia and fibula. It forms a strong
tendon which passes beneath the flexor longus digitorum on its ventral side, and enters the ventral
division of the groove on the outer surface of the distal extremity of the tibia.
In Phoca, near the tubercle of the scaphoid, it gives off a slip which becomes the middle slip of
the abductor hallucis ; to the inner side of the abductor slip it gives off another in which the
20G THE VOYAGE OF H.M.S. CHALLENGER.
sesamoid bone is found ; and then is inserted into the tubercle of the scaphoid, spreading over its
plantar surface, and into the entocuneiform bone. A few fibrous bands end upon the proxirna
end of the 1st metatarsal to the inner side of the flexor brevis hallucis. In the large Phoca,
from beneath the inner head of the abductor hallucis, a slip from the tibialis posticus tendon
proceeds backwards to the inner side of the flexor brevis hallucis, and ends on the ventral or inner
surface of the 1st metatarsal. In Macrorhinus the tendon is inserted into the scaphoid tubercle,
into the entocuneiform, and the 1st metatarsal; and gives off a strong slip to the abductor hallucis.
In Arctocefphalus gazella it is of the same size as the flexor longus hallucis. It arises from the
anterior fourth of the ventral border of the fibula, from the inner surface of its head, from
the inner surface of the tibio-fibular fusion, from the anterior three-fourths of the inner surface
of the tibia dorsad to the oblique line, and from the anterior fourth of the interosseous membrane.
About 1 inch from the ankle it forms a tendon, which goes beneath the annular ligament in the
groove near the ventral border of the tibia. After traversing it the tendon expands and is
inserted into the anterior half of the 8th tarsal or entoscaphoid bone.1 On nearing its insertion
it gives off a tendinous slip (slip i.), which crosses the surface of this bone and joins the
plantar fascia. It also sends a strong slip over the inner or ventral half of the bone which
runs along the ventral plantar side of the 1st metatarsal; opposite the middle of the shaft this
slip divides into two (ii. and hi.). The dorsal slip (ii.) is prolonged to the distal plantar surface of
the 1st phalanx of the hallux. The ventral (iii.) is inserted into the distal plantar ventral side of
the 1st metatarsal. In Otaria it is only inserted into the scaphoid; but Murie does not
mention it in Trichcchus. Lucae agrees with me as to its insertion in Phoca, and Humphry
gives the same scaphoid and metatarsal insertions, but states "that a considerable portion
of its tendon extended into the ligaments under the tarsus and into the tendinous structure which
represents the short muscles of the hallux." In the Phocinse it is supplied by a branch of the
great sciatic nerve.
The human tibia upon the posterior surface has a ridge dividing it into two ; the outer
division is for the origin of the tibialis posticus. In all the specimens of Seals there is no ridge, and
the inner surface is for the tibialis posticus. The part of the bone covered by the muscle in the
Phocinas is deeply scooped out, and gutter-like, the convexity being on the outer side, and in most
of the specimens the shaft is semitransparent. This formation gives lightness to the bones of the
leg but little diminution in surface. In Macrorhinus the inner surface is very slightly concave,
and the shaft is triangular in transverse section, the apex of the triangle giving attachment to the
interosseous membrane. In Arctocephcdus the inner surface is only moderately scooped in its
anterior third, and the shaft is triangular like the last. In all, the origin reaches the dorsal
tuberosity of the tibia, for the popliteal line begins at the dorsal side of the ventral tuberosity, but
in man the insertion of the popliteus prevents this.
The ventral surface of the tibia is apt to be included with the inner surface, unless a number
of tibise are examined. The ventral border begins at the internal malleolus, runs along the shaft,
and terminates at the junction of the outer two-thirds and inner third of the ventral tuberosity.
The dorsal border runs from the external malleolus forwards to the junction of the external and
internal tubercles on the outer side of the bone. The space between these two borders is the
1 See Sir W. Turner's Report, p. 50. In the Phocina? I found a sesamoid bone in the tendon of the tibialis posticus,
but there was none in Macrorhinus.
REPORT ON THE SEALS. 207
ventral surface. The popliteal line, which begins about the middle of the shaft in all, runs
forwards and ends at the dorsal side of the ventral tuberosity. A casual look at the bones might
suggest that the concave surface giving origin to the tibialis posticus is the inner surface, but such
is not the case. The small triangular surface for the popliteus also belongs to the inner surface.
This is best seen in Macrorhinus, where the popliteal surface advances upon the inner to a greater
extent than in the rest. In Arctoccphalus the inner surface is more convex, and the popliteal line
stands out like a ridge ; this is still more evident in the Phocinas. The popliteal line in the Seals
only gives origin to the flexor longus digitorum.
In the Phochife and Macrorhinus, where the pes is always in line with the trunk, it will
in the backward and forward motion of the paddle, assist in bringing the pes to the middle
line, i.e., adduct it, and turn the sole a little upwards, i.e., pronate it. In Arctoccphalus in
walking it will extend the ankle, raise the inner side of the pes and the heel, besides giving the
other movements when swimming.
Pes. — The Outer Region in all the specimens has one muscle, the extensor brevis
digitorum.
The Extensor brevis digitorum in the Phocinaj arises from the outer surface of the os calcis,
ventral to the peroneal tendons, from the superior dorsal border of the os calcis, and slightly from
the surface below the latter. It forms three muscular slips which end in two small tendons, the
common extensor of the first running backwards between them. The ventral slip goes between the
1st and 2nd metatarsal bones, and is inserted into the ventral side of the proximal end of the
1st phalanx of the 1st digit. The dorsal slip goes between the heads of the 4th and 5th metatarsals,
and is inserted into the proximal end of the 1st phalanx of the 3rd digit.
In Macrorhinios leoninns it is in two separate slips. The dorsal slip arises from the os calcis
ventral to the peronei, and passes between the 1st and 2nd metatarsals, and is inserted into the
proximal end of the ventral side of the 1st phalanx of the 2nd digit, and, by a small tendon from
the side of this one, into the distal dorsal side of the 1st metatarsal. The ventral slip arises from
the astragalus on its outer surface, and from the outer surfaces of the scaphoid and external cunei-
form bones. The tendon passes back to the interval between the 4th and 5th metatarsals, and is
inserted into the proximal dorsal side of the 4th digit.
In Arctoccphalus gazclla it is in two parts. The dorsal part has two heads of origin. The
larger head arises from the dorsal surface of the os calcis and from the dorsal surface of the cuboid.
The smaller head arises from the same bones, but to the ventral side of the large head. These two
heads unite and are inserted into the proximal end of the 1st phalanx of the 2nd digit. The second
part arises from the adjacent sides of the os calcis and astragalus, and from the cuboid, and is
inserted into the proximal end of the 1st phalanx of the 3rd digit.
The Inner or Plantar Region has adductors, flexores breves of the phalanges, and abductors.
In Macrorhinus the inner head of the flexor of the hallux and the abductor hallucis were the only
two intrinsic muscles seen ; the rest had evidently decomposed.
The Adductors. — In Phoca vitulina the adductor minimi digiti is found. In Arctoccphalus
there are the adductor hallucis, the adductor minimi digiti, and the adductor ossis metatarsi primi.
The Adductor minimi digiti in Phoca vitulina arises between the bases of the 3rd and 4th
208 THE VOYAGE OF H.M.S. CHALLENGER.
metatarsal bones, crosses the 4th metatarsal, and is inserted by a fine tendon into the tibial side of
the base of the 1st phalanx of the 5th metatarsal.
In Arctocephalus the Adductor hallucis and the Adductor minimi digiti arise by a common origin
from the plantar surfaces of the proximal ends of the 2nd, 3rd, and 4th metatarsal bones, and from
the sheath of the peroneus longus by a tendinous sheet which is semi-circidar. Along the posterior
border of this tendon muscular fibres spring and take the form of a horse-shoe, the two limbs
forming these two muscles. The tibial lirub is the adductor hallucis and is inserted into the fibular
distal end of the 1st metatarsal. The fibular limb is the adductor minimi digiti and is inserted
into the distal tibial side of the 5th metatarsal.
In Otaria these muscles are regarded as superficial interossei, and their origins are more extensive
than in Arctocephalus. In Trichechus (Murie) they are similar to Otaria. In Trichecus Cunningham
says they form the plantar layer, and he figures the adductor hallucis as consisting of two parts —
an adductor obliquus and an adductor transversus, but I did not find the transverse head of the
adductor hallucis, and the adductor rninimi digiti was not a separate fasciculus, as he figures it. The
general plan of origin in the two specimens of Arctocephcdi was like Murie 's drawings, the adductors
being combined at their origins.
The Adductor ossis metatarsi primi in Arctocephalus arises from the anterior third of the tibial
side of the plantar surface of the 2nd metatarsal, crosses to the 1st digit, and is inserted into the
1st phalanx on its fibular proximal side.
The Flexorcs breves. — These are named the deep interossei by Dr. Murie, and form the inter-
mediate layer of Professor Cunningham. In Phoca vitidina and in Arctocephalus the muscles of
the 2nd, 3rd, and 4th digits are double, of the 5th single, and the 1st digit is peculiar.
In Phoca vitidina they are feeble and arise from both sides of the plantar surfaces of the 2nd, 3rd,
and 4th metatarsals, and from the tibial side only of the 5th. The muscle from the tibial side of the
2nd digit also has origin from the fibular proximal end of the 1st metatarsal. They are inserted
into the proximal ends of the first phalanges of the digits, on the same sides from which they spring.
In Arctocephalus they are well developed. The origins and insertion are as in Phoca vitulina,
with one exception — the muscle from the tibial plantar surface of the 2nd metatarsal is only from
the posterior two-thirds of the shaft.
The Flexor brevis primi metatarsi or flexor brevis hallucis. In Phoca vitulina the outer muscle
or outer head arises from the fibular side of the 1st metatarsal, and is inserted into the proximal
fibular side of the 1st phalanx of the 1st digit. The inner flexor or inner head in Otaria is named
the adductor hallucis, in Trichechus (Murie) the flexor brevis hallucis, and in Trichechus
(Cunningham) this slip was not found. In Phoca it arises beneath the adductor hallucis from the
outer posterior third of the scaphoid bone, upon the tendon of insertion of the tibialis posticus, and
from the anterior outer third of the entocuneiform. It is a glistening tendinous band, with a
reddish tinge at the anterior end, which passes backwards and inwards and is inserted into the
inner side of the base of the 1st metatarsal.
In Macrorhinus the inner head arises from the outer posterior half of the scaphoid bone in
common with the outer head of the abductor hallucis, over the middle of the entocuneiform bone
the tendon splits into two equal portions — the inner is the abductor hallucis, the outer forms the
flexor brevis hallucis. It is inserted into the proximal tibial plantar surface of the 1st metatarsal
to the outer side of the abductor hallucis.
REPORT ON THE SEALS. 209
In Arctocephalus the outer muscle is the same as in Phoca ; the inner arises from the tendon of
the tibialis posticus, which is attached to the outer side of the anterior end of the entocuneiform
bone, and from the anterior outer half of the same, and is inserted as in Phoca.
The flexores breves in Otaria consist of one single and four double muscles ; as also in Trichechus
(Cunningham). While Professor Cunningham alludes to no differences in their insertions, Dr.
Murie gives the insertion in Otaria of the first interosseus into the fascia covering the metacarpo-
phalangeal joint of the hallux, which is very like what I have stated. Excluding Phoca, we agree
as to some change in the tibial side of the 2nd metatarsal. Murie in Otaria derives the tibial head
of the 2nd muscle from the proximal ends of the 1st and 2nd metacarpals. The smaller moiety
of this muscle, that next the hallux, has also a partial origin or attachment to the superficial layer
of the interosseous fibres and hallucial metacarpal. Professor Cunningham, in describing the flexor
brevis indicis, gives the origin of the tibial head from the base of the 1st metatarsal. In my
account of this digit in Arctocephalus I describe an adductor from the anterior tibial third of the
2nd metatarsal.
Tlic Abductors. — In Phoca vitulina these are the abductor hallucis, the abductor minimi digiti,
and the abductor tertius quinti digiti.
In Arctocephalus the abductor hallucis, the abductor minimi digiti, the abductor tertius quinti
digiti, the abductor ossis metatarsi quinti, and the abductor ossis metatarsi primi are found. In
Macrorhinus the abductor hallucis only is described.
The Abductor hallucis in Otaria is named the flexor brevis hallucis, in Trichechus (Murie) the
abductor hallucis, and in Trichechus (Cunningham) the inner head of the flexor brevis hallucis.
In Phoca vitulina it originates by three separate slips, which are close to each other and attached
posteriorly. The outermost arises from the scaphoid bone upon the tendon of insertion of the
tibialis posticus, a little to the outer side of the inner head of the flexor brevis hallucis, which lies
at a greater depth in the sole, and from the adjacent posterior surface of the os calcis ; the middle
from the tendon of the tibialis posticus before it reaches the scaphoid just anterior to its insertion ;
the inner by a slip which comes from the outer posterior side of the sesamoid bone of the tibialis
posticus tendon. These three slips unite a little posterior to the sesamoid bone, forming a strong
tendon, which is inserted into the inner distal plantar side of the 1st metatarsal. On both sides of
the slip which comes from the tibialis posticus tendon, and on the outermost side of the middle
two-thirds of the outermost tendon, there are a few muscular fibres.
In Macrorhinus it arises, in common with the inner head of the flexor brevis hallucis, from the
outer posterior half of the scaphoid bone upon the tendon of insertion of the tibialis posticus before
it reaches the scaphoid. It is a strong fibrous band which is directed backwards ; midway
between its origin and insertion, it is joined on the outer side by the outer head, and the two
together are inserted into the proximal tibial plantar surface of the 1st metatarsal.
In Arctocephalus it arises from the tubercle on the posterior end of the sesamoid bone of the
tibialis posticus, and is closely united with its tendon. It courses backwards along the tibial side of
the 1st metatarsal, and is inserted into the distal tibial side of the 1st metatarsal and the proximal
end of its 1st phalanx, receiving some fibres from the tibialis posticus, which pass over the
sesamoid bone into it. Its almost tendinous nature, its close association with the tendon of the
posticus, and its arising from the sesamoid bone, show that it has a similar function to the posticus.
In Otaria Murie does not describe this muscle.
(ZOOL. CHALL. EXP. — PART LXVIII. — 1888.) Yyy 27
210 THE VOYAGE OF H.M.S. CHALLENGER.
In their descriptions of Trichcchus Dr. Murie and Professor Cunningham differ. The former
states that it arises by a long narrow belly, by a tendon from the extra bone outside the cuneiform,
and is fleshy three-quarters the length of the hallucial metacarpal, being inserted by tendon and
fascia over the metacarpophalangeal joint ; the latter that it arises from a sesamoid bone which
slides upon the tibial side of the internal cuneiform, and is inserted into the inner side of the base
of the 1st phalanx of the hallux. Murie writes that it is fleshy, and Cunningham that it is
tendinous, which he considers is probably owing to his specimen being a pup ; and although Murie
explains that this muscle in the Seal is entirely tendinous, I found muscular fibres in Phoca.
The Abductor tertius quinti digiti in Phoca vitulina is exposed after reflecting the tendinous
structure concealing it. Murie has classed it as the 2nd head of the abductor ossis metacarpi
quinti. It arises from the adjacent sides of the bases of the 4th and 5th metatarsal bones, and
the tendinous structure covering these phalanges ; after crossing the 5th metatarsal it is inserted
into the fibular side of the head of this bone.
In Arctocephalus it is 1 inch long, with no fibres, and arises from the fascia completing the tunnel
for the peroneus longus tendon. This origin is a small round tendon at right angles to the plantar
surface. It is inserted into the tendon of origin of the abductor ossis metatarsi quinti. In Otaria
it is one of the heads of origin of the abductor ossis metatarsi quinti.
In Trichcchus (Murie) it is also named the abductor ossis metatarsi quinti. Its origin is the
same as in Arctocepihcdus and Otaria, but the insertion is into the base of the 5th metatarsal bone.
In Trichcchus (Cunningham) the origin and insertion are similar to the former.
The Abductor minimi digiti in Phoca vitulina arises between the proximal ends of the 3rd and
4th metatarsal bones, and crosses obliquely outwards to the tibial proximal plantar side of the 1st
phalanx of the 5th digit. In Arctocephalus it has two bellies. The first arises from the dorsal half
of the plantar surface of the os calcis by muscular fibres, and extends longitudinally from the
insertion of the gastrocnemius to the posterior tendon of this bone. The second belly arises from
the base of the 5th metatarsal bone. The first belly is inserted, after forming a flat tendon, into
the base of the 5th metatarsal beneath the second belly, which goes to the dorsal distal end of
the 5th metatarsal, and is inserted into the outer or dorsal side of the flexor brevis minimi digiti.
Murie's description of this muscle in Otaria differs from the above.
In Trichcchus Murie states that it comes from the outside of the os calcis and not from the
plantar fascia, while Cunningham says that it arises from the fascia covering the outer surface of
the abductor ossis metatarsi minimi digiti, and is inserted into the outer side of the base of the
1st phalanx of the minimus.
The Abductor ossis metatarsi quinti in Arctocephalus is the flexor brevis minimi digiti (Murie),
and the abductor ossis metatarsi minimi digiti (Cunningham). It is a small muscle, and arises
from the os calcis to the outer side of the origin of the abductor minimi digiti by a slender
elongated tendinous slip, which is closely united with the muscle just mentioned, and lies along its
fibular edge. It is inserted into the distal fibular side of the 5th metatarsal to the fibular side of
the abductor.
In Otaria the origin is double ; the outer head is the same as described above ; the inner I
regard as the 3rd abductor of the 5th digit which is found in Phoca. In Trichcchus (Cunningham)
it exhibits the usual attachments, but Murie gives a different description.
REPORT ON THE SEALS. 211
The Facial Muscles of Expression.
These are arranged in the following groups : — The occipito-frontalis, the muscles of the ear,
the muscles of the nose, the muscles of the eyelids, the muscles of the orbit, and the muscles of
the mouth.
The Occipito-frontalis in Phoca vitulina. — The specimen from which the cervico-scapular
panniculus is described had two V haped prolongations over the frontal region, which represented
the occipito-frontal muscles. In another Phoca these fibres extended towards the middle line and
touched each other, forming a complete occipito-frontalis. It is supplied by the facial nerve which
ascends from the ear over the temporal muscle. In Otaria it is imperfectly formed.
TJic Muscles of the Ear. — There are three small muscles to the cartilaginous meatus in Phoca
vitulina, two small pale fasciculi forming the protractors, and one a retractor.
The Internal protractor or Attollcns aurem arises from the skin above the middle of the orbit and
passes outwards and backwards, and is inserted into the inner and under surface of the cartilaginous
meatus of the external ear at the junction of the skin with the cartilage. It is recognised by
Humphry ; in Otaria it is indistinct ; in Trichechus the muscle is well developed, but arises
posterior to the orifice of the meatus, and is therefore a retractor.
The External protractor or Attrahens aurem arises from the fibrous tissue over the articulation of
the malar with the zygoma, and is inserted into the outer and under surface like the former. It is
not noted by Humphry ; in Otaria it is present but undescribed ; in Trichechus it is distinct.
The Retractor or Retrahens aurem arises from the superior border of the zygoma extending from
the osseous meatus, to midway between the articulation of the zygoma with the malar bone anteriorly
and the osseous meatus posteriorly. It is inserted into the inner and under surface of the
cartilaginous tube of the external ear. It is named the attrahens by Humphry ; in Otaria it is
feeble ; in Trichechus it is a strong muscle.
The Cartilaginous meatus is 1 inch long and S-shaped, the attollens pulls the tube forward
and opens it, the attrahens draws it forward and outwards, also opening it. The retrahens retracts
the tube and flexes the anterior bend of the cartilage, thereby closing the meatus. They are
supplied by the facial nerve.
The Muscles of the Nose. — The combined Dilator et depressor nasi in Phoca vitulina is rectangular.
It arises from the fossa of the superior maxilla on the outer side of the infraorbital foramen,
and from above the foramen, extending forwards to the third last molar tooth. The inferior
division of the 5th nerve pierces it, and the fibres above the nerve form the dilator nasi, those
below the depressor nasi. The former, after crossing the levator anguli oris and the constrictor
nasi, is inserted into the side of the nose. The latter is inserted into the side of the nose inferior
to the dilator, and into the upper Up, beneath the septum ; some fibres join those of the
opposite side, while a few are attached to the skin of the upper lip, coming off from the depressor
portion in small slips. The nerve of supply is the facial.
In Otaria they are described separately.
The Constrictor nasi (named the compressor in Otaria) arises in Phoca vitulina superiorly
from the whole length of the nasal cartilage, and is inserted into the premaxillary bone and
muscle of the other side, partly under cover of the levator anguli oris. It is supplied by the facial
nerve.
212 THE VOYAGE OF H.M.S. CHALLENGER.
The Levator labii superioris et aim nasi in Phoca vitulina is a rectangular muscle. It arises
from the dorsal surface of the frontal bone which lies between the orbits, the same surface of
the nasal bone, and the superior maxilla, nearly reaching the nasal orifice anteriorly. The fibres
proceed downwards and outwards, and are inserted into the muzzle from the nose to near the
angle of the mouth. It is supplied by the facial nerve.
The Muscles of the Eyelids. — The Orbicularis palpebrarum in Phoca vitulina arises from the tendo-
palpebrarum inferior to the pulley for the superior oblique, from the palpebral ligament superior
to it, from the frontal bone, and from the superior maxilla. The orbicular portion blends with
the corrugator supercilii and the occipito-frontalis fascia, and is attached to the superior maxilla,
to the ligament completing the orbit, and to the malar bone. The palpebral portion is feeble and is
attached to the malar bone.
The Tendo-palpebrarum arises from the nasal process of the superior maxilla, and ends in the
orbicularis palpebrarum, lying along the inferior surface of the tendon of the superior oblique.
The Superior palpebral ligament goes from the superior maxilla above the tendon of the superior
oblique to the orbicularis palpebrarum.
The Corrugator supercilii arises from the frontal bone and is inserted into the under surface of
the orbicularis and the occipito-frontabs.
The Tensor tarsi arises from the orbital surfaces of the frontal and superior maxilla three-fourths
of an inch below the nasal process of the latter. It ascends and is inserted into the tendo-palpe-
brarum at the junction of the lid and the tendo-palpebrarum. It blends with the orbicularis.
The Levator palpebrm superioris in Phoca vitulina arises from the upper margin of the optic
foramen external to the superior oblique. It passes forwards, expands, and is inserted into the
upper eyebd. It is supplied by the 3rd nerve.
The Muscles of the Orbit} — Besides the four recti and the two oblique muscles, there are two
retractors and two depressors of the third eyebd.
The Superior, Inferior, External, and Internal recti muscles resemble the corresponding human
muscles.
The Superior oblique arises from the inner, and sUghtly from the upper, surface of the optic
foramen. It passes forwards along the inner wall of the orbit beneath the bgament for the upper
eyelid through the pulley attached to the superior maxilla, and goes outwards to the eyelid and is
inserted into the eyeball on the inner side of the insertion of the superior rectus. The tendon after
passing through the pulley lies between the tendo-palpebrarum and the superior palpebral bgament,
and pierces the upper eyelid to gain its attachment. The pulley is attached to the superior
maxilla upon the margin of the orbit close to the articulation with the nasal process of the
frontal bone, posterior to the nasal process of the superior maxilla. It is supplied by the 4th nerve.
The Inferior oblique arises from the orbital surface of the superior maxilla to the inner side of
the inferior orbital foramen. It goes upwards and outwards round the eyeball, and is inserted into
it inferior to the external rectus attachment. It is supplied by the 3rd nerve.
The Superior external and Superior internal retractor muscles arise from the outer side of the
optic foramen, run along the optic nerve to the eyeball, and are inserted on the corresponding sides
of the sclerotic beside the optic nerve.
The Deprrcssors of the third eyelid arise together from below the optic foramen, widen out as
1 Rosenthal's description of the muscles of the orbit differs considerably from mine.
REPORT ON THE SEALS. 213
they near the inferior margin of the third eyelid, pass on each side of the Harderian gland,
and are lost in the substance of this lid. The cartilage in this eyelid is strongest in the centre,
where it forms a strong vertical rod. The eyeball is surrounded by a fibrous case which lines
the wall of the orbit, forming the periosteum, and on the outer side, where there is no osseous
protection, the periosteum is continuous with the fibrous case along the sharp edge of the
pterygoid and palate bones inferiorly, and the nasal and frontal supeiiorly.
The Orbital ligament completes the break in the circumference of the orbit. It is attached to the
malar and temporal bones at their zygomatic articulation, and to the frontal bone above the orbit
The Muscles of the Mouth. — The Orbicularis oris in Phoca vitulina surrounds the mouth, and,
where it passes from jaw to jaw, is indistinct. It arises from the under surface of the inferior
maxilla by fine fasciculi as far back as the 4th molar tooth. The fibres ascend round the angle of
the mouth, and are inserted into the superior maxilla from the 4th molar to the articulation of the
premaxilla ; some of the fasciculi next the symphysis and the premaxilla circle round the mouth.
It is supplied by the facial nerve.
The Levator labii supcrioris proprius arises from the superior maxilla, forming the margin of the
orbit, and is inserted into the upper lip. It is supplied by the facial nerve.
The Buccinator muscle in Phoca vitulina is very small and composed of feeble muscular fibres.
It arises from the superior maxilla from a line extending from the last molar backwards along the
edge of the palate bone to midway between the root of the zygoma and the hamular process of the
pterygoid, from the inferior maxilla, from a linear origin from above the inferior dental foramen
to the last molar, and from the pterygo-maxillary ligament. It is inserted into the orbicularis
oris. It is supplied by the facial and the 5th nerves.
The Levator anguli oris in Phoca vitulina, arises from the junction of the nasal bone with the
premaxilla, and from the protuberance of the superior maxilla. It is inserted into the canine fossa
and into the skin of the mouth anteriorly beneath the infraorbital nerve and the dilator nasi. It
is supplied by the facial nerve.
The Muscles of Mastication.
The Masscter in Phoca vitulina and in Arctoccphalus arises from the whole of the zygomatic arch,
and is inserted into the fossa of the lower jaw below the coronoid root to where the alveolar
margin commences. It is supplied by the inferior maxillary nerve.
In Otaria there are two layers of fibres, a superficial and a deep set.
The Temporal muscle in Phoca vitulina and in Arctocephalus covers the side of the cranium, below
the temporal ridge which traverses the parietal bone, running obliquely forwards from the middle of
the occipital ridge to the root of the nasal process of the frontal bone. It lies between this line
superiorly and the zygomatic arch inferiorly, and arises from the lower half of the parietal, from the
squamous surface of the temporal bone, from the outer surface of the frontal inferior to the
oblique line, from the outer half of the anterior orbital surface of the same, and from the superior
tip of the alisphenoid. It converges and is inserted into the outer border, anterior border, and
internal surface of the coronoid process of the lower jaw, coveiing the internal surface above a line
drawn from the condyle to 1 inch behind the last molar. It is supplied by the inferior
maxillary nerve.
214 THE VOYAGE OF H.M.S. CHALLENGER.
In Otaria the fibres have two directions.
The Pterygoideus intemus (Lucae's pterygoideus and Humphry's pterygoid) in Phoca vitulina and
Arctocephalus is a strong muscle. It arises from the external surface of the pterygoid bone, and from
the fossa on the outer side of the hamular process in Arctocephalus only, for this is absent in Phoca.
It is inserted into the subcondyloid process of the lower jaw in Arctocephalus and in Phoca, beside
the inner side of the ramus below the condyle, to midway between it and the angle of the jaw.
The subcondyloid process is feeble in Phoca, but extensive in Arctocephalus. It is supplied by the
inferior maxillary nerve.
In Otaria and Trichechus it has not been described.
The Pterygoideus extemus in Phoca vitulina and Arctocephalus is a very small cylindrical bundle,
and arises in the former from below the foramen rotundum, and in the latter from the bridge
of bone connecting the alisphenoid with the external pterygoid plate over the foramen rotundum.
In both it crosses transversely outwards and is inserted into the inner side of the condyle of
the lower jaw. It is supplied by the inferior maxillary nerve.
This muscle has not been described by Vrolik, Humphry, Lucae, nor by Murie. I found it in
all the specimens by dividing the symphysis and pulling the jaw gently outwards, when the bundle
of fibres attached as above was seen.
The Muscles of the Neck.
The only superficial muscle is the Stcrno-mastoid ; in Phoca vitulina it is a riband-shaped
muscle, and arises from the under surface and side of the anterior third of the presternum. It is
joined to its fellow for one inch and a half anterior to the presternum by a fine aponeurosis,
ascends to the mastoid process and is inserted into it at the root of the zygoma behind the insertion
of the trachelo-mastoid. It is supplied by external branches of the cervical plexus and a twig from
the spinal accessory nerve.
In Arctocephalus it is triangular with the base resting on the fascial slip representing the
clavicle. It arises from the dorsal surface of the presternum and cartilage of the 1st rib, from the
deltoid ridge of the humerus between the insertions of the pectoralis major on the inner side and
the deltoid on the outer, blending with the origin of the inner part of the brachialis anticus below;
and from the fascial slip representing the clavicle. This last origin is thin, and midway between
the sternal and humeral origins is almost devoid of fibres, the deficiency being filled in with fibrous
tissue. The humeral part blends with the cephalo-humeral muscle along its outer edge and the
pectoral along its inner. The muscle runs forwards, narrows, and is inserted into the occipital ridge
near the external auditory meatus anterior to the splenius. It is supplied by twigs from the
external branches of the cervical plexus.
In Otaria the sternal end of the muscle represented in Dr. Mime's Memoir (pi. lxxiii. fig. 33) is
like what I have described in Arctocephalus, and, as it extends outwards to the shoulder, it must have
other attachments than the manubrium, which is all that Murie gives in his description.
In Trichechus it is as in Otaria, but a division into two parts is not described.
The Infea-hyoid Region includes the sterno-thyro-hyoid and the thyro-hyoid.
The Sterno-thyro-hyoid in Phoca vitulina is called the costo-thyreo-hyoideus by Lucae, the sterno-
REPORT ON THE SEALS. 215
hyoid by Humphry, and the sterno-hyoid and sternothyroid by Murie. It is a long band with a
triangular expansion from its outer side opposite the head of the humerus, the apex lying upon it.
It is situated along the side of the neck upon the carotid artery and pneumogastric nerve, &c, and
arises by muscular fibres from the outer surface of the cartilage of the 1st rib, from the outer
surface of the lesser tuberosity of the humerus, from the transverse ligament going between the
two tubers of the humerus, and from the fascia binding together the great vessels and nerves going
to the nipper from the thorax and stretching from the 1st rib to the lesser tuber. At the level of
the thyroid gland it splits into two parts, an anterior and a posterior. The deeper or posterior is
inserted into the thyroid cartilage, the superficial or anterior into the hyoid bone ; in the large
specimen, the division was about midway between the origin and insertion. The part from the
lesser tuberosity may be named the omo-hyoid. It is supplied by the communicans noni nerve.
In Arctocephalus it arises from the tip of the dorsal surface of the presternum, proceeds for-
wards, and about 1 inch posterior to the thyroid cartilage divides into a dorsal and ventral band.
The dorsal is the sterno-thyroid, and is inserted into the thyroid cartilage, the ventral is the sterno-
hyoid and is inserted into the hyoid bone. The nerve was destroyed.
The Omo-hyoid in Plioca vitulina is the part of the sterno-thyro-hyoid attached to the humerus,
in Arctocephalus it is the outer margin of the sterno-mastoid. The Sterno-thyro-hyoid in Arcto-
cephalus has an origin somewhat like the sterno-mastoid in Plioca. Humphry does not refer to the
part forming the sterno-hyoid.
The Thyro-hyoid in Phoca vitulina and in Arctocephalus arises from the posterior part of the
oblique ridge of the thyroid cartilage, and is inserted into the hyoid bone.
The Supra-hyoid Region. — In this region are the digastric, stylo-hyoid, mylo-hyoid, and genio-
hyoid.
The Digastric in Phoca vitulina and Arctocephalus arises from the mastoid hollow and the
tympanic bulla. About its middle, in Phoca, a superficial transverse tendinous division exists. It
is inserted into the inferior surface of the angle and inferior border of the lower jaw to opposite the
last molar tooth on the inner surface. It is supplied by the facial and by the mylo-hyoid branch of
the inferior dental nerve.
In Otaria there is no tendinous intersection, but it is present in Trichcchus.
The Stylo-hyoid in Phoca vitulina and Arctocephalus is a narrow transverse band, and arises
from below the external auditory meatus, and is inserted into the hyoid bone. Humphry
considers it to be a part of the digastric, and Murie does not describe it. It is supplied by the
facial nerve.
The Mylo-hyoid in Phoca vitulina- and Arctocephalus is a triangular muscle, which, with its fellow,
fills in the intermaxillary space. It arises from the inner surface of the lower jaw above and a
little behind the inferior dental foramen, and from the alveolar margin until opposite the last molar
tooth, where the line of origin turns suddenly to reach the inferior border of the lower jaw. Thus
far the origin is muscular, but at the symphysis it is tendinous. The fibres are inserted into a
median fibrous raphe" and into the body of the hyoid bone. The digastric intervenes between it
and the lower jaw at the angle. It is supplied by the mylo-hyoid branch of the inferior dental
nerve.
The Gcnio-hyoid in Phoca vitulina and Arctocephalus is a small rectangular muscle situated
21(5 THE VOYAGE OF H.M.S. CHALLENGER.
immediately beneath the inner border of the stylo-hyoid. It arises from the inner surface of the
symphysis, and from the inferior margin of the lower jaw opposite the 2nd, 3rd, and 4th incisors.
It takes a small turn to the middle line and meets its fellow, and is inserted into the inferior
surface of the body of the hyoid bone. It is supplied by the hypoglossal nerve.
The Muscles of the Tongue.
The extrinsic muscles are the stylo-glossus, hyo-glossus, genio-hyo-glossus, and palato-glossus
(see Soft Palate).
