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THE 


JOURNAL 


OF 


THE LINNEAN SOCIETY. 


ZOOLOGY. 


VOL. XXVII. 


LONDON: 
SOLD AT THE SOCIETY’S APARTMENTS, BURLINGTON HOUSE, 
PICCADILLY, W., 
AND BY 
LONGMANS, GREEN, AND CO., 
AND 
WILLIAMS AND NORGATE. 
1899-1900. 


Dates of Publication of the several Numbers included in this Volume. 


No. 173, pp. 1- 59, published April 1. 1899. 


» 174, ,, 60-209, 5 July 1, 1899. 

» L75, ,, 210-268, » duly 1, 1899. 

55 WES 5, we ms November 1, 1899. 
» 177, ,, 345-458, i December 30, 1899. 
» 178, ,, 454-583, r April 24, 1900. 


FRiINTED BY TAYLOR AND FRANCIS, 


RED LION COURT, FLEET STREET, 


LIST OF PAPERS. 


Page 
BERNARD, Henry M., M.A.Cantab., F.L.S. 
Recent Poritide, and the Position of the Family in the Madre- 


porarian System. (With 2 illustrations.)............ 127-149 
On the Structure of Porites, with Preliminary Notes on the Soft 
Parts. (Plate 35:and 5 diagrams.) ........0ss0ce0: 487-503 


BovutENGER, G. A., F.R.S., F.Z.S. 
Reptilia and Amphibia of Lake Urmi. (See GutnrTuEr, 
IRGrsmaeH! 1) Gboeoonacoqod po doo boo oD DbOo boDE CEC 378-381 


Brive, Prof. T. W., Sc.D., F.L.S., Mason University College, 
Birmingham. 
The Air-bladder and its Connection with the Auditory Organ 
in Notopterus borneensis. (Plates 36 & 37) ........ 503-540 


Burr, Matcoum, F.Z.S., F.E.S. 
Orthoptera of Lake Urmi. (See GintHER, Ropert T.). 416-418 


Butier, A. G., Ph.D., F.L.S. 
Insecta (Lepidoptera Rhopalocera) of Lake Urmi. (See 
(Croagumenoin,, Taoist AN)! Beeboooseoduacqodueunooben 408-411 


Cartman, W. T., B.Sc., University College, Dundee. 
On the Characters of the Crustacean Genus Bathynella, Vej- 
dovsky. (Communicated by Prof. D’Arcy W. Thompson, 
(Os, JAS) CRD AO) oooossobccnceoo bu s0od00 338-344 


IV 
Page 
CAMBRIDGE, F. O. PicKARD. 
On some Spiders from Chili and Peru collected by Dr. Plate of 
Berlin. (Communicated by Prof. G. B. Howes, F.R.S., 
SecwerS yn Ge late 2) eae sce. leei wis le. ote exes leprae 15-22 


Coz, F. J., University College, Liverpool. 
On the Discovery and Development of Rhabdite-“ Cells” in 
Cephalodiscus dodecalophus, McIntosh. (Communicated by 
Prof. G. B. Howes, Sec.L.8.) (Plate 17) .......... 256-268 


Oricx, G. C., F.G.8., of the British Museum (Natural History). 
Note ona Jurassic Ammonite. (See GUNTHER, RopERT T.) 418-419 


Durrpen, J. E., Assoc. Roy. Coll. Sci. (Lond.), Curator of the 
Museum of the Institute of Jamaica. 
The Edwardsia-stage of the Actinian Lebrunia, and the Forma- 
tion of the Gastro-ccelomic Cavity. (Communicated by Prof. 
G. B. Howes, Sec.L.8.) (Plates 18 &19) .......... 269-316 


Exiwes, H. J., F.R.S., F.L.S. 
On the Zoology and Botany of the Altai Mountains. (With 
Grillustra tions.) Wisi epe sete ee ads cee eeicle atereeeetne ei eiseee 25-46 


Greeory, J. W., D.Sc., F.G.S. 
Fossil Echinoidea of Lake Urmi. (See GunrHEeR, RoBERT 


WV aes a's. joreie a javeiaoe aeuetanstie wose evens veusnaiighsustenenerctor eas bae -, 419-424 
Fossil Corals of Lake Urmi. (See GUNTHER, RopEerT T.) 424-480 


GunTuER, Dr. A. C. L. G., M.A., F.R.S., P.L.S. 
The Wild Sheep of the Urmi Islands. (See GuinruHEr, 
PROBE TL (Li.)i a revere aioe e's .dsu re aye lonolmereuctetleRome era fee ote 374-375 
Fishes of Lake Urmi. (See GunTHER, Ropert T.).... 381-391 


GinTHeR, Ropert T., M.A., F.R.G.S., Fellow of Magdalen College, 
Oxford. 
Contributions to the Natural History of Lake Urmi, N.W. Persia, 
and its Neighbourhood. (Communicated by the President, 
Dr. A. C. L. G, Giinther, M.A., F.R.S., F.L.S.) (With Map 
Plate 21 and Plates 22-30) ............c.seeseeee 345-453 


Hamesoy, Sir G. F., Bart. 
Insecta (Lepidoptera Phaleenz) of Lake Urmi. (See GUNTHER, 
OBER LPs) meee oeielereieveleleceeere terre eitenereiernerhsns te 411-414 


Page 
JOHNSTONE, JAMES, Fisheries-Assistant, University College, Liver- 


pool. 
On the Gastric Glands of the Marsupialia. (Communicated by 
Prof. G. B. Howes, Sec.L.S.) (Plate 1 and 1 woodcut.).. 1-14 


Kye, H. M., M.A. 

On the Presence of Nasal Secretory Sacs and a Naso-pharyngeal 
Communication in Teleostei, with especial reference to Cyno- 
glossus semilevis, Gunther. (Communicated by Prof. G. B. 
JsOwas, SRUIS,)) (EES SS) h ose aseoonsouncsoneoo. 541-555 


Luspock, The Rt. Hon. Sir Joun, Bart., M.P., F.R.S., LL.D., F.L.S. 
On some Australasian Collembola. (With 7 illustrations.) 334-338 


Metvitt, James Cosmo, M.B., F.LS., F.Z.S., and STanpen, 
RosBeERrr, Assistant-Keeper, Manchester Museum. 
Report on the Marine Mollusca obtained during the First Expe- 
dition of Prof. A. C. Haddon to the Torres Straits, in 1888-89. 
in (latest LOY aL Weer ier txoriiciictrsiel sioverol sic: salen citispal 150-206 


MicHakrt, ALBERT D., F.L.S., F.Z.S., F.R.MS. 
Acari of Lake Urmi. (See GUNTHER, ROBERT T.) ........ 407 


MircHELL, P. Cuaumers, M.A., F.LS. 
On so-called ‘ Quintocubitalism ” in the Wing of Birds; with 
special reference to the Columbe@, and Notes on Anatomy. 
(Plates 12 & 13 and 7 illustrations.) ................ 210-236 


Newron, R. Buuren, F.G.8., of the British Museum (Natural 
History). 
Marine Tertiary [Miocene] Mollusca of Laké Urmi. (See 


GUNTER PROBE R TMs) Winer. srs colerduatten ate notices oral arate 430-452 
Note on a Paleozoic Limestone of Lake Urmi. (See GUNTHER, 
FRVCOB ER Tye les) arene tegrey eter stei dv vor aiel eco ater ool: okel aver Aistepaierssovers 452-453 


Parsons, F. G., F.R.C.S., F.L.S., Lecturer on Comparative Ana- 
tomy at St. Thomas’s Hospital, and Hunterian Professor at the 
Royal College of Surgeons. 

The Position of Anomalurus as indicated by its Myology. 
QWathyoplllstratioms,)) par cet sisters clei! eevee soit) 317-334 


vi 
Page 
Pocock, R. 1., of the British Museum (Natural History). 
Chilopoda and Arachnida of Lake Urmi. (See GUNTHER, 
ROBE RTE) yeni cet lelotvensta ccs ele acs oNerslcharte Siete eee omen 399-406 


PycrarFt, W.P., A.L.S. 
Note on the External Nares of the Cormorant. (With 2 
HUIS TRMMONS,)) S15 66666006 a:séyaliniseie.roletayeng etomtireuane axa cattle 207-209 
Some Facts concerning the so-called “ Aquintocubitalism” in 
the Bird’s Wing. (Plates 14-16 and 2 illustrations.).. 286-256 


Rivewoonp, W. G., D.Sc., F.L.S., Lecturer on Biology at St. Mary’s 
Hospital Medical School, London. 

Some Observations on the Caudal Diplospondyly of Sharks. 
(Wath teil istration) pereiektete tit ntit tri rek teenie ierererene 46-59 

On the Hyobranchial Skeleton and Larynx of the new Aglossal 
Toad, Hymenochirus Boettgert. (Plate 31).......... 454-460 

Note on the Carpus of the new Aglossal Toad, Hymenochirus 
Boettgert. (Plate 31. fig.5.) ...... err ener uchen ee 460-462 


Scorr, Tuomas, F.L.S., Naturalist to the Fishery Board for Scotland. 

Report on the Marine and Freshwater Crustacea from Franz- 

Josef Land, collected by Mr. William S. Bruce, of the Jackson- 
Harmsworth Expedition. (Plates 3-9) .............. 60-126 


Smit, Epa@ar A., F.Z.8. 
Land and Freshwater Mollusca of Lake Urmi district. (See 
(Cripamenn, Inverse Wb) oso aogaocoaasoondeoGs00000C9 391-395 


STANDEN, Ropert, Assistant-Keeper Manchester Museum. 
Report on the Marine Mollusca obtained during the First Expe- 
dition of Prof. A. C. Haddon to the Torres Straits, in 1888-89. 
(S22 Mibsigyanvin, datans| OSIM.) onococcdctsoaca00008 150-206 


THomson, GrorGE M., F.L.S. 
On some New Zealand Schizopoda. (Plates 33 & 34) .. 482-486 


Wacer, Harozp, F.LS. 
On the Eye-spot and Flagellum in Euglena viridis. (Plate 
SH) GuocoeoasoasuogueageoduGo OE GO COD CoS oodaD aac 463-481 


Vil 


EXPLANATION OF THE PLATES. 


Illustrating Mr. W. T. Calman’s paper 
on the Characters of the Crustacean 
Genus Bathynella. 


PLATE ! 
eee Mr. Jas. Johnstone’s paper on the Gastric Glands of the 
: Marsupialia. 
2. { Tllustrating Mr. F. O. Pickard-Cambridge’s paper on Spiders from Chili 
and Peru. 
3.) 
4. | 
| Illustrating Mr. Thos. Scott’s paper on the Crustacea of Franz-Josef 
6. Ff Tand. 
7. 
2. | 
9. / 
10. Meee Messrs. J. C. Melvill’s and R. Standen’s paper on the 
11. Mollusca of Torres Straits. 
12. ae Mr. P. Chalmers Mitchell’s paper on so-called “ Quinto- 
13. cubitalism ” in the Wing of Birds. 
es pee Mr. W. P. Pycraft’s paper on so-called “ Aquintocubitalism ” 
16. in the Bird’s Wing. 
17 [pees Mr. F. J. Cole’s paper on the Development of Rhabdite- 
“cells” in Cephalodiscus. 
a Mr. J. H. Duerden’s paper on the Hdwardsia-Stage of 
19. Lebrunia. 


20 BATHYNELLA NATANS, ae 


ANASPIDES TASMANIA, Thoms. 


21. Map of Lake Urmi Basin. 

22, { OVIS OPHION, var. URMIANA, Illustrating Dr. A. Gimther’s paper on the 
Wild Sheep of the Urmi Islands. 

ABRAMIS URMIANUS. 

GoBIO PERSA. Illustrating Dr. A. Giinther’s paper on the 

LEUCISCUS ULANUS. Fishes of Lake Urmi. 

LEUCISCUS GADERANUS, 


23. { 
25 eae URMIANA. Illustrating Mr. R. T. Giinther’s paper on the 


24. 


Crustacea of Lake Urmi. 

Illustrating Mr. R. I. Pocock’s paper on the Arachnida of Lake Urmi. 

Insecta and Acarip from Lake Urmi. Illustrating Mr. R. T. Giinther’s 
paper. 

Illustrating Dr. J. W. Gregory's paper on the Fossil Hchinoidea and 
Corals of Lake Urmi. 


126. 
27. 


28 


Vili 


PLATE 
29. { Illustrating Mr. R. Bullen Newton’s paper on the Miocene Mollusca of 
30. Lake Urmi. 


1. eo Laryngeal, and Carpal Skeletons of Hymenochirus 
Boettgeri. Illustrating Dr. W. G. Ridewood’s papers. 
32. { Evetena viripis. Illustrating Mr. Harold Wager’s paper on the Hye- 
spot and Flagellum in Huglena viridis. 
1-5. SIRIELLA DENTICULATA. Illustrating Mr. G. M. Thomson’s 
Ss: les TENAGOMYSIS NOVE-ZEALANDIA. ! paper on some New Zealand 
34. 'TENAGoMysIS NOVH-ZEALANDIA, Schizopoda. 
35. Illustrating Mr. H. M. Bernard’s paper on the Structure of Porites. 
36. NotoPrerus BoRNEENSIS. Illustrating Prof. T. W. Bridge’s paper on 
37. | the Air-bladder and its connection with the Auditory Organ in 
Notopterus borneensis. 
38. ee Mr. H. M. Kyle’s paper on the Presence of Nasal Secretory 
Sacs and a Naso-pharyngeal Communication in Teleostei, &c. 


ERRATA. 


Page 367, line 8 from bottom, Bathyoscopus poceilus, H.-Sch., read Bythoscopus 
pecilus, Herr.-Schatf. 

Page 416, line 10 from top, Bathyscopus pocillus, read Bythoscopus pecilus, 
Herr.-Schaff. 


THE JOURNAL 


OF 


THE LINNEAN SOCIETY, 


On the Gastric Glands of the Marsupialia. By James Joun- 
sTonE, Fisheries-Assistant, University College, Liverpool 
(Communicated by Prof. G. B. Howes, Sec. Linn. Soc.) 


[Read Ist December, 1898.4 
(Puate 1.) 


W. A. Fores in 1881, in an account of an investigation of the 
anatomy of a Koala, was led, by a consideration of many points 
of similarity between this animal and the Wombat, to deduce a 
closer degree of relationship between the two forms than was 
then generally admitted. The common possession of a “ gastric 
gland,” a structure which Forbes considered to be so peculiar as 
- to render it highly improbable that it should have been inde- 
pendently acquired in two forms unrelated to each other, afforded 
him a “convincing token of their affinity.” Whether or not a 
fuller knowledge of the nature of this structure than Forbes 
possessed materially strengthens his contention, does not seem 
certain, but this investigation into the minute structure of the 
gland,—an investigation suggested by reading Forbes’ paper,— 
seems to disclose certain points worth noting. 

The first mention of the presence of a gastric gland in the 
stomach of Phascolomys seems to be that made by Home (4) in 
1808, who refers to it as closely resembling that of the Beaver, 
and ‘“‘ forming a very extraordinary peculiarity.” Home gives 
a very correct figure of the external appearance of the gland, a 
figure which Owen (9) repeats. Owen knew of the existence 
of the gland in Phascolarctus, and described it as closely 

LINN. JOURN.— ZOOLOGY, VOL. XXVII. 3 


2 MR. JAMES JOHNSTONE ON THE 

resembling that found in both Phascolomys and the Beaver. 
Later references to the gastric gland of the Marsupials in the 
literature are those of Huxley, 1871 (6), Flower, 1872 (2), Forbes, 
1881 (1), Fleischman, 1891 (3), and Oppel, 1896 (8). Forbes 
had the opportunity of examining the fresh stomach of a newly- 
dead Koala, and described the gastric gland as red and vascular, 
while the surrounding mucous membrane was pale ; he suggested 
a histological comparison of the glands of Phascolarctus and 
Phascolomys, with a view to finding whether the resemblance was 
more than an external one. — 

Oppel, in his important work on the comparative microscopic 
anatomy of the stomach, gives a very short description of the 
stomach of the Koala (8. pp. 291-2) and a figure of the gastric 
gland in section. The glands of the greater curvature and 
of the gastric gland itself he describes as “ Fundusdriisen.” 
There is no exact account of the limits of the gland regions, 
but a reference is given to a paper by Edelmann in which the 
absence of the peculiar “ Cardiadriisenregion”” in the Koala is 
described. Oppel’s work (8. p. 298) contains no account of the 
histology of the gastric gland in Phascolomys. 

Fleischman has a criticism of Toepfer’s work (11) on the 
comparative anatomy of the stomach in the Rodentia, and the 
author makes some interesting remarks on the parallelism in 
structure of the stomach in the Rodents and in the Diprotodont 
Marsupials, which lead him to a belief in a close genetic 
relationship of these two groups. 

“‘ Gastric giands”’ in the Mammalia puta Marsupials occur 
only in Manis among the Edentates, and in Castor among the 
Rodents. The glandular appendages on the stomach of Manatus 
evidently belong to a distinct category. The structure in the 
stomach of the Beaver (fig.1, I., p. 4), which seems to have been 
first mentioned by Schmidt (10)in 1805, was figured and described, 
so far as external characters are concerned, by Home (4), and its 
minute anatomy was more exactly described by Toepfer (11) in 
1891. The stomach of Castor is a simple one, lined throughout 
by a glandular epithelium. The cuticular lining of the esophagus 
ceases at the opening of that organ, and the gastric gland is 
situated to the pyloric side of it on the surface of the lesser 
curvature. Home gives a figure of a hand-section through one of 
the openings of the gland, which shows a number of short tubules 
opening into a short terminal duct, and forming a structure 


GASTRIC GLANDS OF THE MARSUPIALIA. 3 


more closely resembling the gastric gland of Phascolomys than 
the corresponding structure in Phascolarctus. But the gland 
_ differs from that in the Wombat and Koala in that the openings 
have a serial arranzement; though, from the accounts given by 
various authors, the number and precise disposition of these 
openings is irregular. Such variation also occurs in the two 
Marsupials. 

Manis javanica also possesses a complex glandular apparatus 
which may be termed a gastric gland, using the term in the 
same sense as in connection with the stomachs of Castor and the 
Marsupiais. But the stomach of Manis (tig. 1,1V., p. 4), which 
was minutely described by Weberin1891(12),is a very specialized 
one ; and the gastric gland is certainly morphologically a different 
structure from the “ gastric glands” of other Mammalia. The 
epithelium of the stomach, with the exception of certain patches, 
is a non-glandular one, cuticular in nature. Three groups of 
opevings, leading into much branched gland tubules, oecur— 
one at the pyloric aperture, one on the lesser curvature, and one 
on the greater near the orifice of the gastric gland. The latter 
is a prominent pad on the region of greater curvature, projecting 
into the cavity of the stomach and opening into it by a single 
opening. Within there is a system of complex foldings, lined 
with an epithelium, which consists of closely set gland tubules 
possessing the characteristic central and parietai cells. At the 
openiag of the gastric gland there is a portion of the cuticular 
epithelium covered over with small horny processes. A similar 
structure (Triturationsorgan) is found on the greater curvature 
at the opening of the gastric gland. 

The “ Vormagen” of Myoxus avellanarius (fig. 1, V., p. 4), and 
teh glandular appendage on the cardiac portion of the stomach 
of Manatus australis, which Leydig (7) compares with the gastric 
gland of the Beaver and the “ Vormagen ” of Myoxus, are evi- 
dently only analogous structures. 


My material consisted of the stomachs of specimens of Phasco- 
lomys and Phascolarctus. The Wombat was a full-grown animal, 
and the area on the smaller curvature occupied by the glandular 
thickening measured 2°8 cm. along the longitudinal axis, and 
3°4 em. on the shorter axis of the stomach. The gland was 
situated closer to the esophagus than in the case of the Koala, 


and its thickened rim partially embraced the lat‘er, several of the 
1* 


4. ; MR. JAMES JOHNSTONE ON THE 


openings being situated laterally to the ostium of the cesophagus. 
The number of these openings was about thirty, but it was difficult 
to count them exactly, as occasionally several seemed to open out 
from the bottom of little gutters, and very short ducts frequently 
branched quite near to the surface into several smaller tubules. 
Their arrangement was an irregular one, but they occupied a central 


Weise N 
el, ty aN 
AUTRE HAY 
\ \ hn) 


WiMWZ 


FA 
q 


AU 


Schematic representations of the stomachs of—I. Castor (after Home), II. 
Phascolomys (Home), III. Phascolarectus (Oppel), IV. Manis (Weber), 
V. Myoxus (Toepfer). I. and II. have their inner surfaces everted ; III. 
and IV. are in sagittal section. All the figures reduced.—é.v., bulbus 
ventriculi; duo, duodenum; g.g., gastric gland; @, cesophagus; py, 
pylorus ; ¢r, “ Triturationsorgan.” 

position on the thickened area, leaving an annular space free. The 

Koala was a young one, measuring about 26 em. from the snout 

along the back to the root of the tail. The gland on the lesser 

curvature was almost circular in shape and measured about 


GASTRIC GLANDS OF THE MARSUPIALIA. 5 


10 mm. in diameter. Although it had not attained its greatest 
development in point of size, I have no reason to suspect that its 
anatomical details differed in any essential respects from those 
characteristic of the gland in the fully-grown Koala. The-openings 
were much smaller than those in the Wombat, 25 in number, and 
occupied a central portion of the glandular thickening, round 
which was a part of the thickened area of mucous membrane free 
from openings. Unlike Phascolomys, the gland was situated 
nearly midway between the cesophagus and pylorus. 


PHASCOLOMYS. 


A section of the gland in the Wombat, taken parallel to the 
surface of the stomach, some little distance below the surface but 
before the bifurcation of the tubules has taken place, shows a 
number of tubules of varying diameter and of irregular distribu- 
tion. The area of the gland is sharply bounded by a circularly 
running tract, which contains the cardiac glands of the surround- 
ing epithelium of the stomach, and in which those glands are 
cut somewhat obliquely: this appearance is due to the plane 
of the section passing through the thickening of the gastric 
gland and surrounding epithelium. The gland itself is a some- 
what lenticular shaped pad, due entirely to the complex folding 
which the mucous membrane has undergone. The surface of 
this thickening, facing the interior of the stomach, is slightly 
depressed. ‘Towards the external surface the gland presents a 
convex border. In relation to each tubule there is a closely 
investing sheath of muscularis mucose, which accompanies it in 
its evagination outwards. This sheath consists mostly of a 
Jayer of plain muscle fibres, running transversely in relation to 
the long axis of the tubule, and, external to this transverse layer, of 
a very meagre, and in many places discontinuous, sheath of fibres 
running in the direction of the long axis of the tubule. Between 
these tubules is a space which is an extension of the submucosa of 
the general gastric epithelium, and which is occupied by areolar 
tissue containing blood vessels, lymph spaces, and muscle fibres. 
The latter are of two kinds—bundles of plain muscle fibres 
and of less abundant striated fibres. These are almost entirely 
derived from the muscular coats of the stomach, but probably 
also to some extent from the muscularis mucose. The primary 
tubules in the epithelium lining the evaginations which make up 
the gland are continuous with those m the cardiac region of the 


6 MR. JAMES JOHNSTONE ON THE 


stomach, and, like these, contain the typical central and parietal 
cells. In sections taken parellel to the surface of the gland, 
these tubules are for the most part cut longitudinally, though 
many are obliquely and even transversely cut. 