The Stylo-glossus both in Phoca vitidina and Arctocephalus is a small muscular band which arises
from the ventral surfaces of the stylo-hyal and epi-hyal, and goes obliquely forwards over the end
of the hyo-glossus, ending at the tip of the tongue. It is supplied by the hypoglossal nerve.
The Hyo-glossus both in Phoca vitulina and Arctocephalus is a quadrilateral band, and arises from
the thyro-hyal, slightly from the basi-hyal and cerato-hyal. It extends along the under surface of
the tongue to its tip beneath the stylo-glossus. It is supplied by the hypoglossal nerve.
The Genio-hyo-glossus both in Phoca vitulina and Arctocephalus is thin and triangular. It arises
by muscular fibres from the inferior margin of the lower jaw, reaching from the symphysis to opposite
the second last molar tooth, and is blended near the symphysis with its fellow. The posterior fibres
go to the body of the hyoid bone and some to the pharynx ; the middle to the middle of the
tongue ; the anterior to the front of the tongue. It is supplied by the hypoglossal nerve.
The Muscles of the Pharynx.
These are as usual, the inferior, middle, and superior constrictors, and the stylo-pharyngeus
(for the palato-pharyngeus and salpingo-pharyngeus, see the Soft Palate).
The Inferior constrictor in Phoca vitulina arises from the side of the cricoid cartilage, &c, as in
human anatomy, and almost hides the middle constrictor. It is supplied by the pharyngeal plexus,
and the external and recurrent laryngeal nerves.
The Middle constrictor in Phoca vitulina arises from the posterior cornu of the hyoid bone,
along its anterior surface, and slightly from the cerato-hyal, and is inserted as usual. It is supplied
by the pharyngeal plexus.
The Superior constrictor in Phoca vitulina arises from the inner surface of the stylo-hyal and
epi-hyal, and from the fibrous tube of the pharynx between the two tympanic bulla?. It is inserted
into the median raphe of the pharynx and to the vertebral column between the two recti antici
majores. It is supplied by the pharyngeal plexus.
The Muscles of the Soft Palate.
These are the levator palati, tensor palati, palato-glossus, palato-pharyngeus, azygos uvulae, and
salpingo-pharyngeus.
The Levator palati both in Phoca vitulina and in Arctocephalus arises from the tympanic bulla below
and a little to the inner side of the Eustachian tube, and from the inferior surface of that tube. It
REPORT ON THE SEALS. 217
passes beneath the salpingo-pharyngeus, and is inserted into the posterior part of the soft palate
between the palato-pharyngei.
The Tensor palati both in Phoca vitulina and Arctocephalus is a round muscular bundle arising
from the tympanic bulla on the outer side of the Eustachian tube, and from the outer side of this
tube. It runs along the outer side of the pterygoid plate, and turns round the anterior aspect of
the hamular process ; then it spreads out as a fine tendon, fan-like, upon the aponeurosis of the
palate anteriorly. It is supplied by the otic ganglion.
The Palato-glossus both in Phoca vitidina and Arctocephalus arises by a few scanty fibres from
the anterior surface of the soft palate, and blends with the stylo-glossus.
The Palato-pharyngeus both in Phoca vitulina and Arctocephalus arises beneath the levator
palati from the posterior surface of the soft palate by one head. It is inserted as in man.
The Azygos-uvtdm both in Phoca vitidina and in Arctocephalus arises from the aponeurosis of the
soft palate, and is distributed as usual.
The Salpingo-pharyngeus both in Phoca vitidina and Arctocephalus arises from the hamular
process of the pterygoid, which is feebly developed in the former, but strongly in the latter. It
takes a backward course to blend with the stylo-pharyngeus.
Prevertebral Muscles. ■
The prevertebral Cervical Region contains the rectus capitis anticus major and minor, rectus
lateralis, and the longus colli.
The Pectus capitis anticus major both in Phoca vitidina and in Arctocephalus is a long slip
arising by three fasciculi from the ventral division of the transverse processes of the 3rd, 4th, 5th,
and 6th cervical vertebra?. Its origins are between the inner slips of origin, and the outer
slips of insertion of the longus colli. The anterior parts of the origins from the vertebrae are
tendinous. It runs forwards, and is inserted at the inner side of the foramen lacerum posterius,
and to the anterior three-quarters of the fossa on the ventral surface of the basi-occipital, anterior
to the rectus capitis anticus minor. It is supplied by an anterior branch of the suboccipital
nerve, and by the internal branches of the cervical plexus.
The Rectus capitis anticus minor in Phoca vitulina and in Arctocephalus is a small slip arising
from the atlas behind the condyle, and to the inner side of its foramen at the anterior border of the
lamina. It is inserted into the posterior three-quarters of the fossa on the ventral surface of the
basi occipital. In Arctocephalus it also has an origin from the tip of the transverse processes of
the axis, but the fossa is much deeper in Arctocephalus than in Phoca. It is supplied by the sub-
occipital nerve, and by the deep internal branches of the cervical plexus.
The Rectus lateralis in Phoca vitulina arises from the anterior surface of the transverse process
of the atlas outside the foramen, and is inserted into the inferior termination of the occipital ridge,
into the paramastoid process, and into the outer quarter of the fossa to the inner side of this process.
In Arctocephalus it arises as in Phoca, but on the inner side of the foramen, and is inserted
posterior to the foramen lacerum posterius and the origin of the digastric into the exoccipital bone.
It is supplied by the suboccipital nerve.
The Longus colli in Phoca vitulina is a long muscular roll situated upon the anterior surface
of the thoracic and cervical vertebra. It consists of two parts, an anterior and a posterior. The
(zool. chall. exp — part Lxvin. — 1888.) Yyy 28
218 THE VOYAGE OF H.M.S. CHALLENGER.
posterior arises from the bodies of the 1st to the 7th dorsal vertebrce, from the intervertebral discs,
from the ventral surfaces of the rib joints, and from the body of the last cervical vertebra. It
ascends and lies to the outer side of the rectus capitis anticus major, and is inserted into the
ventral divisions of the cervical transverse processes of the 2nd to the 6th cervical vertebra;, to the
outer side of the origins of the rectus capitis anticus major. The anterior part arises from the
inner sides of the ventral divisions of the transverse processes of the 3rd to the 6th cervical
vertebra;, the origins from the anterior ends being tendinous. It lies to the inner side of the
rectus capitis anticus major, and is inserted into the bodies and intervertebral discs of the vertebra;
reaching to the atlas, the fibres from the 6th going to the 5th, from the 4th to the 3rd, from the 3rd
to the 2nd and 1st cervical vertebra;. It is supplied by branches of the brachial plexus.
In Arctocephalus there are three parts. The posterior oblique arises from the bodies of the 1st
to the 4th dorsal vertebra?, from the intervertebral discs, from the ventral surfaces of the rib
joints, and from the body of the last cervical vertebra; and is inserted into the outer side of the
ventral division of the 6th cervical vertebra. The anterior oblique lies on the outer side of the
rectus capitis anticus major, and arises from the outer sides of the ventral divisions of the transverse
processes of the 3rd, 4th, 5th, and 6th cervical vertebra; ; and is inserted by three slips into the
dorsal tubercles of the 3rd, 4th, and 5th cervical vertebra;, and the outer half of the ventral surface
of the wing of the atlas. The vertical part arises from the inner surface of the ventral divisions
of the transverse processes of the 2nd to the 6th cervical vertebra;. The fibres go forwards, and are
inserted into the hypapophyses of the 2nd to the 6th cervical vertebrse.
In Otaria and Trichechus Dr. Murie describes two parts.
The Lateral Vertebral Region includes three muscles in Phoca vitulina — the scalenus
anticus, medius, and posticus. In Arctocephalus the scalenus medius is wanting.
The Scalenus anticus in Phoca vitulina is a short band of muscle, and arises from the anterior
and outer surface of the 1st rib at its junction with its cartilage, and proceeds forwards to be inserted
into the antero-posteriorly elongated hatchet-shaped ventral divisions of the transverse processes of
the 4th and 5th cervical vertebra; by tendinous slips. A fasciculus from it blends with the tendon
of insertion of the scalenus medius into the 3rd cervical. It is supplied by the branches of the
brachial plexus.
In Arctocephalus it lies between the longus colli ventrally and the serratus posticus dorsally. It
arises from the anterior border of the 3rd rib, anterior to the origin of the digitation of the serratus,
from the same border of the 2nd and 1st ribs, but to the inner side of the digitations of the
serratus. The muscle forms a flattened band, and is inserted into the tip of the ventral division of the
transverse process of the 7th cervical, and into the ventral sides of the dorsal divisions of the
transverse processes of the 3rd, 4th, 5th, and 6th cervical vertebrse.
The Scalenus medius in Phoca vitulina is larger than the last, and arises from the anterior
triangular surface of the 1st rib near its vertebral end, lying behind and a little to the outer side
of the anticus. It is inserted by a tendon into the ventral hatchet-shaped division of the 4th cervical
vertebra, by a fasciculus into the under surface of the tendon of the posticus going to the 4th
cervical vertebra, and into the posterior surface of the tendon of the same muscle going to the
3rd cervical vertebra by the same fasciculus, which is continued forwards from behind the tendon
of the posticus to the 4th vertebra. In the large Phoca vitulina the muscular arrangement
REPORT ON THE SEALS. 219
differs from the above : on the right side it arises from the anterior and outer surface of the 1st rib,
extending from its junction with the cartilage nearly to the inner third of the rib. It runs
forwards and is inserted into the root of the outer side of the ventral division of the transverse
process of the 6th cervical vertebra by muscular fibres, into the posterior end of the ventral
division of the 5th cervical vertebra by a tendinous slip, and likewise into the corresponding part
of the 3rd and 4th cervical vertebra. It is supplied by the brachial plexus.
The Scalenus 2)ostic2is in Phoca vitulina is much the largest of this group, and arises from the
posterior border of the 4th rib at the junction of the cartilage with the rib ; and similarly from
the posterior border and outer surface of the 3rd rib. The digitation of the serratus coming from
the 3rd rib lies between these two points of origin. It is inserted into the ventral hatchet-shaped
division of the transverse processes of the 3rd and 4th cervical vertebrae by two strong tendons.
In the large Phoca vitulina the origin is similar, but the insertion is by tendons into the posterior
of the ventral divisions of the transverse processes of the 3rd and 4th cervical vertebras, these
tendons being common to the scalenus secundus. It is supplied by the brachial plexus.
In ArctocepJuilus it lies in the cervical region and is of the same shape as the anticus, but
smaller. It arises from the ventral tip of the transverse process of the 7th cervical vertebra, and
is inserted into the ventral border of the transverse process of the atlas, dorsal to the anterior
oblique portion of the longus colli, into the dorsal tip of the transverse process of the 3rd cervical,
and into the dorsal divisions of the same processes of the 4th, 5th, and 6th cervical vertebrae.
The Muscles of the Thorax.
In Phoca vitulina we find the sterno-costalis anterior and posterior, internal intercostals, external
intercostals, scalenus lumborum, levatores costarum, and triangularis sterni. In Arctoceplialus the
sterno-costalis posterior is wanting, otherwise the muscles are alike.
The Sterno-costalis anterim' is Lucae's transversus costarum latus and Murie's supracostal. In
Phoca vitulina it lies next the sternum, and is a flat slender layer of muscle which arises from the
side of the presternum and mesosternum as far back as the 6th rib, and from the cartilages of the
1st to the 6th ribs at their junction with the sternum. It remains tendinous to 1 inch from the
side of the sternum, then it becomes muscular, and ultimately divides into three digitations, the 1st
being the longest. It is inserted into the outer surface and cartilage of the 1st rib, extending
outwards to the digitation of the serratus, into the posterior border of the 2nd rib close to the
digitation of the serratus, and into the same part of the 3rd rib.
In Arctoccphalus it is a small triangular muscle, and arises from the sides of the 2nd, 3rd, and 4th
sternebrae, from the cartilages of the 2nd, 3rd, 4th, and 5th ribs by a fine aponeurosis. The fibres
run forwards and outwards, and are inserted into the posterior border of the cartilage of the 1st rib,
and into the 2nd at the junction of the cartilage with the rib.
Lucae figures it attached to the sternum and the ribs, covering the tendon of the rectus (pi.
vi. fig. 1), but describes it as going from the 4th rib to the 1st cartilage and rib, with its tendon
united to that of the rectus.
In Otaria it lies to the sternal side of the scalenus anticus between the cartilages of the 3rd
and the 1st ribs.
In Trichechus it extends from the 4th costal cartilage to the 2nd rib and 1st intercostal space.
220 THE VOYAGE OF H.M.S. CHALLENGER.
The name supracostal given to this muscle by Dr. Murie is confusing, for thus we should
understand that the muscle is the musculus rectus thoracis of Professor Sir Wm. Turner, and the
supracostal of Mr. Wood and others.1 In Phoca the rectus tendon is unbroken and reaches the
cartilage of the 1st rib beneath the sterno-costalis ; in Arctocephalus the rectus ends posterior to the
origin of this muscle, but the fibres of the sterno-costalis anterior, as in Phoca, are obliquely directed
outwards and not antero-posteriorly. In Murie 's dissections it does not touch the sternum, but the
situation and direction of the fibres in his plates are the same as in Phoca and Arctocephalus ; so
I do not regard Murie's supracostal as a part of the rectus which the name he uses indicates.
The Sterno-costalis posterior is the transversus tenuis of Lucae. It is only found in Phoca vitu-
lina, and arises from the mesosternum at its junction with the cartilages between the 3rd and 7th
ribs, and from the tendon of the rectus which passes forwards beneath it. On the right side it
receives a muscular slip from the xiphisternum and then divides into three slips, which are inserted
into the outer surfaces of the 1st, 2nd, and 3rd ribs. The first digitation is fixed outside the origin
of the sterno-thyro-hyoid ; the second and third beside the origins of the serratus, these two being
crossed by the origin of the scalenus posticus. This part may not be quite separate from the
anterior.
The External intercostals in Phoca vitulina commence close to the side of the vertebral column,
and leave only a hollow for the levatores. They arise from the posterior part of the rib joints
by a few fibres, at the outer side of the levatores they thicken, and then pass from the ribs in
front backwards and outwards to the ribs behind. They extend from end to end of each rib.
In Arctocephalus they extend from the heads of the ribs to the middle of the costal cartilages.
The Internal intercostals in Phoca vitulina are quite close to the vertebra?, where the subcostals
are wanting. They arise from the anterior part of the rib joints, and from the anterior borders of the
ribs, and are inserted into the posterior borders of the ribs sternad to their origins, the fibres of
the muscles almost meeting over the ribs. There is an aponeurosis between the two muscles,
strongest near the column and almost disappearing about the middle of the bony ribs. In the last
space the muscle lies only on the outer half of the rib, and begins at the outer edge of the scalenus
lumborum.
In Arctocephalus they arise from the whole anterior borders of the ribs to their necks, and from
the same borders of the cartilages. The fibres are inserted into the posterior borders of the ribs
sternad to their origins, and end at the sternebrae.
The Scalenus lumborum is so named by G. H. Meyer in the human body.2 In Phoca vihdina
it arises from the transverse processes of all the lumbar vertebrae by an aponeurosis, and forms a
triangular muscle with the apex posterior. It is inserted in its outer half into the posterior
border of the outer half of the last rib ; the inner half crosses over this rib and is strengthened
by a few fibres from it ; whilst the outer half of this portion goes into the 14th rib, the inner crosses
over to the 13th rib, and ends on its posterior border.
In Arctocephalus it arises from the transverse processes of the 1st, 2nd, and 3rd lumbar verte-
brae, forming a thin triangular sheet with the base forwards. It is inserted into the last three ribs.
A portion of its fibres from the outer side is attached to each of the last two ribs, the remainder to
the 13th rib, but the proportion in which the fibres are distributed differs from that in Phoca —
1 Sir Wm. Turner, Proc. Roy. Soc. Edin., vol. vi. pp. 268, 270, 1869.
- Lehrbuch der Anatomie, 1855.
REPORT ON THE SEALS. 221
over the ventral surface of the 15th rib the outer third of the muscle terminates, over the 14th
rib the half of the remaining fibres, and over the 13th the rest.
Lucae makes no distinction between the true subcostals and the scalenus lumborum, but
describes both under the name subcostalis vertebralis. The drawing by Meyer shows the scalenus
lumborum terminating at the last rib, but in Phoca and Arctoccplialus it is prolonged forwards to
the under surface of the third last rib. In Otaria and Trichech/as it has not been figured or
described.
The Subcostales in Phoca vitulina usually commence at the 7th or 8 th intercostal space ; there
are six or seven of them, and they lie next the vertebral column. Each arises from the ventral
surface of the rib joint, from the posterior part of the vertebra in front of the one with which the
rib articulates, and from the rib close to the joint. The fibres pass over the rib in front of their
origin, and are inserted into the posterior border of the next. The last two muscles have the
scalenus lumborum along their outer borders.
In Arctocephcdus there are twelve of these ; all arise from the posterior part of the body of a
dorsal vertebra, and are inserted as in Phoca vitulina, having a rib intervening between the origin
and insertion. The muscles are much narrower than in Phoca, and commence in the 3rd intercostal
space.
In Otaria and Trichcchus they are not mentioned.
The Lcvatorcs costarum both in Phoca vitulina and in Arctocephalus are a series of small triangular
muscles, with their bases directed away from the spinal column. There are fifteen on each side of
the back ; they arise from the dorsal tips of the dorsal divisions of the transverse processes of the 7th
cervical and the anterior ten dorsal vertebras, and from the under surfaces of the anapophyses
of the 11th, 12th, 13th, and 14th dorsal vertebrae. The anterior muscles are small and narrow, the
posterior short and broad, and the intermediate much the longest. They are inserted into the dorsal
surface of the rib below the originating point, then into the anterior border of the same rib for a
little distance beyond this. In the large Phoca vitulina I found only fourteen.
The Triangularis stcrni is Lucae's subcostalis sternalis; in Phoca vitulina it lies upon the inner
side of the sternum and covers the internal mammary artery. It arises from the side of the
manubrium reaching as far forwards as the anterior border of the 4th rib, from the junction of the
cartilages with the sternum from the 4th to the 10th ribs, and from the anterior half of the side of
the ensiform cartilage. The muscle is divided into a number of serrations; in this dissection there
were nine on each side, six arising from the manubrium and three from the ensiform cartilage ; those
from the manubrium were much larger than those from the ensiform. Each serration frorn the
manubrium extended from the posterior border of one rib to the back of the next behind. The
posterior fibres are not so obliquely directed forwards and outwards as the rest ; the anterior
fibres are most oblique. The last serration from the ensiform cartilage crosses the cartilage of the
11th rib, and is inserted into the lower border of the cartilage of the 10th rib; the penultinate
crosses the cartilages of the 10th and 11th ribs, and is inserted into the lower and inner surface of
the cartilage of the 9th rib ; the last but two crosses the cartilages of the 11th, 10th, and 9th ribs,
and is inserted into the lower border and inner surface of the 8th cartilage. The other serrations
are inserted into the under and inner surfaces of the cartilages of the 2nd to the 7th ribs, each
serration crossing two ribs before reaching its' attachment.
In Arctocephalus it arises from the anterior half of the sternum by fibres, and from the ventral
222 THE VOYAGE OF H.M.S. CHALLENGER.
sides of the 3rd, 4th, 5th, and 6th sternebrae by a narrow tendon. The fibres pass transversely
outwards posteriorly, and slightly outwards and forwards anteriorly, and terminate laterally 1 inch
to the outer side of the sternum. It is inserted into the cartilages of the 3rd to the 8th ribs, and
into the fascia between them upon the intercostal muscles. These muscles are supplied by the
intercostal or lumbar nerves.
The Muscles of the Abdomen.
In Phoca vitulina and in Arctoccphalus these are the external oblique, internal oblique, trans-
versalis, rectus abdominis, and cremaster. The pyramidalis and quadratus lumborum are wanting.
The Obliquus externus abdominis in Phoca vitidina is much the strongest of the abdominal group.
It has fourteen digitations. The first arises from the outer and posterior surface of the 4th rib, inter-
digitating with the serratus magnus, and, after mingling with a few of the fibres of the first or
anterior head of the scalenus posticus, is partially crossed by part of the same muscle from the 5th
rib. The second digitation from the outer and posterior surface of the 5th rib interdigitates with the
serratus and touches the origin of the posterior slij) of the scalenus posticus, which passes beneath
the digitation of the serratus from this rib. The third to the seventh spring from the outer and
posterior surfaces of the 6th to the 10th ribs, interdigitating with the serratus magnus only. The
eighth to the thirteenth digitations spring from the outer and posterior surfaces of the 10th to the
15th ribs, interdigitating with the latissimus dorsi. The digitation from the last rib is the largest of
the series from ribs. The fourteenth digitation has no bony origin, but comes from the lumbar fascia,
interdigitating with the latissimus dorsi, and is still larger than the thirteenth. The fibres from the
first and second digitations run backwards and inwards and terminate near the inner edge of the
rectus muscle, and are continuous with the fascia covering its posterior surface ; those from the
third and fourth have the same course, but form an aponeurosis near the inner edge of the rectus, and
join the aponeurosis of the other side. From behind the ensiform cartilage to the symphysis pubis
the muscle terminates in an aponeurosis upon the outer border of the rectus, the fibres of one side
decussating with those of the other. The anterior digitations run obliquely backwards and inwards,
the middle more so, and those fibres coming from the posterior ribs and the dorsal fascia are nearly
antero-posteriorly directed. The muscle is thinnest at its anterior end and thickest at its posterior
and dorsal side, and does not thin off at the side of the rectus but ends abruptly as a tendon. The
dorsal border behind the last digitation overlies the lumbar fascia, and is kept in its position by the
fascia over the surface of the muscle passing on to it. The dorsal edge of the muscle runs directly
backwards to a little to the outer side of the patella, whence the posterior border runs obliquely
backwards and inwards to the symphysis in a direct line from the patella. From the posterior
border, midway between the outer edge of the patella and the symphysis, two strong flat tendons
are developed ; the outermost goes behind the cord and is attached to the brim of the pelvis a
little more than 1 inch to the anterior side of the symphysis, its posterior border being posterior
at the attachment. From the anterior border of this a thin triangular aponeurosis is attached
to the brim of the pelvis, extending from the anterior part of the outer pillar to an inch and a
quarter posterior to the pectineal eminence.
In ArctoccpJialus it arises by ten digitations from the posterior and outer surfaces of the 6th to
the 15th ribs, each digitation ending about half an inch from the junction of the ribs with their
REPORT ON THE SEALS. 223
cartilages. The last digitation also has origin from the lumbar fascia; the direction of the fibres is
much the same as in Phoca. As most of the aponeuroses of the trunk had lost distinctness owing to
the length of time the specimen had been in salt, I cannot enter into a description. From
the last digitation the fibres pass directly backwards and reach the ventral anterior spine of the ilium,
and are attached to it. The mesial fibres terminate upon the middle of the rectus muscle in the
anterior two-thirds, and upon its inner edge in the posterior third. From the ventral anterior
spine of the ilium the insertion remains muscular until it reaches the pectineal eminence, where it
becomes tendinous, and is inserted into the pelvic brim, reaching the outer side of the rectus abdominis
and forming the outer pillar of the external abdominal ring. The rest of this aponeurosis crosses
over the cord and terminates upon the origin of the rectus. The five anterior slips interdigitate with
the serratus magnus, the sixth, seventh, and eighth with the latissimus dorsi.
The Obliquus interims abdominis in Phoca vitulina is hidden by the external oblique, and has
a slight resemblance to a quadrilateral figure. It arises by muscular fibres from the ventral border
of the ventral anterior spine of the ilium to the inner side of the origin of the sartorius ; from the
brim of the pelvis by muscular fibres from the front of the attachment of the inner pillar of the
external abdominal ring to the posterior part of the pectineal eminence ; from the lumbar fascia
between the last rib and the crest of the ilium by an aponeurosis; from the inner surfaces and tips of
the cartilages of the 13th, 14th, and 15th ribs; and from the adjacent sides of the 11th and 12th ribs
an inch from their terminations. The fibres of the muscle midway between the anterior ventral
spine of the ilium and the last rib are almost all transverse ; anterior to this they are directed
upwards and inwards, and anteriorly end opposite the posterior end of the xiphisternum. The fibres
are grouped into bands which are closely united, and between these bands the arterial supply for
the abdominal walls penetrates. The muscle ends anteriorly and mesially as an aponeurosis and by
fibres along its dorsal border ; the anterior part of the aponeurosis crosses over the posterior half
of the xiphisternum, and is inserted into the side of its anterior half and into the cartilages of
the 10th and 11th ribs. The two strong broad anterior bands, arising from the lumbar fascia
between which the transversalis partially arises, are inserted into the posterior border of the last
rib, and into the inner surface and tips of the cartilages of the 13th, 14th, and 15th ribs, and by
muscular fibres into the inner surfaces only of the 11th and 12th ribs. Behind the ensiform
cartilage and anterior to the level of the 14th rib the tendinous termination passes behind the
rectus and unites with the internal oblique of the other side ; posterior to this and anterior to the
posterior seventh of the rectus, the tendon passes over the rectus, and unites with the tendon of
the external oblique above it ; and at the posterior seventh of the rectus the muscle ends upon
the rectus by small muscular digitations. The posterior border of the muscle crosses over from the
anterior ventral spine of the ilium to the outer pillar of the external oblique, crossing the middle of
the thigh ; and the fibres rising from the brim of the pelvis turn outwards upon the thigh, and then
curve to the middle line. The most posterior fibres turn over the cord and are attached to its
outer fourth ; and four fasciculi descend upon the testicle, and form the cremastcr, being prolonged
from the inner and curved side of the fibres which arch over the cord.
In Arctocephalus it arises by muscular fibres from the pelvic brim, beginning midway between
the symphysis and the pectineal eminence; the fibres between the anterior part of the pectineal
eminence and the anterior ventral spine of the ilium arise from the transversalis fascia ; between
this spine and the last rib the fibres anterior to the 4th lumbar spine spring from the lumbar fascia ;
224 THE VOYAGE OF H.M.S. CHALLENGER.
posterior to the 4th lumbar vertebra and anterior to the anterior ventral spine it has two fascial
origins, from the dorsal surface of the erector spina?, and from the tips of the transverse processes of
the 5th and 6th lumbar vertebra?. The dorsal border has an oblique direction, sloping gradually
from the 1st lumbar vertebra to the anterior ventral spine. The fibres springing opposite the 1st
lumbar vertebra pass transversely beneath the digitations of the external oblique from the 13th
rib, and are inserted into the posterior border of the 13th rib beneath that muscle; the fibres
from the 2nd lumbar spine are similarly inserted into the 14th rib, and those from the 3rd
lumbar spine like the last into the 15th rib. The greater portion of the last digitation of the
external oblique is blended with the transversalis between the fibres from the 3rd and 4th lumbar
spines. Posterior to the 3rd lumbar vertebra the fibres ascend and are inserted into the posterior
border of the cartilages of the 13th to the 15th ribs and into the 11th and 12th cartilages by fascia.
From the xiphisternum to midway between it and the pubes the fibres end upon the under surface
of the rectus ; posterior to this they turn over its ventral surface to end upon it ; and the fibres
from the pubes arch over the round ligament and also end upon the rectus.
The Transversalis abdominis in Phoca vitulina arises from the transversalis fascia between the
front of the outer pillar and the anterior ventral spine of the ilium, from the lumbar fascia to
the last rib, and from the inner surface of the cartilages of the 10th to the 15th ribs. The fibres are
strongest anteriorly and thinnest posteriorly. It is inserted by muscular fibres into the side of the
anterior surface of the ensiform cartilage, and blends over its posterior half with its fellow. Behind
the ensiform cartilage it forms a tendon which unites with its fellow to the posterior third of the
rectus, behind this it ends on the outer side of the rectus. Near the symphysis it terminates upon
the transversalis fascia, and near the outer side of the same fascia it crosses over the cord near
the internal ring.
In Arctocephalus it arises from the inner surface of the cartilage of the 11th rib in front of the
tip of the 12th rib, from the same part of the 12th to the 15th ribs, from the lumbar fascia coming
from the tips of the transverse processes of the lumbar vertebra? ; and between the anterior ventral
spine and the pectineal eminence it lies on the transversalis fascia. The fibres are far apart,
especially near the pubes, and are inserted into the xiphisternum and the liuea alba; midway
between the xiphisternum and the pubes the fibres gradually shorten, and at the posterior fourth of
the linea alba they just cross the infundibulum.
The Rectus abdominis in Phoca vitulina arises from the symphysis and slightly from the brim of
the pelvis a little anterior to this. It rims anteriorly over the 11th rib and then along the side of
the manubrium. Over the 5th rib it forms a broad thin tendon, which is inserted into the junction
of the cartilages with the manubrium from the 1st to the 5th ribs. On the right side the tendon
of the rectus was prolonged outwards from the junction of the 1st cartilage with the sternum to the
humerus.
In Arctocephalus it is narrow anteriorly, and arises from the symphysis and the adjacent pubic
bar. At the posterior fourth of the muscle it passes between the transversalis and the internal
oblique, and midway between the xiphisternum and the pubes between the internal and external
oblique. It is inserted by tendinous slips into the outer surfaces of the cartilages of the 5th to
the 10th ribs. In Phoca and Arctocephalus the inscriptiones tendinea? are wanting.
This set of muscles is supplied by the ilio-inguinal, ilio-hypogastric, dorsal, and 1st lumbar
nerves.
REPORT ON THE SEALS. 225
The Diaphragm.
In Pkoca vitulina the diaphragm has a costal and a vertebral origin. The costal portion
arises by a broad fleshy slip from the dorsal surface of the xiphisternuni which stretches across
its terminal expansion, from the posterior surface only of the cartilage of the 10th rib, and from
the posterior and inner surfaces of the cartilages of the 11th to the 15th ribs. These costal origins
interdigitate with the transversalis. The vertebral portion arises by two crura, the left crus by
two tendinous slips, the posterior from the ventral surface of the posterior margin of the body of
the 2nd lumbar vertebra, the anterior from the same part of the 1st lumbar ; the posterior slip
joins the outer side of the anterior opposite the middle of the 1st lumbar vertebra. The right
crus also arises by two slips, the posterior from the ventral surface of the posterior part of
the 2nd lumbar vertebra, the anterior from the back of the 1st lumbar exactly opposite the
anterior slip of the opposite side, reaching as far forward as the back of the last dorsal ; this
slip is much larger than the slip of the opposite side. The lumbar artery of the right side has just
sufficient space to pass between the slips, while the corresponding vessel for the left side is in
the same relation to the short anterior slip but far removed from the posterior one. In addition
the left crus also takes origin from both sides of the transverse process of the 1st lumbar vertebra
by tendinous fibres, and both crura from the posterior border of the last rib. It has three openings
which are nearly in the middle line. The one for the vena cava lies a little to the right of the
mesial plane and most anterior, and is in the tendon of the diaphragm, and therefore is surrounded
by fibrous tissue. The middle opening is for the oesophagus and is elliptical, the long diameter
being antero-posterior ; a small portion of the anterior end is tendinous, the rest is muscular. The
third opening gives passage to the aorta, and is also elliptical, the long sides of the ellipse being
parallel with the aorta ; it is formed by the crura of the diaphragm which meet over the aorta,
opposite the posterior part of the last dorsal vertebra, the left crus slightly overlapping the right, but
the fibres not crossing each other. The central tendon is a large V-shaped slip of fibrous tissue.
It begins on each side of the back about the middle of the penultimate rib as two fine fibrous
streaks, which widen as they near the vena cava. These two streaks meet around this vessel, forming
a fibrous ring which fills in the space between the opening for the oesophagus and the vena cava.
On the dorsal margin of the diaphragm there are two V-shaped slips of fibrous tissue let into
its substance on both sides ; the outer arches over the psoas secundus, the inner is between the tip
of the transverse process of the 1st lumbar vertebra and the last rib. The apices of these V's are
directed forwards, the abdominal fascia forming one side of these slips and the pleura the other.
The muscular fibres from the xiphisternuni go straight to the tendon. The costal fibres run
into the front side of the lateral slips of the central tendon, those from the ribs being most
oblicpie, and those next the xiphisternuni most transverse. The dorsal fibres take a straight
course to the posterior side of the tendon ; those from the transverse processes of the lumbar
vertebras pass towards the middle of the last rib, where they reach one side of the central tendon
opposite where the last costal fibres reach its other side ; between these two is one of the fibrous
V's whose base is between the middle of the last rib and the transverse process of the 2nd lumbar
vertebra.
The fibres are thus distributed round the openings : — Around the aorta on the under surface
the crus of the left side slightly overlaps the right ; on the upper surface they touch and run on.
(zool. chall. exp. — pabt Lxvin. — 1888.) Yyy 29
226 THE VOYAGE OF H.M.S. CHALLENGER.
Around the oesophageal opening upon the under surface the fibres from the crura meet at its
posterior part, and pass on to the central tendon on each side of the vena cava ; upon the upper surface
the fibres from the right crus divide and run along each side of it, and end behind the vena cava.
The vena cava is fibrous on the under surface of the diaphragm ; the upper is not, but it receives
a few fibres from the left side.
In Arctoccphalus as in Phoca the diaphragm has a costal and a vertebral origin. The former
arises by fleshy slips from the ensiform sternebra and not from the spade- shaped cartilage attached
to it, posteriorly from the posterior surface of the 8th rib, and from the posterior and inner
surfaces of the 9 th to the 14th ribs. These also interdigitate with the transversalis. The latter
origin consists of two crura, the left crus arises by muscular fibres from the body of the 1st and 2nd
lumbar vertebra; and the disc between, and by tendon from the 2nd and 3rd lumbar vertebras
and the disc between. The right crus is larger than the left, and arises by muscular fibres from
the 1st, 2nd, and 3rd lumbar vertebras and the discs between, and by tendon from the 3rd and 4th
lumbar; the crura expand and form an oval slip which fits into the back of the central tendon.