In a nearly sagittal section (Pl. 1. fig. 1), taking in pyloric 
and oesophageal orifices, the extent of the gland exhibited 
is rather less than in one passing to one side of the cesophagus. 
This is due to the extension of the gastric gland laterally 
to the cesophageal opening. All the layers of the stomach- 
wall are present. The serosa (ser.) is scanty in the middle line, 
but can be recognized ; the muscularis (muse.l., muse.t.) is rather 
reduced ; the submucosa (sm.) is largely encroached on by the 
folded mucosa and the presence of muscle bundles between these 
folds. The mucosa itself (muc.), to which the great thickness ot 
the wall is due, is seen to have undergone an elaborate folding. 
At the pyloric extremity there is a gradual transition between 
tubules containing the parietal cells, which are present in the 
gland, and the epithelium immediately adjoming on the pylorus 
(muc.py.), in which the primary gland-tubules contain only 
central cells. At the msophagus there is a sharp transition 
between those parietal-celled tubules which are present on the 
epithelium to the left of the former and the stratified cesophageal 
lining. 

In any one section in such a plane (PI. 1. fig. 1) one or more 
of the openings (0) of the gland are cut through. These are 
then seen to lead into a more or less complex system of cavities, 
into which the mucosa is prolonged. There appear, also, isolated 
portions of the mucosa often containing a lumen, and in which 
the primary gland tubules are cut in all possible planes. 
Accompanying each of the secondary tubules making up the 
gastric gland is a sheet of muscularis mucose (PI. 1. fig. 3, mm.). 
As stated above, this consists of a tunic containing fibres mostly 
running transversely to the long axis of the tubule. 

The muscularis is present over the whole surface of the 
gland. Passing from the pyloric to the cardiac extremity, the 
strongly developed layer of transversely running fibres forming 
the pyloric sphmeter (m.py.) thins out with the commence- 
ment of the gland-thickening, and is almost entirely replaced 
by a layer of striated muscle fibres (Pl. 1. fig. 1, mwse.t.). The 
course of these is both transverse and longitudinal to the lone 
axis of the stomach; and the arrancement is such that the 


GASTRIC GLANDS OF THE MARSUPIALIA. 7 


longitudinal bundles (fig. 1, musc./.) are external and the trans- 
verse ones (musc.¢.) internal. But many are cut obliquely, and 
over the area occupied by the gastric gland (2. e. almost the 
whole of the lesser curvature) the oblique and longitudinal 
bundles form the greater part of the muscularis. Mixed with 
these striated muscle bundles there are strands of plain muscle 
fibres (PI. 1. fig. 3, mp.), the number and masses of which diminish 
towards the cesophagus. Striated muscle fibres are found over 
the region of the pyloric glands, and indeed form the muscu- 
lature of the gastric-gland thickening. Delicate strands of both 
plain and striated (Pl. 1. fig. 3, m.cnt.) muscle fibres penetrate 
into the submucosa, between the secondary tubules of the gland, 
although most of these are unstriated fibres, and are derived 
probably both from muscularis and muscularis mucose. 

The course taken by the secondary tubules or involution of 
the gastric gland is, in Phascolomys, a comparatively simple one. 
Many openings on the surface of the gland lead into simple pits, 
but others are more complex, and a single tubule divides into a 
small number (2-6) of brauches. But the length of these side 
tubules relatively to their diameter is much less than in Phasco- 
larctus. The lumen is always a narrow one; and the thickness 
of the epithelium lining these tubules is generally greater than 
that on the free surface of the stomach. 


PHASCOLARCTUS. 


The most striking differences in the structure of the gastric 
gland of the Koala and that just described for the Wombat, lie in 
the greater compactness of the former and greater complexity in 
the ramifications of the tubules, and in the nature of the muscular 
coat. The latter is arranged in an external longitudinal (PI. 1. 
fig. 2, musc.J.) and an internal circular layer (muse.¢.). But the 
musculature over the gastric gland, which in Phascolomys was 
composed predominantly of striated fibres, is here made up entirely 
of nonstriated fibres (Pl. 1. fig. 4, muse.l., musc.t.). The muscu- 
laris is less strongly developed than in Phascolomys, and the 
transverse bundles are the more numerous. At the cesophagus 
striated muscle fibres (Pl. 1. fig. 2, m.e@.) are present; and the 
transition from these to plain fibres is, on the pyloric side of the 
cesophageal opening, a sharp one. On the cardiac side, however, 
the longitudinal musculature of the cesophagus extends for a short 
distance unmixed with nonstriated fibres over the surface of 


8 MR. JAMES JOHNSTONE ON THE 


the stomach. The striated, transverse cesophageal musculature 
is gradually replaced by the nonstriated musculature of the 
stomach. On the pyloric side, the distribution of striated muscle 
fibres is coterminous with that of the oesophageal epithelium. 
On the cardiac side this double line of demarcation between 
cesophagus and stomach is not so clear, but exists to a large 
extent. 

The primary gland tubules, as in the case of Phascolomys, are 
made up of central and parietal cells. But the tubules which in 
the Wombat are straight and unbranched, are here more complex. 
A single tubule (Pl. 1. fig. 6) is lined at its opening on the 
surface of the stomach with columnar cells (en.), which in the 
neck are replaced by clear cubical cells. At the first bifurcation 
the parietal cells (cp.) appear. In the tubule figured a group of 
eight lesser tubules is brought about by a triple bifurcation. 
Other of the glands are simpler or more complex, but this type 
seems to be the more general. 

Hach opening (0) on the surface of the gastric gland leads 
into a relatively wide tubule which, after remaining undivided 
for a length equal to twice or more than twice its own diameter, 
branches into two or more divisions, from which other branches 
are given off, either laterally or terminally. These branches 
end blindly ; in diameter they are about 0°5 mm. ‘There are no 
anastomoses. In the end a very complex cluster of short 
tubules is produced. The course of these branch tubules is 
generally transverse to the long axis of the stomach, so that 
in sections taken through the pylorus and cesophagus they 
are mostly cut transversely. In sections in the same plane, 
passing through the more peripheral portion of the glandular 
thickening, where there are no openings on to the internal 
surface of the stomach, the whole gland-pad consists of a closely 
packed mass of these branched tubules bound together by 
muscular and connective tissue. I have counted as many as 35 
of them cut transversely in a section passing through the peri- 
pheral portion of the gland. 


As might have been expected from their external form and 
situation, the gastric glands of Phascolomys, Phascolarctus, and 
of Castor do not exhibit any essential points of difference in their 
minute anatomy. From Toepfer’s account (11), the gland in the 


GASTRIC GEANDS OF THE MARSUPIALIA. 9 


Beaver, except in the arrangeinent of its openings on the internal 
surface of the stomach, is not dissimilar from the structures 
I have described in the Koala and Wombat. In each case 
the cavity of the stomach is prolonged by means of a variable 
number of evaginations into short tubular extensions which, 
branching frequently, end in series of tubules which do not anas- 
tomose but end blindly. The mucous membrane covering the 
general surface of the stomach is prolonged without interruption 
into, and forms the walls of, these systems of branching cavities. 
But whereas in Phascolomys the ramifications of the proximal 
portions of the evaginations are comparatively simple, the 
terminal portions short, the branches few, and the lumen a con- 
tracted one, in Phascolarctus each separate evagination forms a 
very complex system of tubules communicating with the stomach, 
which simulate the form of a true racemose gland. The ter- 
minal portions are relatively long and their lumina conspicuous. 
The primary gland tubules making up the gastric epithelium of 
the gland in both cases are cardiac glands similar to those found 
on the cardiac portion of the stomach and contain parietal cells. 
In Phascolomys these glands (Pl. 1. fig. 5) form a close-set 
series of straight unbranched tubules, in length about 0°56 mm., 
in which parietal cells extend from near the neck to the base 
of the gland, although they are rather concentrated at the 
middle of the tubule; while in Phascolarctus each gland (PI. 1. 
fig. 6), which is about 0°3 mm. in length, consists of a terminal 
neck portion, lined with columnar or cubical cells (en.), which 
branches several times, forming a series of long tubules opening 
through a common orifice at the lumen of the gastric-gland 
involution. Asin Phascolomys and the Beaver, the gland contains 
both central (cn.) and parietal (ep-) cells. 

Only the mucosa and muscularis mucose of the various coats 
of the stomach take part in the formation of the gastric gland. 
Submucosa and muscularis are only passively affected. In 
Phascolarctus the musculature over the thickened area is the 
layer consisting of outer longitudinally and inner transversely 
disposed fibres, which is characteristic of the other parts of the 
stomach. But in Phascolomys the musculature consists almost 
entirely of bundles of striated fibres; and the division of these 
into external longitudinal and internal transverse layers is not 
so evident as in the case of Phascolarctus, most of the bundles 
pursuing a more or less oblique course. 


10 MR. JAMES JOHNSTONE ON THE 


The great development. of striated muscle fibres over the 
gastric giand seems remarkable, but since the posterior limit of 
the striated musculature characteristic of the upper part of the 
cesophagus seems to vary in different animals, this extension into 
the region of lesser curvature of the stomach is probably without 
any special significance. But it suggests a comparison with the 
elandular “ bulbus ventriculi ” of Myoxrus avellanarius—an organ 
which, though not strictly homologous with, probably belongs to 
the same category of structures as, the gastric glands of the 
other mammals mentioned, and in which the musculature is a 
striated one. This organ was first described by Home (5), 
who compared it with the gastric gland of the Beaver, with 
which he found it to correspond “ very minutely in its internal 
structure.” It appears as a bulb-like enlargement of the lower 
portion of the cesophagus resting on the stomach, from which it is 
separated by a deep constriction. The cuticular liming of the 
cesophagus ceases at its anterior extremity, and is replaced by a 
very thick layer of glandular epithelium containing glands made 
up of central and parietal cells. Home describes these glands as 
having an arrangement similar to those in the glandular pad of 
the Beaver’s stomach. “ Hach orifice,” he says, “ exposes three 
small openings, these again lead to smaller processes, as has been 
described and delineated in the glandular structure of the Beaver.” 
Leydig (7), speaking of the glandular appendage on the cardiac 
part of the stomach of Manatus, where a compound tubular 
gland is found, refers to the “ Vormagen” of Jyorus as some- 
thing similar. Meckel regarded it as a truly avian structure. 
Toepfer (11), who investigated tke structure of this organ, found 
it to be provided with a thick layer of striated muscle-bundles 
which are a direct continuation of those found in the upper 
partof the cesophagus; and, in a discussion as to its morphological 
nature, he regards the presence of the epithelium rich in glands as 
affording a more reliable test of the morphological nature of the 
organ than the presence of an csophageal musculature. If the 
bulbus is gastric and not cesophageal in origiv, then the whole 
stomach of Myoaus—i. e. the true stomach and bulbus ventriculi— 
is homologous with the simple stomach of Castor provided with a 
concentration of cardiac glands on the region of its smaller 
curvature, since in A/yozus such a concentration occurs round the 
entrance of the cesophagus. 

In the Beaver, as in Phascolomys and Phascolarctus, there is the 


GASTRIC GLANDS OF THE MARSUPIALTA. sgl 


important difference in the disposition of the gland regions of 
the stomach as compared with Myoaws, that the cardiac glands of 
the latter, that is the glands composed of parietal and central cells, 
are restricted to the bulbus ventriculi, and the rest of the stomach 
contains only pyloric glands; while in the three first mentioned 
forms the whole stomach is glandular, and cardiac glands, although 
predominantly crowded together in the gastric-gland thickening, 
are not wholly restricted to it as they are to the bulbus of 
Myoxus, but are found to a certain extent over the region of the 
fundus. The area on the stomach of the Beaver occupied by 
cardiac glands is therefore to be regarded (on Toepfer’s view) 
as homologous with the “ Vormagen” of JZyoxus. The extent to 
which these glands extend over the lateral wails and region of 
ereater curvature of the stomach has not been investigated, nor 
have the precise limits of the pyloric glands been determined, 
either for Castor or for the Marsupials referred to. 

In Manis, although the gastric gland on the greater curvature 
of the stomach cannot be regarded as homologous with those in 
other mammals, it is more comparable to the bulbus ventricu1 
of Myoxus than to any of the others, for here there is a definite 
restriction of the cardiac glands to a portion of the stomach- 
wall, where they are arranged as a complex glandular organ, 
the rest of the stomach being non-glandular, with the excep- 
tion of certain patches which contain glands probably homologous 
with the pyloric glands of the other mammalia. Here, too, it 
seems that a direct correlation with the nature of the food and 
the other parts of the alimentary canal and disposition of the 
mucous membrane of the stomach exists; a proposition that can 
hardly be made with regard to the gastric glands of either the 
Marsupialia or the Rodentia. There are simple stomachs un- 
provided with any complex glandular apparatus in the Phalangers, 
as in the Myoxide other than JL. avellanarius. 