The tendon is V-shaped, and the crura are attached to its dorsal side. The fibres from the ensiform
cartilage and the ribs pass towards the anterior part of the tendon, those from the 14th rib meet the
central tendon midway between the opening for the vena cava and this rib, and the gap between is
filled in by fibrous tissue. The oesophagus is in the apex of the central tendon, the vena cava to
the right, and the aorta between the crura.
In both specimens the left phrenic nerve pierces the diaphragm half an inch to the right of the
vena cava ; the right goes through the same spot on the other side, which is one and a half inch to
the left of the vena cava, and they supply the muscle.
The Deep Muscles of the Back.
The muscles may be considered in the following groups : — The serratus posticus, splenius,
erector spinas, complexus and transverso-spinales, interspiuales, intertransversales and interzyga-
pophyses, and the short postero-cranio-vertebral muscles.
The Serratus posticus in Phoca vitulina is a very thin muscular band, and arises from the 2nd
to the 5th dorsal vertebras. The fibres course downwards and backwards, and are inserted by a
short aponeurosis into the lower borders of the 5th to the 9th ribs, outside the tendons of the ilio-
costalis. In the large Phoca vitulina it arises by thin tendons from the same vertebras and from
the intervals between the vertebras by thin aponeuroses, and is inserted into the posterior borders
of the 6th to the 10th ribs by fine tendons. As it is not mentioned by Murie in Otaria and
Trichechus, I conclude it is absent. It is supplied by the external branches of the dorsal spinal
nerves.
The Splenius in Phoca vitulina, is not a double muscle as in human anatomy, neither is it strap-
shaped, but triangular. It is hidden by the cephalo-humeral, rhomboideus-capitis and cervicis,
and arises from the ligamentum nuchas, by muscular fibres in its posterior part, and by a fine
aponeurosis in its anterior. It extends posteriorly from where the fibres of the rhomboideus
cervicis begin to take a transverse course from the middle line of the neck, which is about one
inch anterior to the vertebral anterior angle of the scapula, and terminates anteriorly at the back
REPORT ON THE SEALS. 227
of the interparietal bone. A small fasciculus is continuous with the rhomboideus cervicis. The
ill ires course outwards and forwards, and are inserted partly by muscular fibres and partly by
aponeurosis into the occipital ridge. It is supplied by the external division of the great occipital
nerve, by a branch of the external division of the 3rd cervical, and by a branch from the suboccipital.
In Arctocephahis it has the same shape and relations as in Phocn vitulina, and arises from the
ligamentum nucha?, the 7th cervical vertebra, and the three anterior dorsal spines, and is inserted into
the occipital ridge from the posterior termination of the sagittal suture, to the posterior margin
of the external auditory meatus. It is blended with the trachelo-mastoid near its insertion. In
Otaria and Trichechus a splenius capitis and colli are described. The former is the same as the
splenius in Arctocephahis, and the splenius colli is the trachelo-mastoid in Arcloccphalus.
The Erector spinm in Phoca vitulina divides into the sacro-lumbalis, longissimus dorsi, trans-
versalis colli, and trachelo-mastoid. In Arctocephahis, in addition to the above, the spinalis dorsi
and colli are found.
In Phoca vitulina it lies between the caudal region and the last rib. Its aponeurosis extends
from the sacral to the dorsal region as far as the 14th dorsal spine, crosses over the multifidus,
forming its dorsal covering, and ends laterally on the middle of the dorsum of the erector spinas
posterior to the last rib. It is a massive roll of muscle anterior to the ilium, but posterior to this
is in two small but distinct parts, corresponding to the sacro-lumbalis and the longissimus dorsi.
Anterior to the ilium there is an indication of the existence of two muscles, for a partial fibrous
partition is found running for a short distance into the fibres from the anterior dorsal surface of the
sacrum. The erector arises in two parts, the division ultimately forming the longissimus dorsi from
the rudimentary zygapophyses of the 3rd, 4th, 5th, and 6th caudal vertebras, from the transverse
process of the 1st caudal, from the ligamentous structures covering the dorsum of the sacrum
between the zygapophyses and the transverse processes, and from the ligamentous partition on each
side. It runs forward as the erector spins, lying next the zygapophyses of the lumbar vertebra?,
and turns over the last rib. The division joining the fibres of the erector, and forming the
sacro-lumbalis, arises from the transverse processes of the 1st, 2nd, and 3rd caudal vertebras, and
from the zygapophyses of the 1st and 2nd caudal vertebra?. This origin is bound to the outer side
of the posterior sacro-iliac ligament, and runs into the erector anterior to the ilium, forming the
partial septum already mentioned. This structure is supplemented by a tendon from the dorsal
surface of the ilium, and by fibres from its anterior surface, from the same surface of the sacrum, and
from the ligament between them, the three last origins being ventral to the septum. The
longissimus portion of the erector from the caudal region is inserted into the dorsal posterior borders
of the anterior zygapophyses of the 2nd, 3rd, 4th, and 5th lumbar vertebras, into the anapophyses of
the last dorsal and 1st lumbar vertebra', into the outer surface of the lumbar vertebras, by muscular
fibres extending from the dorsal tips of the anterior zygapophyses to midway between these and the
tranverse processes, and into the inner posterior third of the last rib. The sacro-lumbalis portion
is inserted into the ventral halves of the outer surfaces of the lumbar vertebras, into the outer sur-
faces of their transverse processes, and into the outer two-thirds of the posterior border of the last
rib to within 1 inch from its outer end. The erector is under cover of the lumbar fascia.
In Arctocephahis the longissimus dorsi is not separable from the sacro-lumbalis as in Phoca,
yet the formation of the two is partly evident in the lumbar region. The erector arises in the
sacral region by an aponeurosis, from the spines of some of the caudal and all the sacral vertebrae,
228 THE VOYAGE OF H.M.S. CHALLENGER.
which passes forwards, and ends opposite the 8th dorsal vertebra, and covers the whole of the
longissimus division in the lumbar region, and 1 inch of the sacro-lumbalis division next the
ilium ; also from the neural spines and zygapophyses of the caudal vertebra;, from the neural
spines, zygapophyses, and transverse processes of all the sacral vertebra?, from the inner surface of
the ilium, from the anterior surface of the sacrum, from the ligament between the ilium and sacrum,
from all the lumbar vertebra? between the neural spines and the zygapophyses, and from the sides
of these vertebras between the zygapophyses and the ventral tips of the transverse processes.
The Eio-costcdis or Sacro-lumbalis in Phoca vitulina is the outer division of the erector spina?.
It is a long band running along the back, broadest and strongest at the posterior end, narrowest
and tendinous at the anterior. Along its outer margin is a series of serrations, the seven posterior being
muscular, the nine anterior tendinous. It is chiefly adherent to the outer surfaces of the ribs over
which it lies, especially along their posterior and anterior borders. It is inserted by the digitations
along its outer edge into the outer surfaces of the posterior seven ribs by muscular fibres, into the
posterior borders of the anterior seven ribs by tendinous slips, and into the dorsal tubercle of the
transverse process of the 7th cervical vertebra. In the large Phoca vitulina it also arises by
tendinous slips from the angles of the 14th to the 5th ribs on their anterior borders. It is supplied
by the posterior primary division of the spinal nerves.
In Arctocejthahis it is an offshoot from the erector spina?, and its origin can be partly traced
to 1 inch anterior to the crest of the ilium. It is very narrow posteriorly, and expands gradually
as it approaches the last rib, where it covers its inner two-thirds, while the anterior two-thirds of
the muscle is narrow and tendinous on the dorsal surface, and terminates by giving off long tendinous
slips. It is inserted into the dorsal surfaces of the 6 th to the 15th ribs. Along its outer margin
it has a number of tendinous digitations which are long at the anterior end, and short posteriorly.
The most anterior goes into the dorsal tip of the transverse process of the 7th cervical, the rest
pass to the dorsal borders of the 1st to the 12th ribs. The posterior slips are half tendinous and
half muscular.
The Musculus acccssorius ad ilio-costalcm and the Ccrviccdis ascendens are wanting in both Phoca
vitulina and Ar otocephalus.
The Longissinms dorsi in Phoca vihdina lies to the inner side of the sacro-lumbalis, but is not
quite so large or long, and is under cover of the fascia lunibo-dorsalis. It arises from the under
surface of this fascia out of the erector spina?, from the dorsal tips of the zygapophyses of some
of the lumbar vertebra?, and from the anterior zygapophyses of all the dorsal vertebra?. It lies
along the outer side of the zygapophyses, and is inserted by muscular fibres into the outer surfaces
of the posterior five ribs, by tendons into the anapophyses of some of the dorsal vertebra? (posterior
six), into the dorsal tips of the transverse processes of all the other dorsal vertebra?, which are
homologous to the anapophyses, into the dorsal division of the transverse process of the 7th cervical
vertebra, and by tendinous slips along its inner border into the dorsal surfaces of the anterior
borders of the 6th to the 11th ribs. From the anterior zygapophyses of the 11th and 12th dorsal
vertebra? two strong tendons arise, which divide equally between the multifidus and this muscle.
Opposite the 6th rib a large piece of this muscle goes into the transversalis cervicis. In the large
Phoca vitulina it is inserted in addition into the 9th. 10th, and 11th ribs on their posterior surfaces.
In Arctoccphalus it is a long narrow band covered by dense fascia in its posterior half. This
division of the erector runs into the neck, giving off a number of serrations from its under surface, the
REPORT ON THE SEALS. 229
anterior ones long and chiefly tendinous, the posterior more muscular. They are inserted into the
posterior borders of the 11th to the 15th ribs, into the zygapophyses and anapophyses of the 11th
to the 15th vertebras, into the anapophyses of the 1st to the 10th dorsal vertebrae, and into the
tips of the dorsal tubercles of the transverse processes of the 3rd to the 7th cervical vertebrae.
Some fibres of this muscle seem to take origin along the internal borders of the vertebrae.
The Transversalis ecrvicis or colli in Phoca vitulina is in part a continuation of the longissimus
dorsi, and branches off from it at the 6th rib. It also arises by muscular fibres from the dorsal
surfaces of the transverse processes of the 1st to the 5th dorsal vertebras, to the inner side of the
tendons of insertion of the longissimus dorsi into the same processes, which if developed would be
the anapophyses, and from the anterior halves of the anterior zygapophyses of the 5th, 6th, and 7th
cervical vertebras and the 1st dorsal. It forms in conjunction with the part coming out of the
longissimus dorsi a muscular band which advances towards the cervical region, and there lies
between the digitations of the serratus magnus on its outer side and the trachelo-mastoid on its
inner, and is inserted by five tendinous slips into the dorsal tubercles or divisions of the transverse
processes of the 3rd to the 7th cervical vertebrae. In the large Phoca vitulina it arises in addition
from the transverse processes of the 6th to the 10th dorsal vertebras.
In Arctoccphalus it is a prolongation of the longissimus, and can only be regarded as the
part of the longissimus which lies in the region of the neck and is described with it, because there
is no slip given off from it as in Phoca.
The Trachelo-mastoid in Phoca vitidina lies between the complexus and the transversalis cervicis.
It arises from the anterior and posterior zygapophyses of the 3rd to the 7th cervical vertebras, and
the surfaces of the vertebras between the articular surfaces. It divides into two parts ; the portion
arising from the 3rd, 4th, and 5th vertebras is inserted into the posterior edge of the transverse process
of the axis ; the rest goes to the cranium, and is inserted into the mastoid process. It is supplied
by the external division of the 2nd and 3rd cervical nerves.
In Arctoccphalus it is called by Murie splenius colli, and arises from the 3rd, 4th, 5th, 6th, and
7th cervical vertebras between the posterior zygapophyses and the hyperapophyses, and from the dorsal
surfaces of the laminas between each vertebra, and from the sides of the roots of the 1st, 2nd, 3rd,
and 4th dorsal spines. The last two cervical origins blend with the complexus. It courses anteriorly
below the outer border of the splenius, and is inserted by a narrow tendon into the inferior part of the
occipital ridge behind the external auditory meatus posterior to the insertion of the splenius.
The Spinalis dorsi is only found in Arctocephalus. It lies between the neural spines and the
longissimus dorsi; the dorsal surface is muscular as far back as the 12th rib, from this to the 14th
rili it is tendinous and appears continuous with the longissimus; but, by scraping away the muscular
fibres which arise from the neural spines, a set of tendons is reached which appears to be the
aponeurosis of the longissimus, but can with a little care be parted from it. It arises by muscular
fibres and by long tendinous slips ; the fibres spring from the sides of the neural spines from the
9th to the 12th dorsal vertebras, the long tendinous slips from the metapophyses of the 11th to the
14th dorsal vertebras. The muscular fibres are sparse and thin at the posterior extremity, but
deepen and expand anteriorly; the tendons of origin lose themselves on the under surface of the
muscle. It is inserted by small tendinous slips into the sides and tips of the neural spines from
the 1st to the 8th dorsal vertebras ; but the 1st, 2nd, and 3rd vertebras also have fibres inserted
into them. A large muscular slip is continued into the cervical region, forming the spinalis colli.
230 THE VOYAGE. OF H.M.S. CHALLENGER.
The Spinalis colli in Arctocephalus branches off from the spinalis dorsi between the 2nd and
3rd dorsal vertebrae, passes forwards, receives additional fibres from the posterior cervical spine,
surmounts them, and ends on the neural spine of the axis. It is inserted into the anterior cervical
spines as far forwards as the axis.
The Complexus in Phoca vitulina lies above the rectus capitis posticus major, and beneath the
splenitis capitis and trachelo-mastoid. It is a fleshy band, and arises from the zygapophyses of the
3rd to the 7th cervical vertebrae by fleshy digitations. The fibres proceed to the occipital region, and
are inserted by tendon in its inner three-fourths and by muscular fibres in its outer fourth into
the back of the interparietal element of the occipital bone, and into the occipital ridge, reaching
nearly to the root of the zygoma. It is supplied by the internal branches of all the cervical, and
a branch of the suboccipital nerve.
In Arctocephalus it arises from the hyperapophyses of the 3rd, 4th, 5th, 6th, and 7th cervical
vertebras, and the metapophysis of the 3rd cervical. It courses forwards upon the cervical laniinas,
and is inserted by tendon into the occipital ridge to the outer side of the biventer cervicis.
The Biventer cervicis in Phoca vitulina arises by fibres from the anterior zygapophysis of the
7th cervical vertebra, from the posterior zygapophysis of the same, from the anterior and posterior
surfaces of the 1st dorsal vertebra, and from the anterior part of the 2nd dorsal vertebra. There
are three digitations of origin; at the spine of the axis it unites with the inner border of the
complexus and is inserted with it. In the large Phoca vitulina it arises from the zygapophyses of the
1st to the 4th dorsal vertebras, and joins the complexus opposite the spine of the axis. In a small
male Phoca vitulina it was absent. The nerve supply is the same as that of the complexus.
In Arctocephalus it is long and riband-like. It arises from the roots and sides of the neural spines
of the 2nd, 3rd, and 4th dorsal vertebrae to the inner side of the trachelo-mastoid, with which it is
blended at its origin. It passes forwards to the inner edge of the complexus, and is inserted by
fibres into the occipital ridge, between the complexus on its outer side and the sagittal suture
on its inner.
The Oblique Rotator Muscle of the Spincd Column} — This muscle is in two layers in Phoca vitulina
and in Arctocephalus, and lies between the neural spines and the zygapophyses. These layers are
of a totally different formation in these animals. In Phoca the superficial layer is an extensive
muscular bundle extending from the caudal to the cervical region, and the deeper layer forms a set
of triangular imbricated muscles. In Arctocephalus the superficial layer resembles the deeper layer
in Phoca, and the deep layer is similar of the rotatores muscles in human anatomy.
The superficial layer of fibres of the oblicpue rotator in Phoca vitulina lies in the hollow
between the neural spines and the zygapophyses, stretching from the caudal region into the
cervical under cover of the lumbo-dorsal fascia. In the caudal region the origins are tendinous
slips ; the first slip arises from the rudimentary zygapophyses of the 4th and 5th caudal vertebras
and the dorsal surfaces of the laminas between them, and is inserted into the side of the neural
spine of the 4th sacral vertebra, the corresponding parts of the 1st, 2nd, and 3rd caudal, and also
into the posterior part of the zygapophyses of the 3rd caudal ; the second slip arises from the
zygapophysis of the 3rd caudal vertebra, and is inserted into the sides of the neural spines of the
2nd and 3rd sacral vertebra3, and the laminae between the 2nd and 3rd, and 3rd and 4th sacral
vertebras ; the third slip arises from the zygapophysis of the 2nd caudal vertebra, and is inserted
1 See for the use of this term Sir Win. Turner's Introduction to Human Anatomy, revised edition, p. 76, 1882.
REPORT ON THE SEALS. 231
into the sacral portion of the multifidus. In the sacral region they arise by a tendinous slip from
the zygapophysis of the 4th sacral vertebra, from the zygapophysis of the 3rd sacral by a very short
tendinous slip, from the dorsal surface of the sacrum between the neural spines and the zygapo-
physes, and from the fascia covering it. The mass thus formed and strengthened by the caudal
slip joins anteriorly the lumbar portion, and is inserted by muscular fibres into the neural spines
and the contiguous dorsal surfaces of the sacral laminae, and by tendinous slips into the neural
spines of the 4th and 5th lumbar vertebra;. In the lumbar region the fibres arise by broad
tendinous slips from the anterior zygapophyses of the 1st sacral and all the lumbar vertebrae,
from the posterior zygapophyses and dorsal surfaces of the laminas of all the lumbar vertebra?,
and from the fibrous covering above the muscles. These fibrous slips from the anterior zygapo-
physes join the muscle on the under surface of its outer border. They are inserted by tendinous
slips along its inner border into the posterior aspect of the dorsal tips of the neural spines of the
12th to the 15th dorsal vertebras, into the corresponding part of the 1st, 2nd, and 3rd lumbar
vertebras, and by muscular fibres into the sides of the neural spines of the same vertebras. In
the dorsal region the fibres arise from the metapophyses of the 8th to the 15th dorsal vertebrae,
from the dorsal surfaces of the laminae, as far forwards as the 8th or 9th dorsal vertebra, and are
inserted in the same way as the lumbar portion, into the 1st to the 11th dorsal vertebrae. From
the 10th to the 15th dorsal spines the fibres of the superficial layer are intimately connected with
the longissimus dorsi, and have origin from the under surface of the dorso-lumbar fascia, as far-
forwards as the 14th dorsal vertebra. The tendons of origin from the 6th to the 15th vertebrae
are shared between the longissimus and the multifidus ; the cervical portion is an offshoot from the
dorsal about the level of the 8th rib, and forms long slips which are inserted into the neural spines
of the 3rd to the 7th cervical vertebrae.
The deep layer of fibres of the oblique rotator in Phoca vitulina. The first muscle arises from
the 3rd to the 5th cervical vertebrae from the posterior zygapophyses, and is inserted into the
posterior zygapophysis of the axis. The second arises from the posterior zygapophyses of the 4th
to the 6th cervical vertebrae to the inner side of the 7th, and is inserted into the posterior border
of the lamina of the 3rd cervical vertebra. The third arises from the same parts of the 5th to
the 7th, and is inserted into the corresponding part of the 4th cervical vertebra. The fourth arises
from the laminae of the 6th and 7th cervical and the 1st dorsal vertebra3, and is inserted into the
dorsal surface of the same portion of the 5th cervical vertebra. The fifth arises from the posterior
zygapophysis of the 1st dorsal vertebra, from the dorsal surface of its lamina, and from the dorsal
surface of the lamina of the 7th cervical, and is inserted into the dorsal side of the spine of the 6th
cervical vertebra. The sixth arises from the transverse process of the 2nd dorsal, from the posterior
zygapophysis of the 1st dorsal, and is inserted into the spine of the 7th cervical. In the dorsal
region they are not so oblique as in the cervical region. One arises from the zygapophyses of the
2nd to the 4th dorsal vertebrae, and is inserted into the spine and lamina of the 2nd dorsal vertebra,
and so on to the 10th dorsal vertebra, the last muscle only having origin from one vertebra. The
cervical muscles are all oblique, the dorsal are oblique and transverse, and all are imbricated.
'The superficial layer of the oblique rotator muscles in Arctoceplialus is well developed. The
cervical muscles are much longer than the dorsal, the first and most anterior a/rises by three slips,
from the dorsal surface of the lamina and the posterior zygapophysis of the 3rd cervical
vertebra, from the hyperapophysis of the 4th cervical, and from the same part of the 5th cervical
232 THE VOYAGE OF H.M.S. CHALLENGER.
vertebra. It is inserted into the back of the hatchet-shaped neural spine of the axis, into the
posterior border of its lamina, and into the neural spine of the 3rd cervical vertebra. The second
arises from the hyperapophyses of the 5th, 6th, and 7th cervical, ascends, and is inserted into the
neural spines of the 4th and 5th cervical. The third arises from the hyperapophysis of the 7th
cervical and the posterior zygapophysis of the 1st dorsal vertebra, and is inserted into the back
of the neural spine of the 5th and the neural spine of the 6th cervical. The dorsal muscles
are well marked in the anterior dorsal region, but are feeble near the lumbar, for the space between
the transverse processes and the neural spines narrows considerably near the lumbar region. They
are all formed on the same plan, and arise from the inner sides of the transverse processes of these
vertebrae, and are inserted into the side of the neural spine opposite the most anterior vertebra
from which they spring.
The deep layer of the oblique rotators in Arctocephalus arises from the anterior of the transverse
processes, passes inwards, and is inserted into the roots of the neural spines and posterior borders
of the lamina? of the vertebra? anterior to their origin.
The erector spinas, sacro-lumbalis, longissimus dorsi, transversalis colli, trachelo-mastoid, and
spinalis dorsi are supplied by the external branches of the posterior divisions of the cervical, dorsal,
and lumbar nerves. The rotator muscles and spinalis colli by the internal branches of the posterior
divisions of the cervical, dorsal, and lumbar nerves.
The Supraspinales were not seen in Phoca vitulina or Arctocephalus.
The Intcrspinales in Phoca vitulina and in Arctocephalus are most distinct in the cervical
region.
The Intcrtransvcrsalcs in Phoca vitulina and in Arctocephalus are found in the cervical and
dorsal regions. In the latter they are between the dorsal divisions of the transverse processes.
The Intcrzygapophyscs in Phoca vitulina and in Arctocephalus, in the lumbar region, are strong
muscular slips. The supraspinales and interspinales are supplied by the internal divisions of the
cervical, dorsal, and lumbar nerves. The intertransversales and interzygapophyses by the external
divisions of the cervical, dorsal, and lumbar nerves.
The Rectus capitis posticus major in Phoca vitulina is riband-shaped and arises from the posterior
three-fourths of the side of the tip of the neural spine of the axis by muscular fibres. It courses
forward, and is inserted by a short tendon into the lambdoidal suture between the rectus capitis
posticus minor and the rectus capitis posticus major accessorius behind the occipital ridge beneath
the complexus. It is supplied by the suboccipital nerve.
In Arctocephalus it arises from the anterior two-thirds of the side of the hatchet-shaped neural
spine of the axis, and slightly from the adjoining surface. It is inserted into the lambdoidal suture.
The Rectus capitis p>osticus major accessorius is present in all the specimens. It is a narrow mus-
cular slip arising by muscular fibres from the anterior fourth of the side of the tip of the neural
spine of the axis. This small bundle takes a turn outwards and ascends to the occiput between the
rectus capitis posticus major and minor. It is inserted into the occipital bone a little posterior to
the lambdoidal suture, to the inner edge of the rectus capitis posticus major, between the insertion
of the rectus capitis posticus minor and the insertion of the superior oblique, and slightly into
the back of the condyle of the occipital bone, posterior to the major. It is supplied by the
suboccipital nerve.
In Arctocephalus it arises from the anterior and under surface of the neural spine of the axis, and
REPORT ON THE SEALS. 233
is inserted into the occipital bone posterior to the insertion of the complexus, and between the
superior oblicpae and the rectus capitis posticus minor.
The Rectus capitis posticus minor in Phoca vitidina is nearly rectangular, and its anterior end
is the broader. It arises from the tubercle on the tip of the neural arch of the atlas, and from the
thin dorsal surface of its lamina anterior to the foramen. It ascends, and is inserted into the
whole of the surface of the occipital bone, behind the insertion of tne major rectus accessorius in
front of the foramen magnum, and as far out as the inner side of the condyle of the occipital bone.
This insertion is extensive. It is supplied by the suboccipital nerve.
In Arctoeephalus it arises from the anterior dorsal half of the atlas, between the neural spine and
its foramen and the articular surface for the occipital condyle. It is inserted into the supraoccipital
bone, posterior to the biventer cervicis internally, and the rectus capitis anticus major externally.
It is bounded by the rectus capitis posticus and the complexus, and posteriorly by the foramen
magnum.
The OUiquus capitis inferior in Phoca vitidina is a short rectangular muscle, and arises from the
side of the neural spine of the axis beneath the major and accessorius muscles, from the whole of
the dorsal surface of its lamina, and slightly from the dorsal surface of the posterior zygapophysis
of this vertebra. It passes outwards and forwards, and is inserted into the concave posterior surface
of the transverse process of the atlas ventral to its foramina. It is supplied by the suboccipital and
the great occipital nerves.
In Arctoeephalus it arises from the outer side of the neural spine of the axis, and the dorsal
surface of the lamina to the inner side of the hyperapophysis. It is inserted into the concave sur-
face on the posterior dorsal half of the wing-like transverse process of the atlas.
The Obliquus capitis superior in Phoca vitidina is the same shape as the last ; it arises from the
dorsal anterior surface of the condyle of the atlas, and from the dorsal edge of the transverse pro-
cess of the same. It is inserted into the middle of the occipital ridge between the rectus capitis
posticus major, and the rectus lateralis beneath the complexus. It is supplied by the suboccipital
nerve.
In Arctoeephalus it arises from the anterior surface of the atlas outside the foramen, and is
inserted into the lower half of the occipital ridge, into the upper half of the paramastoid process,
and the exoccipital bone.
The Muscles of the Tail.
I have only observed one muscle arising from the caudal region in Phoca and in Arctoeephalus.
This is named in the text of Lucae the abductor caudae, while Murie calls it the levator caudaa
externus. The levator caudas of Lucae, and the levator caudse internus of Murie, are simply prolon-
gations backwards of the erector muscles of the back into the caudal region, and are described as
part of these muscles. The ventrales caudae of Lucae are the same as the pubo-, ilio-, sacro-, and
infra-coccygeus of Murie, and are included in my description of the levator ani.
The Abductor caudse in Phoca vitidina arises from the dorsal surface of the dorsal sacro-iliac
ligament, and from the under surface of the transverse processes of all the sacral vertebra?, and is
inserted into the same parts of the caudal vertebrae. It is supplied by the caudal nerves.
In Arctoeephalus it arises from the dorsal anterior spine of the ilium, from the dorsal surface of
(ZOOL. CHALL. EXP. — PART LXVIII. — 1888.) Yyy 30
234 THE VOYAGE OF H.M.S. CHALLENGER.
the dorsal sacro-iliac ligament, and from the transverse processes of all the sacral vertebrae, and is
inserted into the transverse processes of all the caudal vertebra, partly by tendon and partly by
muscular fibres.
Some of the Perineal Muscles.
The Sphincter ani in the female Phocinse and the female Arctocephcdus is a broad, strong band.
It arises from the ventral mesial caudal region, and encircles the posterior end of the rectum and
vagina, but in the male it only winds round the rectum.
The Levator ani in Plwca vitidina is a triangular muscle. It arises from the anterior inner
wall of the pubes, ending half an inch anterior to the posterior margin of the obturator foramen ;
from one inch of the pubic bar at the anterior margin of the obturator foramen ; between these two
points of origin from the obturator fascia close to the ventral margin of the obturator foramen :
from the internal surface of the innominate bone dorsal to the obturator nerve, and posterior to the
sacro-iliac articidation ; and from the lateral and ventral surfaces of the 2nd, 3rd, and 4th sacral, and
1st and 2nd caudal vertebra?. It forms several tendons and these proceed backwards ; the inner-
most is inserted into the ventral mesial surface of the 5th caudal vertebra, the outermost into the
ventral surface of the transverse process of the same vertebra, and the other three tendons into the
ventral surfaces of the last caudal vertebra by passing backwards between the other two insertions.
From the middle of the 2nd caudal vertebra to the back of the 4th sacral, many fibres pass
down around the rectum and vagina, and proceed backwards beneath the sphincter for the vagina
and rectum. The combined levators form a funnel-shaped tube which passes through the pelvic
outlet surrounding the rectum and vagina.
In Arctocephalus it arises from the inner surface of the pubic bar above the pelvic brim, between
the pectineal eminence anteriorly and the side of the symphysis posteriorly; from the inner surface
of the ilium anterior to the obturator foramen ; and from the ventral sides of the sacral and caudal
vertebrae. The levators form a muscular tube as in Phoca, the posterior pubic fibres proceed back-
wards and encircle the vagina and rectum, but principally the former. Then they turn inwards
upon the ventral side of the vagina, and end posteriorly among the fibres of the sphincter for the
vagina and rectum outside the pelvis. The rest of the fibres run backwards along the caudal
region, and are inserted into the ventral surfaces of the transverse processes of the caudal vertebrae.
The Protractor of the prepuce, hi the Phoeinre between the symphysis pubes and the umbilicus, is
a muscular band arising from the outer border of the rectus by three slips. These soon unite,
proceed backwards, and are inserted into the side of the prepuce around the orifice, the fibres of
both muscles meeting on its ventral surface.
The Retractor vagina? in Phoca is a quadrilateral muscle. It arises half an inch ventral to the
ischial spine from the posterior borders of the ischium and pubes, and descends upon the side of the
vagina, being dorsally blended with the levator ani.
INDEX TO REPORTS ON MARINE MAMMALS.
This Index refers both to the Keport on the Bones of Cetacea (Zool. Chall. Exp., part iv., 1880) and to the
Keport on the Seals. The references to the Keport on the Bones of the Cetacea are distinguished by
the Eonian numeral i. ; the references to the Seals by the Konian numeral ii.
Alactherium, ii. 69.
Allen, J. A., on Macrorhinus leoninus, ii. 5, 19.
on Otariidee, ii. 35, 73.
on Trichechus, ii. 69, 71.
Anterior extremity, of Arctocephalus australis, ii. 45.
Leptonychotes weddelli, ii. 25.
Macrorhinus leoninus, ii. 15.
Muscles of, iL 142.
Apes, brain of, compared with Seals, ii. 121.
Arctocephalus, characters of, ii. 82.
vertebra? of, ii. 24.
argentatus, ii. 54, 87.
australis, described, ii. 39.
compared with A. ursinus, ii. 86.
delalandii, ii. 84.
elegans, ii. 36, 87.
forsteri, characters of, ii. 87.
gazella, described, ii. 36, 83.
gillespii, ii. 77.
gracilis, ii. 83.
hookeri, ii. 78.
lobatus, ii. 63.
monteriensis, ii. 76.
nigrescens, ii. 82, 88.
philippii, characters of, ii. 87.
pup referred to, ii. 54.
pusillus, characters of, ii. 84.
schisthyperoes, ii. 85.
sp. incerta, described, ii. 52.
ursinus, characters of, ii. 86.
pup of, ii. 52.
size of, ii. 52.
Arctophoca falclandica, ii. 82, 83.
gazella, ii. 83.
Badger, brain of, ii. 118.
Balsaia antipodarum, i. 33.
australis, bones of, i. 32, 40
lalandii. See Megaptera, i. 30.
Balsenoptera antarctica, ear-bone, i. 34.
huttoni, ear-bone, i. 35.
rostrata, ear-bone, i. 35.
sibhaldii, ear-bone, i. 34.
Baleenopteridffi, ear-bones, i. 35.
Ballast bag of Seals, ii. 136.
Bardeleben, K., on scaphoid bone, ii. 50.
Bear, Brown, brain of, ii. 131.
Polar, brain of, ii. 117.
Beluga catodon, stones in stomach of, ii. 137.
Berardius arnouxii, teeth, i. 21.
Bibliography of —
Brain of Carnivores, ii. 90.
Myology of Seals, ii. 139.
Black, A. M., specimen presented by, i. 2.
Bonner, John, specimen presented by, i. 2.
Brain, comparison of convolutions of, ii. 113.
Nomenclature of convolutions of, ii. 96.
of Elephant Seal, ii. 91.
of "Walrus, ii. 102.
Broca, Paul, on Brain of Primates, ii. 126.
on cerebral nomenclature, ii. 96.
Buenos Ayres, Epiodon australis from, i. 7.
Burmeister, H., on Epiodon, i. 27.
on Otariidse, ii. 73, 83.
Callidon giintheri ( — Mesopdodon laijardi), i. 3.
Callocephalus vitalinus, ii. 58.
Callorhinus ursinus, pup of, ii. 53.
size of, ii. 52, 86.
Cape of Good Hope, specimen, i. 3.
Carcharodon, teeth from deep sea, i. 42.
Cat, brain of, ii. 120.
236
THE VOYAGE OF H.M.S. CHALLENGER.
Cerebellum of Elephant Seal, ii. 100.
Walrus, ii. 111.
Chatham Islands, Cetacean from, i. 3, 27.
Chincha Islands, Seal from, ii. 30.
Clark, J. W., on Eumetopias hookeri and E. cinereus, ii.
78, 80.
on Arctocephalus schisthypierdes, ii. 85.
Classification of Pinnipedia, ii. 55.
Coati, brain of, ii. 119.
Convolutionsof Brain in various groups compared, ii. 1 1 3.
Crozets, Seals from, ii. 3, 36.
Cunningham, D. J., on muscles of pes, ii. 208.
Cunningham, R. O., skulls collected by, ii. 30, 41.
Cuvier, F., on Macrorhinus leoninus, ii. 5.
on Megaptera lalandi, i. 30.
Cystophora, characters of, ii. 67.
compared with Macrorhinus, ii. 69.
with Ommatophoca, ii. 66.
crisfata, characters of, ii. 68.
uterus of, ii. 136.
CystophorinEe, characters of, ii. 67.
Dean, Mr., presented a specimen, ii. 29.
Delphinus, ear-bones, i. 37, 39, 40.
Distribution, Geographical — Mesoplodon layardi, i. 3.
Ziphioids, i. 39.
Dolichodon layardii ( = Mesoplodon), i. 3.
Doran, A. G., on auditory ossicles, ii. 11.
Ear-bones — Balxnoptera, i. 34.
Cetacean, i. 33.
MegapAera, i. 31.
Mesoplodon and Ziphius, i. 8.
Eared Seals, ii. 29, 73.
Ehlers on Epiphysis cerebri of Plagiostomata, ii. 109.
Elephant Seal, ii. 3, 69.
Elliott, H. E., on Alaska Seals, ii. 52, 53, 76.
on stones in stomach, ii. 137.
Entoscaphoid bone, ii. 50.
Epiodon australis = Ziphius cavirostris, i. 27, 39.
chathamiensis, specimen of, i. 11.
Hector on, i. 27.
identical with Ziphius cavirostris, i. 29.
novx-zealandiee = Ziphius cavirostris, i. 29.
Erignathus barbatus, ii. 60.
Eubalxna austral is, i. 33.
Eumetopias, characters of, ii. 76.
californianus, characters of, ii. 77.
cinereus, characters of, ii. 79.
tarsal bones of, ii. 50.
hookeri, characters of, ii. 78.
lobatus, ii. 81.
Eumetopias stelleri, characters of, ii. 76.
Euotaria nigvescens, ii. 41, 82.
External characters of —
Arctocephalus australis, ii. 39.
Macrorhinus leoninus, ii. 3.
brain of, ii. 91.
Otaria jubata, ii. 29.
Extremities, bones of anterior, ii. 15, 25, 45.
posterior, ii. 17, 26, 48.
Falkland Islands, Whale from, i. 2.
Seals from, ii. 29.
Felis domesticus, brain of, ii. 120.
tigris, brain of, ii. 120.
Ferret, brain of, ii. 119.
Ferrier, Dr., on carnivorous brain, ii. 96.
on experiments on brain, ii. 125.
Fissure, of Rolando, ii. 1 26.
of Sylvius, ii. 95, 122.
parieto-occipital, ii. 131.
prrecentral and prsesylvian, ii. 130.
Fleming, J., on genus Monachus, ii. 66.
Flesch, M., on Brain of Ursus arctos, ii. 131.
Floe rat ( = Phoca hispida), ii. 59.
Flower, W. H., on cerebral nomenclature, ii. 95, 96.
on Macrorhinus leoninus, ii. 5, 19.
on Otariidae, ii. 73.
on Ziphioids, i. 21, 22, 26, 29.
Fossil Seal, ii. 60.
Fur-Seals, ii. 73.
Globiocephalus, ear-bones, i. 37, 39, 40.
inelas, kidney of, ii. 136.
Golspie, Seal from, ii. 62.
Gray, J. E., figures by, ii. 63.
on Macrorhinus leoninus, ii. 5.
on Megaptera, i. 31.
on Otaria, ii. 33.
on Otariidce, ii. 73.
Haast, Sir J. von, on teeth of Mesoplodon, i. 1 7, 22.
on Ziphius cavirostris, i, 8.
Hair-Seals, ii. 73.
Halichcerus, characters of, ii. 61.
grypus, characters of, ii. 62.
pup of, ii. 53.
uterus of, ii. 136.
Hapale jacchus, brain of, ii. 133.
Heard Island, Seals from, ii. 3, 19.
Hector, Sir J., on teeth of Mesoplodon, i. 16.
on Megaptera, i. 31.
on Ziphius cavirostris, i. 27.
presents skull of Stenorhynchus, ii. 20.
REPORT ON THE SEALS.
237
Histriophoca fasciata, ii. 61.
Homologous cerebral convolutions and fissures, ii. 134.
Humpback Whale of New Zealand, specimen examined,
i. 1.
Humphry, G. M., on Myology of Seals, ii. 140, e. s.
Hyoid Bone, Mesqplodon layardi, i. 26.
Inaccessible Island. See Tristan da Cunha, ii. 3, 36.
Juan Fernandez, Seals from, ii. 3, 36, 52.
Kerguelen Island, Seals from, ii. 3, 29, 36, 83, 91.
Kogia macleayi, ear-bone, i. 37, 41.
Krueg, J., on brain of Seal, ii. 114.
Lamna, teeth from deep sea, i. 42.
Langley, J. N, on cerebral nomenclature, ii. 96.
Lankester, E. R., on tooth of Mesoplodon, i. 18, 20.
Leptonyclwtes, characters of, ii. 64.
specimens of, ii. 20.
weddelli, characters of, ii. G5.
described, ii. 20.
Leptonyx, ii. 63.
weddelli, ii. 20, 63, 65.
Leuret, on Mammalian brains, ii. 113.
Lobodon, ii. 20.
carcinophaga, ii. 63, 64
Lucae, J. C. G., on Myology of Seals, ii. 140, e. s.
Lutra vulgaris, brain of, ii. 119.
McBain, Jas., on skull of Otaria, ii. 30, 77.
M'Kellar, John, specimen presented by, i. 2.
Macrorhinus, external characters of, ii. 68.
skeleton of, ii. 12.
skull of, ii. 5.
specimens of, ii. 3.
tarsalia of, ii. 50
anguirostris, ii. 69.
leoninus, brain of, ii. 91.
characters of, ii. 69.
described, ii. 3.
reference to figures of, ii. 63.
viscera of, ii. 135.
Magellan Strait, Seal from, ii. 30, 36, 41.
Maldonado, Seal from, ii. 30.
Man, brain of, compared with Seals, ii. 121.
Manganese, on Cetacean bones, i. 33.
Marion Island, Seal from, ii. 3.
Marmoset monkey. See Hapale jacckus, ii. 133.
Maxilla of Cetacean from deep sea, i. 38.
Medulla oblongata of Elephant Seal, ii. 101.
Walrus, ii. 112.
Megaptera lalandi, account of, i 30
ear-bone of, i. 40.
longimana, i. 30.
Megaptera nov&zealandim, i. 31.
Helen taxus, brain of, ii. 118.
Hellivora indica, brain of, ii. 118.
Mesoplodon australis, mentioned, i. 4.
vertebrae of, i. 22.
ftoweri ( — 31. layardi), i. 3.
grayi, mentioned, i. 4.
vertebras of, i. 22.
giintheri ( = H. layardi), i. 3.
hectori, mentioned, i. 4.
layardi, characters of, i. 2.
ear-bones of, i. 36, 37, 39.
geographical range of, i. 3.
occurrence recorded, i. 3.
specimens examined, i. 1.
longirostris ( = M. layardi), i. 3.
sqwerbyi, skull of, i. 4.
vertebrae of, i. 22.
Messier Channel, Seals from, ii. 36, 39, 41.
Meynert, T., on Mammalian brain, ii. 128.
Miller, W. C. S., Appendix on Myology of Pinnipedia,
ii. 139.
Mivart, St. G, on Macrorhinus leoninus, ii. 5.
on Otariidre, ii. 74.
on Ursine lozenge, ii. 95, 116.
Monaclnis, characters of, ii. 66.
albiventer, ii. 67.
monaclnis, characters of, ii. 67.
Montrose, Fossil Seal from, ii. 62.
Moon, Charles, presented brain of Walrus, ii.
Morse. See Trichechus rosmarus, ii 61.
Morunga elephantina, ii. 69.
Moseley, H. N, Notes by, i. 2, 16.
Murie, J., on Brain of Otaria, ii. 114, 132.
on Myology of Otaria and Trichechus, ii. 140, e. s.
on Otariidae, ii. 74.
on Sea Lion, ii. 29.
Murray, John, on Dredging of Cetacean bones, i. 33.
Muscles — ■
Abdominal, ii. 222.
Back, ii. 226.
Dermal, ii. 140.
Diaphragm, ii. 225.
Facial, of Expression, ii. 221.
Fore Limb, ii. 142.
brachial — ■
anterior, ii. 158.
posterior, ii. 160.
extensor, ii. 169.
first layer, ii. 142.
238
THE VOYAGE OF H.M.S. CHALLENGER.
Muscles —
Fore Limb —
rnauus, ii. 168.
seeond layer, ii. 145.
shoulder, ii. 153.
thoracic —
lateral, ii. 151.
ventral, ii. 148.
Hind Limb, ii. 176.
femoral —
internal, ii. 185.
ventral, ii 181.
fibular, ii. 196.
gluteal, ii. 188.
ilio-femoral, ii. 176.
leg, ii. 194.
pelvis to leg, ii. 191.
pes, inner or plantar, ii. 206.
outer, ii. 205.
tibio-fibular —
inner, ii. 198.
outer, ii. 194.
Masticatory, ii. 213.
Neck, ii. 214.
infia-hyoid, ii. 214.
supra-hyoid, ii. 215.
Perinseal, ii. 234.
Pharynx, ii. 216.
Praevertebral, ii. 217.
cervical, ii. 217.
lateral, ii. 218.
Soft Palate, ii. 206.
Tail, ii. 233.
Thoracic, ii. 216.
Tongue, ii. 216.
Mustela furo, brain of, ii. 119.
pennanti, cranial variation in, ii. 80.
vulgaris, brain of, ii. 119.
Nasua rufa, brain of, ii. 119.
Nehring, A., on Seals from Brazil, ii. 83.
New Zealand, Cetacea from, i. 3, 30.
Nightingale Island. See Tristan da Cunha, ii. 36.
Odobaenidae. See Trichechida?, ii. 69.
Odobeenus. See Trichechus, ii. 70.
(Edi/puB, brain of, ii. 133.
Ogmorhininas, characters of, ii. 62.
Ogmorhinus, ii. 20.
characters of, ii. 63.
carcinophagus, characters of, ii. 64.
leptonyx, characters of, ii. 64.
Ommatophoca, characters of, ii. 65.
rossi, characters of, ii. 64.
mandible of, ii. 23.
reference to figure, ii. 63.
Otaria, characters of, ii. 29, 75.
albicollis, ii 81.
argentata, ii. 54, 87.
australis, ii. 82.
californiana, ii. 77.
cinerea, ii. 79.
falMandica, ii. 82.
forsteri, ii. 87.
gazella, ii. 83.
gillespii, ii. 77.
godeffroyi, ii. 35, 75.
hookeri, ii. 78.
jubata (Forster), characters of, ii. 75.
described, ii. 29.
reference to figure of, ii. 63.
tarsalia of, ii. 50.
varieties of, ii. 35.
leonina, ii. 35, 75.
minor, ii. 35.
pygmxa, ii. 35.
pusilla, ii. 84.
stelleri, ii. 76.
ullox, ii. 30, 35, 75.
ursina, ii. 86.
weddelli, ii. 20, 55.
Otariidce, characters of family, ii. 56.
classification of, ii. 73.
Otter, brain of. See Lutra vulgaris, ii. 119
Owen, Sir E., on Epiphysis cerebri, ii. 110.
on cerebral nomenclature and homologies, ii. 96,
113, 128.
on Ziphioid Whales, i 3, 5, 13.
Oxyrldna, teeth from deep sea, i. 42.
Pansch, Adolf, on cerebral nomenclature, ii. 96.
on homology of cerebral fissures, ii. 128.
Pelagios monaclms, ii. 67.
Pelagius, ii. 66.
Pelvis, Arctocephalus australis, ii. 47.
Leptonychotes weddelli, ii. 26.
Macrorhinus leoninus, ii. 16.
Peters, W., on Arctocephalus, ii. 36.
on Macrorhinus leoninus, ii. 19.
on Otaria, ii. 34.
on Otariidoa, ii. 73.
Petrorhynchus capeims ( = Ziphius cavirosiris), i. 27,
29, 39.
REPORT ON THE SEALS.
239
Philippi, R. A., on a Fur-Seal, ii. 54.
Phoca, characters of, ii. 58.
albioenfer, ii. 67.
annellata, ii. 59.
antarctica, ii. 84.
australis, ii. 39, 82.
harbata, characters of, ii. 64.
carcinophaga, ii. 64.
caspica, ii. 61.
cristata, ii. 68.
elephantina, ii. 69.
equestris, ii. 61.
falklandica, ii. 82.
fcetida, ii. 59.
groenlandica, characters of, ii. 58
gryphus, ii. 62.
grypus, ii. 62.
hispida, characters of, ii. 59.
jubata, ii. 29, 75.
leonina, ii. 3, 69.
leptony.c, ii. 64.
monachus, ii. 67.
pusilla, ii. 84.
rosmarus, ii. 71.
siberica, ii. 61.
ursina, ii. 86.
vitulina, hrain of, ii. 115.
characters of, ii. 5S.
myology of, ii. 139.
pup of, ii. 53.
tarsalia of, ii. 50.
Phocardos elongatus, ii. 7S.
hooketri, ii. 79.
Phocidoe, characters of family, ii. 55.
PhocinEe, characters of, ii. 57.
Physeter macroeephalus, ear-bone, i. 37.
absence from dredgings, i. 41.
Pineal body of Elephant Seal, ii. 99.
Walrus, ii. 109.
Pineal eye in Lacertilia, ii. 110.
Pcescopia lalandii = Megaptera, i. 31.
Polar Bear. See Ursus maritimus, ii. 117, 125, 131.
Posterior extremity —
Ardocephalus australis, ii. 48.
Leptonyehotes weddelli, ii. 26.
Macrorhinus leoninus, ii. 17.
Muscles of, ii. 176.
Pup of various Seals, ii. 53.
Ratel. See Mellivora indica, brain of, ii. 118.
Red Crag, Baleenoptera from, i. 35.
Ribs of Ardocephalus australis, ii. 44.
Leptonyehotes weddelli, ii. 24.
Macrorhinus leoninus, ii. 14.
Mesoplodon layardi, L 25.
Right Whale of New Zealand, vertebrae of, i. 1, 32.
Rio de Janeiro, Seal from, ii. 83.
Rolando, Homology of Fissure of, ii. 126.
Rorqual du Cap, i. 30.
St. Andrews, Seal from, ii. 68.
Scammon, Capt., on Macrorhinus leoninus, ii. 19.
Sea Bears, ii. 73.
Sea Elephant, ii. 3, 69.
Sea Horse. See Tricheehus rosmarus, ii. 71.
Sea Lions, ii. 29, 73, 75.
Seals —
Auckland Island Hair-, ii. 78.
Bearded, ii. 60.
Californian Sea Elephant, ii. 19, 69.
Common Harbour, ii. 58.
Crab-eating, ii. 64.
Crested, ii. 68.
Elephant, ii. 3, 69.
False Leopard, ii. 20.
Fur-Seals, of —
Australia, ii. 87.
Cape of Good Hope, ii. 84.
Crozets, ii. 84.
New Zealand, ii. 87.
North Pacific, ii. 86.
Grey, ii. 62.
Grey Sea Lion, ii. 79.
Harp, ii. 58.
Kerguelen Island Fur-Seal, ii. 36.
Leopard, ii. 64.
Lion, ii. 29.
Monk, ii. 67.
Ross's Large-eyed, ii. 23, 65.
Saw-toothed, ii. 64.
South American Fur-Seal, ii. 39, 82.
Southern Sea Lion, ii. 75.
Steller's, ii. 76.
Weddell's, ii. 20, 65.
Shark's teeth on floor of ocean, L 41.
Shetland, Cetacea from, i. 27.
Seal from, ii. 62.
Skeleton —
Ardocephalus australis, ii. 41
gaxella, ii. 37.
Leptonyehotes 'weddelli, ii. 20.
Macrorhinus leoninus, ii. 5.
240
THE VOYAGE OF H.M.S. CHALLENGER.
Skull—
Ardocephalus austraMs, ii. 41.
gazella, ii. 37.
sp. incerta, ii. 53,
Leptonycliotes weddelli, ii. 20.
Macrorhinus leoninus, ii. 5.
Mesoplodon layardi, i. 3.
Otaria jubata, ii. 29.
Smith, E. T., presented a specimen, ii. 29.
Spinal column —
Ardocephalus australis, ii. 43.
Balxna australis, i. 32.
Leptonycliotes weddelli, ii. 23.
Macrorhinus leoninus, ii. 12.
Mesoplodon layardi, i. 21.
Megaptera lalandi, i. 30.
Stenorhinque, ii. 63.
Stenorhynchus, ii. 20, 63.
leptonyx, ii. 63, 64.
serridens, ii. 64.
Sternum —
Ardocephalus australis, ii. 45.
Leptonycliotes weddelli, ii. 25.
Macrorhinus leoninus, ii. 15.
Mesoplodon layardi, and other sp., i. 26.
Stones in Stomach of Seals, ii. 136.
Sydney, specimen from, i. 3, 18.
Sylvius, fissure and convolution of, ii. 94, 95, 104, 122,
124.
Tables of—
Dimensions of —
Brain of Elephant Seal, ii. 92.
Walrus, ii. 103.
Cast of Cranial Cavity of Elephant Seal, ii. 92.
Crania of Ardocephalus australis, ii. 41.
gazella, ii. 37.
Eumetopias californiamis, ii. 77.
cinereus, ii. 80.
Macrorhinus leoninus, ii. 6.
Leptonycliotes and Stenorhynchus, ii. 21.
Otaria jubata, ii. 30.
Fur-Seal from Juan Fernandez, ii. 52.
South American Fur-Seal, ii. 39.
Teeth of Elephant Seal, ii. 7.
Table of homologous Fissures and Convolutions, ii. 134.
Tay, Seal from, ii. 62.
Teeth—
Ardocephalus australis, ii. 42.
gazella, ii. 37.
Leptonycliotes iveddelli, ii. 20.
Macrorhinus leoninus, ii. 7.
Mesoplodon layardi, i. 10.
soiverbyi, i. 20.
Otaria jubata, ii. 31.
of Sharks, i. 41.
Thomas, Oldfield, on Mustela pennanti, ii. 80.
Thomson, Allen, on ossification of digits, ii. 51.
Tiger, brain of. See Felis tigris, ii. 120.
Trichechida?, characters of family, ii. 56.
extinct, ii. 69.
Tricliechodon, ii. 69.
Trichechus, characters of, ii. 70.
obesus, ii. 71.
rosmarus, characters of, ii. 61.
Tristan da Cunha, Seals from, ii. 3, 36.
Tympanic bulla, i. 8, 34 ; ii. 11, 23, 33, 43.
Ursine lozenge, ii. 95, 116.
Ursus arctos, brain of, ii. 131.
marinus, ii. 86.
maritimus, brain of, ii. 117, 125, 131.
Van Beneden, P. J., on Ziphius, i. 29.
on Megaptera, i. 31.
Vertebrae. See Spinal column.
Viscera of Elephant Seal, ii. 135.
"Walrus, brain of, ii. 202.
Weasel, brain of, ii. 119.
Wellington, Cetacea from, i. 27.
Seals from, ii. 21.
Zalophus gillespii, ii. 77.
Ziphius, beak from deep sea, i. 38.
cavirostris, specimen examined, i. 1.
skull measurements, i. 4, 8.
general sketch, i. 27.
ear-bone, i. 36, 39, 41.
chathamiensis, identical with Z. cavirostris, i. 29.
indicus, identical with Z. cavirostris, i. 27, 29.
novx-zealandix, identical with Ziphius caviros-
tris, i. 29.
PLATE I.
(zool. chall. exp. — part Lxvm.— 1888.)— Yyy.
PLATE I.
Elephant Seal.
Fig. 1. Profile view of head and anterior part of body of well-grown female Mavrorhinus
leoninas, from Christmas Harbour, Kerguelen.
Fig. 2. Dorsal aspect of the hind limbs and caudal region of the same animal.
Fig. 3. Ventral aspect of the hind limbs and vent of the same animal.
I am indebted to Arthur Thomson, M.B., for the drawings from nature from which
these figures were taken.
The Voyage of H.M S'Challenger"
Seals PI I
ATthur Thomson M H pinxit
ELEPHANT SEAL ?
FHtiUi.Lilh' Kdm'
PLATE II.
PLATE II.
Elephant Seal.
Fig. 1. Profile of skull of adult male (h) Macrorhinus leoninus, from Heard Island.
Fig. 2. Profile of skull of well-grown male (e), from Kerguelen Island. The skeleton
of this animal has been described in Part I.
Fig. 3. Profile of skull of a large female (/), from Kerguelen Island.
Fig. 4. Profile of young skull, apparently a female (h), from Kerguelen Island.
Fig. 5. Anterior surface of atlas vertebra. This and the other vertebra are from a male
skeleton.
Fig. 6. Anterior surface of axis vertebra.
Fig. 7. Anterior surface of 3rd cervical vertebra.
Fig. 8. Anterior surface of 7th cervical vertebra.
The Voyage of H.M.S."Challenger.'
Seals
ELEPHANT SEAL.
■
PLATE III.
(ZOOL. OHALL. EXP. PART LXVIII. — 1888.) — Yyy.
PLATE III.
Elephant Seal.
Fig. 1. Vertex view of skull of adult male (h) Macrorhinus leoninus.
Fig. 2. Vertex view of skull of a well-grown male (e).
Fig. 3. Vertex view of skull of a large female (f).
Fig. 4. Vertex view of a young skull, apparently a female (h).
Fig. 5. Side view of pelvis of a female (c), from Kerguelen Island.
The Voyage of HM.S.'Thalleiiger
Seals PI III.
V.
SteS;
r*
""^SsaSSS*- ~ "
■^
*
From I
ELEPHANT SEAL
yHuth.LitKEdin*
PLATE IV.
PLATE IV.
Elephant Seal.
Fig. 1. Dorsum of the left scapula of a well-grown male Macrorhinus Iconinus, from
Kerguelen Island ; the cartilage is still attached to the vertebral border.
Fig. 2. Anterior aspect of the right humerus of the same animal.
Fig. 3. Dorsal surface of the bones of the left fore-arm and hand of the same animal ;
si, scapholunar ; c. cuneiform ; p, pisiform ; tr, trapezium ; v, unciform ; P,
pollex ; V, minimus.
Fig. 4. Front of the right femur of the same animal.
Fig. 5. a, Articular surface of the right patella ; b, the same bone in profile.
Fig. 6. The ventral surface of the left tibia and fibula of the same animal.
Fig. 7. Dorsal surface of the right tarsus and metatarsus of the same animal ; a, astra-
galus, with c, its calcanear process ; cl, os calcis with its calcanear process ;
cu, cuboid bone ; sc, scaphoid bone ; en, opposite the three cuneiform bones ;
H, hallux ; II., III., IV., V., metatarsal bones of four outer toes.
The Voyage of H M.S. "Challenger."
tr si
Fyr.S.
Fig. 2.
Fig. I.
Fie,.*.
Fig. 5
Fig. 6.
From Photo" and 'Nature.
F Hull, Li
ELEPHANT S EAL
PLATE V.
(ZOOL. CHALL. EXP. — PART LXVIII. — 1888.) Vyy.
PLATE V.
Weddell's Seal.
Fig. 1. Dorsal surface of the skull of Leptonychotes weddelli ; f, supraoccipital venous
foramen ; reduced.
Fig. 2. Profile view of the lower jaw of the same skull ; reduced.
Fig. 3. Series of upper and lower post-canine teeth of the left side ; natural size.
Fig. 4. Dorsal surface of the bones of the right anterior extremity of the same animal ;
p, the pisiform bone ; si, scapholunar bone ; tr, trapezium ; P, pollex ; V, the
minimus.
Fig. 5. Left side of the pelvis with the last lumbar and the more anterior caudal
vertebrae.
Fig. 6. Anterior surface of the left femur.
Fig. 7. a, Articular surface of the left patella ; b, the same bone in profile.
Fig. 8. Ventral surface of the left tibia and fibula.
Fig. 9. Dorsal surface of the left pes of the same animal ; a, astragalus with its calcanear
process ; cl, calcanear process of os calcis ; cu, cuboid ; sc, scaphoid ; en,
opposite the ento-cuneiform ; H, the hallux; II., III., IV., V, the four
outer toes.
The Voyage of H.M.S."Challen|er:'
.
S
Fig. 9.
Fig. 6.
Fig. I.
y
Fig. 8.
Fig. J.
Fig. 5.
\
Fig. ;
Front Photo* and Nature
FHolh,:illi'Eiinr
WEDDELL'S SEAL
PLATE VI.
PLATE VI.
Fur-Seals, South America and Kerguelen Island.
Fig. 1. Profile view of skull of adult male Arctocephalus australis, from the Messier
Channel.
Fig. 2. Profile view of skull of Arctocephalus gazella, from Kerguelen Island, probably a
male, but not adult.
Fig. 3. Vertex view of the skull of the same Arctocephalus australis.
Fig. 4. Vertex view of the skull of the same Arctocephalus gazella.
Fig. 5. Inferior surface of the skull of the same Arctocephalus australis.
Fig. 6. Inferior surface of the skull of the same Arctocephalus gazella.
The Voyage of H.M.S"Chal] i
From Photo" and Nature.
FUR SEALS, SOUTH AMERICA & KERGUELEN ISLAND.
F Huth. I
PLATE VII.
(zool. chall. exp. — paut Lxvin. — 1888.) — Yyy.
PLATE VII.
Fur-Seal, South America.
Fig. 1. Profile of left side of the series of cervical vertebrae of male Arctocephalus
australis.
Fig. 2. Dorsal surface of right scapula.
Fig. 3. Front of right humerus.
Fig. 4. Radius aud ulna of right fore-arm.
Fig. 5. Palmar surface of right carpus and metacarpus ; P, pollex ; V, minimus ;
p, pisiform ; si, scapholunar bone ; tr, trapezium ; c, cuneiform.
Fig. 6. Left side of pelvis with three of the caudal vertebras.
Fig. 7. Anterior surface of right femur.
Fig. 8. a, Articular surface of left patella ; b, profile view of same bone.
Fig. 9. Tibia and fibula of left leg.
Fig. 1 0. Plantar surface of left tarsus and metatarsus ; H, hallux ; V, minimus ;
a, astragalus ; cl, os calcis ; c,m, cuboid ; en, entocuneiform ; es, entoscaphoid.
The Voyage of H.M S.'Xbtllenger."
Seals. PI VII
Fig. 7-
Fig. 2.
... •
.
Fig. 8.
Fig. 9.
m
r %
Fig. 6.
Fig. 10.
Fig. 3.
From Phoios and Nature
F Huth.L'
FUR SEAL, SOUTH AMERICA.
PLATE VIII.
PLATE VIII.
Brain of Elephant Seal.
The fissures, convolutions, and other divisions of the brain, both in this and the succeeding Plates, are lettered as below :
Fissures.
s.
Sylvian.
c.
crucial.
pc.
prsecruciate.
ps.
prsesylvian.
CO.
coronal.
ml.
medilateral.
1.
lateral.
ss.
suprasylvian.
ssp.
posterior suprasylvian.
rh.
rhinal.
■ r.
postrhinal.
.
intraorbital.
ol.
olfactory.
h.
hippocauipal.
sp.
splenial.
sps.
suprasplenial.
ph.
postero-horizontal.
psp.
postsplenial.
V.
vorticose.
I. olfactory.
II. optic.
III. motor oculi.
IV. trochlearis.
Convolutions, &c.
re.
gyrus rectus.
isc.
internal supraorbital.
esc.
external supraorbital.
ur.
ursine lozenge.
sac.
sagittal or 1st external.
mlc.
medilateral or 2nd external.
ssc.
suprasylvian or 3rd external.
syc.
Sylvian or 4th external.
sgc.
sigmoid gyrus.
cc.
callosal gyrus.
he.
hippocampal gyrus.
Ih.
lobus hippocampi or uncinatus.
spc.
splenial convolution.
sspc
suprasplenial convolution.
pre.
prorean convolution.
to.
tuber olfactorium.
ob.
olfactory bulb.
P.
pineal body.
H.
hypophysis cerebri or pituitary body
eel.
corpus callosum.
cs.
corpus striatum.
oth.
optic thalamus.
eh.
choroid plexus.
km.
hippocampus major.
f.
fornix.
th.
taenia hippocampi.
a.
1st cervical spinal nerve.
ncrals
'.-XIL inclusive indicate the cranial nerves, as follows : —
V.
trigeminal.
IX. glosso-pharyngeal.
VI.
abducens.
X. pneumogastric.
VII.
portio dura or facial.
XI. spinal accessory.
VIII.
portio mollis or auditory.
XII. hypoglossal.
Fig. 1. Superior surface of the brain of Macrorhinus leoninus. The clotted outline
represents the form of the brain as taken from a cast of the cranial cavity.
Fig. 2. Inferior surface of the same brain.
Fig. 3. Profile view of the right hemisphere of the same brain.
The above figures were drawn from nature by T. W. Dewar, M.B.
Fig. 4. Superior surface of the brain of a fcetal Phoca grcenlandica, with the pia mater
not stripped off.
Fig, 5. Vertical transverse section through the right hemisphere and lateral ventricle
of the cerebrum of brain a of the Walrus, ecl, corpus callosum forming the
roof of the lateral ventricle ; cs, corpus striatum.
Figs. 4 and 5 were drawn from nature from my dissection by Professor
Eichard Caton.
The Voyage of H.M.S."Challei^
Seals. PI. VIII.
£
>=>
^
*
iwar M.B. del
F Huth, 1
ELEPHANT SEAL
PLATE IX.
(ZOOL. CHALL. EXP.— PART LSVIII.— 1888.)— Yyy.
PLATE IX.
Elephant Seal and Walrus.
Fig. 1. Anterior end of the cerebrum of Macrorhinus leoninus. The olfactory bulb is
turned down on one side to show the olfactory and prsecruciate fissures and
ursine lozenge. Drawn from nature by T. W. Dewar, M.B.
Fig. 2. Tentorial and inner surfaces of the left hemisphere of the same brain, obtained
by making an antero-posterior mesial section through the corpus callosum.
Drawn from nature from my dissection by K. M. Scott, M.B.
Fig. 3. Tentorial and inner surfaces of the left hemisphere of the Walrus (brain a.)
This and fig. 4 were drawn from nature from my dissection by Professor
Richard Caton.
Fig. 4. View of the floor of the lateral ventricle and the descending horn in the right
hemisphere of the cerebrum of the Walrus, obtained by removing the corpus
callosum.
The Voyage of H M S "Challenger"
Seals [X
S
KM.Scott4T.YI
ELEPHANT SEAL & WALRUS,
PLATE X.
PLATE X.
Walrus.
Fig. 1. Superior surface of the brain (a) of Trichechus rosmctrus ; the ends of the
olfactory bulbs are seen anteriorly, and the pineal body is visible posteriori)'
between the two diverging hemispheres. The pineal body has been filled in
from brain c, by K. M. Scott, M.B.
Fig. 2. Inferior surface of the same brain from which the pituitary body drawn in
fig. 6 had been removed. The sensory root of the right 5th nerve has the
Gasserian ganglion connected with it. The vorticose fissure in the hemi-
sphere of the cerebellum is well seen. The cranial nerves behind the 8th
had been torn off. The shape of the medulla oblongata was restored from
brain c.
Fig. 3. Profile of the left hemisphere of the same brain.
The above figures were drawn from nature by Professor Richard Caton.
Fig. 4. Anterior end of the left hemisphere of the cerebrum of brain c, to show the
crucial fissure.
Fig. 5. Tentorial surface of the left hemisphere of the same brain, reduced. When
compared with fig. 3, PL IX., several modifications in the splenial fissure
and adjacent convolutions will be seen.
Fig. 6. Inferior surface of the pituitary body divided into four lobes.
Fig. 7. Profile view of the pineal body, epiphysis cerebri.
The Voyage of H.M.S."Challenge
PI X
R
WALRUS
THE
VOYAGE OF H.M.S. CHALLENGER
ZOOLOGY.
REPORT on the Actiniaria dredged by H.M.S. Challenger during the
years 1873-1876. By Prof. Richard Hertwig.
SUPPLEMENT.
INTRODUCTION.
After I had concluded my Eeport on the Actiniaria of the Challenger Expedition,
a number of additional specimens were sent to me, on which I now present a short
Supplementary Report. Unfortunately the work has been delayed longer than I could
have wished, partly on account of a series of experimental investigations upon the
fertilisation and segmentation of the ovum, which I had undertaken in concert with my
brother, but mainly owing to the claims on my working-time caused by my trans-
ference from Konigsberg to Bonn, and from Bonn to Munich.
Amongst the material occurred several specimens of species which have been previ-
ously described and can therefore be treated in few words ; besides these, there are also
several new forms, representing new and interesting genera, which require a detailed
description, and which are, for the sake of clearness, designated by an asterisk. At the
end (p. 54) will be found a list of those Actinias of which a systematic study was
impossible, either because they were not sufficiently well preserved, or because their
appearance was no longer characteristic owing to the absence of sculpturing and colour,
the necessary result of the method of preservation.
Since the publication of the earlier part of this Report, the great monograph of
Angelo Andres on the Actiniaria has appeared.1 During the progress of his work this
1 Fauna und Flora des Golfes von Neapel, Le Attinie, 1884.
(ZOOL. CHALL. EXP. — PAET LXXIII. 1888.) lUldd 1
2 THE VOYAGE OF H.M.S. CHALLENGER.
author was acquainted only with the short preliminary notice of my researches published
in the Jena Proceedings,1 not with the Eeport itself ; a fact easily understood when one
considers how long before the date of publication a monograph constructed on such a
plan must be completed. In his comprehensive revision of the Actinise, and re-definition
of families and genera, he has been prevented from referring to my contemporaneous
attempt at revision, since this first appeared in the detailed Report. As it is most
desirable that two systems, appearing within a short time of one another, should be
brought into such relation as to avoid future discordance and mistake, I accept with
pleasure the opportunity of a critical utterance on their mutual relations.
As against the six chief divisions into which I divide the Actinise (Hexactinise,
Paractinise, Monaulese, Edwardsise, Cerianthese, Zoanthese), Andres erects seven, viz.