The formation of the “‘ gastric gland”’ as we find it in the 
Marsupials seems to be explicable in one of two ways. First, as 
the result of a tendency to localization of the gland regions,—a 
tendency which finds its expression alike in the structures I have 
described, in the glandular cardiac appendage of Manatus, and 
more completely in the “ gastric gland”’ of Afanis and the bulbus 
ventriculi of yoxvus. But we may also with Oppel (8. pp. 402-3) 
in the case of the Beaver and the Marsupials, regard it as 
due to the necessity for increase in the area of the gastric 


12 MR. JAMES JOHNSTONE ON THE 


epithelium containing cardiac glands (“ progressive development’’), 
and in Manis as due to the restriction of that area (‘‘ retrogressive 
development”). In the first case it is related to the necessity of 
digestion of an exceeding large amount of food-material ; and 
in the second to the digestion of a small quantity of easily 
assimilated food. 

The terms ‘“ gastric gland” of English authors and “ grosse 
Magendriise” employed by the Germans, are alike misnomers. 
The organ, as both Oppel and Toepfer have pointed out, is not 
a gland in the sense in which that term is legitimately employed, 
but a complex evagination of the gastric wall, bearing, in common 
with the rest of the stomach, true glands; andas such it does not 
seem as if its value can be very great as affording any trust- 
worthy indication of the phylogenetic history of the animal in 
which it occurs. In the case of two forms like Phascolarctus 
and Phascolomys, known, from other considerations, to be nearly 
related to each other, the common presence of a “ gastric gland ” 
may indeed afford reason for their closer approximation; and 
in these animals the organ does seem to be homologous, although 
in the Koala its structure is more specialized than in the case 
of the Wombat. But statements as to an organ in the Rodentia 
resembling the “‘ gastric gland’” of Marsupialia, and conclusions 
as to a closer degree of genetic relationship of these two 
groups, deducible therefrom, in view of the great specialization 
of the stomach occurring in some Rodents, must, I think, be 
received with hesitation. 

I am indebted to Prof. G. B. Howes for supplying me with 
the material reported on, and for kindly furnishing me with 
many of the works cited. Part of this investigation was done 
while I was still a student under him inthe Research Laboratory 
of the Royal College of Science, London. 


BIBLIOGRAPHY. 


1. Forprs, W. A.—“On some points in the Anatomy of the 
Koala (Phascolarctus cinereus). Proce. Zool. Soc. London, 
1881, pp. 180-195. 

2. Frowrr, W. H.—Lectures on the Comparative Anatomy of 
the Mammalia. Med. Times and Gazette, vol.1. pp. 215-673 
& vol. i. pp. 1-945. 1872. 


GASTRIC GLANDS OF THE MARSUPIALIA. 13 


3. Fierscuman, Dr. A.—“‘ Bemerkungen uber d. Magen d. 
Rodentia.” Morphologisches Jahrbuch, Bd. xvii. 1891, 
pp. 408-416. 

Discussion of Toepfer’s results (11). 

4. Homer, E.—“ An account of some peculiarities in the Ana- 
tomical Structure of the Wombat, with observations on the 
Female Organs of Generation.” Phil. Tians. Roy. Soe. 
1808, Pt. IT. pp. 304-811, pl. ix. 

5. Howe, E.—‘“ Observations on the Structure of the Stomachs 
of different Animals, with a view to elucidate the Process of 
converting Animal and Vegetable substances into Chyle.” 
Phil. Trans. Roy. Soc. London, 1807, Pt. II. pp. 189-178, 
pis. V.—xiil. ; 

Huxiey, T. H.—Anat. Vertebrate Animals. London, 1871, 
pp: 327 & 436. 

7. Leypie, Dr. F.—Lehrbuch d. Histologie d. Menschen u. d. 
Thiere (pp. 551). Hamm, 1857. 

Oppet, A.—Lehrbuch d. vergl. mikrose. Anat. d. Wirbelthiere. 
Teil I. Der Magen. Jena, 1896. 

9. Owen, R.—Comp. Anat. and Phys. of Vertebrates, 1868, vol. 

i. pp. 412-3, fig. 310. 

10. Scumrpr, Fr. Adolf—De Mammalium esophago atque 
ventriculo. Spec. inaug. medicum. Hale ex offic. Bath., 
pp: 28. 1805. 

11. Torprer, K.—‘‘ Die Morphologie der Magens der Rodentia.” 
Morphologisches Jahrbuch, Bd. xvii. 1891, pp. 380-406, 
Taf. 2 A. 

12. Wezer, M.—“ Beitr. z. Anat. u. Entwickl. d. genus Manis.” 
Zool. Ergebnisse einer Reise in Niederlandisch-Ostindien, 
Bd. II. Taf. i-ix., pp. 116. Leiden, 1891. (Abstract in Oppel 
(8), pp. 305-311, figs. 211-220.) 


o 


so) 


EXPLANATION OF PLATE 1, 


Fig. 1. Verticai section through the gastric gland of Phascolomys parallel to the 
long axis of the stomach, but passing to one side of the esophageal and 
pyloric apertures. X 6 diameters. 

Neither in this nor in figs. 2, 3, & 4 has any attempt been made 
to represent the histological details of the mucous epithelium. But 
broken lines and circles are employed to indicate the direction in 
which the primary gland-tubules are cut. 


14 GASTRIC GLANDS OF THE MARSUPIALIA. 


Fig. 2. Vertical section through the gastric gland of Phascolarctus parallel to 
long axis of the stomach but passing to one side of the cesophagus. 
x 8 diameters. 

3. Portions of three contiguous tubules of the gastric gland in Phascolomys 
from its pyloric extremity. Vertical section, parallel to longitudinal 
axis of stomach. > 20 diameters. 

Fig. 4. Three contiguous tubules of the gastric gland of Phascolarctus. Vertical 

section, parallel to longitudinal axis of stomach.  X 380 diameters. 

Fig. 5. Transverse section of a primary gland-tubule from the gastric gland of 

Phuscolomys. xX 694 diameters. 

Fig. 6. A primary tubule from the gastric gland of Phascolarctus, seen in optical 

section. X 266 diameters. 


Fi 


= 
oa 


Reference Letters. 


bv. Blood-vessels. 
cn. Neck-cell in primary gland tubule. 
cp. Parietal cell. 
ce, Central cell. 
ct. Connective tissue between primary gland tubules. 
lp. Lumen of primary gland tubule. 
ts, Lumen of gastric-gland tubule. 
musc.l. Longitudinal layer of muscularis. 
musc.t. Transverse layer of muscularis. 
m.int. Nonstriated nuscle fibre between gastric-gland tubules. 
mp. Nonstriated muscle fibres. 
mst. Striated muscle fibres. 
mm. Muscularis mucose. 
m.e@. Cisophageal musculature. 
m.py. Pyloric sphincter. 
muc.. Mucosa. 
muc.p. Pyloric mucosa. 
» Opening of the gastric-gland evaginations into the stomach. 


oS 


0 

ser, Serosa. 

sm. Submucosa. 
C. Cardiac extremity of gastric gland. 
P. Pyloric extremity of gastric gland. 


Bp. _— 


Johnstone. Lrnw. Soc. Journ. Zoot. Vou. XX VIL Pr. 1. 


J. Johnstone del. 
Parker & Percy lith. 5 Geo West k Sons imp. 


GASTRIC GLANDS OF MARSUPIALIA. 


ON SPIDERS FROM CHILI AND PERU. 15 


On some Spiders from Chili and Peru collected by Dr. Plate of 
Berlin. By F. O. Picxarp-Campriner. (Communicated 
by Prof. G. B. Howes, F.R.S., Sec.L.S.) 


|Read 17th November, 1898. ] 
(PuaTE 2.) 


THE present paper contains a list of the Spiders collected by 
Dr. Plate on a voyage extending from T'umbez in North Peru, 
down the coast of Chili, to Cape Horn. The collection contained 
nineteen species, of which seven are new toscience. One species 
has been described in MS. for some years by Mr. R. I. Pocock, 
of the British Museum of Natural History, and this description 
has recently been published. There is nothing very striking or 
noteworthy in this collection, although all spider-forms are 
interesting from whatever part of the world they come. 

There is one curious point to be noted. Two species taken in 
the island of Juan Fernandez, 500 miles west of Valparaiso, are 
identical with examples which are indigenous to Northern Europe, 
namely Teutana grossa and Dysdera crocota. ‘These spiders are, 
however, found almost all over the world, so that their presence 
in Juan Fernandez is not perhaps so astonishing, when we find 
them also in the Sandwich and Cape Verde Islands, separated by 
thousands of miles of ocean and continent. 


List of Species. 
THERAPHOSID ZH. 
Paraphysa manicata, E. Simon. Coquimbo, Chili. 
Phryxotrichus roseus (Walck.) ? Corral. 


Citharoscelus Kochi, Pocock. Coquimbo, Chili. 
ScyTopipz. 

Stcartus thomisoides, Walck. Coquimbo and Iquique, Chili 
DyspDERID&. 

Ariadna marima (Nic.). Juan Fernandez. 

Dysdera crocota, C. Koch. i ns 
CLUBIONID&, 

Sparassus bombilius, sp. n. Amciven,} 

Gayenna maculatipes, Keys. Juan Fernandez. 
DicryNipm™. 


Amaurobius Plater, sp. n. Tumbez, Peru. 


16 MR. F. O. PICKARD-CAMBRIDGE ON 


ARGIOPID&. 
Meta nigrohumeralis, sp. 1. Juan Fernandez. 
Tmeticus Defoer, sp. 0. Dp ” 
4 Platei, sp. n. 9 op 
THERIDIID®. 
Teutana grossa, O. K. 99 - 
Theridion tepidariorum, C. Kk." Cavan. 
PHOLCIDE. 
Pholcus americanus, Nic. Chili. 
AGELENIDA. 2 
Rubrius annulatus, sp. n. Corral. 
Lycosipa. 
Lycosa implacida, Nic. Coquimbo. 
»  fernandezt, sp. Nn. Juan Fernandez. 
» australis, K. Simon. Tekenilla, Cape Horn. 


Family THeraPHosip#. 


Parapnysa MANICATA, Sim., Hist. Nat. Ar. i. p. 166. 
A single adult female was taken at Coquimbo by Dr. Plate 
in Sept. 1893. 


PHRYXOTRICHUS ROSEUS ( Walck.)? 
An adult male and female of this species were taken by 
Dr. Plate in Corral, Dee. 1894. 


CrrHarosceLus Kocutt, Poc. Ann. Mag. Nat. Hist. April 
1899. (Pl. 2. figs. 13-15.) 

A single female of this species was taken by Dr. Plate at 
Coquimbo. 


An adult female Hurypelma, which I hesitate to assign to any 
species, as well as several immature forms, occurred in Dr. Plate’s 
collection. 


Family ScyTrop1p#. 

SICARIUS THOMISOIDES, Walck. 

Thomisoides terrosa, Nic., Gay, Hist. de Chile, vol. iii. p. 350, 1849. 

Walckenaer forestalled Nicolet in publishing a description of 
this species and used that author’s generic name as the specific. 
The genus and species belong undoubtedly to Walckenaer, who 
obviously saw Nicolet’s work in print, though not published. 

The type of the genus can scarcely be terrosa, Nic., for this was 
an unpublished name. To the single species which Walckenaer 


SPIDERS FROM CHILI AND PERU. 17 


recognized, the name thomisoides was given when he gave 
the name Svcarius to the genus. SS. thomisoides, Wlk., is there- 
fore the type of the genus Sicarius. If we regard Sicarius as the 
proper generic title on the ground that, at the time of its publi- 
cation, Thomisotdes existed only in manuscript, or, if in type, as 
an unpublished name, as M. Sition has done in Hist. Nat. Ar. 
2 ed. p. 271, we must also regard thomisoides as the proper 
specific title, which M. Simon has however not done in the place 
mentioned. 