Edwardsinse, Actininae, Stichodactylinse, Thalassianthinse, Zoanthinse, Cerianthinse,
Minyadinse. With regard to three chief groups we are in complete accord (Edwardsise,
Cerianthese, Zoanthese), except for the fact that Andres, in my opinion, relies on too
inconstant and unimportant external characters ; while, as I have shown, these groups,
at least, admit of anatomical characterisation by the arrangement of their mesenteries,
and thus can be far more clearly and sharply circumscribed. If the reader compare in
this connection the definitions of the Zoanthese furnished by myself and by Andres, it
will be readily admitted that none of the characteristics of the latter author, such as
colony-formation or incrustation, are constant within the group ; that, on the other
hand, all the forms follow one and the same law of mesenterial arrangement, first
recognised by G. von Koch.
If we carry the comparison further, we find that Andres places beside the Actininse,
as separate groups, the Thalassianthinse, the Stichodactylinse, and the Minyadinse ;
though with a certain caution, as having himself studied no representative of them.
I believe that he has here exceeded the systematic value which can be safely assigned
to the form of the tentacles and their distribution on the mesenterial chambers. I have
studied certain Stichodactylinse (Corallimoiyhus rigidus, Corallimorphus profundus,
and Heterodactyla hemprichii), and of the Thalassianthinse, Thalassianthus aster,
and can assert, as the result of a thorough examination of their structure, that in all
important points they agree with the hexamerous Actinise ; nor have I any doubt that
these forms, even if united into separate families, must be ranged among the Hexactinise.
Finally, the group of Minyadinae has for many reasons, which I entirely recognise,
undergone at the hands of Andres so sharp a criticism, that one can hardly see why he
retains it, or why at least he does not allow it to rank merely as a subdivision of
Hexactinise, until the necessity of its removal from that group is rendered apparent by
anatomical investigation.
From the point of view explained, I am of opinion that all the forms referred to
1 Jtnaische Zeitschr., Bd. xv. p. 10, 1881.
REPORT ON THE ACTINIARIA. 3
by Andres must be comprehended in the four divisions, Edwardsiae, Hexactiniae,
Zoantheae, and Cerianthese, and accordingly hold to the systematic classification which
I have published. The groups of Paractiniae and Monauleae are in all respects natural,
and would also certainly be retained by Andres had representatives of them been
known to him.
Even greater discordance than that of which I have hitherto spoken, between the
classifications of Actiniae followed by Andres and myself, presents itself when the
determination and nomenclature of families and genera are regarded. Independently
of each other, and from different standpoints, we have taken in hand a systematic
revision of Actiniae : Andres starting; with the advantage of a richer material, and
studying species with which earlier publications are especially concerned, and which he
could command in a living condition ; while my qualification for a systematic classifica-
tion was that afforded by close anatomical investigation, namely, that I relied for
systematic characteristics upon such weighty differences as the structure of the
sphincter, the arrangement of the mesenteries, the structure of the musculature and of
the oral disc, etc., points which Andres has, hitherto at any rate, entirely left out of
consideration. Thus it has resulted that in the determination of families and genera,
and also in the value assigned to existing names, we have in many cases taken up a
totally different attitude ; and as, in consequence of this, no inconsiderable confusion
has arisen in the method of diagnosis, I hold it advisable to inquire critically what
must be retained of the system of the Italian observer.
Of least importance are our differences of opinion relating to those Actinias which
possess acontia. Andres has here adopted the separation, instituted by Verrill, into
Sagartidse and Phellidae. Having regard to his wider acquaintance with the species, I
agree with him in accepting as a distinctive character the chitinous covering extending
over two-thirds of the body-wall ; and for clearer characterisation of both families, the
following marks not mentioned by Andres should be included in the diagnosis, — a
mesodermal sphincter, and a differentiation of the mesenteries into sterile complete
primary mesenteries, and incomplete secondary mesenteries provided with generative
organs. Of the Challenger Actinia?, there would belong to the Phellidae only Phellia
pectinata ; to the Sagartidse, Sagartia sp., Cereus spinosus, Calliactis polypus,
Bunodes minuta. Of these, the two latter require an alteration of name ; Calliactis
polypus must be termed Adamsia polypus,1 and Bunodes minuta be known as Cylista
minuta, since it has been shown by Andres that the typical Bunodes possesses no
acontia, and therefore cannot belong to the Sagartidae.
Andres has incorrectly allowed the generic name Cereus (Oken) to drop, and has
1 The specific name Tlondeletii has heen -wantonly substituted by Andres for the older polypus, the former being
used for the first time by delle Chiaje in 1825, while the latter was already instituted by Forskal in 1775. Milne-
Edwards is therefore correct in calling the animal Adamsia polypus.
4 THE VOYAGE OF H.M.S. CHALLENGER.
introduced in its place the more recent name Heliactis, for Sagartidae with numerous
large papilla; ; although Oken adduces Cereus bellis as the type form, which stands in
the same relation to the genus Heliactis. The papillate Sagartidae are of two kinds, the
one having a soft surface, while in the other the body-wall is covered as far as its upper
edge with a bark -like cuticle which recalls the Phellidae ; it is therefore advantageous to
confine to the former the name Heliactis, applied, though unjustifiably, by Andres, and
for the latter to restore Cereus, the old designation of Oken, a representative of the
newly characterised genus being Cereus spinosus.
In discussing the families instituted by Andres, we next come to the Paractidae.
As I understand the diagnosis given for this family, — " margine tentaculato, non
rilevato e privo d' acroragi," — the tentacles spring at the edge where body-wall and
oral disc pass into one another, just as is the case both in the Corallimorphidse and
Antheomorphidae, which I have described in more detail, and, generally speaking, in
such Actinias as are devoid of a circular muscle. But this relation also holds good in
Actiniae with a weak sphincter, as, for example, in Anemonia cereus (to which Andres,
strange to say, ascribes a " margine rilevato ") ; and, finally, in Actiniae, in which the
sphincter is developed at some distance outwards from the upper edge of the body-wall.
The facts adduced are sufficient to prove that this characteristic is systematically
useless ; and in addition to this I insist that the few forms grouped in the family do
not appear to agree with the diagnosis. The tentacles of an Anemonia are, according
to Andres, formations placed more at the edge than are those of a Paranthus or a
Paractinia. On the contrary, the Paractis peruviana, which Andres adduces as the
type of the family, seems to me to have no tentacles which would be marginal. Indeed,
it agrees so entirely with a Challenger form, Paractis excavata, that I long doubted
whether it were not right to unite the two. In Paractis excavata, I am certain that a
strong mesodermal sphincter is present, and, corresponding to this fact, body-wall and
oral disc are sharply marked off from each other, whence I conclude that the same holds
for Paractis peruviana. Since I have thus apod ground for holding; unsuitable the
methods by which Andres has instituted his family Paractidae, and can, in addition,
claim the right of priority, I adhere to the definition which I previously published,
leaving only to future investigators to decide upon the advisability of erecting Actiniae
with marginal spherules, sucking-papillae, and papillae into a family separate from the
Paractidae (sensu stricto) with smooth body-wall. ^
The next family in the system of the Italian naturalist is formed by the Actinida?,
and corresponds to the Antheadae and Actinidae of Gosse. I formerly followed Gosse
in separating these two families, but had previously maintained that anatomically they
are closely related, and should perhaps on that account be united. I have therefore
nothing to adduce against this proceeding of Andres, though the detailed investigation
of the Actinidae, which I recommended, has not vet been carried out. It is also
REPORT ON THE ACTINIARIA. 5
correct to replace the name Anthea by the older Anemonia, and to range the genus
Comactis under it. On the other hand, my Comactis jlagelUfera is not identical with
Anemonia sulcata [Anthea cereus), and should therefore be referred to as Anemonia
fiacjellifera.
In the system of Andres the Bunodidae bear the closest relation to my family
Tealidae. I was unacquainted with any typical Bunodes, and had supposed (ef. supra)
that they possessed acontia. This supposition is, according to Andres, incorrect ; and
the close relationship to Tealia is thus anew proven. Accordingly I withdraw the
name Tealidae in favour of the older designation Bunodidae ; but, now as formerly, the
endodermal sphincter must occupy the first place in the diagnosis. I relinquish, how-
ever, to future observers, as with the Paractidae, the decision whether forms with
smooth and with papillate body-wall should be separated from one another, or not.
A last point of dispute with Angelo Andres lies in the fact that I reckon the
Halcampce among the Ilyanthidae, while he erects them into a separate family. I will
not decide in this place either for the one opinion or the other, but will discuss merely
the point of view, which, as it seems to me, must be of importance for a decision.
The more we have learnt in late years of the structure of these forms, the
more has it become apparent that Actinias, which are rounded posteriorly and
devoid of pedal disc, exhibit in most cases a sort of ancestral character ; eminently
primitive forms are, above all others, the Edwardsiae. Among such forms is the
genus Halcampa, from which again the genus Halcampella is a transition to the
remaining Actiniae, in virtue of its numerous tentacles, and of its commencing to
exhibit accessory mesenteries. I opine that the genus Uyanthus stands in close
relation to the Halcampcllce ; the regular increase of the mesenterial pairs by multiples
of six, which is commencing in the one case, is in the other clearly expressed,
as may be inferred from the presence of the numerous longitudinal furrows of the
body-wall ; while the siphonoglyphes (ciliated grooves), the hinder edge of the body,
and the sphincter, are obviously of weak development, as among the Halcampce.
Possibly a study of the mesenteries may yield further points of agreement, but,
unfortunately, nothing is accurately known of these important features in the structure
of Uyanthus ; and so long as this is the case, no conclusion can be certain. If my
expectations be confirmed, a union of the Halcampce with the Ilyanthidae would be
desirable ; the latter would form a transitional family placed at the top of the
Hexactiniae, and bridging the gap between them and the Edwardsiae ; while, as a
peculiar and aberrant branch of the Actiniae, would be ranged near them the
Siphonactidae. the forms possessing a conchula.
All the forms of which we have as yet spoken possess the typical digitate
or tubular Actinian tentacles, so arranged that one tentacle corresponds to each
radial chamber ; there are, however, two variations of this arrangement. In the one,
6 THE VOYAGE OF H.M.S. CHALLENGER.
the tentacles are replaced by appendages of a different value, for instance, by stomidia
in the Liponemidse which I have described, or by bushy or arborescent growths in
the families Sarcophianthidre and Thalassianthidse erected by Andres. On the other
hand, there are forms in which more tentacles than one correspond to a mesenterial
chamber ; accessory tentacles, placed on the oral disc, being present in addition to the
primary tentacles. This is conclusively proved only for species of Corallimorphus, but
Andres has rendered it excessively probable also for species of Corynactis (compare the
account of Corynactis ? sp. ? p. 10, infra). For such forms I have instituted the family
Corallimorphidse, Andres the family Corynactidse. I believe that my designation
deserves preference, because it is the older, and because my diagnosis of the family
alone insists upon the important anatomical characteristic ; on the other hand, I concede
to the Italian naturalist that the family may be restricted to species with knobbed
tentacles, and that all Actinias with modified tentacles, of which an accurate investigation
is still required, may be brought under a series of further families.
For a comprehension of the above discussion, I give a view of that arrangement of
Hexactinian famdies which I hold the most advantageous, in the form of a synoptic
table.
A few changes have been made iu the English terminology used in the former part
of this Report : " oesophagus " has been replaced by " stomatodseum," " mesoderm "
by "mesoglcea," and "oesophageal groove" by "siphonoglyphe."
REPORT ON THE ACTINIARIA.
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DESCRIPTION OF GENERA AND SPECIES.
Tribe I. Hexactini^e.
Family 1, Corallimorphid.e, R. Hertwig.
Genus Corallimorplms, Moseley.
Corallimorphus rigidus, Moseley.
Amongst the supplementary material I have found the original specimen on which
Moseley formerly founded the species Corallimorphus rigidus. I had already mentioned
this on Moseley's authority in my earlier Report, though I had not myself seen it,
and had described from my own observation four more specimens, of which one, from
Station 157, agreed in all essential particulars with the three others from Station 146.
I am now in a position to confirm the statement that the three latter agree with
Moseley's specimen in form, in colour (of which traces only remain in spirit specimens),
and in the condition of the body-wall, — they exhibit no thickenings, but merely forty-
eight longitudinal furrows corresponding to the insertions of the mesenteries. Another
specimen, from Station 299, also agreeing with Moseley's type, is of interest, since, of
the twenty-four tentacles on the oral disc, one accessory tentacle of the first order is
duplicated, two little tentacles being planted close together. I have already described
a similar, though more strongly expressed, development of supernumerary tentacles
in Corallimorphus "profundus, so that it appears probable that the law of increase in
the tentacles of Corallimorphidse is not yet so definite as among other Actinias, and
allows of more variation than in other cases.
Corallimorphus obtectus, n. sp.
While the five last-named specimens agree with one another, that from Station
157, on which I chiefly based my former description, demands a separate position, so
that I now account it the representative of a new species to which I give the name
Corallimorphus obtectus, having regard to the buckle-like thickenings which cover the
insertions of the mesenteries. A further difference lies in its disc-like shape, due to the
relations of size between pedal and oral disc. Both are in this case of the same size, but
in Corallimorphus rigidus the former is considerably the smaller, producing a saucer-
shaped profile. The two species may be differentiated by the following diagnosis : —
1. Corallimorphus rigidus. — Twenty-four tentacles are planted on the oral disc,
(ZOOL. CHAI.L. EXP. — PART LXXIII. — 1888.) DdJJ 2
10 THE VOYAGE OF H.M.S. CHALLENGER.
and forty-eight at its edge ; the insertions of the mesenteries are recognisable by
longitudinal furrows ; the oral disc essentially larger than the pedal.
In addition to the examples referred to above, there belongs to this species one
specimen from Station 299, December 14, 1875 ; lat. 33° 31' S., long. 74° 43' W.; depth,
2160 fathoms. — Dimensions, height, T3 cm.; breadth of the oral disc, 4 cm., of the
pedal disc, 17 cm.
2. Corallimorphus obtectus. — Twenty-four tentacles are situated on the oral disc,
and forty-eight at its edge ; the mesenterial insertions are covered, in the lower third of
the body-wall and the peripheral third of the pedal disc, by cylindrical thickenings ;
the pedal and oral discs of approximately the same size.
To this species belongs only the example from Station 157, with which my former
description was concerned.
Genus Corynactis, Allman.
Corynactis (?) sp. (?)*
The tentacles, both on the disc and at its edge, are knobbed ; those on the disc are
arranged in several circles, so that more than one tentacle communicates with each
intra-mesenterial chamber.
Habitat.— Station 219, March 10, 1875 ; depth, 150 fathoms.
Dimensions.— Diameter of the oral disc, 2-5 cm. ; height of the column, 0'8 cm. ;
greatest length of the tentacles, 1*6 cm.
Angelo Andres gives in his monograph a description of the genus Corynactis,
based partly on personal observation, partly on the account of Allman, from which I
infer that, with one and the same radial chamber communicate one of the marginal
tentacles and, in many radii, several of those placed on the disc ; five cycles are present,
of which the first contains four tentacles, the second sixteen, the third, fourth, and
fifth twenty-four. Gosse records other numbers, namely, four rows with sixteen, twenty-
four, thirty -two, and thirty-two tentacles respectively. In such remarkable contra-
diction, one may well doubt whether one has any right to deduce a law of position from
either account ; and the descriptions of the manner of distribution of the tentacles
are so inadequate, that it is impossible to conjecture how many of the tentacles placed
on the disc correspond to a radial chamber.
Amongst the Challenger material was an Actinian which I originally took for a
Corallimorphus, till I recognised that on four radii of the body two tentacles on the disc
and one at its edge proceed from one and the same radial chamber. This is in contra-
diction to the law of the position of tentacles in Corallimorphus, but on the other hand
is related to that in Corynactis ; to the latter genus I therefore provisionally refer it, even
though many characters do not agree in the two forms. Especially is its shape divergent,
being saucer-like as in Corallimorphus, and not elongated as in Corynactis. Further,
REPORT ON THE ACTINIARIA. 1 1
the animal is very irregularly developed ; the number of marginal tentacles amounts to
fifty-six, larger and smaller generally alternating ; two cycles, each of twenty-eight, might
thus be recognised, did one not consider that the tentacles of each cycle differ markedly
and somewhat irregularly in size. One is compelled to rank under the primary circlet,
tentacles which in diameter are far short of tentacles of the second order. Even more
irregular is the arrangement of those tentacles which are situated on the disc : their
total number, twenty-three, falls into three cycles, six tentacles being placed near the
mouth (oral tentacles), ten near the edge (peripheral tentacles), and seven intermediately.
Despite these apparently irregular numbers, I have noticed the complete validity of a
law in one-half of the animal, and it is of importance that this regular half commences
with the one pair of directive mesenteries and reaches to the other, thus just completing
one side of the animal. In the half in which a regular arrangement is followed, we
have three oral, six intermediary, and six peripheral, accessory tentacles. The six inter-
mediary alternate with the six peripheral, three of them standing on the same radius
as the oral tentacles. If we compare with these the marginal tentacles, the larger
twelve are on the radii already occupied by the tentacles on the disc, while the smaller
twelve are placed on the intermediate radii. At each of two points two tentacles are
present, a larger and a smaller ; and, being out of accord with the law which governs
Actinias, are either a token of the commencement of further growth, or constitute a case
of those numerous abnormalities which occur in the group.
In the other half of the animal occur important gaps in the ground plan just
quoted. The three oral tentacles are in the same place as in the other half, (one is over
the chamber bounded by the directive mesenteries), but five of the intermediary
tentacles and two of the peripheral are wanting. The single intermediary tentacle
occurs in the region bordering on the directive mesenterial chamber just mentioned ;
this region is normally arranged, the peripheral tentacles being also present on it. As
with the tentacles on the disc, so also the marginal ones exhibit great irregularities ; their
number amounts to twenty-eight ; in size their relations are also variable, so that the
rule, that larger and smaller tentacles alternate, is in places infringed.
The peculiar results of a macroscopic examination induced me to cut out a sextant
of the animal for a closer study by means of sections, choosing that sextant of the
normal side which contained the directive septa, and which only departed from the
regular scheme of the Hexactinise in the presence of two supernumerary tentacles. The
results were, that the mesenteries are grouped in pairs by the arrangement of their
muscles thus, — one pair of directive mesenteries, and four other pairs, all of which reach
to the stomatodseum. Of these four pairs I reckon one in the second cycle, two in the
third, the remaining pair being developed asymmetrically and repeating the irregularity
already noticed in the tentacles.
From the intra-mesenterial chamber of the directive septa are evaginated three
12 THE VOYAGE OF H.M.S. CHALLENGER.
tentacles ; besides the marginal, one oral tentacle and one intermediary are placed
on the disc. In the intra-mesenterial chamber of the second order the former (oral) was'
wanting ; in those of the third order, the intermediary tentacle was also wanting, or
rather was replaced by a peripheral tentacle. From all the inter-mesenterial chambers,
and also from the supernumerary intra-mesenterial chamber, spring only marginal
tentacles.
Histologically our Corynactis is closely related to the Corallimorphi. The
mesogloea is homogeneous, branched stellate cells are richly scattered in it, while the
modified bladder cells, which occur in Corallimorphus obtectus, are wanting. Beneath
the endoderm runs a fibrous layer, sometimes closely under it, sometimes separated from
it by a homogeneous layer, giving off bundles which run to the endodermal surface.
The musculature of the oral disc and tentacles is weak and ectodermal ; there is no
special sphincter, and the mesenteries are provided with only weak muscles on both sides.
On the other hand, I was surprised at the occurrence of longitudinal muscles on the
outer side of the body-wall. They are not very strong, and are mostly composed of
short spindle-shaped fibres, the lamella being always slightly pleated here and there.
This discovery made it necessary to study Corallimorphus obtectus anew, with reference
to the body-wall. The epithelium having been preserved only at exceedingly few spots,
constituted the reason why I had not previously observed the muscle, but a renewed
study yielded figures by which I arrived at the following definite opinion, based on
numerous preparations from different parts of the body.
At the basis of each epithelial cell lies a small body, staining in carmine, and
resembling, in sections accurately transverse to it, a muscle fibril. If the section be
taken at an angle of about 30°, these bodies appear elongated and somewhat spindle-
shaped ; but I have seen no such obvious longitudinal fibres as in Corynactis. I am
therefore of opinion that Corallimorphus possesses longitudinal muscles, but that they
are extremely rudimentary.
The observation of ectodermal longitudinal muscles on the body-wall of Corynactis
is an exceptionally interesting discovery. Among all Anthozoa, we know of a similar
condition in Cerianthus alone, and, as I may here mention, anticipating future
investigation, in Arachnactis, a genus very closely allied to Cerianthus: while in
the typical Anthozoa the ectodermal musculature is confined to the tentacles, the oral
disc, and the stomatodseum. On the other hand, all Hydroids in the hydra-form (i.e.
Hydroid-polypes and Scyphostoma;) possess ectodermal longitudinal muscles of the
body-wall, which are prolonged directly into the tentacles and oral disc (peristome).
We have here, throughout the whole body, circular muscles on the endodermal side, and
longitudinal, i.e. radial, on the ectodermal.
On the ground of previous researches on the sexual organs, I have published the
view, since defended by Gotte, that the Scyphomedusse are ancestral forms of the
REPORT ON THE AGTINIARIA. 13
Anthozoa, the development of radial (mesenterial) folds which commences in the
former being further advanced in the latter. In this case the ectodermal longitudinal
musculature of Corynactis and the Cerianthi would be, as it were, heirlooms from the
Scyphostoma?. Both genera would thus retain an ancestral character no longer to be
found elsewhere among Anthozoa, with which would agree that both genera must on
other grounds be placed near to the original ancestor of the group. Of all Hexactinise,
the Corallirnorphidse are, next to the Halcampce,, the most primitive ; the Cerianthidse,
again, must be derived from the extremely primitive Edwardsise.
Family 2, Antheomorphid^e, Hertwig.
Genus Ilyanthopsis, n. gen.
Antheoinorphidse with the tentacles in several rows ; body-wall smooth ; body
goblet-shaped, broadening upwards from the small pedal disc to the broad oral disc.
Ilyanthopsis longifilis* n. sp. (PI. II. fig. 2).
Tentacles very long, pointed, with an obvious terminal pore, ranged in four circlets,
increasing in length from the centre outwards.
Habitat. — Reef of the Bermudas, June 1873. One specimen.
Dimensions. — Diameter of base, 4 cm., of oral disc, 7 cm. ; height, 3"5 cm.
The single specimen, which was well preserved but strongly contracted, in its
shape occupies a middle position between Aiptasia and Ammonia. The base is
relatively small, the body not very high, but broadening out conically towards the
mouth. The body-wall being raised in goblet shape over the edge of the oral disc, the
animal possesses a "collar" in the sense of Angelo Andres, and consequently, owing
to the absence of cinclides and acontia, must be reckoned near the Ilyanthidse. From
these it differs in the presence of a well-developed pedal disc, by which it undoubtedly
attaches itself to rocks.
The thin body-wall is smooth, except for transverse wrinkles due to the strong
contraction of the mesenteries. No sphincter is present. The circular muscle-lamella
is, in all parts of the body-wall equally, pleated into muscular lamina?, which are low,
and either not at all or only slightly arborescent.
The tentacles are very numerous, and are arranged in four rows, the oral disc being
free from them in the immediate neighbourhood of the mouth. Since I counted but
160, not all the tentacles of the sixth order can as yet have been developed. The
longest of them were some 4 cm. in length, and 0-5 cm. broad at the base ; the slightly
truncated tip possessed a small pore. In studying the ectodermal muscle-lamellag,
peculiarities presented themselves which suggested the longitudinal muscles on the
outer surface of the body-wall in Cerianthus. The muscular pleats are generally
14 THE VOYAGE OF H.M.S. CHALLENGER.
slightly arborescent, as is shown in PI. II. fig. 2, and arranged close to one another
like the leaves of a book. At the free edge of the pleat the musculature is
interrupted, since here the fibres of the mesoglcea, which serve as foundation for
the muscle-pleat, radiate into the epithelium. For some distance they are united
in a bundle ; they then part, and each fibre individually tends in the direction of the
epithelial surface. The nerve-fibre layer is consequently pierced by fine fibrils, arranged
parallel to and at equal distances from one another. I would have gladly determined
how far the connective tissue fibres reach, and whether they are connected with indi-
vidual epithelial cells or not ; but in thin sections I could only follow them into the
dim granular striated layer of epithelial cells, in which they were no longer distin-
guishable from other fibres. Attempts to exhibit the isolated fibres by brushing and
agitating thin sections, or by maceration in alkali, yielded no result ; and staining with
picrocarmine was also unsuccessful. The latter generally stains the mesoglceal struc-
tures of a deep red, and is therefore peculiarly adapted for exhibiting the mesoglceal
lamina which carries the muscles, but it refuses to differentiate the fibrils. The red
tint is therefore only seen to extend so far as is expressed in the figure by shading ;
the fibrils probably do not stain, but only the cement substance uniting them. The
condition here described may be followed on to the oral disc, inasmuch as the sup-
porting laminae of the muscle pleats here also run out in fibres, and the individual
fibres radiate to the epithelium. I have only further to remark that radial furrows,
shallow and slightly expressed, run from the edge of the oral opening towards the
tentacles.
The stoniatodaeum, in the only specimen which I could examine, was evaginated,
and consequently so tightly stretched that even the siphonoglyphes were almost
smoothed out, and hardly recognisable.
The mesenteries agree in number with the tentacles ; all reach the stomatodseum,
and bear generative organs. The younger mesenteries touch the stomatodseum some-
what further back, and are in other respects less developed than the older ; but their
generative organs are more voluminous than those of the first and second orders.
Stomata in the mesenteries, and acontia, I have not been able to recognise.
Family 3, Actinidje, A. Andres.
Antheadw, Herhvig.
Genus Hormathia, Gosse.
Actinias with broad diffuse endodermal sphincter ; smooth thin body-wall, and
parietal spherules (i.e. marginal spherules placed on the body-wall).
REPORT ON THE ACTINIARIA. 15
Ilormathia delicatula* sp. n. (PI. II. figs. 1, 3, IV. fig. 9).
More than 160 tentacles; parietal spherules tentacle-like, one of the latter to
about every four tentacles.
Habitat. — (?) (Inscription on the label conrpletely soaked away.) Two specimens.
Dimensions. — Diameter of the nearly spherical body, 2'5-3'0 cm.
Gosse has conferred the name of Ilormathia margarithce on an Actinian brought
up by the line of a deep-sea fishing-boat. Having obtained but one specimen, and that
not till some time after death, he could give but an incomplete description ; the most
imjiortant point of which is that on the delicate body-wall, at some distance from its upper
edge, are placed prominences resembling marginal spherules, the number of which is
about ten, and is essentially less than that of the tentacles. In his monograph treating
of the Actinias, Angelo Andres has included the animal among the doubtful genera, as
being of uncertain systematic position. It was therefore very agreeable to me to find
in the Challenger material two Actiniae obviously belonging to the genus Ilormathia,
by the study of which I am enabled both to justify the creation of a new genus, and
also to define accurately its systematic position.
Both specimens were so strongly contracted as to resemble an apple in shape.
The upper part of the body-wall, the pedal disc and the mouth being entirely drawn
in, and the latter covered over, one saw at first only the lower part of the body-wall,
the smooth surface of which was so little characteristic that I came near to ranking the
animal among the undeterminable forms. Only after dividing a specimen longitudinally
did the circlet of parietal spherules come into view, their position being characteristic
of the genus Ilormathia.
The pedal disc is strongly constricted and pleated by the violent contraction.
The body-wall is exceptionally delicate, so that the septa are plainly visible through it,
and is quite smooth. By a circular fold, which recalls to mind the boundary between
body-wall and oral disc, and marks the limit of retraction in a withdrawn specimen, is
bounded a separate invaginable region of the body-wall ; close up to this fold, and on
the side nearest to the oral disc, is placed a circlet of 42 knobs, which are hollow and
beset with nematocysts, and which therefore recall the structure of the marginal vesicles
or "bourses marginales" (PI. IV. fig. 9). They are of different sizes, the largest
generally longer than the marginal spherules ; and are curved in a digitate manner at
the end, so as to present some resemblance to tentacles. The number of tentacles and
mesenteries being about 160, the parietal spherules, as I term these structures, are not
placed, like the marginal spherules, one on each inter- and intra-mesenterial chamber ;
but there is one spherule to about every four chambers, with one of which it is
always in communication, leaving the remaining two or three free.
The marked retractibility of the animal is effected by a sphincter muscle in a
definite region of the body-wall, which, commencing at some little distance from the
1G THE VOYAGE OF H.M.S. CHALLENGER.
parietal spherules, and reaching to the origin of the tentacles, extends therefore over a
airly wide belt. Correspondingly to this broad extension, it is nowhere strongly
developed, and falls under the category of " diffuse " endodermal muscle, the lamella
being most markedly pleated in the centre. Its arrangement is very characteristic, as
a transverse section presents the appearance of numerous closely-packed acinose glands,
excavated in the mesoglcea. In the more central parts — to continue the comparison —
the gland-like crypts are longer and more closely packed than in the upper and
lower parts (PL II. fig. 1).
The strongest development of muscles occurs on the ectodermal side of the disc,
where the supporting lamina rises into high plates, covered by strong fibrils and
richly arborescent. Here and there I have also noticed the plates fusing together, with
a resultant mesodermal inclusion of the muscle fibrils (PI. II. fig. 3). Towards the
tentacles the muscles become weaker.
The tentacles are of a medium length, broad at the base, and drawn out to a fine
point, which is probably not provided with an opening at the tip. The siphonoglyphes
are hardly marked on the stomatod?eum. To the latter, besides the mesenteries of the
first cycle, those of the second and third cycles at least are attached. Their musculature
is in no region strongly developed ; in the specimen investigated nearly ripe testicular
follicles occurred on them.
Family 4, Bunodid^e.
Genus Aulactinia, Verrill.
Aulactinia, sp. (?) *
Habitat. — Simon's Bay, Cape of Good Hope, December 1873; 10-20 fathoms.
One specimen.
Dimensions. — Height in a strongly contracted condition, 2 cm. ; breadth of pedal
disc, 3 cm.
In this place I will devote only a few words to a Bunodidan, of which I reserve
a detailed description till I shall have reviewed a rich supply of species of this
family which has been forwarded to me. The body-wall of the sole specimen lying
before me is thickly beset with thin-walled vesicular outgrowths, which are about
1 mm. in size, show a tendency towards arrangement into transverse and longitudinal
rows, and are so thickly set that the intermediate stouter parts of the body-wall have
a reticulate appearance. The three upper rows of these vesicles (about seventy in
number) are closely packed with nematocysts, and so take on the character of marginal
spherules ; they may be distinguished into a stalk, and a branching head like a cauli-
flower. They recall somewhat those external appendages of species of Oulactis, which
REPORT ON THE ACTINIARIA. 17
have been termed, — I Jo not know for what reason, — tentacles. The tentacles are
arranged in three rows, and more than 200 are present. The endodermal sphincter
is extraordinarily strongly developed, in the form of a ridge projecting into the
ccelenteron.
Family 5, Paraotid^.
Genus Dysactis, Milne-Edwards.
Dysactis crassicornis, R. Hertwig.
Two additional examples of this Actinian have reached me, dredged from a depth
of 55 fathoms at Station 313. One had died in an expanded condition, so that the
tentacles wrere in better preservation than in the specimens previously studied ; from
this I am enabled to determine some further characters of these organs.
In many cases terminal pores, which I was before unable to discover, were easily
recognised on a surface view ; I have therefore re-investigated the older material, and
was able with some trouble to prove the existence of openings by injecting air into
them under water.
Further, in the well-preserved tentacles, comes strongly into view a characteristic
which I had previously figured (former Report, pi. vii. fig. 12), but had not introduced
into the text ; the tentacles are longitudinally striated, so covered with longitudinal
ridges and furrows as to recall a fluted pillar ; in section this is still more prominent.
At tolerably regular intervals the mesogloea rises in high ridges (PI. II. figs. 6, 7),
and at these points the mass of muscle lying in it is correspondingly increased.
The muscles therefore form in transverse section a continuous ring, which in the
region of the ridges of mesogloea is drawn out into cusps. At the base of an especially
strong tentacle I counted twenty-two longitudinal ridges, of which, however, some only
reach to the tip.
Family 6, Liponemid.e, R. Hertwig.
Genus Liponema, R. Hertwig.
Liponemidse with weak endodermal sphincter ; the body-wall marked by longi-
tudinal furrows, without marginal spherules ; stomidia very numerous.
Liponema multiporum, R. Hertwig (PI. I. fig. 13, PI. II. fig. 4).
Stomidia, several hundreds in number, distributed in several cycles, and scattered
over the whole oral disc ; body apparently cup-shaped, broadening out from the small
pedal disc upwards to the wide oral disc.
Habitat. — (a) Station 305a, January 1, 1876; 120 fathoms. One specimen.
(b) Station 147, December 30, 1873 ; 1600 fathoms. One specimen.
(ZOOL. C'HALL. EXP. — TART LXXIII. — 1888.) Dddd 3
18 THE VOYAGE OF H.M.S. CHALLENGER.
Among: the Actinia? with degenerate tentacles. I described in my former Challenger
Report a new species, in which the extent of retrogression of the tentacles can be
recognised in a degree attained by no other form. I was compelled to dispense with a
detailed description of its structure, since the only specimen at my disposal was on
the one hand much mangled, and on the other rendered so brittle by preservation in
chromic, acid that it could not be methodically investigated. I am glad to be in a
position to fill up the deficiency by means of two specimens found in the supplementary
material, both well preserved, although considerably altered in shape by violent con-
traction. In both cases, as in the example previously described, the stomatodaeum is
so much evaginated as to take the place usually occupied by the oral disc, the latter
falling outwards from this point like a body-wall (PI. I. fig. 13). On the other hand,
pedal disc and body-wall are alike deeply retracted on the lower side. The body-wall
forms a cup like the shell of a Patella, the pedal disc projecting into the cup somewhat
like the body of the Patella. In so marked a de-formation, dimensions can with
difficulty be given, and can serve only for approximate orientation. In the larger of
the two specimens (from 120 fathoms at Station 305a), the pedal disc had a diameter
of about 2 cm., the distance between the edge of the oral disc and the mouth reached
2-5 cm. ; the length of the stomatodseum was at most places l-5 cm., and at the
siphonogiyphes more than 2 cm. The corresponding dimensions of the smaller
example (Station 147 ; depth, 1G00 fath.) are essentially less, — diameter of pedal disc,
0'07 cm. ; radius of oral disc, T2 cm. ; length of the stomatodseum, TO cm. From the
nature of the contraction may be inferred that in both cases the dimensions of oral
disc and stomatodaeum are excessive, as the result of evagination, while those of the
pedal disc are too small.
On the pedal disc are about 160 radial furrows, of which, however, only a
proportion reach the centre, the rest dying out sooner or later. The ridges between
the furrows are somewhat toothed, in the manner formerly described by me as occurring
in Polyst&midium and Polysiphonia. In the centre of the pedal disc lies a pit about
the size of a pin's head, which cannot be proved to be an opening.
On the exterior of the body-wall also, similar ridges, alternating with furrows,
run longitudinally from pedal to oral disc ; their number is greater, being close on 400 ;
they differ in size, some few of less considerable development rising between every two
of the stronger ridges. At the edge of the oral disc they all pass into a strong circular
ridge, which forms the sharp boundary between body-wall and oral disc.
The pedal disc and body-wall possess on their inner surfaces the circular muscle-
fibre layer occurring in all Actinias ; on the body-wall this is strongly pleated, and the
more so, the nearer we approach to the upper edge. In the immediate neighbourhood
of the edge the pleating is so marked that one may term it a sphincter ; it causes here
the circular ridge mentioned above as occurring at the upper edge of the body-wall
REPORT ON THE ACTINIARIA. 19
(PL II. fig. 4). This circular ridge appears in transverse section as a pushing out of
the body-wall, the circular muscle exhibiting a very different structure in the different
regions. At the base of the organ the pleating of the muscle-lamella is insignificant,
indeed weaker than at other points of the body-wall, but at both edges of the
evagination it is exceptionally strong, and more especially 30 at the boundary of the
oral disc. When the section comes to the actual spot on which one of the stomidia
is set, the inner sphincter — as we call the nearest muscular pleating — is beautifully
recognisable as a ridge projecting inwards, into the axis of which protrudes a mesoglceal
ingrowth. From this axial ingrowth are given off on both sides richly branching
mesoglceal lamellae, clothed by powerful muscle-fibres in transverse section. At the
remaining points, where the oral disc presents no stomidia, the sphincter is less clearly
bounded, and resembles more the outer sphincter, which is essentially nothing but an
approximation of muscular folds at two closely-adjacent points.
Relatively to the size of the animal, both sphincters are weak ; a consequence of
this is the fact that they have not drawn up the body-wall over the mouth disc, but
that stomatodaeum and oral disc have rather been pressed outwards.
The oral disc recalls in appearance a toadstool, having a faintly flesh-coloured
surface, covered by whitish, slightly elevated spots. These spots are the stomidia or
tentacles, which are distributed nearly up to the mouth, leaving but a narrow strip
free. Between the stomidia the radial furrows run in undulating lines. Their number
is difficult to determine, but may amount to about 400.
The stomidia are openings in the oral disc, surrounded by a slightly developed
ridge, and projecting a little above the surface ; roughly speaking, they are distributed
uniformly over the oral disc, or allow only of a vague distinction into several zones.
Of these zones one is peripheral, set close to the edge of the oral disc ; one is
central, not far from the oral opening ; and two intermediate zones are placed betwTeen
them. The openings increase in size from without towards the centre, and at the same
time undergo an alteration of shape ; in the peripheral zone they are like radially-set
slits, with a long axis of 0"7-l"5 mm. ; in the intermediate zones they are circular,
with a diameter of 1-2 mm. ; and in the central they again form slits of 2 "0-2 "5 mm.
in the longer diameter, but are here placed at right angles to the radii.
The structure of the stomidia can best be exhibited by figures of transverse
sections. Each stomidium completely occupies the intermediate space between two
neighbouring mesenteries, and forms a tube, opening peripherally by a wide mouth.
The walls of the tube appear in section to be direct continuations of the adjacent septa ;
morphologically their lower part is to be regarded as oral disc, their ujiper part as
rudimentary tentacle; accordingly, they exhibit below the numerous muscular pleatings
which at other points cause the radial ridges on the oral disc, while above these pleat-
ings are absent. A remarkable structure is a small circular fold projecting below into
20 THE VOYAGE OF H.M.S. CHALLENGER.
the lumen of the tube, and constricting it like a diaphragm. This doubtless serves to
close the tube, since it is covered by a marked layer of muscle fibres, running circularly
round the opening.
Where no stomidium is placed, the oral disc exhibits on its ectodermal side a
thick layer of radial muscle fibres, arranged in simple lamella?, which, at most, branch
but once. The lamellae being higher midway between two mesenteries than elsewhere,
the radial ridge-like thickenings of the oral disc are the result.
With reference to the relations between the stomidia and the inter- and intra-
mesenterial chambers, in my former publication I expressed the opinion that an
intra-mesenterial chamber might carry more than one stomidium, Liponema thus
approximating to the Corallimorphidae ; an opinion which I can now designate as
erroneous, on the ground of more accurate investigation. Each intra-mesenterial
chamber possesses but one stomidium, which is the more closely approximated to the
centre of the oral disc, in proportion as its adjacent septa are of older formation.
The stomatodseum is brownish-violet in tint, and 2 cm. long ; on it are placed
the marked siphonoglyphes, about 1*5 cm. in breadth, projecting considerably at the
lower part of the tube, where they pass into the boat-shaped stomatodseal cone. They
are bounded by two stout, transversely pleated, lips. Further, the stomatodaaum is
marked by about 200 longitudinal folds, of which some 80, by their stronger build,
deserve the name of primary folds. Between every two primary folds lie, in many
cases, two secondary folds ; but at some places one only may occur, or they may be
entirely wanting.
The number of mesenteries was determined by the method before mentioned, that of
cutting out a sextant of the animal and studying it closely anatomically. I found six
cycles, in all therefore 192 pairs of mesenteries. In the first four cycles all the.
mesenteries reach the stomatodseum, though those of the first two cycles only are
attached to it for its whole length ; they all possess wide openings near the edge of
the lip (internal mesenterial stomata), and their muscular nature so far preponderates
that only those of the fourth cycle carry generative organs. In this respect these
mesenteries of the fourth cycle agree with those of the fifth and sixth, but the muscular
development of the latter is considerably inferior to that of the others. The mesenteries
of the sixth cycle are practically nothing else than small genital folds, projecting but
slightly into the ccelenteron, and never provided with mesenterial filaments.
Of the generative organs I found exclusively the testicular follicles, containing
spermatozoa in parts ripe, in parts only commencing to develop.
It is possible that in this animal a further growth takes place, with the formation
of new mesenteries ; this I infer from the great number of stomidia. In the sextant
investigated they amounted to about 120, or to 700-800 for the whole animal. Since
only about 196 intra- and inter-mesenterial chambers are present, and each of the
REPORT ON THE ACTINIARIA. 21
former possesses but one stomidium, the latter apparently must be provided each with
two or three, — an inference confirmed by dissection. Since it is the rule amongst
Actinia? that the development of tentacles precedes that of mesenteries, we can also
infer in this instance from the plentiful development of stomidia, an imminent addition
to the mesenteries.
Genus Aulorchis, n. gen.
Liponemidae, whose generative organs are modified into a tube perforating the oral
lip ; gonidial grooves on both sides drawn out into a long ear-like cone.
Aulorchis paradoxa* sp. n. (PI. I. figs. 9, 10 ; PI. III. figs. 2-6 ; PI. IV. figs. 1-6).
Stomidia arranged in two alternating rows, approximately sixty in number.
Habitat— Station 299, December 14, 1875; lat. 33° 31' S., long. 74° 43' W.;
depth, 2160 fathoms. One specimen.
Dimensions. — Height, 4 cm. ; greatest breadth (measured about half-way up the
animal), 3 cm.
Among the accessory Challenger Actinias occurs this form, of great interest as
enlarging by a new genus and new species the group of forms devoid of tentacles.
Unluckily, I have had but the one solitary specimen for study, and even this was badly
preserved, and had apparently suffered much from the dredge. It was exceedingly
contracted ; oral and pedal discs were externally unrecognisable, since both ends of the
body-wall were closely drawn together. As a natural result of this condition, I have
not been able to clear up many important points of the organisation so well as I could
have wished. For investigation, I divided the specimen longitudinally, and dissected a
sextant with scalpel and scissors, arriving at the following results.
The strongly contracted, and therefore small, pedal disc exhibits indistinct radial
brownish wrinkles and furrows, and is sharply marked off from the body-wall, the
surface of which is smooth. The latter is of a whitish tint, and of inconsiderable thick-
ness, only here and there becoming more powerful, but never forming hooks or papillae.
Its consistence is less firm than that of cartilage, but considerably more so than that of
Medusan mesogloea. The tissue is of a fibrous nature, composed of very fine fibrils,
which are generally interlacing and reticulate. At many points, however, they are
thicker and bound together in more parallel series, so that cords and lamellae are
formed, which, though staining brilliantly with carmine, are not sharply differentiated
from their surroundings. These lamellae are ranged parallel to the two surfaces, and
run constantly closer to one another till a firmly united mass of fibres is formed just
below the epithelium. At other points, however, the fibres are more loosely plaited, so
that spaces remain between them, which are filled up by homogeneous mesoglcea.
In some places I detected hollow spaces in the tissue, which were devoid of an epithelial
22 THE VOYAGE OF H.M.S. CHALLENGER.
lining. They occur also in the mesenteries, the stomatodseum, and the oral disc, and
may perhaps be caused by inadequate preservation.
In the upper part of the body-wall lies, close under the endoderm, a mesodermal
sphincter muscle, its length amounting to about 1 cm., while its greatest breadth
reaches 5 mm. at the upper end, from which point it gradually thins out. It is of
interest from several points of view ; in the first place, the muscle-fibres are abnormally
strong ; consequently the muscle-bundles are formed of but few elements, and consist
in many cases only of two to four. Again, the individual tracts are so far from running
parallel to one another, that in a longitudinal section many bundles are cut absolutely
transversely, others obliquely, and others for long stretches siqjerficially ; thus an
appearance of extremely entangled fibres is presented (PL III. fig. 3a).
Finally, Atrforchis affords proof of the endodermal origin of the mesogloeal muscle-
bundles, as we find on the endodermal side every transition from the mesogloeal bundles
to the endodermal layer of circular fibres ; in one place the bundles lie close under the
fibrous layer, at another are in communication with it by a broader or narrower band ;
finally, we find slight infoldings of the endodermal muscle-layer (PI. III. fig. 3b).
The stomidia lie in two alternating rows between the edges of the mouth and of the
body-wall, somewhat nearer to the former ; they are about sixty-four in number (thirty-
two between two pairs of directive mesenteries). The stomidia of the inner row are
larger than those of the outer ; the smallness of the latter producing the impression,
that they have just been formed, and that a further increase of their number is taking
place. Radial ridges on the oral disc start at the edge of the body-wall and run up
to the individual stomidia.
Transverse sections through the oral disc exhibit a strong mesodermal musculature ;
this is interrupted along the lines of mesenterial insertion, and falls therefore into
marked radial bands which cause the radial ridges of the oral disc. The individual
muscle-bundles contain a few strong fibres, and are so separated from one another by
mesogloeal sheaths, stout or slight, that the lines of mesoglcea form dendritic figures
springing now from the ectodermal, now from the endodermal side (PI. III. fig. 2).
The mesoglcea sends into the ectoderm arborescent supporting offsets, on which
to my surprise I was unable to find muscle-fibres. It seems as if in Aulorchis
the ectodermal musculature is completely wanting ; I would gladly have expressed
something definite on this point, had the histological condition of the animal not
been so indifferent ; but the ectoderm, where present, was unfortunately reduced to a
detritus, in which no structure could be detected.
In order to demonstrate how the stomidia penetrate the thickness of the oral disc,
I have drawn two figures, in the one of which (PL III. fig. 4) are seen the openings of
the tube to the exterior and to the coelenteron ; in the other (fig. 5) the section passes
through a spot where the stomidial tube is closed at both ends, whence it may be
REPORT ON THE ACTINIARIA. 23
inferred that its diameter is here considerably greater than that of the two openings.
The radial mesodermal muscle-fibres pass into its walls with a longitudinal trend.
On the stomatodamm are placed the two siphonoglyphes, which are of a very
characteristic appearance, as being more powerfully developed than in any Actinia
which I have as yet seen ; each projects over the mouth edge and upwards with
two long ear-shaped cones. The groove itself is correspondingly deep and broad,
pleated, and of a cartilaginous consistence. Between the two siphonoglyphes run
on each side about ten strongly-marked longitudinal ridges, terminating in rounded
knobs on the lip.
In investigating the mesenteries, I could at least prove their arrangement in pairs,
but could not convince myself that the Hexactinian symmetry was carried out. Neither
by microscopic preparations of a sector, nor by dissection of individual mesenteries, could
I arrive at a definite law of arrangement ; this point therefore requires investigation.
The mesenteries dissected bore no generative organs ; these appeared to me to be
confined entirely to one mesentery, and to possess a tubular structure unparalleled in the
whole class of Anthozoa, a fact which decided me to choose for the genus the name
Aulorchis. Even before dissection it had struck me that at a spot on the edge of the
lip, and by a pore specially present for the purpose, was the openiug of a cylindrical
organ ; this organ had obviously once been longer, as at its end a fracture was clearly
recognisable. By splitting up the opening and the adjacent stomatodseurn, the organ,
which I will term in future, for reasons to be mentioned, the genital tube, could be
clearly followed into an inter-mesenterial chamber (PI. I. fig. 9). It meets one of the
complete mesenteries, lies at this point embedded in the tangle of mesenterial coils,
and, as appeared later from sections, ends at the mesentery in a horseshoe-shaped curve.
The curved portion was firmly united with the mesentery. Transverse sections yielded
further conclusions relative to its structure ; but, unfortunately, owing to bad preserva-
tion, no exhaustive account of this is possible. For instance, I have not been so
fortunate as to determine how far the structure of the genital tube can be referred to
that of the ordinary Aetinian ovary (PL IV. figs. 1-6).
The genital tube is superficially clothed by epithelium, which is limited exter-
nally by a border resembling a cuticle, but perhaps produced only by mucous
secretion; then follows the mesogloea with the ova embedded in it; internal to these
lies a cavity, more or less spacious according to the mass of the ova. The mesogloea is
divided by a narrow granular layer into inner and outer zones, which here and
there, by failure of the intermediate layer, join together. The outer zone is narrow,
and exhibits what appear to me to be circular muscle -fibres referable to the
epithelium, which in longitudinal sections through the organ (fig. 3) resemble narrow
laminae placed close together. The state of preservation was inadecpiate for the
determination of the histological character of the granular median layer ; in transverse
24 THE VOYAGE OF H.M.S. CHALLENGER.
section it gave the impression of a disintegrated epithelium, in longitudinal it resembled
a loose connective-tissue. This layer is important as containing small, spherical, deeply-
staining cells, which I regard as young ova. The masses of ova are in parts so con-
siderable as to present the appearance of mosaic, if part of the wall of the genital tube
be cut out, stained, and viewed from the surface (fig. 2). Next comes the second zone
of mesoglcea, the layer of most importance, since ova of various sizes are embedded in
it. Some of these are certainly connected with the superficial epithelium ; this con-
dition, I believe, occurs in all ova, and is effected by the fibre-arrangement characteristic
of Actinian ovaries, of which remnants only could here be detected (fig. 1).
The lumen of the tube was mostly filled by a cell-detritus, but at some points was
lined by a clearly ciliated epithelium (figs. 4, 6) ; I reckon therefore the lumen as a
ciliated canal, serving for the transit of ripe ova and perhaps also of embryos, and
opening to the exterior outside the oral disc. The ripe ova appear to lie on the floor of
the tube, since here I found compact masses of a finely granular substance, appearing
to me to resemble ova.
As to the distribution of the ova in the genital tube, I have the following facts to
add : the smaller ovules are met with in sections through the upper part of the tube,
forming a ring, on the one side of which the generative elements are more closely
packed than on the other. This lop-sided development of sexual cells is expressed
more obviously lower down, where on one side of the section they are entirely
wanting, the ripening ova being only present in the other half.
With regard to the connection of the genital tube with the body of the Actinia, I
have arrived at no positive results. At the pore, the organ merely perforates the oral
lip without being attached to it, as I can assert both from macroscopic dissection and
transverse sections ; while at the lower end I have discovered no intimate connection
with the mesentery ; what I saw there was only an epithelial adhesion, not a transition
from the mesoglcea of the mesentery into that of the genital tube. Such a connection,
however, must certainly occur at this point.
From my description it may be recognised that Aulorchis is one of the most
interesting Actiniae, and that it would be very desirable that a richer material of it
should be acquired by fresh Deep Sea investigations.
Family 7, Phellid.e.
Genus Phellia, Gosse.
Phellia spinifera, n. sp. (PI. II. figs. 8, 9).
The bark-like part of the body-wall is bedecked with thorn-like pointed knobs,
distributed more richly on the upper than on the lower parts.
REPORT ON THE ACTINIARIA. 25
Habitat. — (a) Station 311, January 11, 1876 ; depth, 245 fathoms. Three speci-
mens, (b) Station 320, February 14, 1876 ; depth, 600 fathoms. One specimen.
Dimensions. — Length of the contracted animal, 2-5-3-2 cm. ; breadth, 2-5-3'5.
At first I was inclined to refer the three specimens from Station 311, which were
seated on Molluscan shells, and the single specimen from Station 320, to Phellia
pectinata ; for they possessed the characteristic appearance of the body- wall, resembling
the tunic of Cynthice, while the upper indrawn part of the wall presented the ridged
surface which has been already figured. I was, however, persuaded to a closer study
by observing some points of divergence in the structure of the peripheral region of
the body-wall. The transverse and longitudinal ridges are wanting, instead of which
occur knobs, resembling those of Cereus spinosus ; these start with a broad base, and
terminate in a slightly truncated tip ; they are distinguished from the body- wall, which
is nearly white, by a brownish tint, and may amount to 200 in number, distributed
more abundantly on the upper than on the lower regions of the body-wall. The upper
knobs are as much as 0"25 cm. long, and are more strongly developed than the rest;
they become gradually smaller below, and finally appear only as fine grains. Such an
arrangement of the knobs in series, as exists in Bunodes, does not occur.
The mesoglcea of the body-wall is so extraordinarily stiff as to cause some trouble,
before good sections of the sphincter can be effected. The latter is essentially constituted
as in Phellia pcctinata, so that reference to the description given under that species is
.sufficient. In position it is considerably nearer to the ectoderm than to the endoderm.
The oral disc and stomatockeum are of a brownish violet (partially altered in the
alcohol), the former lighter in tint than the latter. On the stomatodaeum the two
siphonoglyphes, which are not pigmented, and are consequently of a whitish yellow,
strike the eye on opening the animal as two broad, sharply-marked, stripes. They are
only distinguished from their surroundings by this difference of colour, since they are
flush with the rest of the stomatodgeum. They are crossed by transverse folds regularly
arranged, which are continuous over the rest of the stomatodasum. Further, the
stomatodgeal cone is hardly expressed at all, and the longitudinal furrows, which so
commonly run parallel to the siphonoglyphes between the mesenterial insertions, are
wanting.
For the characterisation of the species the condition of the musculature of the oral
disc is also of importance ; it exhibits two methods of formation. In the one case it is
purely ectodermal and markedly pleated, the pleats running parallel to one another,
and only slightly arborescent (PI. II. fig. 9). At other points (fig. 8) the arborescence
is very considerable, the individual branches anastomosing with one another ; the
musculature thus becomes partly mesoglceal, and a very obvious and stout muscle-
layer arises. The muscle-fibres are here, as in the sphincter and the powerfully
developed laminae of the retractors, of exceptional thickness.
(ZOOL. CHALL. EXP. — PART LXXIII. — 1888.) Dlllld i
26 THE VOYAGE OF H.M.S. CHALLENGER.
All the mesenteries are unusually muscular ; the primary mesenteries are sterile,
and reach to the stomatodaeum, while the secondaries are incomplete but bear generative
organs. I observed a few acontia ; cinclides, on the other hand, are wanting.
In conclusion, I might refer to the possibility that Phellia spinifera may be only a
variety of Phellia pectinata. In the sole example from Station 320, the spinose knobs
were developed only on the upper part of the bark-like body-wall, and even here not
abundantly ; so that its appearance is intermediate between the characters of Phellia
pectinata and Phellia spinifera. In spite of this, I have retained the separation of the
two species, because the musculature of the oral disc of Phellia pectinata does not yield,
on further study, the characteristic appearance drawn in PI. II. fig. 8. In this respect,
the transitional form agrees with the type of Phellia spinifera.
Family 8, Amphianthid.^, E. Hertwig.
Genus Amphianthus, E. Hertwig.
Amphianthus ornatum* n. sp. (PL I. fig. 8).
Body-wall beset with numerous (about 26) longitudinal rows of papillae ; the latter
are for the most part recognisable by the naked eye, and are not arranged in transverse
series.
Habitat. — (a) Station 56, May 29, 1873 ; depth, 1075 fathoms. One specimen.
(b) Station 241, June 23, 1875; depth, 2300 fathoms. Three specimens, (c) Station
244, June 28, 1875 ; depth, 2900 fathoms. One specimen.
Dimensions.- — Height, 0*2-0-5 cm. ; length of the pedal disc, 0"3-2 cm.
The five specimens which I describe under the name of Amphianthus ornatum
have on the one hand many points of resemblance to Amphianthus bathybium, on the
other to Cylista (Bunodes) minuta ; with the latter they agree in the form of the
papillae, but differ from it in the divergent shape of the body and in characteristics of the
family Amphianthidae, as also in the absence of acontia ; with the former, on the other
hand, they tally in general habits, but exhibit a divergent condition of the body-wall.
Amphianthus bathybium possesses small papillae, recognisable only with the aid of a
lens, and arranged in small groups, with a tendency to transverse series. In Amphi-
anthus ornatum, however, they are large, and comparatively isolated in position ; they
form about 20-30 longitudinal rows, which die out sooner or later at some distance
from the lower end of the body-wall. The papillae are not all of one size ; indeed, it
even happens that rows of larger and smaller alternate.
In the very young specimen from Station 244, only twelve rows of papillae were
present, all most regularly distributed on the periphery of the body, and all of essentially
similar structure, since both in the size and number of the papillae the individual
REPORT ON THE ACTINIARIA. 27
rows were closely identical. The number of the papillae varies between six and seven.
From this observation it may be inferred that with increasing growth an addition to the
rows of papillae occurs, proportional to the additions to the pairs of mesenteries. In
connection with this is the fact that the rows of papillae correspond to the intra-
mesenterial chambers.
Two examples of the present species were taken from the same locality as the one
example of Amphianthus bathybium, Station 241. This renders it necessary to weigh
the possibility that the differences which have been made of importance, are perhaps
only of secondary significance, and that all the specimens may be referred to one species.
Owing to the limited material, the cpiestion could not well be decided.
From the minuteness of the organism, anatomical investigation could only be
effected by means of longitudinal and transverse sections ; to this purpose I devoted
two complete examples, the one from Station 241, the other from Station 244, besides
quadrants of specimens from Stations 241 and 56. It resulted that the papillae were
proved to be solid outgrowths of the body-wall, and, like it, consist of an extremely
fibrous mesogloea. The fibres are generally interlacing, as is for the most part normal
among Actiniae, so that the tissue appears finely granular ; they also here and there
show a tendency to arrangement into bundles. In transverse sections, therefore, a
reticulate figuring appears round the endodermal lining ; this can be rendered clearer
by staining, when it appears that small branches of the fibrils cross the course of the
rest of the fibrils in a longitudinal direction. Similarly, one sees numerous radial fibres
also in the peripheral parts of the body-wall, and a corresponding radial striation is
thus produced.
The sphincter is completely embedded in the connective-tissue of the body-wall,
and consists of small mesoglceal muscle-bundles composed of few, but powerful, fibres.
In some places only two or three fibres are united in a bundle, or a single fibre even
may run in the connective-tissue. The individual bundles are enclosed in such numbers
in the mesogloea as to be separated from the two epithelial surfaces by only a narrow
layer. In transverse section, the muscle in most cases' forms a club-shaped figure, being
of weak development below and broadening out strongly upwards ; this increase in
breadth is so considerable that the whole upper end of the body-wall is strongly thickened.
Even in the youngest specimens the sphincter was completely formed, and inclosed in
the mesogloea. As it is separated from the endodermal circular musculature by the
insertion of a layer of connective-tissue, it seems that in the course of further growth
the bundles can only increase by division of the bundles of fibrils.
The musculature of the oral disc and tentacles is purely ectodermal, but very
markedly pleated. The number of tentacles corresponds to the number of mesenteries,
and this is different in the different individuals investigated. In the youngest specimen
from Station 244, the two first cycles were already formed, and of the third traces
28 THE VOYAGE OF H.M.S. CHALLENGER.
seemed to me to be recognisable in that inter-mesenterial chamber which is equidistant
from the ends of the transverse and sagittal axes. With this agreed the distribution
of the tentacles ; they were about equal in number to the mesenteries, and amounted to
more than twenty-four, i.e. the first three cycles and some tentacles of the fourth were
present, and those of the fourth cycle lay at points corresponding to the inter-mesenterial
chambers above mentioned. In the other specimens, between forty and forty-eight
mesenteries were present in the whole circuit of the body-wall, so that here the fourth
cycle was nearly complete. The number of mesenteries was still greater in the angle
between pedal disc and body- wall, the point where mesenterial growth is first recognisable
in Actiniae. This part being very transparent, the number could be determined with
approximate accuracy, and reached to nearly a hundred.
From the fact that in places the mesenteries were discontinuous in transverse
section, I infer the existence of mesenterial stomata. On the other hand, I could not
demonstrate acontia ; generative organs (testicular follicles) I saw only in one specimen,
and, as they were at all points adherent to the mesenteries, I could not determine
whether they were present on all mesenteries, or were wanting on the primaries.
Directive septa and siphonoglyphes were distinguishable on all four specimens, but
only in two examples, namely the smallest and largest, could I accurately determine the
position which agrees with the typical attitude of Amphianthidge. The sagittal axis
of this Actinian is at right angles to the long axis of the body on which it has fixed
itself, or, in other words, the lengthening of the animal takes place in the direction of
the transverse axis.
With tolerable certainty I can at length assert that only the mesenteries of the
first order are complete.
With the sole example of Amphianthus ornatum from Station 56 was associated
another Amphianthidan, externally so little characterised, that I decided not to describe
it. It possessed a smooth body- wall, which was pleated only as the result of contraction ;
the pedal disc was 1'5 cm. long, and the total height 0'4 cm.
Family 9, Ilyanthidje, Gosse.
Genus Halcampa, Gosse.
Halcampa kerguelensis* n. sp. (PI. II. fig. 5).
Tentacles devoid of longitudinal furrows, pointed ; circular muscles of the body-
wall weak ; retractor muscles of the mesenteries powerfully formed.
Habitat. — (a) Station 149 a, Betsy Cove, Kerguelen, January 10, 1874; depth,
25 fathoms. Ten specimens, (b) The same locality ; 25-30 fathoms. One specimen.
(c) Station 149 G, off London River, Kerguelen, January 29, 1874; 110 fathoms.
REPORT ON THE ACTINIARIA. 29
Three specimens, (d) Station 149 J, off Cumberland Bay, January 29, 1874; 105
fathoms. Three specimens, (e) Station 149 H, off Cumberland Bay ; January 29, 1874 ;
127 fathoms. One specimen.
Dimensions. — Length, T5-2-5 cm. ; greatest breadth, 0-7-l'0 cm.
On an external inspection I was inclined to identify this species with Halcampa
clavus, which it strongly resembles. The preparation of transverse sections, however,
caused me to abandon this view, and a more accurate study produced a number of
points of divergence, which I will briefly enumerate.
1. The tentacles, though twelve are also present in this species, are essentially
longer than in the other, and end in a fine point. The two longitudinal furrows which
occur on them in Halcampa clavus, can be recognised neither superficially nor in
transverse section.
2. The circular muscles of the body- wall are weakly developed ; the laminaa which
they form are not so striking as in Halcampa clavus; and they project into the
coelenteron at greater distances from each other. The sphincterdike enlargement of
the circular muscledayer is wanting.
3. On the stomatodseum the marked projections, which designate the insertions of
the mesenteries, are absent.
4. In the mesenteries the muscle-lamina is pleated in a most complicated manner,
so that in transverse section it exhibits an abundant arborescence. The centre of the
muscle forms a sort of tree (PI. II. fig. 5), a thin lamina starting outwards from the
mesentery, and branching like the top of a tree. This whole region is usually marked
off by an indentation from the adjacent parts, the mass of muscle being thus divided
into three sections.
Genus Halcampella, Angelo Andres.
Ilyanthidse with six powerfully developed pairs of mesenteries, but with numerous
rudimentary mesenteries, and numerous tentacles.
Halcampella maxima* n. sp.
Tentacles small, approximately 46 ; body devoid of longitudinal furrows ; its
surface partly bark -like, partly somewhat incrusted ; the polyp of considerable size.
Habitat. — Station 209, Zebu, Philippine Islands, January 22, 1875 ; 95 fathoms.
Six specimens.
Dimensions. — Length, 8-15 cm. ; greatest breadth, 2-3 cm. ; breadth at narrowest
point (near the pedal disc), 0-4-r2 cm.
In all the specimens the body is a lax thin-walled sack ; its diameter is least at the
posterior end, which is stalk-like and rounded off, but anteriorly it bellies out, contract-
ing again in the region of the oral disc. With the exception of the largest, all the
30 THE VOYAGE OF H.M.S. CHALLENGER.
specimens are so contracted that tentacles, oral disc, and upper part of body-wall were all
drawn inwards together; in the largest, however, part of the tentacular crown protruded.
The surface of the body-wall is incrusted with sand-grains, so that at first sight
I was inclined to take the animal for a Sphenopics. The sand-grains are not, however,
embedded in the mesoglcea, but adhere to the cuticle of the ectodermal epithelium, so
that they can easily be removed by scraping. At the anterior end they are more
sparse, and are practically absent on the upper third. This part of the body-wall
assumes a different appearance to the rest, being more leathery or bark-like, and
traversed by rough longitudinal furrows. The bark-like appearance is produced by
the cuticle, which is strongly developed, and of a brownish tint, resembling that of
Phellia pectinata and Tealia hunodiformis. A fairly sharp boundary marks off from
the rougher part of the body-wall a strip about 1 cm. wide, which adjoins the oral
disc and wreath of tentacles, and which has a completely smooth surface. One can thus,
as in Halcampa claims, recognise three regions of the body, — capitulum, scapus, and
physa ; but only the capitulum is marked off from the rest with any degree of sharpness.
Histologically the body-wall is composed of a strong fibrous connective-tissue.
The individual fibres are extremely fine, and are united in great numbers into tracts ;
they are not so sharply bounded, as, for example, in the connective-tissue of Verte-
brata, but, like them, have a curving course. Generally they cross one another and
interlace in every direction, and only under the endodermal surface does a longitudinal
arrangement preponderate, parallel to the endoderm. Here the fibres stain exceedingly
deeply in picrocarmine, while at all other points fine cords alone retain the stain
after washing.
The endodermal circular muscle-layer is formed into lamellar pleats, arranged
closely like the leaves of a book, and seldom showing arborescence in section. A
muscular region specially developed for a sphincter is not present.
The tentacles are small conical stumps, measuring in the contracted condition about
0"5 cm., and devoid of the two longitudinal ridges occurring in Halcampa clavus. On
the other hand, the terminal pores are obvious, and in many cases are recognisable
with the naked eye. The tentacles are arranged in several rows ; their number in one
case amounted to forty-six, and was perhaps increasing, as I found several small tentacles
among the larger. The longitudinal muscle lamella is ectodermal, and but little pleated.
The oral disc is very small, and presents twelve radial ridges, produced at the
edge of the mouth into the longitudinal ribs of the stomatodasum ; the latter are
sharply-angled, with deep furrows between them. A specially differentiated siphono-
glyphe is not present. The length of the stomatodgeum in the largest example
amounts to nearly 2 cm. Correlated with the absence of a siphonoglyphe is that of
a stomatodeeal cone. The boundary between oral disc and stomatodreum is sharply
marked by the lip being elevated into a circular fold.
REPORT ON THE ACT1NIARIA. 31
The radial muscles of the oral disc are ectodermal, and form a slightly pleated layer ;
a notable point is the presence of small bundles irregularly embedded in the mesoglcea.
The number of mesenteries appears on macroscopic examination to be confined
to twelve, set a.t equal distances on the periphery of the stomatodaeum ; they are so
grouped in pairs according to the muscular distribution, that one can distinguish two
pairs of directive and four pairs of intermediate mesenteries. They resemble thin
veils stretching between body-wall, oral disc, and stomatodseuru, unusually delicate, and
tearing at the slightest strain ; below, they reach nearly to the posterior pole of the
body, but are here so weakly developed as to hardly project at all into the coelenteron.
In these veil-like mesenteries are recognisable, as special thickenings, the following
organs: — 1, the muscle pennons or retractors; 2, the muscles of the edge; 3, the
generative organs ; 4, the digestive filaments.