Several females and a male of this species were taken by 
Dr. Plate at Coquimbo and Iquique. 


Family DyspERrIDs. 


ARIADNA MAXIMA (Wicolet). 

Dysdera maxim1, Nic., Gay, Hist. de Chile, Zool. iii. p. 541, plate ii. 
fig. 6. 

Two examples were taken by Dr. Plate in the island of Juan 
Fernandez. 


Dyspera crocota, C. K., Die Arachniden, Bd. v. p. 81. 
Three adult males of this species were taken in J uau Fernandez 
by Dr. Plate. 


Family CLuBronips. 


SPARASSUS BOMBILIUS, sp. n. (Pl. 2. figs. 1 & 2.) 

3g. Total length 15 mm.; carapace 7X7; legs, i. 34— 
ii. 80—ill. 27—1iv. 28. 

Colour. Carapace mahogany-brown clothed with golden-grey 
hairs. Mandibles black, clothed with long golden-grey hairs. 
Fang-groove fringed with ferruginous-red hairs. Legs very dark 
mahcgany, almost black, hirsute with stiff golden-grey hairs; 
abdomen clothed with rough golden-grey hairs, a transverse band 
across the shoulders black, continued indistinctly down the dorsal 
line, with a second narrower transverse dark band just before the 
middle, followed towards the spinners by a narrow central band 
having very short indistinct oblique lateral rays. Lateral area 
dark brown; sub-ventral area golden-grey, embracing the spinners; 
central ventral area black. Sternum deep brown, clothed with 
short golden-grey hairs. Palpi: femora and patelle orange, tibiz 
and tarsi deep brown or black. 

Structure. Carapace almost circular, convex. Anterior row of 
eyes procurved, posterior row straight. Central antericrs 

LINN. JOURN.— ZOOLOGY, VOL. XXVIL. 2 


18 MR. F. 0. PICKARD-CAMBRIDGE ON 


slightly larger than laterals: central posteriors slightly smaller 
than posteriors ; central quadrangle slightly broader than long. 
Tibial apophysis of palpus S-shaped, falciform, its apex directed 
outwards and slightly dilate, compressed, spiraliform. Palpal 
bulb surrounded by a stout circular spine, lying in a groove in 
the surface of the bulb. : 

A single adult male of this beautiful species was taken by 
Dr. Plate at Amciven. 


GAYENNA MACULATIPES, Keys., Brasilianische Spinnen, p. 141, 
pl. iv. 97 & 97a. 

Although the adult male example from Juan Fernandez is 
considerably larger than the type in Keyserling’s collection, yet 
I am unable to find any difference in the structure. 

An adult male and an immature female were taken by 
Dr. Plate in Juan Fernandez. 


Family Dictynip#. 


AMAUROBIUS PrATEI, sp.n. (PI. 2. fig. 3.) 

3. Total length 12 mm.; carapace 5°5 x 3°5; leg i. 21. 

Colour. Carapace pale orange-yellow, suffused with sootv- 
black on the margins of the caput. Mandibles deep mahogany- 
brown. Legs pale yellow, deepening to mahogany-brown towards 
the extremities. Legs i—femora mahogany-brown, with two 
indistinct dark transverse spots beneath; patella, tibia, pro- 
tarsus, and tarsus deep mahogany-brown. Legs ii—femora pale 
yellow, with two transverse spots beneath, the rest asin i. Legs 
iu. and iv. paler; tibiz and protarsi indistinctly annulate with 
sooty-brown. Abdomen black, with pale anterior dorsal area, 
containing four ill-defined white spots, followed to the spinners 
by a series (3 or 4) of narrow white A-shaped bars. Apex of 
abdomen deep black. Sternum pale orange-yellow. 

Structure. For figure of palpus see Plate 2. fig. 3. 

A single adult male was taken by Dr. Plate at Tumbez. 


Family ArGropips. 


Mera NIGROWUMERALIS, sp. n. (PI. 2. fies. 4 & 5.) 

2. Total length 15 mm. 

Colour. Carapace pale orange-yellow ; the caput, central stria, 
and a submarginal line rich brown. Mandibles rich mottled 
brown. Legs pale orange-yellow ; femora suffused with brown 
towards the apex; patelle, tibia, and pretarsi of i. and ii. 


SPIDERS FROM CHILI AND PERU. 19 


suffused and indistinctly annulated with brown ; those of iii. and 
iv. more definitely annulate. Abdomen pale clay-white with 
deep black shoulder-spots—or a large black spot, bilobate and 
margined with pure white behind; a central brown band runs 
to the spinners, giving off lateral sooty-brown oblique lines, 
blending with the mottled brown and black lateral areas; 
ventral surface paie yellow-brown, margined on each side with a 
straight, narrow, broken white band. Sternum rich brown. 

Structure. A figure of the vulva appears on the Plate (fig. 5). 

Six adult females were found on the island of Juan Fernandez 
by Dr. Plate, but no males. This species bears a general 
resemblance to the European Meta Menardi, which is also found 
in North America, but is none the less quite distinct from it. 


Tmeticus Deroet, sp.n. (PI. 2. figs. 6 & 7.) 

2. Total length 7-5 mm. 

Colour. Carapace, sternum, and legs rich orange-mahogany- 
brown. Abdomen black, with a pale yellow curved band on each 
side of the anterior dorsal area, enclosing a large black trian- 
gular patch; bebind these longitudinal bands come two short 
transverse pale bands on each side. The lateral area has two 
longitudinal indistinct pale bands, and the ventral surface bears 
in the centre a dull pale blotch. 

Structure. The mandible exhibits on the outer side the usual 
transverse strie ; and a figure of the vulva will be found on the 
Plate (fig. 7). The inferior margin of the fang-groove bears a 
row of five short cusps; the superior a row of seven longer teeth. 
The tibie of the legs bear a pair of short erect dorsal bristles. 

Two adult females were taken by Dr. Plate on the island of 


Juan Fernandez. 


Tmericus Pratret, sp.n. (Pl. 2. figs. 8 & 9.) 

©. Total length 85 mm. 

Colour. Carapace, mandible, legs, and sternum dull orange- 
brown, more or less suffused with darker brown. The base and 
apex of the femora are suffused with darker brown. The tibie 
and protarsi have each an indistinct broad dark annulation 
towards the base, and another narrower one towards the apex. 
Abdomen olive-brown; dorsal area mottled with dull yellow- 
white blotches, the best defined being a series of five transverse 
A-shaped bars, their apices not contluent, extending from the 


middle to the spinners. ; 
9% 


20 MR. F. O. PICKARD-CAMBRIDGE ON 


Structure. Mandibles very stout, with two rows of teeth, the 
inferior with 9 small conical cusps, the superior with 11 long 
stout teeth increasing in length, the longest being the fifth to 
the tenth. Outside the superior row of teeth lies an irregular 
row of stiff bristles. yes. Posterior row straight ; eyes subequal, 
centrals slightly closer together, three fourths of a diameter apart; 
anterior row slightly reeurved; centrals small, one fourth smalier 
than the laterals, half a diameter apart, over one diameter from the 
laterals: lateral eyes equal and in contact. Tibiz of legs with 
two erect bristles above, one near the base, the other towards 
the apex. The mandibles bear a series of strie on the outer 
side, and the coxe of the palpi a few stiff bristles on the inner 
side. 

A single adult female was taken on the island of Juan 
Fernandez by Dr. Plate. 


Family TuEeripiip#. 


TEUTANA GROSSA, C. K. 

Theridion grossum, C. K., Die Avachniden, iv. p. 112. 

Two adult females were taken on the island of Juan Fernandez 
by Dr. Plate. 


THERIDION TEPIDARTORUM, C. K., Die Avachniden, viii. p. 75. 
A single adult female was taken at Cavan by Dr. Plate. 


Family Prorcips. 


PHOLCUS AMERICANUS, Wc. 


Two examples, which probably belong to this species, occurred 
in Dr. Plate’s collection from Chili. 


Family AGELENID&. 


RUBRIUS ANNULATUS, sp.n. (PI. 2. fig. 10.) 

©. Total length 15 mm. 

Colour. Carapace, sternum, and legs pale orange-yellow, man- 
dibles black, maxille and labium deep brown. The sides and 
anterior margin of the caput are deep brown, besides a narrow 
central and two narrow lateral brown bands converging at the 
central stria. An irregular brown band runs midway between 
the central stria and the margin, consisting in reality of a serics 
of blotches on the lateral strie which are confluent. The margin 
of the sternum is suffused with dark brown. The femora of 


SPIDERS FROM CHILI AND PERU. Zyl 


the legs bear three indistinct dusky annulations; the patelle are 
dusky along their sides, while the tibie have two deep brown 
annulations cn the upperside, obsolete below. The protarsi also 
exhibit two dark annulations. The abdomen, which hag some- 
what lost its colour, is dull white mottled with dark grey. 

Structure. Carapace long, narrow; caput raised, convex, with 
three rows of stiff hairs along the three brown bands. Posterior 
row of eyes slightly procurved, eyes equal, centrals rather closer, 
one and an eighth diameter apart. Anterior row straight, centrals 
slightly smaller than the laterals, three fourths a diameter 
apart, the same distance from the laterals. Clypeus equal to a 
diameter of the anterior centrals. Mandibies stout, triangular, 
conical, gibbous above (as in Cotes). The margins of the fang- 
groove bear, each, 5 teeth, and the upper margin bears also a 
fringe of long incurving hairs. Legs spinose, femora with five 
or six spines above, tibiz with three pairs beneath and one on each 
side, toward the apex, and often one towards the base. Protarsi 
with three or fuur pairs of stout, long spines beneath. Anterior 
tarsi with a double series of stiff bristles beneath ; posterior with 
some spines towards the apex, amongst the bristles. Tarsal 
claws three. 

I refer this spider to the genus Rubrius, E. Simon, although 
the character assigned to it, wita several other genera, ‘“ che- 
larum margo inferior dentibus, 3 or 4 armatus,” does not strictly 
apply. The species is undoubtedly closely allied to R. sub- 
fasciatus, Sim., Mission Sci. du Cap Horn, p. 14 (1887), but still 
distinct from it. The central anterior eyes are not very much 
smaller than the laterals, certainly not “ plus duplo ” as sM. Simon 
says of subfasciatus. 

A single adult female was taken by Dr. Plate at Corral. 


Family Lycosrpz. 


Lycosa IMPLACIDA, Vic. 
Gay, Hist. de Chile, Zool. iii. p. 358, pl. ii. fig. 10. 
A single specimen (2 adult) from Coquimbo, Chili. 


LYCOSA FERNANDEZI, sp.n. (PI. 2. figs. 11 & 12.) 
3. Total length 16 mm.; carapace 8 x 6; legs, i. 26—H, 23:5— 
il. 22°5—1v. 29:5 


2. Total length 25 mm. ; carapace 10 x7; legs, i. 27—ii. Op 
il, 24—iy. 33. 


DO ON SPIDERS FROM CHILI AND PERU. 


3. Colour. Carapace mahogany-brown, clothed with fine 
golden-brown pubescence. Legs clothed with golden-brown pu- 
bescence. Anterior tibie and tarsi sooty black. Abdomen: 
shoulders black ; anterior half with a central dorsal brown bar, 
its three angles picked out with black, followed towards the 
spinners by two or more small triangular black marks; on each 
side is a row of three or more black spots, contrasting with 
some white ones lying adjacent. Ventral surface unicolorous 
pale golden grey. 

©. Colour similar to that of the male, except that the golden 
pubescence has a deeper yellow-olive tinge. Abdomen entirely 
clothed on the upper side with golden brown pubescence, with a 
central dorsal suffusion of rust-red. Shoulders slightly tinged 
with brown, anterior dorsal mark very obscure, followed towards 
the spinners by two or three very obscure pale A-shaped marks. 

3. Structure. Palpus & organs. See Plate 2. fig. 11. 

OF i Vulva. See Plate 2. fig. 12. 