The retractors are powerful swellings about 1-2 mm. wide, which are tolerably
sharply bounded, and appear as if glued to one side of the mesentery ; they commence at
the angle where oral disc and stomatodasum are continuous, and run from this point in
a slight curve outwards and downwards to the boundary between the first and second
thirds of the body-wall, where they terminate, thus dying out disproportionately soon,
far sooner than even in Ilalcarrvpa claims. Transverse sections exhibit their structure
in greater detail ; in the region of the muscle the supporting lamina is strongly
thickened, and is elevated, together with the muscle-layer resting on it, into lamellae
which are long, thick, and parallel to one another, but which either do not branch at
all, or only slightly. An arborescent or bush)^ appearance is occasionally produced by
a ridge of the mesogloeal mesenterial lamina bearing on both sides a complete series
of muscular lamellae. The sharp boundary of the muscular masses is referable to the
circumstance that on both sides the pleating of the muscular layer ceases abruptly.
The edge-muscles form a band of tendinous appearance running close along the
body-wall, and are most clearly expressed in the posterior parts of the body. Here
they constitute nearly the whole of the mesentery, and the mesenterial filament is
affixed almost directly to them.
The mesenterial filament is fairly obvious for the first two centimetres below the
stomatodaeum, and is arranged in a few coils. Afterwards it becomes finer, but is
wound into a mass of twisted loops, continuing thus for about the next four centi-
metres. The contortions then become gradually less marked, till, sooner or later, the
whole filament dies out ; in one mesentery it could be followed to within two centimetres
of the posterior pole. The first section of the filament is trilobate, possessing one
glandular and two ciliated lobes ; lower down it undergoes, as in other cases, a simpli-
fication of structure by the dying out of the ciliated lobes.
Both the glandular and the ciliated lobes are of exceptionally strong development ;
the continuations of the mesogloeal lamina entering them broaden out in the shape
32 THE VOYAGE OF H.M.S. CHALLENGER.
of a wing, so that an accurately transverse section of the trilobate filament exhibits
the mesoglcea in form of a cross, the arms of which are broad and wing- shaped.
The generative organs lie in the thin septum which is intercalated between the
retractor and the mesenterial filament, and were male in one specimen investigated, in
the other female. The testes are 1*5 cm. long, 0"2 cm. broad, composed of separate
follicles which are arranged in about thirteen transverse swellings. At the edge of the
organ occur small bodies, recognisable only in transverse sections, which I take to be
the first commencements of the follicles ; the supporting lamina widens out, enclosing
a space in which are included roundish cells (spermatoblasts ?), fewer (five) or more
numerous according to the size of the cavity. The latter always opens towards the
epithelium by a small but obvious pore. The latter would argue, if there were any
question here of stages of development of the testis, for its derivation from endoderm ;
unfortunately, however, the mesenteries were not sufficiently well preserved for a close
histological investigation.
In the female organ the conditions were similar ; the ova are irregularly scattered
in the mesentery as larger or smaller grains ; those of fair size project above the surface,
while the largest of all stand out markedly beyond its plane, and are connected with
the mesentery only by means of a fine pedicle. The pedicle passes into a chorion which
surrounds the ovum on all sides, the latter being about 1 mm. in diameter. In this
condition the ovum appears to be already in segmentation.
In the mesenteries occur, finally, external stomata ; they are oval, about 0-5 cm.
long, and occur rather to the outer side of the great mesenterial muscles, on a level
with the wreath of tentacles. Whether also internal mesenterial stomata exist just
below the oral lip, remains doubtful.
From the type of the true Halcampw, this Actinian diverges in exhibiting a
commencement of additional mesenterial cycles, although these are extremely weakly
developed. The accessory mesenteries are small projections, which, in the upper part
of the body alone, emerge from the angle between body-wall and oral disc ; here
there occur pairs of mesenteries both of the second and third orders, readily dis-
tinguishable by difference of size. Since, as we have seen above, the number of the
tentacles also is larger than in the true HalcampcB, the genus Halcampella leads up
to the remaining Ilyanthidae, and through them to the true Actiniae.
Halcampella, sp. (?)*
Habitat. — Shallow water; St. Vincent, Cape Verde Islands, July 1873.
To the genus Halcampella doubtless belongs another Ilyanthidan with numerous
tentacles, although too much mutilated for close investigation or systematic determina-
tion. It is to be distinguished from Halcampella maxima at once, by the absence of
incrustation on the body-wall.
REPORT ON THE ACTINIARIA. 33
II. Paractini^e.
Family 10, Sicyonid^e.
Genus Sicyonis, E. Hertwig.
Sicyonis elongata* n. sp.
The animal is elongated, with about 54 tentacular papillae ; the genital mesenteries
project into the ccelenteron from between the oral disc and the body-wall.
Habitat. — Station 244, June 28, 1875 ; 2900 fathoms. One specimen.
Dimensions. — Height, 7 cm. ; breadth about 3 "5 cm. ; diameter of the pedal
disc, 2 cm.
The sole specimen at my disposal was so strongly contracted that one could hardly
find the entrance to the oral disc. The pedal disc was also exceedingly small, due
partially, no doubt, to contraction. Had the specimen, which in other respects also
was but poorly preserved, not been so compressed in the packing, it would have had
the shape of a long sack sewn up at both ends.
The external appearance of the animal is therefore essentially different from that of
Sicyonis crassa, the body of which is flattened like a cake ; but in the internal structure
there is considerable agreement between the two. The sphincter, the muscles of the
tentacles and oral disc, the cuticular consistence of the mesoglcea, the differentiation of
muscular and genital mesenteries, the enormous folding of the siphonoglyphes, the
radial striation of the oral disc, the shape and arrangement of the tentacles, are in both
cases identical. I was therefore inclined to regard it as a new specimen of Sicyonis
crassa, had I not lighted on one distinguishing characteristic of great importance.
The genital mesenteries in Sicyonis crassa are thin lamellae, which bear only the
generative organs, and spring in the angle between pedal disc and body -wall ; but in
this new specimen the muscles are obvious, and are arranged in " muscle-pennons ; "
the most noteworthy point, however, is, that the genital mesenteries belong to the
upper section of the body, lying in the angle between oral disc and body-wall ; on the
former they reach as far as the oral opening, and on the latter, in the form of slight
folds, up to the pedal disc. Mesenterial filaments do not occur on them. Since the
specimens of both Sicyonis crassa and Sicyonis elongata were males, the different
position of the mesenteries cannot be due to the difference of sex.
Part of the animal was anatomically investigated with reference to the arrange-
ment of the mesenteries, and part of the body-wall, with the mesenteries in the
neighbourhood of the stomatodeeum, was utilised for transverse sections. I was able
to prove the normal arrangement of the mesenteries in pairs at some points ; but at
certain spots irregularities occur, owing to the alternation of isolated genital mesenteries
(ZOOL. CHAIX. EXP. — TART I.XX1II.— 1888. ) DcUld 5
34 THE VOYAGE OF H.M.S. CHALLENGER.
with isolated complete ones. It is probable that, here and there, the one mesentery
of a pair is formed, the other arrested. I was compelled to relinquish the determina-
tion of the number of the mesenteries, in order to spare the specimen. I counted,
however, the number of tentacular papillas, amounting to fifty-three ; some of these, in
the neighbourhood of the single siphonoglyphe, were very small. I infer from this
that increase of the number of the tentacles was not yet concluded.
III. Edwakdsia
Family 11, Edwardsid^e.
Genus Edwardsia.
Edwardsia, sp. (?).*
Habitat. — Station 168, July 8, 1874 ; 1100 fathoms. One specimen.
The sole example of the genus Edwardsia which I met with in the Challenger
material, and which came from a depth of 1100 fathoms, was so strongly contracted
that the capitulum was concealed within the scapus, and in the posterior section was so
completely crushed that it was difficult to detect the rounded hinder pole.
The surface is extraordinarily rough and bark-like, probably in consecpience of an
incrustation of mud on the cuticular layer ; at the anterior end the entrance to the
mouth is visible, and round it are eight radial furrows, which, owing to the indifferent
preservation, could be followed only for a short distance upon the body-wall. The
opening is slit-like ; the wedge-shaped regions bounded by the furrows at the anterior
pole are dissimilar in size, and are so arranged that the broadest is at one end of the
slit, the smallest at the other, while the remaining six are symmetrically arranged right
and left. At the posterior end of the animal, only seven of these furrows, which
correspond to the mesenterial insertions, can be recognised.
I attempted to investigate the structure further by means of transverse sections,
but was reluctantly forced to the conviction that nothing remained of the mesenteries
and stomatodseum.
IV. ZOANTHEvE.
As the result of researches instituted by G. von Koch and myself, I have in my
former Report separated from the hexamerous Actinias, the sharply marked group of the
Zoanthese, and have described as their representatives the genera Sphenopus, Zoanthus,
and Epizoanthus.
I conceived it to be eminently inappropriate that such discordance should exist
in the nomenclature of the individual species and genera of Zoantheae, a discordance
REPORT ON THE ACTINIARIA.
35
referable chiefly to the fact that the forms described had been quite insufficiently
studied, and that consequently the systematic characters had been referred to points of
secondary moment only. In this condition of affairs no alteration has been effected by
the monograph of Angelo Andres ; the great abundance of forms cannot be compressed,
as he has attempted to compress them, into the three genera, Zoanthus, Palythoa, and
Sphenopus (the genera Verrillia, Bergia, and Antinedia having but a doubtful position,
so long as we possess such scanty information about them as at present).
I have therefore requested Dr. Erdmann, one of my students in Bonn, to undertake
a revision of the Zoantheaj with reference to the following important anatomical
characters : — (l) condition of the ccenenchyme ; (2) arrangement of the mesenteries ; (3)
structure of the sphincter; (4) condition of the integument; (5) colony - formation.
His conclusions are as follows : — The Zoanthese may live solitary (Sphenopidse), or may
form colonies (Zoanthidse) ; in the latter case the ccenenchyme may either consist of
basal stolons more or less branching, sometimes even anastomosing, or of a connecting
lamella, or of a mass which unites the polyps almost for their whole height. The
integument either consists merely of an epithelium and cuticle, or else there occur
on it foreign bodies, which penetrate the mesoglcea of the body-wall, and more or less
fill it. In the arrangement of the mesenteries two points are of importance : (l) that
the pairs of mesenteries, with the exception of the directives, consist of a macro- and
a micro-mesentery ; (2) that a dorsal and a ventral zone of mesenteries must be
distinguished. The two zones may approximate either with small (Microtype) or with
large mesenteries (Macrotype). Finally, the sphincter exhibits three modes of forma-
tion; it may be (l) endodermal; (2) mesoglceal ; (3) it may be mesoglceal, but
distinguished by a muscle-free region into upper and lower portions.
AVith reference to the points above mentioned, Erdmann has distinguished five
genera in the colonial Zoanthidse, the characteristics of which may be followed without
further comment in the accompanying table : —
Genus.
Mesenterial
arrangement.
Sphincter.
Ccenenchyme.
Integument.
Generative
organs.
Zoanthus.
Mammilifera.
Epizoanthvs.
Palythoa.
Corticifera.
Microtypal.
Microtypal.
Macrotypal.
Macrotypal.
Microtypal.
Mesodermal,
duplex.
Mesodermal,
simple.
Mesodermal,
simple.
Endodermal.
Mesodermal,
simple.
Stolonar.
Stolon-like, with a
tendency to form
lamellae.
Connective, lamellar.
Resembling a ribbon
or tongue.
Polyps sunk in the
ccenenchyme to
their upper ends.
Soft.
Soft.
Incrusted.
Incrusted.
Incrusted.
Hermaphrodite.
(1)
Dioecious.
Dioecious.
(?)
36 THE VOYAGE OF H.M.S. CHALLENGER.
The material which I was able to place at Dr. Erdmann's disposal was derived
partly from the Bonn Museum, partly from the Triton expedition, but chiefly from
the Challenger collection. For the descriptions of the Challenger Zoanthese I
o-ive here short extracts from his Memoir,1 for the accuracy of which I can vouch, as
the whole investigation was carried out under my direction. I have achieved, what he
omitted, in identifying as far as possible the forms obtained with species previously
described, and, where that was impossible, have introduced new names, and have
reduced the diagnoses of species to shorter and more precise terms.
Family 12, Zoanthid^:.
Genus Zoanthus, Cuvier (pro parte).
Integument not incrusted ; ccenenchyme stolonar, with an occasional tendency to
lamellar extension ; sphincter differentiated into upper and lower sections ; mesenteries
arranged on the microtype.
Zoanthus dance (?), Le Conte (PI. I. fig. 1).
Polyps with fleshy body-wall, the larger borne on a stalk-like extension, and
arranged closely together on reticulately branching stolons ; approximately fifty
tentacles arranged in two cycles.
Habitat. — Bermuda Islands ; shallow water.
Dimensions. — Of the individual polyps — height, 0-5-2'5 cm. ; breadth, 0 3-0 "5 cm.
This animal, which I refer with considerable reserve to Zoanthus dance, is identical
with the Zoanthus which I have already described. To that description I can add the
following points, based on Erdmann's researches : —
1. The colony grows on a foundation of rock in such a manner that the upper ends
of all the polyps lie in the same plane. As the foundation is irregular, the individual
polyps must be of unequal lengths, a result of wdiich is that those animals which
correspond to hollows in the foundation are produced posteriorly into a kind of stalk,
distinguished from the body proper by a constriction, and by the thinner consistence of
the body-wall.
2. A peculiar attachment of the cuticle to the body-wall, and one perhaps more
widely distributed among the Zoanthea3, is effected by mesoglceal processes which
perforate the epithelium and are inserted on the cuticle.
3. The colony investigated by Erdmann was sexually mature ; ova and testicular
follicles occurred in the same mesentery.
1 Erdmann, Ueber einige neue Zoautheen. Ein Beitrag zur anatomischen uud systematischen Kenntniss der
Actinien, Jenaische Zeitschr., Bd. xix. pp. 430-188, pis. iv. v.
REPORT ON THE ACTINIARIA' . 37
Zoanthus confertus* Verrill (PI. I. fig. 12).
Polyps with thin transparent body-wall, so closely packed as to be polygonally
flattened.
Habitat. — Simon's Bay, Cape of Good Hope ; 10-20 fathoms.
Dimensions. — Of the individual polyps — height, 06-0"8 cm. ; breadth, 0"3-0'4cm.
The species is in general structure very close to the preceding, but differs in the thin
consistence of the body-wall, through which may be seen the mesenteries, and in the
compact arrangement of the polyps. The latter being consecpiently compressed
polyhedrally, a character of importance is afforded for the species, which is further
marked off by the transparence and delicacy of the body-wall.
Genus Epizoanihus, Verrill.
Integument incrusted, ccenenchyme (mostly ?) lamellar ; sphincter simple, meso-
gloeal ; mesenteries arranged on the macrotype ; colonies (mainly V) parasitic.
Epizoanthus thalamophilus* n. sp. (PL I. fig. 3 ; PI. IV. figs. 7, 8).
Incrustation scanty, exclusively composed of Foraminiferal shells, which are
arranged on the individual polyps into 15-20 longitudinal rows, bifurcating
downwards ; body-wall transparent ; tentacles 30-40, very long, and arranged in
two rows.
Habitat. — Station 299, December 14, 1875 ; 2160 fathoms; on Gastropod shells.
Dimensions. — Height of the contracted individuals, 0-2-l-3 cm. ; diameter at the
base, 0"9-l'5 cm.
" The colony of seventeen individuals has settled on a deserted Fusus shell about
8 cm. long. The polyps are principally situated on the back of the shell, and only the
five young individuals at its apex are arranged in a whorl round it. The region round the
aperture of the shell is free from polyps ; they rise with elliptical bases from a common
ccenenchyme, and arch upwards like a dome. The largest specimens have a base of
10-15 mm. in diameter, and are 13 mm. high ; but we find every transition to the smallest
specimens, which appear as flat elongated projections with a base of 5-9 mm., and a
height of 1*5-3 mm. The ccenenchyme is a continuous sheet, 0*3-0*5 mm. in thickness,
which covers the shell as far as the colony reaches. Towards its termination it becomes
constantly thinner and more transparent, till it ends as a very delicate pellicle, which
may be easily rubbed off. All the polyps were in a highly contracted condition ; and
at the dome-shaped summit lies, on a prominence which is bounded by a circular furrow,
the entrance to the interior ; it is hardly recognisable as an opening, and is formed by
the indrawn parts of the body-wall. The latter is of slight thickness, so that the
38 THE VOYAGE OF H.M.S. CHALLENGER.
mesenteries may be seen through it as clear stripes. In the external zone of its mesoglcea
lie the deposits above mentioned, consisting exclusively of Foraminiferal skeletons.
They are evenly distributed over the ccenenchyme ; but on the body- wall are ranged in
a most regular and elegant manner, the following facts being recognisable with the
aid of a lens. From the apex outwards run, in a well-grown individual, fifteen to twenty
looping rows of Foraminifera in clear elevated lines. Where the body-wall bends
downwards at right angles, each row bifurcates, and each branch so produced runs
downwards on the body-wall in a straight line ; a single row of Foraminifera is thus
situated over each mesentery, the insertion of the latter being externally clearly
recognisable, owing to the thinness of the wall. While therefore, from the apex of the
polyp outwards, the ridges agree in number with the pairs of mesenteries, in the lower
part of the body-wall there are present as many rows of shells as there are individual
mesenteries. Towards the base these become less plain, so that at the lowest part
of the polyps, as on the ccenenchyme, the Foraminiferal coating is evenly distributed
all over " (Erdmann). The rows of shells are continued on to that region of the body-
wall which has been drawn inwards ; and their arrangement can here be only understood
by referring to the point of transition from body-wall into oral disc. This occurs along
an undulating curve, since at one point the oral disc with its outer circlet of tentacles,
at another the body-wall with its rows of shells, projects the farthest. A horizontal
section therefore, through the region under discussion, meets alternately with rows of
Foraminifera and the origins of tentacles (PI. IY. fig. 8). Further, at the point of
junction, the body-wall forms a strongly projecting fold in which lies the greater part
of the sphincter (PL IV. fig. 7). The horizontal section represented in fig. 8 exhibits
this fold on the inner side, while on the outer lie the body-wall and oral disc, united
by mesenteries.
The fold of the body-wall bears, on both sides, rows of Foraminiferal shells,
supported on ridge-like processes of the body-wall, and appearing therefore in trans-
verse section as coronets ; they are, as we learn from longitudinal sections, discontinuous
at the free edge of the fold, so that the outer and inner rows of shells do not pass into
each other.
The sphincter embedded in the fold of the body-wall is mesoglceal and simple, and
forms here an evenly distributed complex mass of muscle-bundles, the latter being
variously shaped. It also overlaps a small strip of that region of the body-wall which
is not drawn inwards.
The tentacles are, as in other cases, in two alternating circlets, and are in part
produced into long pointed filaments, in part contracted into short stumps. Their
muscles are ectodermal and slightly pleated ; the mesoglceal supporting lamina lying
at the base of the pleats sends processes into the epithelium.
The stomatodseum is oval, and the siphonoglyphe only slightly expressed.
REPORT ON THE ACTINIARIA. 39
The number of mesenteries varied in the three specimens investigated between
twenty-eight and thirty-six, according to their size. The dorsal and ventral zones of
mesenteries approximate always with macromesenteries.
No channel filled with cells is present at the bases of the mesenteries ; the muscle-
pennons indistinct ; the generative organs so abundantly developed as to fill the
greater part of the ccelenteron. These latter occur only on the macromesenteries,
and consisted of testicular follicles in the three specimens studied.
The coenenchyme is extremely thin, and possesses internally smooth connecting-
tubes lined by endoderm ; on the upper surface Foraminiferal shells are sparsely
embedded ; while on the other side, which covers the Gastropod shell, these are entirely
absent.
The name thalamophUus was chosen with reference to Thalamophora and Poly-
thalamia, names which have been applied to the Foraminifera.
Epizoanthus stellaris* n. sp. (PL I. fig. 4).
" Polyps of inconsiderable height, nearly saucer-shaped; body-wall vertical at the
sides, but strongly flattened above ; on its horizontal upper surface are numerous radial
ridges, separated by furrows, 15-20 in the adult animal ; colour of the colony dark
greyish-brown ; deposits very various."
Habitat. — Station 201, off Samboangan, Philippine Islands; 82 fathoms.
Dimensions. — Of the individual polyps— height, 0'05-0"4 cm. ; diameter, 0'15-0-7 cm.
" Of this species I possess a colony, covering the rooting spicules of a Hyalonema for
a distance of about 14 cm., and consisting of about 100 individuals. The coenenchyme
forms a tube open at both ends, and surrounds like a sheath the bundle of spicules, the
latter being about 5 mm. thick. The individuals spring from it at longer or shorter
intervals by an elliptical base, measuring in the largest polyps (3-4 mm. high) about
5-7 mm. in diameter. From these to the smallest, which hardly project above the
coenenchyme, and are 1-5-3 mm. broad, by 0'5-l mm. high, every transition is found.
All the animals are strongly contracted ; on the strongly flattened, discoidal, horizontal
surface of the body -wall may be dimly seen the entrance to the interior by a circular
pit. From this point outwards radiate over the surface of an adult specimen, about
15-20 ridges separated by furrows.
" The colour of the colony is a dirty dark-grey. The body-wall is of considerable
thickness, caused by the strongly developed mesoglcea. The exterior surface of the
latter is charged with various deposits, consisting of irregular grains of sand and lime,
sponge spicules of very varied origin, and finally of the small dark crystalline bodies
which cause the dark tint of the colony. These deposits occur in additional quantity
on the radial ridges before mentioned. They are continued inwards as elevated ridges
over the edge of the covering fold without a break, and run even further, on the inner
40 THE VOYAGE OF H.M.S. CHALLENGER.
face of the indrawn part of the body-wall. Sections through the upper region of the
polyp yield appearances similar to those described under the preceding species, though,
owing to the abundant and various deposits enclosed, they are not so regular and
elegant.
" In those inner parts of the mesoglcea which are free from adventitious accretions
there lie embedded in the homogeneous matrix — 1. fine radial fibres, penetrating
the whole thickness of the soft mesoglcea, provided here and there with nuclei ; 2. round
mesoglcea-cells containing a large nucleus ; 3. round or oval spaces packed with cells.
Hertwig, who has observed similar structures in the JEpizoanthus parasiticus described
by him, conjectures that these oval cell-islets are produced only by indifferent preserva-
tion, and result from the breaking down of a system of anastomosing cords, such as
the mesoglcea of Zoanthus exhibits. I [Erdmann] am inclined to regard these roundish
heaps of cells as primary structures, like the canals of Zoanthus, since I have been
able to recognise them in almost all my species of Epizoanthus, which were without
exception in a very good state of preservation. As to their origin I have no data ; but
there is no reason why they should not be referred to an ectodermal origin as well as
the cell-canals of Zoanthus, the derivation of which from ectoderm is indisputable ;
besides, many of these cell-islets clearly exhibit an elongate outline, with here and
there even a slight tendency to branch, by which an external approximation to
Zoanthus is effected.
"The mesoglcea of the mesentery is well developed, and on its inner edge is
thickened like a club. The micromesenteries project only slightly into the interior,
but, like the macromesenteries, clearly present marked muscle-pennons. On these
mesenteries there springs on the side opposite to the muscle-pennons a mesoglceal
lamella, which is considerably elongated in order to carry the generative organs
and to form, centrally to these, the mesenterial filaments. The former are present
in considerable numbers ; and, being cut more or less superficially owing to the con-
torted course of the mesentery, may be recognised in transverse section as roundish
balls enveloped in a thin mesoglceal lamella, pressed against the body-wall and generally
filling the adjacent chamber. All the specimens which I investigated were female, the
generative balls consisting of a large number of ova closely appressed together, but
separated by a fine mesoglcea lamina.
" The body-wall is deeply drawn inwards, and conceals in this region a strongly
built sphincter, which has the shape described for the preceding species, but which
is distinguished by a greater complication in the branching of the bundles of fibrillas.
" The stomatodseum is oval, with a clearly defined siphonoglyphe. The ensheathing
ccenenchyme measures 1-1 "3 mm. in thickness ; in its interior run longitudinally
numerous connecting tubes. The mesoglcea carries on its surface foreign deposits of
the same character and quantity as those on the body-wall, but the inner face, which lies
REPORT ON THE ACTINIARIA. 41
ou the foundation, is completely free from incrustation. The soft mesoglcea of the
ccenenchyme is, with reference to histological differentiation, in the same relation to the
body-wall as it is in Zoanthus, since here also, in addition to the other points of marked
agreement, the nucleated fibres are supplanted by mesogloeal cells.
" With a view to observing the mesenterial arrangement, I studied two examples,
one of medium size, and one fully grown ; both exhibit the regular macrotype. In the
•younger specimen occurred a symmetrical arrangement of the pairs of mesenteries ;
of these there were sixteen, seven being regularly distributed on each side of the
■directives. The other polyp possessed nineteen pairs, of which nine were situated on
the one side, and eight on the other."
Epizoanihus elongatus* n. sp. (PI. I. fig. 2).
" The individual polyps form elongated cylindrical tubes, the body-wall is flattened
'above, with a marked indentation, but terminates without radial furrows ; colour of
the colony a yellowish-grey."
Habitat. — Station 322, off Monte Video; February 26, 1876 ; 21 fathoms.
Dimensions. — Height of the polyps, 0-05-l'0 cm. ; breadth, 0- 15-0 '4 cm.
" This species can only be externally distinguished from the preceding. The
colony is 10 cm. high, consisting of about 100 individuals, and lives on a bundle of
the siliceous threads of a Hyalonema, about 3 mm. only in thickness. The largest
polyps are long cylindrical tubes, about 8-10 mm. high and 3-4 mm. broad ; in their
neighbourhood occur gradations to the youngest buds, which are small warts projecting
from the ccenenchyme, of 0"5-2 mm. in height, l"5-2"5 mm. in breadth. All the
animals are in a state of the most marked contraction ; the horizontal upper surface of
the body-wall is more or less flattened, and exhibits a circular indentation. This part
of the body-wall is entirely free from radial ridges and furrows. The colour of the
colony is a greyish-yellow.
" The body-wall is thinner than in the preceding species, and possesses in its
outer zone the same deposits, though in smaller quantity. The remaining anatomical
and histological relations agree closely with those of the former species, but it is
important to observe that the sphincter is less strongly developed. The body-wall is
drawn inwards less deeply ; its sphincter is in transverse section correspondingly short,
but curved, and pointed at both ends. The generative organs consisted of ova in the
five specimens investigated."
Epizoanthus cancrisocius* Studer (PI. I. fig. 15).
Colony much incrusted, and consequently so brittle as to break readily in pieces ;
individual polyps slim, body-wall at the upper end bent outwards in the contracted
(ZOOL. CHALL. EXP.— PART LXX1II.— 1888.) Dddd 6
42 THE VOYAGE OF H.M.S. CHALLENGER.
condition into a plate-like expansion, from the indented centre of which run 15-20
furrows towards the thickened edge.
Habitat. — Station 49, May 20, 1873; 85 fathoms, upon a Gastropod shell
tenanted by a Pagurus, the shell entirely dissolved away by the coenenchyme.
Dimensions. — Length of the polyp, 0'6-rO cm.; breadth, 0-3-0 '5 cm. ; colour,
greyish-yellow.
"This species forms a colony of eleven individuals, on a shell some 2'0 cm. high.
The calcareous substance of the latter is completely absorbed, and at all points replaced
by the coenenchyme, the latter having obviously taken its place, while preserving its
external form. Only the anterior side of this ccenenchymatous structure, i.e. the part
directed forwards in movement of the Crustacean, possesses polyps ; the free posterior
side allows the coils of the former Gastropod shell to be clearly recognised. Of the
eleven individuals, eight large mature polyps occupy the edge of that side which is
directed anteriorly in the movement of the crab. They form long cylindrical tubes,
G-10 mm. high and 3-5 mm. broad. In the median space which they bound, stand
three very young polyps, projecting as vertical cylindrical warts from the coenenchyme,
with height and breadth alike of 1-5-2 mm. One may remark that the large polyps
bend forwards, i.e. their oral discs face upwards, in the direction corresponding to the
locomotion of the Pagurus, so that they are most favourably placed for the reception
of the food matters which stream against them. Owing to the curving just mentioned,
the large polyps are above strongly compressed laterally.
" The whole colony has a rough shagreen-like exterior, of a grey colour. The
otherwise smooth body-wall forms above a horizontal plate, which not only projects
like the capital of a column over the vertical part, but has also a characteristic sculp-
ture, and the appearance of a plate with raised edges and indented centre ; in the
middle of the latter lies the entrance to the interior, which is slit-like, corresponds
to the lateral compression, and is always recognisable as an obvious opening. From
this median point outwards radiate over the plate-like surface 15-20 radial furrows,
which are continued outwards for a short distance over the marginal thickening,
appearing on it as deep notches.
" When a polyp is opened with scissors, one remarks that the mesenteries run
down the whole length of the body-wall, but do not pass over on to the horizontal
floor of the ccelenteron. In the lowest parts of the polyp, the mesenteries are visible
as slightly projecting ridges, striking the eye by their clear colouring ; at about one-
fourth of the total height, the macromesenteries form filaments ; these are yellowish-
white contorted coils, which completely obscure the micromesenteries. One can
without damage remove the mesenteries from the body-wall, and study them indepen-
dently. The supporting lamina of the mesenteries is very thin, and runs simply to
the base without any excavation ; the mesenterial filaments are of the customary
REPORT ON THE ACTINIARIA. 43
structure. I have been unable to detect generative organs in any specimen
investigated.
" Owing to the abundant incrustation, the body-wall becomes as hard and brittle
as stone, and does not permit therefore of investigation by means of sections. In this
case therefore, and in the remaining forms with similarly strong incrustation, I made
use of the method of grinding tested and recommended by G. v. Koch in his researches
on Tubipora.
" The body- wall is of considerable thickness ; its mesoglcea exhibits a structure very
different from the remaining species of Epizoanthus, as being penetrated by deposits
throughout its whole depth. These deposits consist of particles of sand with irregular
angles, and are set in a strong circular fence, reducing the mesoglcea to thin lamellae ; but
there persists a very narrow internal lamella bounding the endoderm all round. In the
homogeneous lnesoglcea-lamellas are situated roundish cells which give off fine radiating
processes, and fine fibres provided with nuclei ; the presence of the cell-heaps, which
are to be met with in the remaining species of Epizoanthus, I was unable to
demonstrate in this case. A transverse section through the wall of the shell exhibits
a similar condition in the ccenenchyme. This latter is also of considerable thickness,
and is internally traversed by the large endodermal tubes which connect the various
ccelentera together.
"The body-wall is, as has been already mentioned, bent above at a sharp angle,
thus forming a plate-like surface. In contrast to the remaining members of the
genus, where it turns deeply inwards vertically, it is here only slightly invaginated, a
difference resulting from the slighter development of the sphincter. The latter com-
mences to a certain extent on the horizontal part of the body-wall, and then thickens
gradually into a truncated muscular mass, which appears fusiform in section, and is
only slightly curved inwards. It lies enclosed in the innermost lamella of mesoglcea ;
the latter is thus much thickened, and is free from adventitious deposits. The
sphincter is on both sides bounded by a layer of mesoglcea, which extends inwards to
the commencement of the oral disc, is charged with the usual accretions, and is a direct
continuation of the outer sandy layer."
So much for the anatomical description given by Erdmann, which sufficiently
proves that Epizoanthus cancrisocius must be separated systematically from Epizoanthus
parasiticus, the latter possessing larger and coarser polyps and far less incrustation. I
have identified the animal with the Epizoanthus cancrisocius of Studer, as he records
for his specimens similar dimensions, and a marked incrustation, at least for the basal
membrane.1 In other points his description is not sufficiently exhaustive, and this is
still more true of Gray's account.2 Only the statement of the latter that the large
1 Monatuber. d. k: Akad. d. Wiss. Berlin, 1878, p. 547.
s Proc. Zool. Sue. Lond. 1867, p. 237.
44 THE VOYAGE OF H.M.S. CHALLENCxER.
polyps break up easily, and the reference to a figure of Gosse's which recalls our
Epizoanthus cancrisocius, make it probable that his Epizoanthus papillosus and the
Epizoanthus cancrisocius are identical.
Erdmann refers it in his Memoir to the expedition of H.M.S. "Triton." I find,
however, his specimen in a bottle from the Challenger collection, with the label given
above ; some mistake must therefore have occurred in his manuscript.
Genus Corticifera, Lesueur.
Ccenenchyme extending from the base upwards between the individual polyps, and
uniting them together almost as far as the upper edge of the body-wall ; integument
incrusted ; sphincter mesoglceal ; mesenteries arranged on the micro type.
On the above diagnosis I may remark that, on the body-wall of each polyp may
be distinguished two regions, the one surrounded by ccenenchyme, the other projecting
freely above it. When the animal contracts, the latter is drawn inwards to the level
of the ccenenchyme as in Madreporaria ; it partly serves to close over the anterior end,
and partly is invaginated inwards. A colony in contraction consequently forms a crust-
like covering, in which the individuals are only indistinctly marked off from each other.
Corticifera lutea* Quoy and Gaimard (PL I. fig. 6).
Individual polyps marked off by fairly obvious stripes on the ccenenchyme, and
recognisable as annular ridges on the common surface of the colony ; they differ but
little from each other in size.
Habitat. — Bermuda, June 1873; shallow water.
Dimensions. — Height, 1 cm. ; breadth, 0-4-0"5 cm. ; colour, j'ellowish-white.
"The colony at my disposal consists of a flat, quadrangular, crust-like structure,
about 16 cm. long and 7 cm. broad. It does not present a complete whole, but is
merely a piece torn off from a larger mass, carrying about 400 individuals ; the latter
reach a height of 10-15 mm., and are in diameter 4-5 mm. It must be insisted
that this external height of the polyps in no way corresponds to the internal, since the
ccenenchyme forms on the under side so thick an investment that of the total height
only about two-thirds belong to the ccelenteron, the other third to the ccenenchymatous
layer beneath. All the individuals are strongly contracted, and the body-wall is drawn
deeply inwards. The edge of the body-wall projects above the general surface as an
annular depressed ridge, in the centre of which lies, always clearly open, the aperture
to the interior. At the unmutilated edge the individuals stand out as slight swellings
" In that part of the ccenenchyme which borders on the ectoderm, are present
numerous accretions, producing a firm pellicle. The main bulk of the incrustation
consists of irregularly-shaped calcareous bodies ; besides these, occur more sparingly
REPORT ON THE ACTINIARIA. 45
Foraminiferal and Radiolarian skeletons, and finally, numerous Sponge -spicules of
various kinds. In the ccenenchyme between the polyps, the accretions are present only
in small quantity, and fill here simple scattered cavities, which may be recognised after
decalcification as wide lacunae. The rest of the ccenenchyme is soft ; and in its
homogeneous matrix we meet with large canals, lined by pigmented epithelium and
traversing the ccenenchyme in every direction ; they are especially numerous in the
lower ccenenchymatous investment, which consequently presents a reticulate spongy
texture. As appears from longitudinal sections, these canals are direct continuations
of the ccelenteron from the base of the polyp outwards, and extend from this point
upwards through the whole of the cosnenchyme ; they may consequently be homologised
with the endodermal connecting tubes to be found in all Zoanthidae. The mesoglcea of
the ccenenchyme exhibits also numerous roundish cell-islets lined by epithelium, in which
we may perceive the origin of such ectodermal cell-heaps as have been described for
Epizoantlms. The whole of the endodermal epithelium is pigmented by dark granules,
as are also the large endodermal connecting-tubes. On the other hand, the roundish
cell-aggregations just mentioned are free from pigment granules ; this difference of
condition affords an indirect proof that the latter are by no means of endodermal
origin, but are purely ectodermal structures. Finally, the soft ccenenchyme exhibits
fine nucleated fibres starting from the endoderm, and, as is usual, numerous mesoglceal
cells provided with fine processes.
" The main bulk of the whole colony is to be regarded as ccenenchyme ; the
individual polyps consist merely of a mesoglceal cylinder lined internally by endoderm,
of moderate thickness and homogeneous consistence. The supporting lamina of the
mesenteries is of similarly weak development. Below, the latter enclose a canal filled
with cells, which in the case of the macromesenteries is frequently divided up by cross
anastomoses. The muscle-pennons are well developed, and appear, especially in the
larger mesenteries, as branching processes, which extend over a wide stretch of the
mesentery. Nothing of interest can be said about the mesenterial filaments. In none
of the specimens investigated could I find generative organs. The stomatoda3um is
pear-shaped in section, with a well-marked siphonoglyphe."
The sphincter is mesodermal, simple, and only slightly developed. It begins early,
as a narrow strip, in that part of the body-wall which is drawn horizontally inwards,
and extends without any thickening to the edge of the invaginated part. The number
of the mesenteries, which are arranged on the microtype, varied in five individuals
between thirty-four and forty.
Corticifera tuberculosa* Klunzinger (PI. I. fig. 5).
Individuals closely appressed together and flattened polygonally, generally separated
by a deep furrow, and of very dissimilar sizes, so that the surface of the contracted
46 THE VOYAGE OF H.M.S. CHALLENGER.
colony appears to be irregularly covered with knobs. These knobs exhibit radial
furrows which run outwards from the indistinct opening.
Habitat. — Simon's Bay, Cape of Good Hope ; 10-20 fathoms.
Dimensions— OS-OS cm. in height; diameter, 0"2-0"5 cm.
Colour. — Brownish.
The small colony of about forty individuals differs essentially from the above
described Corticifera lutea in its external appearance. From the small development of
ccenenchyme, it results that the individual polyps press closely on one another, and
frequently become polyhedrally, generally hexagonally, flattened. They are separated
by deep grooves on the surface, which, at few points only, become shallower or dis-
appear altogether. The absence of the groove between two polyps possibly signifies a
genetic dependence, the one having arisen by gemmation from the other ; and smaller
individuals are "frequently adjunct to the larger polyps in this fashion.
The individuals of the colony are of most varying size ; from the large dome-
shaped convex animals with a diameter of 0'6 mm. those of intermediate size lead to
the smaller, which measure only 0"1 mm. in the one direction and 0'2 mm. in the
other. Since the surface therefore exhibits smaller and larger knobs, I refer the species
to the Palythoa tuberculosa of Klunzinger, and have therefore retained the well-chosen
specific name.
In length there is but little difference between the larger and smaller animals, the
former measuring 0 6 cm., the latter 0"4 cm. As they all diminish downwards in a
wedge-shape, the lower side of the colony is so much narrower that the polyps on the
edge are nearly horizontal.
All the polyps are so strongly contracted that the entrance to the interior is
recognisable only as an indistinct indentation, from which radiate outwards numerous
shallow furrows.
With reference to the finer anatomy, what has been said for Corticifera lutea
holds good in this species. In the two specimens investigated there were respectively
thirty-four and thirty-six mesenteries, which followed the microtype.
Genus Palythoa, Lamouroux.
Integument strongly incrusted ; ccenenchyme little developed, ribbon- or tongue-
like ; mesenterial arrangement on the macrotype ; sphincter endodermal.
Palythoa anguicoma* Norman (PL I. fig. 7).
Incrustation superficial, so that a thick layer of mesoglcea remains free of deposit ;
ccenenchyme tongue-shaped ; individuals, when in a contracted condition, long, with a
terminal capitular enlargement, on which run 15-20 radial furrows.
REPORT ON THE ACTINIARIA. 47
Habitat. — Station 135a, off Inaccessible Island; October 16, 1873; 60-90 fathoms ;
hard ground, shells, and gravel.
Dimensions. — Height of the polyps, 0'4-0-8 cm. ; breadth, 0'2-0'4 cm.
Colour. — Brownish-yellow.
" From the material at my disposal, which appears to have been carelessly detached,
the general form of the present species cannot with certainty be inferred. The greater
part of it consists of single individuals, in which one can recognise the forcible detach-
ment from the colony. One group, which to. all appearance represents a complete
and intact colony, is composed of four individuals ; they are situated, in a row and at
short intervals, on a thin ccenenchyme which is extended like a ribbon ; their dimen-
sions are 4-8 mm. high by 2'5-4 mm. broad. All the polyps are strongly contracted;
the body-wall forms above, in this condition, an obliquely-angled ridge projecting
outwards ; its upper surface presents an elevation, rendered obvious by a circular furrow,
in the centre of which the aperture to the interior is recognisable. From the middle
of this upper surface radiate outwards 15-20 furrows, which are continued over the
projecting ridge on to the vertical body-wall, where they then flatten out. The colour
of the polyps is a dirty yellow.
" The integument is furnished with accretions, and exhibits a rough shagreen-like
exterior. On rubbing awTay the thin sandy layer, there remains the thinner soft part
of the mesogloea, which is excellently fitted for the preparation of longitudinal and
transverse sections with a razor.
" The soft mesoglcea is of considerable thickness, and consists of a homogeneous
matrix, in which come into view the large number of cavities charged with cells.
These may be simple, i.e. preserve their roundish or elliptical outline, or, as in most
cases, may branch to form a system of anastomosing canals which entirely recall
Zoanthus. Below the endoderm such a canal runs in an almost unbroken ring
through the whole of the body-wall ; it lies so close under the epithelium as to be
separated from it only by a narrow lamella of homogeneous matrix. Its diameter is
not constant throughout its whole circuit, but is frequently constricted, and occasion-
ally such constriction produces an actual discontinuity. It is further of importance
that the canal invariably presents a considerable hollow expansion under each mesen-
terial insertion. At many points can be demonstrated a communication between
the smaller branching cell-canals and this large ring-canal, the latter being at such
places apparently expanded into a kind of funnel. Further, there are found in the
mesoglcea numerous mesoglceal cells, giving off fine processes ; and, finally, delicate
nucleated fibres, the course of which, however, is here not radial, but in the main
circular.
" The structure of the ccenenchyme agrees in all respects with that of the body-
wall, except for the fact that it possesses endodermal connecting tubes.
48 THE VOYAGE OF H.M.S. CHALLENGER.
" The mesogloea of the mesenteries is strongly constructed, and on it can be
recognised well-developed muscle-pennons. The generative organs, borne in the
supporting lamina, consisted of ova in the individual which I investigated. The mesen-
terial filaments are of the customary structure.
" The mesenterial arrangement is to be referred to the macrotype. The specimen
investigated possessed thirty-six mesenteries, of which five pairs pertained to the dorsal
zone, and thirteen pairs to the ventral ; in the latter zone were ranged regularly, on each
side of the directives, six pairs, consisting of a macro- and a micro-mesentery.
" The body- wall is drawn inwards at a right angle ; on the inner side of this
region a definite endodermal sphincter may be recognised. The pleatings of the
endodermal muscle-lamina are more clearly marked than in Palythoa axinellce ; and
produce on the mesogloea prominent antler-like prongs. The accretions are continued
on to the inch-awn region of the body-wall, but die out at its lower edge, where the
oral disc commences."
The identity of this animal with Palythoa anguicoma is doubtful, as Norman, who
created the species, gave no figure of it. I was influenced by the circumstance that
eighteen rough radial furrows are ascribed to this form ; besides which the incrustation
on it should be only superficial.
Palythoa, sp. (?)*
Habitat. — (a) Station 135 a, off Inaccessible Island, October 16, 1873; 60-90
fathoms, (b) Station 135 c, off Nightingale Island, October 17, 1873 ; 100-150 fathoms.
In the same bottle with Palythoa anguicoma was another species of Palythoa, which
recurred in a second tube, the contents of which were dredged a day later than the
first, and at a greater depth. The specimens in cmestion could easily be distinguished
from individuals of Palythoa anguicoma by containing black particles of hornblende.
Erdmann attempts to separate the two species, and gives the following description : —
" In this species also the larger part of the material consists of individuals torn away
from the colony ; one colony, which was undoubtedly not mutilated, was represented by
three individuals, ranged behind one another on a ribbon-like ccenenchyme. Externally
this species differs from the preceding in colour only, which is in this case a dull grey-
brown ; besides this, from the greater firmness and unevenness of the body-wall, it may
be recognised that the mass of accretions is greater. The body-wall presents, in contrast
to the former species in which the relations are reversed, a considerable zone charged
with accretions, opposed to a slightly-developed soft zone of mesogloea. In the latter
there passes close under the endoderm a cell-canal, frequently constricted, but rarely
interrupted ; external to this follow immediately the accessory deposits, so that of
the numerous canals and spaces observed in the preceding species only a few roundish
cell-islets are preserved."
REPORT ON THE ACTINIARIA. 49
Palythoa (?) sp. (?)*
Habitat. — Station 299, west of Valparaiso, December 14, 1875 ; 2160 fathoms.
I found a small Actinia, labelled " Actinia on nodule," which had settled on a piece
of pumice near an Ascidian. The animal, being incrusted with sand particles, probably
belongs to the Palythoce, but its minuteness and the sandy incrustation forbade a
detailed study. The body, not so much as 1 mm. high, was flattened into a disc 5 mm.
broad. The number of mesenteries which, as in the Zoanthese, were very regularly
arranged, amounted to thirty-two.
Family 13, Sphenopid.e.
Genus Sphenopus, Steenstrup.
Sphenopus pedunculatus* n. sp. (PI. I. fig. 11).
Body marked off into an upper swollen trunk, an elongate narrow foot, and a
broad sole-like (?) " clasping-disc ; " from the apex run, over the upper part of the
trunk, about 10-12 indistinct rough furrows.
Habitat. — Station 203, off Panay, Philippine Islands, October 31, 1874; 12-20
fathoms. Three specimens.
Dimensions. — Length, 2-4-3'2 cm. ; breadth, 2-2'4 cm.
Colour. — Grey. i
" This species differs in many respects from the already known Sphenopus marsu-
pialis (Steenstr.) and Sphenopus arenaceus (Hertw.). The fully-grown animal permits
of an external differentiation into three regions. The most obvious part of such a polyp
is formed by the upper bladder-like 'body' (PL I. fig. 11), which conceals within itself
the organs of nutrition and reproduction. On it is marked off, by a more or less
obvious cross-furrow, a hood-shaped anterior region, sculptured by coarse radial furrows.
The body passes into a long narrow ' foot,' from which it is sharply defined by a
marked furrow, and finally the foot broadens out at its base into a kind of ' clasping-
disc' The three animals of this species which were at my disposal represented
stages of different age. In the oldest individual the bladder-like body has been
irregularly contracted by preservation in spirit, its exterior is folded, and exhibits
besides a lateral compression. The head region, defined by an obvious constriction, is
strongly tuberculate, and marked by twelve coarse radial elevations, separated by
discontinuous and incomplete furrows. The height of the body amounts to 2-5 cm.,
its greatest width to 2 "4 cm. Sharply marked off from it by a circular furrow is the
cylindrical foot, the diameter of which reaches 1*2 cm. Unfortunately this latter has
been broken away, so that I can give no accurate information either about the total
length, or about the clasping-disc of this animal. The second polyp was of medium
(ZOOL. CHALL. EXP. PART LXX1II. — 1888.) Dddd 7
50 THE VOYAGE OF H.M.S. CHALLENGER.
ao-e ; its total length amounted to 3 "2 cm., of which 2-0 cm. belong to the body, and
1'2 cm. to the foot. The former is on one side crushed inwards about the middle,
where it is of the greatest diameter (2 cm.), while on the other it is as strongly swollen
out. Above, it diminishes gradually into the head region, which is indistinctly
furrowed radially ; and below, equally gradually, into the foot. The latter is
cylindrical, and has a diameter of 0"5 cm., while the sole-like clasping-disc has at its
base a breadth of 0'9 cm. The third and still younger polyp consists mainly of the
' body,' which above is flat and discoidal, without differentiation of a head-region, but
is at the periphery pressed into folds ; its height is 2-4 cm., its breadth 2"0 cm. Below
it passes gradually into the foot, which is rudimentary, round, only a few millimetres
high, and ends without a clasping-disc.
" For investigation I made use of the middle specimen, which was completely
preserved. A longitudinal section dividing the polyp into two halves yielded the
following results. The mesenteries run in the foot as clear narrow ridges on the body-
wall, scarcely projecting into the interior ; they extend also on to the horizontal pedal
disc, and appear in this region as radiating lamellae, which meet at the centre of
the flat base. The filaments first appear on the mesenteries at the point of transition
into the broader '_body ; ' they form a thick investment, which nearly fills the whole
ccelenteron and covers the mesenteries completely. The body-wall is fairly thick, and
even with the naked eye can be distinguished into two layers ; an outer, which appears
granular owing to the accretions, and an inner, which is soft, shining, and free from
deposits. It is further noticeable, that the quantitative relations between the incrusted
and the softer layers vary with the height of the part in question, and in such a
manner that, at the upper part of the body, both parts are about equally strongly
developed, while with increasing depth the harder constituents become more numerous,
till at last, in the foot, a complete obliteration of the softer zone is produced. Above,
the body-wall is drawn rather deeply inwards at a sharp angle. On to this infolded
region the accretions are uninterruptedly continued as far as the point of origin of the
oral disc, the latter being inserted just at the inner edge of the fold. The stomatodseum
reaches far downwards, and is characterised by a siphonoglyphe of considerable depth.
" A transverse section in the region of the stomatodaaum allows the mesenterial
arrangement to be recognised even by the naked eye. The longitudinal section having
been carried midway between two mesenteries on both sides, they were completely
intact, and the combination of the two sectional halves yielded a complete picture of
the mesenterial arrangement, which falls under the microtype. Sixty mesenteries in
all are present ; of these, after deduction of the regularly formed dorsal pairs, there fall
into the ventral zone on each side of the directive macromesenteries, twelve pairs,
consisting each of a macro- and a micro-mesentery.
"For a study of the anatomical relations in more detail, I made use of von Koch's
REPORT ON THE ACTINIARIA. 51
method of grinding. The integument is composed, as was stated above, of an interna]
softer zone, and an external zone penetrated by accessory deposits. The latter consist
mainly of clear angular fragments of sand ; but there occur also various indeterminable
mineral splinters of different colours, and finally, more sparingly, Sponge - spicules
and Foraminiferal shells. All these particles lie confusedly mingled, and so closely
together as to form a stout external rind ; between them they allow of oidy thin
mesoglcea-laniellse, in which are embedded fine nucleated fibres, as well as a few
stellate mesoglceal cells. The zone of mesoglcea, which is soft and free from deposits,
consists of a homogeneous matrix, in which sharply circumscribed lenticular cell-
islets are embedded in large numbers and of various sizes. They are especially
plentiful in the neighbourhood of the endoderm ; but, in passing outwards, every
gradation of size, up to fine fusiform structures, is met with. The plane of the
long axis of these cell-islets is always circumferential. The nucleated fibres are
extremely abundant in the mesoglcea ; they extend from the endoderm outwards,
their course being sometimes straight, but more generally undulating, with close coils
almost like a cork-screw. Besides the contents already mentioned, one observes also
the existence of stellate mesoglceal cells, which are sparsely scattered and emit fine
processes into the homogeneous matrix.
" The supporting lamina of the mesentery is well developed, and presents an antler-
like muscle-pennon. At its base passes a canal, filled with cells, and penetrating the
mesenteries for their whole length ; in transverse sections through the micromesenteries
this appears simple and cylindrical, but forms on the macromesenteries a longer cavity
divided up by cross anastomoses. This quite subordinate character accompanies the
microtype through all the genera, however different both externally and anatomically ;
no macrotypal form showing even a trace of this mesenterial canal.
" The sphincter of Sphenopus is mesodermal and simple, and is so far characteristic
that it commences incomparably deeper than in any other known Zoanthean ; it
extends so deeply downwards in the outer part of the body-wall, that, even in the
contracted animal, its lowest point lies in the same horizontal plane as the lower end
of the stomatodaBum. In longitudinal section one can see how, at its deepest point, the
bundles of fibrillre, like small circles, are laid so closely together that they appear
almost to form a continuous line. Above they are more extended, and place themselves
with the long axis perpendicular to the endoderm, from which they are only separated
by a narrow lamina of homogeneous mesoglcea. In this condition the sphincter forms a
system of bacillate fibrillse-bundles, which are arranged extremely regularly in the form
of a palisade. At the edge of the infolding of the body-wall the bundles begin to bay
out irregularly, and finally set themselves, on the indrawn part of the body- wall, to form
the sphincter proper, a plait of delicately branching and anastomosing bundles. This
circular muscle increases in bulk downwards, and terminates below with a rounded
52 THE VOYAGE OF H.M.S. CHALLENGER. :
end. It does not completely traverse the mesoglcea, but leaves free on either side
a homogeneous layer, which in its turn is bounded by a stripe reaching to the com-
mencement of the oral disc, and carrying the usual hard deposits."
Sp)henopus arenacens, R. Hertwig.
Habitat. — Station 187, Torres Strait, Australia, September 9, 1874 ; 6 fathoms.
Two specimens.
Sphenopus marsupialis, Steenstrup.
Habitat. — (a) Station 188, in the Arafura Sea, September 10, 1874 ; 28 fathoms.
One specimen. (6) Station 208, Philippine Islands, January 17, 1875 ; 18 fathoms.
One specimen.
In the Challenger material I have found four further examples of the genus
Sphenopus; two of these I have. determined as Sphenopus arenaceus on account of
their rusty red tint, and other two as Sphenopus marsupialis, in consequence of the
earthy-grey colour and the absence of a stalk. It seems to me, however, desirable that,
with an opportunity of more abundant and fresh material, a renewed study should
be undertaken to decide whether the received specific characters are variable, and
whether all three species should not be united in the single Sphenopus marsupialis.
APPENDIX TO THE ZOANTHE^.
Genus Stephanidium, n. gen.
Among the Zoantheae I include with some reserve a genus which is represented
by a single species, and has thus been insufficiently investigated. It differs from the
characteristic forms of Zoanthese in the absence of incrustations, and the non-formation
of a colony. Both characteristics, however, may be absent in true Zoanthese, e.g. the
soft-skinned Zoanthus and the solitary Sphenopidse. Of more importance is the fact that,
in spite of careful study, I have not yet been so fortunate as to demonstrate beyond all
doubt the decisive characteristic of Zoanthese, namely, the regular distribution of micro-
and macro-mesenteries. I consequently omit to give separate diagnoses of the species and
genus.
Stephanidium scliulzii, n. sp. (PI. I. fig. 14; PI. III. figs. 1, 7).
Habitat. — Station 209, off Zebu, Philippine Islands, January 22, 1875; 95 fathoms.
Dimensions. — Breadth, L5-2-2 mm. ; height, about 1*0 mm.
Some Actinia? were forwarded to me by Prof. F. E. Schulze, found among the
Hexactinellidse entrusted to him for description ; they were mainly small, insufficiently
characterised forms, which I did not care to investigate ; but among them occurred five
REPORT ON THE ACTINIAE! A. 53
specimens of one species, which I will here describe on account of the striking appearance
of the body.
The body of Stephanidium is in diameter l-5-2-2 mm., and about 1 mm. high in
the contracted condition. The epithelium had been stripped off at most points, and
remained only on the lowest parts of the body-wall, the mesoglcea thus being exposed
over a wide extent, and allowing the mesenteries to be seen through it. The resulting
appearance is drawn in PI. I. fig. 14, and was originally interpreted as follows: — I
believed that the surface was indented by deep furrows corresponding to the
mesenteries ; the ridges lying between these furrows become narrower, from a definite
part of the body-wall outwards ; they are extremely unequal in breadth, a broader and
a narrower ridge alternating regularly with one another, and to every broader ridge
corresponds, at the upper edge of the body-wall, a special structure of the following
nature : the edge of the body-wall is elevated into a kind of battlement (PI. III. fig. 7),
on the outer side of which are situated roundish or oval bodies, which call to mind the
marginal spherules of Actinia mesemhryanthemam. The longitudinal ridge of the body-
wall meets the spherule, splits into two forks, and surrounds the structure from below.
Sections through the animal, however, showed that the body-wall is smooth, and
that the appearance of furrows was caused by the insertions of the mesenteries. On
the other hand, the spherules are really present, and form evaginations of the body-wall,
above a spot which is marked by the position of the circular muscle (PI. III. fig. l). The
latter, in spite of the contracted condition of the Actinian, is of weak development,
and is merely a part of that endodermal circular muscle-layer which is at other points
hardly recognisable, but is here elevated into small folds. It is most obvious at those
places where it traverses the thickness of a mesenterial insertion ; here the endodermal
muscle-layer is not recognisable, but mesoglceal muscle-rings are embedded in the
region of the sphincter, largest at the upper end, and becoming gradually less obvious
in a downward direction, till one meets with small groups of only two or three fibres,
or even with completely isolated fibres.
Of the tentacles and oral disc it can only be said that the ectodermal muscle-
layer is strongly pleated.
The mesenteries, the number of which may be learnt even by superficial observa-
tion, amount to twenty-six, and are differentiated, as in the Zoantheae, into macro-
and micro-mesenteries. Of their arrangement, despite much trouble, I have not yet
arrived at a completely clear comprehension, but I could demonstrate the probability
that the directives of the one side are macro-mesenteries, those of the other micro-
mesenteries, that dorsal and ventral mesenterial zones meet with micro-mesenteries, and
that one pair is more developed on the one side than on the other.
The mesenteries (probably only macro-mesenteries) bore ripe male generative
organs. I was unable to recognise a siphonoglyphe.
54 THE VOYAGE OF H.M.S. CHALLENGER.
V. Ceeianthe^;.
Family 14, Cerianthid.e.
Genus Cerianthus, Delle Chiaje.
Cerianthus membranaceus, Spall.
Habitat. — Zebu, Philippine Islands, on the reefs.
Dimensions. — Length, in a contracted condition, 4-5-10"0 cm. ; breadth, I "5-1*8
cm. Two specimens.
UNDETERMINED SPECIES.
1. Station 153, in the Antarctic Ocean, February 14, 1876 ; 1675 fathoms.
An Actinian, nearly as thin as paper, strongly contracted and folded together,
1*5 cm. in size.
2. Station 173, off Matuku, Fiji Islands, July 24, 1874 ; 310-315 fathoms.
An Actinian, firmly fixed on a Gastropod shell (probably an Adamsia).
3. Station 195, off Banda, October 3, 1874; 1425 fathoms.
An Actinian, about 5 cm. long, very rotten.
4. Station 209, off Zebu, Philippine Islands; 95 fathoms.
An Actinian, 4 mm. broad, 2 mm. high, sessile, strongly contracted.
5. Zebu, 100 fathoms.
An Actinian, 6-0 mm. broad, 2 mm. high, of the usual structure; 12 complete
pairs of mesenteries, two directives, 36 (=12 + 24) small pairs.
6. Station 219, North of Papua, March 10, 1875 ; 150 fathoms.
One small Actinian, 1"5 cm. high, 1*'5 cm. broad, slightly contracted (probably
no sphincter), about 30 short, plump, sack-like tentacles. Preservation
inadequate for study.
7. Station 244, North Pacific, East of Japan, June 28, 1875 ; 2900 fathoms.
One Actinian, 12 mm. broad, 3 mm. high, very strongly contracted.
8. Station 286, South Pacific, between Tahiti and Valparaiso, October 16, 1875;
2335 fathoms.
An Actinian, 1"4 cm. broad, 0"4 cm. high, probably a Phellia.
9. Station 299, off Valparaiso ; 2160 fathoms.
Actinian adhering to a Dentalium; one elongated specimen (3-5 cm.), flattened
by contraction, in shape recalling the Amphianthidse.
TABLE OF CONTENTS.
Introduction,
Description of Genera and Species,
I. HexactinijE, .
Corallimorphida?,
Gorallimorphus,
CoraUimorphus rigidus,
Corallimorphus obtectus,
Corynactis,
Corynadis (?), sp. (?),*
Antheomorphidae, .
Ilyanthopsis,
Hyanthopsis longijilis*
Actinidae, .
Hormatliia,
Hormathia deKratula*
Eunodidse, .
Auladinia,
Auladinia, sp. (?),*
Paractida?, .
Dysadis,
Dysadis erassicornis,
Liponemidae,
Liponema,
Liponema multiporum,
Aulordiis,
Aulordiis paradoxa*
Phellidse, .
Phellia,
Phellia spinifera, .
Amphianthidffi,
Awphianthus, .
A mph ia nfhus 01 11a turn , *
Ilyanthidse,
Halcampa,
Halcampa kerr/uelensis, *
Halcamptella, .
Halcampella maxima*
Halcampella, sp. (?),*
page
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24
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28
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32
5G
THE VOYAGE OF H.M.S. CHALLENGER.
II. Paractinle, .
Sicyonidse, .
Sicyonis,
Sicyonis elongata,*
III. Edwardsue, .
Edwardsidse,
Edwardsia,
Edwardsia, sp. (?),*
IV. Zoanthe.e,
Zoanthidae, .
Zoanthus,
Zoanthus dance, .
Zoanthus confertus,*
Epizoanthus, .
Epizoanthus tJialamqphilu
Epizoanthus st el I arts*
Epizoanthus elongatus*
Epizoanth us cancrisocius,*
Corticifera,
Corticifera lutea*.
Curt icife ra tuber cu losa, *
Palythoa,
Palythoa anguicoma,*
Palythoa, sp. (?),*
Palythoa (?), sp. (?),*
Sphenopidse,
Sphenopus,
Sphenopus pedunculatus, "
Sphenopus arenaceus,
Sphewpus marsupial is,
Appendix to the Zoantheffi,
Stephanidium, .
Stephanidium sch ulzii,
V. Cerianthe^;,
Cerianthidae,
Cerianthus,
Cerianthus ,. embranaceut
Undetermined Species,
page
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PLATE I.
(ZOOL. CHALL. EXF. — PART LXXIII. — 1888.) — Dddd.
ai Cutieula.
ec Ectoderm.
en Endoderm.
g Generative organs.
h Mesenteries.
im Intermediary layer.
m Muscle-fibres.
PLATE I.
The lettering is the same in all the figures.
me Mesoglcea.
ms> Upper circular muscle.
ms2 Lower circular muscle.
o Ova.
r Marginal spherules.
ar Siphonoglyphe (cesophageal groove).
t Tentacles.
All statements given as to magnifying powers have reference to Zeiss's system.
Fig. 1. Zoanthus dance.
Fig. 2. Epizoanthus elongatus.
Fig. 3. Epizoanthus thalamophilus.
Fig. 4. Epizoanthus stellaris.
Fig. 5. Corticifera tuberculosa.
Fig. 6. Corticifera lutea.
Fig. 7. Palythoa anguicoma.
Fig. 8. Amphianthus ornatum ; x 4.
Fig. 9. Aulorchis paradoxa ; genital tube exposed by splitting the lip and the
stomatodseum.
Fig. 10. Aidorchis paradoxa; siphonoglyphe, stomatodseuni, oral disc, and
mesenteries exposed on removal of about one-third of the animal.
Fig. 11. Sphenopus pedunculatus.
Fig. 12. Zoanthus confertus.
Fig. 13. Liponema multiporum ; oral disc and stomatodasum evaginated.
Fig. 14. Stephanidium schulzii ; x 30.
Fig. 15. Epizoanthus cancrisocius.
(Figs. 1-7, 11, 12, 15 are after Erdmann.)
TheVovageofH.M.SThallengei"
Actiniaria (SuppJ PI. ..
•'■' •" •.,.•'.'■: ■; ; ■■■'-:•
v:--'v,:-->v-.:,;y;-'V:v
■
PLATE II.
PLATE II.
The lettering is the same in all the figures.
cu Cuticula.
ec Ectoderm.
en Endoderm.
g Generative organs.
h Mesenteries.
im Intermediary layer.
?/t Muscle-fibres.
me Mesoglcea.
ms1 Upper circular muscle.
m<! Lower circular muscle.
o Ova.
r Marginal spherules.
sr Siphonoglyphe (oesophageal groove V
t Tentacles.
All statements given as to magnifying powers have reference to Zeiss's system.
Fig. 1. Portion of the circular muscle of Hormathia delicatula. D, Oc. 2, somewhat
diminished.
Fig. 2. llyanthopsis longiftlis ; musculature of the oral disc. 1/1S, Oc. 1.
Fig. 3. Hormathia delicatula ; musculature of the oral disc. D, Oc. 2.
Fig. 4. Circular muscle and marginal spherules of Liponema multiporum. A, Oc. 1.
Fig. 5. Muscle-pennon of Halcampa Tcerguelensis. A, Oc. 2.
Fig. 6. Dysactis crassicornis ; transverse section through a tentacle, a3, Oc. 2.
Fig. 7. Portion of the tentacle represented in fig. 6, more strongly magnified.
A, Oc. 1.
Fig. 8. Phellia spinifera ; transverse section through the musculature of the oral
disc. E, Oc. 2.
Fig. 9. The same.
The Voyage of H.M. SXhallenger!
Actiniaria ISupp.l PI. N.
■
-•- -■ .
I
1 — ,;"r -,_
'-XsW-jm -•<>o&ip&&I°
PLATE III.
PLATE III.
The lettering is the same in all the figures.
cu Cuticula.
ec Ectoderm.
en Endoderm.
y Generative organs.
h Mesenteries.
im Intermediary layer.
m Muscle-fibres.
me Mesoglcea,
ins1 Upper circular muscle.
ms' Lower circular muscle.
o Ova.
/■ Marginal spherules.
sr Siplionoglyphe (oesophageal groove).
t Tentacles.
All statements given ns to magnifying powers have reference to Zeiss's system.
Fig. 1. Circular muscle of Stephanidium schidzii ; — la, in the region of an inter-
mesenterial chamber and of a marginal spherule; lb, in the region of the origin of a
mesentery.
Aulorchis paradoxa (tigs. 2-6).
Fig. 2. Transverse section through the oral disc. A, Oc. 1, somewhat diminished.
Fig. 3. Parts of a transverse section through the circular muscle, showing the
connection of the endodermal and mesoglceal muscle-layers : 3a, with D, Oc. 1 ; 3b,
with E, Oc. 1.
Figs. 4, 5. Sections through the stomidia. a3, Oc. 1.
Fig. 6. Transverse section through the mesoglceal musculature of the oral disc.
D, Oc. 1.
Fig. 7. Stephanidium schulzii ; upper edge of the body-wall, x 60.
The Voyage of H.M.S."ChaIIi
Actiniaria (Supp.) I'l III.
7.
PLATE IV
PLATE IV.
The lettering is the same in all the figures.
CM Cuticula.
ec Ectoderm.
en Endoderm.
ij Generative organs.
h Mesenteries.
im Intermediary layer.
m Muscle-fibres.
me Mesogloea.
mnl Upper circular muscle.
ms? Lower circular muscle.
o Ova.
r Marginal spherules.
sr Siphonoglyphe (oesophageal groove).
t Tentacles.
All statements given as to magnifying powers have reference to Zeiss's system.
Aulorchis paradoxa (sections through the genital tube, figs. 1-6).
Fig. 1. Connection of an ovum with the endodermal epithelium, probably by means
of a thread apparatus (" Faden-Apparat"). E, Oc. 2.
Fig. 2. Surface view of the germinal layer. I, Oc. 2.
Fig. 3. Longitudinal section through the germinal layer. E, Oc. 2.
Fig. 4. Transverse section through the lower part of the genital tube. A, Oc. 2.
Fig. 5. Transverse section through the upper part of the genital tube, in the region
of the oral disc ; the central detritus, which is probably produced by degradation of
epithelium, is omitted in the drawing. A, Oc. 2.
Fig. 6. Epithelial layer from the interior of the genital tube (cf. fig. 4). E, .Oc. 1.
Fig. 7. Epizoanthus thalamophilus ; section through the circular muscle (after
Erdmann).
Fig. 8. Horizontal section through the external and the invaginated portions of the
body-wall of Epizoanthus thalamophilus (after Erdmann).
Fig. 9. Hormathia delicatula. Portion of the partly invaginated body-wall cut out
and magnified slightly. The invaginated part bears the parietal spherules.
The Voyage of H.M.Sri
niaria (Suj
1 I 6
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