Four females and one male of this species, belonging to the 
Trochosa group, were taken by Dr. Piate in the island of Juan 


Fernandez. 


Lycosa australis, £. Simon, Bull. Soc. Zool. Fr. 1884, p. 3. 

A single adult male, most probably belonging to this species, 
was taken at Tellenika on the False Cape Horn. M. Simon’s 
type was taken on the le Hoste. 


EXPLANATION OF PLATE 2. 


. Sparassus bombilius, sp.n. Adult male. Right palpus from beneath. 
Full figure. 


08 


1 

2. 39 9 29 99 

3. Amaurobius Platei, sp.n. Right palpus. 
4. Meta nigrohumeralis, sp.n. Adult female. Full figure. 
. 


pear ¥ Vulva. 

6. Tmeticus Defoei, sp.n. Adult female. Full figure. 
7. -s a a x Vulva. 

8. 0 Plate, sp. nu. Adult female. Full figure. 
9 S rf PY Br Vulva. 


10. Rubrius annulatus. Adult female. Vulva. 

V1. Lycosa fernandezi. Adult male. Palpal bulb. 

12. ‘5 es Adult female. Vulva. 

13. Citharoscelus Kochit. Female. Coxa of leg i., inside, showing stridu- 
lating spines. 

14. a ‘ " Coxa of pedipalp, outside, showing 
stridulating spines. 

15. a 3 3 A single stridulating spine, enlarged. 


* 


Pickard-Cambridge. Lim. Soc. Journ. Zoon.Vou.XXVII. PL. 2. 


F.0.Pickard-Cambridge del. et. ith. West, Newman imp 
SPIDERS FROM CHILI & PERU. 


eyes 
sir 
aa 


a 


* 
Sar RETF 

Pah Me 
Rreaiten tori 


er 


fest 
ve 


ZOOLOGY AND BOTANY OF THE ALTAI MOUNTAINS. 23 


On the Zoology and Botany of the Altai Mountains. 
By H. J. Exwes, ¥.R.S., F.L.S. 


[Read 15th December, 1898.] 


Ir would be impossible, within the limits at my disposal, to give 
more than a brief sketch of the results of the journey which I 
undertook during the summer of 1898; but as the Altai Mountains 
are almost unknown to English naturalists, and as I am not 
aware that any Huglishman has previvusly visited or written 
anything about that country, I think it will be of interest 
to point out what a wonderful field for research exists, within 
three wecks’ journey of England, and one which is practically 
less known to naturalists than many parts of Central Africa. 
The country is so extensive, aud the season for travelling so 
short, that I was only able to visit a portion efit. So far as 1 
am aware, the only travellers who have writen on the natural 
history of the country, are Pallas, whose great work is well- 
known, though now rather out of date; Ledebour and Buage, 
who 60 years ago compiled an excellent account of the botany 
of the Altai; Helmersen, who has described the geology of 
the country; and Tchihatcheff, a well-known Kussiau traveller, 
who published an account of his travels in French in 1852. The 
few English travellers who have preceded me, so far as I know, 
are Major Cumberland and Mr. St. George Littiedale, both of 
whom went there solely for sport; and Mr. Rew, who last year 
made a rapid ride vd Kusnetsk, Kobdo, and Uliassutai to Irkutsk. 
Since the first half of this century Iam not aware of any zoologist 
who has written, except in Russian, anything of much importance 
about the Altai; and though no doubt there are many valuable 
memoirs on various paris of the country in the Russian language, 
especially relating to the geology and mineralogy, the majority 
of travellers who have gone to Siberia recently have passed along 
the main high road to Irkutsk, leaving the Altai far to the south 
of them. My late friend Seebohm, who visited Siberia especially 
to study ornithology, confined his exploration to the lower valley 
of the Yenesei river ; and one of the ideas which he expressed to 
me, and which made me specially anxious to visit Siberia, was that 
the Yenesei valley formed probably the most natural boundary 


24 MR. H. J. ELWES ON THE 


between the Eastern and Western Palaretie, or, as it is now more 
properly called, the Holarctic Region. This idea was not con- 
firmed by my own observations, for I found many species of 
butterflies and some birds which were previously only known 
from Dauria and Amurland; and I think that if any boundary 
can be fixed, it must be sought west of the Altai Mountains. 

By the Altai Mountains I understand the western extension of 
the great mountain-range between Semipalatinsk and Irkutsk, 
which is divided from the mountain-ranges of Turkestan by the 
Irtysch river, and from the Himalayas and the great mountain- 
ranges of Central Asia by the Gobi Desert and Mongolia. The 
southern part of this range, known as the Great Altai, is in 
Chinese territory and is at present very little known. I may add 
that the political boundary between Rusria and China follows 
roughly, or is supposed to follow, what I daresay geographers 
thovght was the southern watershed of the Obb and Yenesei 
rivers, but-as a matter of fact the whole of the upper waters 
of the Yenesei are in Chinese territory. One of our objects 
was to visit this great mountain valley, containing the head- 
waters of the Yenesei, which is almost unknown to the Russians 
themselves, though Clements and one or two other travellers 
have passed through parts of it, and it is annually visited by 
a number of fnr-hunters and gold-miners. The upper region of 
the Yenesei, from the sources of the Kemehik to Lake Kossogol, 
including the great valleys of the Beikem and Ulukem between the 
Tannu-ola mountains and the Russo-Chinese frontier, is almost 
uninhabited, and unknown except to those in search of fur and 
gold, though in a general way its outlines are reproduced on the 
map. On reaching St. Petersburg, I made every endeavour to get 
information as to the possibilities of visiting this country, and as 
to what had been done recently by Russian travellers. I was 
introduced to M. Beresowsky, who had accompanied Potanin in 
two of his journeys right through Mongolia to China, and 
had collected what information he could. ‘The result of 
these enquiries, and what I was told by M. P. P. Semenov, 
Vice-president of the Russian Geographical Society, who was 
most friendly to me, tended to show that the greater part of 
these valleys were almost impassable in summer; because the 
mountains were covered with dense forests, and the valleys 
were very marshy, intersected with numerous streams and rivers 
which were difficult to cross with horses. J was also informed, 


q 


ZOOLOGY AND BOTANY OF THE ALTAT MOUNTAINS. 2: 


i) 


and afterwards found it to be the case, that the higher regions 
of the Altai were liable to excessive rains and snow-storms in 
summer, making it a very difficult country for horses to traverse. 
Tchihatcheff (the only writer so far as I know who has crossed 
from the head-waters of the Tchuja to the head-waters of the 
Abakan, the principal western tributary of the Yenesei in 
Siberia) has stated that to cover a distance of about 200 miles 
cost him six weeks’ hard travelling, and the loss of about half 
of the 150 horses with which he started. This will give an 
idea of the difficulties of an explorer in some parts of this 
country. 

But to return to the question of what should properly be called 
the Altat Mountains. I would draw the line eastwards at the 
boundary of the Government district of the Altai, which is some- 
what east of the water-parting between the tributaries of the 
Yenesei on the east and those of the Obb on the west. I do not 
include all those eastern mountains which are known by the 
names of Sayansk, &c., and stretch away as far as the south-west 
end of Like Baikal, though they are from a physical point of view 
part of the Altai. 

Until two years ago the journey to Siberia was one which had 
to be undertaken in the winter, for the roads are almost im- 
passable when the snow is melting in spring. But now the 
railway enables one to do in comfort in ten days what formerly 
entailed three weeks or a month of hard sledging. 

Although my special object was an investigation of the 
Lepidcptera, a subject at which I have been chiefly working of 
late years, I was also anxious to collect birds and plants as well; 
and if M. Beresowsky had not left us just when the real work of 
collecting began, I should have done much more in that direction. 
- My companion, Mr. Fletcher, very kindly assisted me during the 
leisure time he could spare from the pursuit of the Wild Sheep, 
which was the special object of his Journey; and I have in con- 
sequence been able to bring home a very fine and complete 
collection of Butterflies, including specimens of about 200 species. 
I also made a ccllection of Altai plauts, but unfortunately, after 
they were packed and sent off, the horse that carried them 
went down ip one of the numerous rivers the crossing of which 
is the principal difficulty of travelling in the Altai. When I 
subsequently unpacked them, they were almost entirely spoilt ; 
but as Mr. Littledale had made a good collection the year before, 


96 MR. H. J. ELWES ON THE 


which was presented to Kew, I am able, with the permission of 
the Director of the Royal Gardens, and the kind assistance of 
Mr. J. G. Baker, to exhibit specimens of a few of the most 
interesting. 

The language is a difficulty not easily overcome by English 
travellers, for though Russian is of course spoken all over the 
Russian Dominions, yet the inhabitants of the frontier districts 
are Tartars, Mongols, and Kirghiz; aud when all communications 
have to be made through two uneducated interpreters in languages 
which they do not understand perfectly themselves, it is not easy 
to get accurate or full information on any subject. 

There is no difficulty in getting leave from the Russian 
authorities, with proper introductions, to visit any part of 
Western Siberia, and in fact I was seldom asked for a passport 
the whole time I was in the Altai. The Chinese in Mongolia 
are also very civil and friendly to travellers provided with pass- 
ports ; and the only difficulty which prevented us from extending 
our journey far into the Chinese dumiuions was the lack of time, 
and the unwillingness of the Russian subjects to run the risk 
of having their horses stolen by the Kirghiz, who are subject 
only to Chinese authority. 

We left Moscow on the 18th of May, 1898, and, travelling by 
rail without stopping, reached the crossing of the Obb river in 
six days and nights. The whole of our route between the Ural 
Mountains and the Obb lay through an immense flat plain, parts 
of which are marshy and more or less clothed with birch woods, 
and wherever the soil is dry enough a considerable amount of 
cultivation is seen. Large quantitics of wheat were stored at 
the railway-stations for export, but I believe the price is now 
insufficient to enable Siberian wheat to be profitably exported to 
England. 

Spring had hardly set in when we reached the Obb river, 
though it had been quite hot at Moscow; and almost the only 
flowers which I noticed in the Steppe were a blue-and-yellow 
anemone (Anemone patens), closely allied to if not identical with 
A, Pulsatilla, and the brilliant yellow flowers of Adonis vernalis. 
On reaching the Obb river, we had to wait two or three days for _ 
a steamer to take us up to Barnaoul, which is the chief, indeed _ 
I may say the only, town in the whole of the Altai Government. 
From there we drove in two days across the Steppe to Biisk, 
which, although a place of 18,000 inhabitants, is really a large 


ZOOLOGY AND BOTANY OF THE ALTAL MOUNTAINS. DT 


and very dirty village rather than a town. As this is the last 
place, however, of any importance towards the Chinese frontier, 
it is the centre of a large trade in wool, tea, and furs. A Dutch 
fur-trader whom we found there showed us a collection of furs 
he had made during the previous winter. Though in the early 
days of the Russian conquest valuable furs were in such abundance 
in this part of Siberia that all the taxes were paid in them, the 
better kinds have now become very scarce, and the only skins 
which we saw in great quantity were those of the Yellow Marmot 
of Mongolia, which are now exported in enormous quantities at a 
very low cost, and I believe are dyed in Europe to imitate the fur 
of the Mink. We must have passed 400 or 500 horse and camel 
loads of them on the road between Biisk and the frontier. 

All up the valley of the Obb are large villages, some of them 
over a hundred years old, and sometimes two or three miles long, 
and the peasants seem fur the most part prosperous and wealthy, 
according to a Russian peasant’s ideas of wealth; but we were 
informed that the country was so far filled up to the foot of the 
mountains that there was no more room for emigrants on a 
large scale, except in the forest country to the eastward, and 
most of the emigrant trains that we-saw on the railway were 
going farther to the East, into the districts of Krasnoyarsk and 
Irkutsk. 

At Busk, which we reached on the last day of May, there were 
signs of spring. The birch and poplar trees were just opening 
their buds, and here we obtained our first view of the outlying 
spurs of the Altai Mountains ; the country between Barnaoul and 
Busk, which from the map one would suppose to be mountainous, 
being grassy rolling downs of low elevation. After some delay 
in getting horses, we fairly entered the mountains on June 6th, 
aud at once found an immense improvement from a naturalist’s 
point of view in the appearance of the country. Many of our 
well-known old garden flowers such as peonies, erythroniums, 
rhododendrons, and anemones were in full bloom, while in some 
parts the ground was completely covered with the flowers of 
Tris ruthenica. Butterflies also began to appear ; and though I 
did not get any worth mentioning until we reached Ongodai, 
which is four long days’ drive through the mountains, I could see 
that the country was of far greater interest than anything we 
had hitherto passed through. 

The Obb river divides just below Biisk into two great branches— 


28 MR. H. J. ELWES ON THE 


the Bija, which flows out of Lake Teletskoi, and the Katuna, 
which rises on the Chinese frontier in the great mountain called 
Bielucha, which is over 10,000 feet high. 

Ongodai is about 180 miles from Biisk, and is the last place on 
the trade route to Kobdo in Mongolia where Russians are settled. 
It lies in the valley of the Ursul river—a broad valley of steppe- 
like character, whose waters flow into the Katuna, one of the 
two great sources of the Obb river. The character of the 
country thus far is much like that of Colorado. The slopes 
exposed to the south are arid, and covered with dwarf plants and 
dry grass, while the slopes facing north are wooded, in some 
places very thickly, with larch, while spruce grows in the flat, 
marshy bottoms of the valleys. 

We entered the country of the Altai Tartars shortly before 
reaching Ongodai. They are mostly nomads, and from their 
appearance are probably nearly akin to (if not identical with) 
the tribes from whom the Ottoman Turks originally sprung. 
Though Tchihatcheff, who spoke Turkish, says he could not 
understand their language or make himself understood, yet I 
recognized the few Turkish words I know for common objects, 
and the appearance of some of the men is exactly like that 
of the Turks of Asia Minor. They have enormous herds of 
horses, one chief owning as many as six or eight thousand, and 
live in summer in movable ‘ yourts’ or tents, covered with felt on 
wicker frames, like those of the Kirghiz, whilst in winter they 
live in pyramidal huts covered with larch bark, which resemble 
in form the lodges of the North-American Indians. 

From Ongodai we had to carry all our baggage on horseback, 
for beyond this the road is impassable, even for country carts, 
and there was considerable delay in getting horses sufficient for 
our outfit, for the spring had been very severe, and most of the 
working horses had already left with merchandise for Kobdo. 
A day’s journey beyond Ongodai, we crossed the valley of the 
Katuna in a deep rocky gorge, about 3000 feet above the sea, by 
a ferry of log canoes, whilst the horses were all made to swim 
over. After crossing the Katuna we passed over a mountain, 
about 5000 feet high, and descended again into the Katuna valley, 
close to its Junction with the Tchuja river, which we followed for 
six days nearly to its source in a great open valley lymg about 
6000 feet above the sea, and known as the Upper Tchuja Steppe. 
In this Steppe was a sort of frontier market-place, called Kuch 


ZOOLOGY AND BOTANY OF THE ALTAI MOUNTAINS. 29 


Agatch, where the Chinese and Russian traders formerly met 
for purposes of barter, and a small Russian custom-house is 
maintained here, whose superintendent was the only Russian 
official we saw anywhere beyond Ongodai. 

Large numbers of horses and camels and a few sheep and 
yaks are pastured in the Tchuja valley. The yaks that I saw 
appeared to be larger and of a finer breed than those I have seen 
in the Himalayas. The camels were all of the double-humped 
Asiatie variety. 

Having reached Kuch Agatch, we got a fresh lot of horses 
and Tartars to go into the mountains lying south of the Tehuja 
Steppe, where we expected to find the great Wild Sheep (Ovis 
ammon, Linn.), some of whose heads I now exhibit. This is the 
finest and largest wild sheep in the worid, although its horns are 
not equal in spread to those of Ovrs Poli of the Pamir. 

Another animal whose acquaintance I specially wished to make 
was the great Stag of the Altai, known in Asia as the Maral. 

As Mr. Lydekker, when preparing his recently published work 
on the Cervide, had not sufficient material to enable him to 
decide as to the species of Cervus found in Siberia, I have, with 
the kind aid of the Duke of Bedford, brought here for exhibition 
several heads and horns which are of scientific value ; for I believe 
hardly any from Siberia have hitherto been seen in England, and 
some of them would be well worth figuring. First, I show three 
heads of so-called Cervus eustephanus of Blanford; a species which 
was described by him from shed horns from the Thian-shan moun- 
tains, and is well figured by Mr. Lydekker * from a living speci- 
men in the Duke of Bedford’s menagerie, the original painting 
of which His Grace has lent me for exhibition. Though smaller 
than the Wapiti, it is a much larger animal than Cervus maral 
from North Persia, the Caucasus, and Asia Minor, which is rightly, 
I think, regarded by Mr. Lydekker as an Eastern race of Cervus 
elaphus. A head (fig. 1) which was brought by Regel from the 
Thian-shan mts., though not a large one, is typical in character of 
this species ; whilst the other two (figs. 2 & 3), which I procured 
in the. Altai, are not so typical, and, as I thought at first, had 
more resemblance to the horus of the European Red Deer. 
Mr. Blanford, however, has convinced me that they are nearer 
in character to those of Cervus eustephanus. This species has 


* ‘Deer of all Lands,’ 1868, pl. vi. p. 105. 


30 MR. H. J. ELWES ON THE 


now become scarce in a wild state in the Russian Altai owing 
to the number which are shot by the native and Russian hunters, 
who sell their horns, if killed while “in the velvet,”’ at high prices 
to the Chinese. They are, however, kept alive in parks at several 


Fig. 1. 


Cervus asiaticus, var. songarica, Severtzoff. 
= Cervus eustephanus, Blanford. 
From the Kuldja district. (Regel.) 


places in the Altai for the sake of their horns, which are annually 
cut for sale, and which sometimes realize as much as 100 roubles 
a pair at the rate of 10 roubles a pound. 

The killing of these deer has now been prohibited by the 


ZOOLOGY AND BOTANY OF THE ALTAL MOUNTAINS. Bi 


Government in the Altai distr'ct, and we never saw the animal 
in a wild state, and though we picked up horns, shed many 
years previously, in the high treeless mountains south of the 
Tehuja valley (one of which I exhibit to-night ‘to show what a 
large size they attain), I believe that they are now very scarce 
except in the heavily wooded country east of the Katuna. In 


Cervus asiaticus, var. sibirica, Severtzoff. From the Altai. (Elwes.) 


the Yenesei and Abakan valleys this deer, or a nearly allied form 
of it, is much more numerous; and I saw some horns from the 
Yenesei valley iu the St. Petersburg Museum which I thought 
had much more resemblanee to those of Cervus elaphus, having a 
distinct cup or crown of 6 or 7 tives branching from the same 
point on the beam, as in large old specimens of the Red Deer, 


Bi) MR. H. J. ELWES ON THE 


oe 


quite unlike the horns of Cervus eustephanus, in which there is 
no cup-like formation on top, and in which, as in the Wapiii, 
the main beam is deflected backwards; the 4th tine, usually the 


Fig. 3. 


Cervus asiaticus, var. sibirica, Severtz. Bought at Barnaoul. (Elwes.) 


largest, pomts f rwards and all the upper points are nearly in the 


same pline. 
Severtzoff, in the Proceedings of the Moscow Society of 


Naturzlists, vol. vii. p. 2 (1873) (translated from the Russian in 


ZOOLOGY AND BOTANY OF THE ALTAI MOUNTAINS. 30 


Ann. & Mag. Nat. Hist. 4th ser. 1876, vol. xvill. p. 386), was the 
first to call attention to the resemblance between the Stag of the 
Thian-shan, which he calls Cervus maral,and the Wapiti: he divides 
the Asiatic species into two forms, which he calls —“ (a) szbirica,” 
from the Altai, the Yenesei, and the wooded hills of the Sayansk 
and Transbaikal mountains, and “ (b) sexgarica,” from the Thian- 
shan. Mr. Blanford, describing Cervus eustephanus (Proc. Zool. 
Soc. 1875, pp. 637-640), has alluded to Severtzoft’s memoir 
(‘Turkestanski Jevetni,’ p. 108), which being in Russian he 
could not follow, and therefore could not say whether the form 
described as var. sonrgarica is identical with eustephanus or 
not. Now, however, that both he and Mr. Lydekker admit its 
identity, it seems to me that the name soongarica having priority 
should be used, though Mr. Lydekker styles this race -Cervus 
canadensis asvaticus. 

With regard to Severtzoff’s var. sibirica, however, it is impossible 
at present to decide its exact relationship to the others. I am 
able to exhibit four pairs of horns which have been lately sent 
by Herr Hageubeck of Hamburg to the Duke of Bedford, which 
were procured by Herr Dorries in the Chingan mountains and 
from the Sutschan river in Manchuria, which Mr. Lydekker, 
who has seen them, considers to belong to the race which he calls 
Cervus canadensis Luehdorfi. This race he regards as more 
nearly related to the Western Wapiti, from the Pacific coast, 
Washington, and Vancouver’s Island, than to the Thian-shan or 
Altai race. He had seen no adult male or horns of this form when 
he published his work, ‘ The Deer of ail Lands;’ and though all 
these four pairs (evidently those of adult stags) are much smaller 
than any race of the Wapiti with which I am acquainted, they 
certainly to some extent show the horn character ot the Wapit’, 
rather than that of the Red Deer (figs.4.& 5). If it be admitted 
that they belong te a race of Cervus canadensis, we have this 
curious fact in geographical distribution, namely, that the race 
of the Western American coast more nearly resembles the 
Eastern Asiatic race than it does the Rocky Mountain race, 
which latter, on the other hand, has resemblance to the Altai 
and hian-shan race, most widely separated from it in point of 
distance *. 

* Since this paper was read I have received from Herr Hagenbeck another 
head procured by Dérries in the mountains south of Lake Baikal, which 
probably belongs to the same race as the Altai Deer, fig. 6 (p. 35). 

LINN. JOURN-—ZOOLOGY, VOL. XXVII. 3 


3b4 , MR. H. J. ELWES ON THE 


Another point which should be taken into consideration in 
deciding the spec’fic relations of these Deer, is the peculiar call of 
the stags in the rutting time. The Red Deer in all its forms, 
both in Europe and Asia, utters at this season a deep hoarse roar, 
ending in three or four loud grunts; which may be imitated by 
the human voice with the aid of a conch-shell or glass bottle. 


Fig. 4. 


Tf 


Cervus asiaticus, var. Liihdorfi, Bolau. 
From the Sutschan river, Manchuria. (Dorries.) 


On the other hand, the Wapiti in all its races (Asiatic and 
American) has a very different cry, which is described by hunters 
as a whistle. Although I have never listened to this ery myself, 
T have heard hunters in the Altai imitate it with the hollow stem 
of a plant, whilst in America a tin whistle is used for the same 
purpose. Radde who, in his well-known and valuable work on 
the natural history of Amur-land, regards the Stag of the East 
Sayansk mountains and Dahuria as a race of Cervus elaphus, 


ZOOLOGY AND BOTANY OF THE ALTAI MOUNTAINS. 35 


mentions this peculiar cry and reduces it to musical notation *. 
Now though such a fact as this may be looked upon as trifling 
by some naturalists, I venture to think that, as in the case of 
the song of birds, the cry of an animal is a point of material 
value in deciding the question of specific alliance. 

if, on examination, M. Biichner should consider that tie horns 


Fig. 5. 


iS os meee 


Another exaruple of C. Liikdorfi. From the same source as fig. 4. 


, 


from the Yenesei, of which I have spoken, belong to a race 
different from that of the Altai and Thian-shan, and are also 
different from that found on the Amur and in Manchuria, and 
if it should. prove that this race, notwithstanding that its horns 


* Mr. J. E. Harting informs me that the notes indicated by Radde (op. 
ett.) accurately ‘express the call of the Wapiti as heard by him repeatedly 
in the Regent’s Park Zoological Gardens, and are quite unlike the call of 
the Evropean Red Deer, 


3* 


36 MR. H. J. ELWES ON THE 


occasionally assume an elaphine character, is a race of Cervus 
astaticus, then I think the synonymy and distribution of the 
Asiatic raees will stand as follows, though I confess that no 
characters can be given by which these races can be exactly 
defined, and probably they will be found to intergrade. 


Fig. 6. 


Cervus asiaticus, var. sibirica ? 


_ From mountains south of Lake Baikal, (Dorries.) 


CERVUS CANADENSIS ASTATICUS vel Cervus asiaTicus, Severtzoff. 


(a) Var. songarica, Severtzoff, Turkest. Jevotn. p. 109 (1878), 
Cervus eustephanus, Blanford, P. Z. 8. 1875, p. 637. 
O. canadensis asiaticus, Lydekker, Deer of all Lands, 
p: 104 (1898). 
Hab. Thian-shan Mts. of Kuldja; South Altai. 


ZOOLOGY AND BOTANY OF THE ALTAIT MOUNTAINS. on 


(6) Var. sibirica, Severtzoff, 1. e. 
Cervus elaphus, var., Radde, Reisen im Sitiden von Ost- 
Sibirien, 2 vols. (1862-63). 
Hab. Altai; ? Yenesei Valley, E. Sayansk, Transbaikalia ; 
Dauria ;? Upper Amur. 


(c) Var. Luehdorfi.—Cervus Lithdorfi, Bolau, Abhandl. Ver. 
Hamburg, vi. p 33 (1880). 
C. canadensis Luehdorfi, Lydekker, op. cit. p. 102. 
C. isubra, Noack, Humboldt, viii. p. 6, fig. 5 (1889). 
Hab. North and East Manchuria, Sutschan river. 


As Herr Biichner, of the St. Petersburg Museum (where alone 
can be found sutlicient material on which to decide the question 
at issue), has declined at present to express an opinion on a 
subject which he considers should be monographically treated, 
it seems the more desirable to place on record the views above 
suggested. 


The Ibex of the Altai, Capra sibirica, Pallas, the head and 
horns of which I exhibit, is nearly allied to the Himalayan Ibex, 
and is common in some parts of the mountains, though very 
difficult to get atinsummer. The Elk was formerly more numerous 
in the northern districts, but has now become extremely rare ; 
and the single head which I brought back resembles those which 
J have seen from European Russia, differing somewhat in the set 
of the horas from the Elk of Norway. 

The Roe, Capreolus pygargus, Pallas, is very common in some 
parts of the Altai and Sayansk mountains, and is a very much 
larger and finer animal than the European Roe. I was at one 
time under the impression that the wide spread of the horns 
was a peculiarity of this species; but it would appear, from the 
nine heads which I have brought for exhibition—six from the 
Upper Yenesei valley, and three from the Altai—that this pecu- 
liarity is by no means constant, and that there is nothing but 
their size to distinguish them, so far as I see, from the European 
race. 

The Musk Deer is also very abundant near the upper limit of 
forest growth, and is snared in quantities by the natives. We 
saw as many as 200 skins in one merchant’s store. 

Reindeer are said by Radde to occur in some parts of the 
eastern Sayansk range, where they are also kept ina domesticated 


38 MR. H. J. ELWES ON THE 


state, but so far as I could ascertain they do not exist m any 
part of the Alta. 

Birds were not so numerous as I expected, although Cranes 
and Ducks were plentiful in the marshes of the Kurai and 
Tchuja Steppes. I was astonished to find a Scoter breeding 
here, which proves to be the species described as Ordemia 
Stejnegeri, and which is an inhabitant of the N.W. American 
coasts and North Pacific. It has never been hitherto procured, 
as | am informed by M. Alpheraky (who is at present engaged 
on a monograph of the Anatide of the Russian Empire), farther 
west than the Upper Amur. 

Game-birds were very scarce, though I observed Capercailli, 
Ptarmigan, and Quail, and in the highest and barest parts of 
the mountains the magnificent Tetraogallus altaicus was not 
uncommon, though very hard to approach. The only one which 
I got within shot of was a hen bird with a brood of young ones, 
and she fluttered along the ground before me just as a ptarmigan 
would do in similar circumstances. 

I also saw a single pair of Perdix barbata, Pallas, with newly 
hatched young, on July 18in a marshy larch wood at about 6000 
feet elevation—a most unlikely place, as I should have thought, 
in which to find such a bird. I do not think that the Eastern 
Capercaillie (7. wrogalloides) is found in the Altai. The Caper- 
eaillie I saw were apparently the same as those of Europe, 
though in the Southern Ural there is a well-marked variety with 
a white breast, which may be specifically distinct. 

No ornithologist, so far as | know, has yet worked out the birds 
of the Altai, and there are few other regions in Europe or Asia of 
which it can now be said that they are unknown to the members 
of the British Ornithologists’ Union. One fact, however, may 
be mentioned, as it bears upon the question raised recently by 
Mr. Hartert as to the migration of the Siberian Nutcracker, 
which he considers to be a distinct variety from the European 
one. As we rowed down Lake Teletskoi on the 4th of August, 
we saw large flocks of Nutcrackers which were evidently 
migrating, and though their migration may not have extended 
beyond the Altai Mountains, yet from the great abundance of 
edible-seed of Pinus cembra, which were just becoming ripe, I 
could not see any reason to account for this. In Europe the 
Nutcracker is a solitary and not a migratory bird, and yet the 
regular occurrence of Asiatic Nutcrackers in Eastern Europe 


ZOOLOGY AND BOTANY OF THE ALTAI MOUNTAINS. 39 


shows that the Siberian one must be at any rate partially migra- 
tory. Those who wish to study the birds of this country should go 
early and stay late in the year, fur the height of summer is nut 
the time for collecting. I have no doubt, however, that one wio 
would vive up his whole time to it might get very valuable results ; 
and J am certain, that even the plants, which Lave hitherto received 
a greater share of attention than any other branch of Natural 
History, are by no means worked out. 

Next to the Wild Sheep, to whose pursuit I devoted ten days 
of my stay in the mountains, I gave most time to the Butterflies, 
which, though comparatively scarce until about the 5th of July, 
then began to appear in great profusion. I was particularly 
pleased to discover here a number of species hitherto known 
only from the Upper Amur and from Lapland, among them 
some most interesting species of Geis, Hrebia, Argynnis, and 
Lycena. 1 found, however, hardly any of the peculiar types 
which occur in the various mountai.-ranges of Turkestan 
southwards ; and it seems, from a general review of the 
Butterflies of the Altai, that the European element is dominant, 
with a large admixture of species belonging to Hasteru and 
Northern Siberia. I must say, however, that the majority 
of these are confined tv the high bare mountains above 
timber-line. The collections made by Kindermann in 1851-52 
(which have been described by Lederer) and by Ruckbeil in 
the south-western parts of the Altai (which are in Herr Tancré’s 
collection) are of a much more typically European character, and 
do not coutain a large number of the more interesting species 
which I obtained. 

As I have already given some account of my collection of 
Lepidoptera from the Altai at a meeting of the Entomological 
Society, aud propose to publish a full account of them in the 
‘Transactions’ of that Society, I need only remark that the 
number of Butterflies known from the Altai district amounts to 
about 180, of which I took about 140 myself in two months. 
Of these about 40 are not found in Europe. 109 are also found 
in the Amur region. 92 were also taken in the Kentei mountains 
of East Mongolia, which is the only place in that country of 
whose butterflies we have a fair list. 78 were found by 
M. Alpheraky in the mountains of Kuldja, which in point of 
distance are much nearer than the Kentei mountains; and out 
of above 200 species included by Grum-Grishimailo im his list of 


40 MR. H. J. ELWES ON THE 


Butterflies from the Pamir region, only 51, or about one-quarter 
of the species, are also found in the Altai. As compared with any 
area of similar extent in Europe or Asia, this number of species 
is surprisingly large; and as Kindermann in two seasons only 
took about 90 species, and Ruckbeil about the same, as against 
the 140 which I myself collected, it shows that the South- 
western Altai, where they both worked, is not nearly so rich 
and has not nearly so much of an Oriental character as the South- 
eastern Altai, where most of my collection was made. 

In the 20 years during which I have collected butterflies, I have 
never got nearly so many species in so short a time, and must 
attribute my success to the fact that when travelling on horseback 
I always had a net in my band and never passed a likely spot 
without giving it a trial. 

As for Moths, I was not able to do much myself. First, be- 
cause at the high elevatiou at which most of our time was spent 
there were not many night-flying species, or at all events very 
few came to our lights. Secondly, because I was generally too 
tired to sit up at night to collect. But having received since I 
returned home the collection made by M. Beresowsky at 
Ongodai, and having gone through the list of Moths made by 
Kindermann, I do not think they are as numerous in proportion 
to the Butterflies as they are in the Alps or Himalayas. 

Of the plants I cannot say much, because, although they 
have been pretty fairly worked out by Ledebour, yet a very 
intimate knowledge of the Flora of Turkestan and Eastern 
Siberia, as well as that of Europe, would be necessary to erable 
one to say how the distribution of plants coincides with that of 
birds and butterflies. 

The Fauna and Flora are also materially influenced by the 
very peculiar climate of the Altai, which has great extremes of 
heat and cold, and is subject to heavy thunderstorms, which fall 
as snow and hail in the higher regions, almost daily throughout 
the summer. During the whole of the two months we were in 
the mountains, we only had seven or eight days quite free from 
rain or snow. ‘These heavy storms seem mostly to come from 
the eastward, and from the high mountains, at the source of 
the Kemehik river, which is the westernmost tributary of the 
Yenesei. To show what sort of climate it is, 1 may mention 
that there were large beds of unmelted snow close to our camp, 
at about 7000 feet, all through July. On almost every clear 


ZOOLOGY AND BOTANY OF THE ALTAT MOUNTAINS. 41 


night it froze, and on the 17th of July snow fell to the depth of 
8 or 10 inches at this elevation, and though the hot sun and dry 
atmosphere very rapidly melted it, we were assured by the in- 
habitants that the summer was too short to make it worth their 
while to graze sheep there, and that as early as the middle of 
August snow might be expected to fall and lie in the higher 
mountains. Tehikatcheff, in his journey from the Tchuja Steppe 
to the Abakan, the principal northern tributary of the Yenesei, 
also met with heavy snowstorms in June and July. 

When we left England we had the intention of passing through 
North-western Mongolia and returning either by the upper valley 
of the Yenesei river, which is almost entirely unexplored, or, if 
we found that impossible, of going eastwards by the shores of 
the great Lake Kossogol to Irkutsk. But as we found no people 
who could act as guides in either of these directions, we were 
obliged to give up anything like real exploraticn. ‘There is, how- 
ever, no difficulty in travelling from Kuch Agatch through Mon- 
golia, vid Kobdo, Uliassutai, and Lake Kossogol to Irkutsk. 
Mr. Rew informed me that the Wild Horse (Hquus Prejvalskiz) 
was to be found in the desert between Kobdo and Uliassutai, 
and I was in great hopes of procuring a specimen of this animal 
which, so far as 1 know, has not been seen by any Englishman. 
M. Grum-Grishimailo, the celebrated Russian traveller, who has 
spent eight years in exploring Central Asia, and who is the only 
European who has actually seen the Wild Horse in its own 
country, assured me that the animals spoken of by Mr. Rew 
must be the Wud Ass, as the nearest point to the Altai at which 
be found the Wild Horse was about 15 days’ hard travelling 
south from Kobdo, near Guchen, aud as that country is almost 
inaccessible in summer on account of the want of water, time 
would not allow us to visit it. 

Though the geographical results of this journey are therefore 
unimportant, the extra time given to collecting in the Altai 
was of the greatest possible advantage, aud though a great deal 
may be done whilst travelling on horseback, if you stop every 
time you see a new or apparently new species, yet fast travelling 
is incompatible with collecting. 

Our return journey from the Mongolian frontier took us 
through a very different and most interesting part of the Altai. 
Leaving the Tchuja valley by the Kurai pass, we crossed the 
mountains to the north of it into a valley through which flows a 


42 MR. H. J. ELWES ON THE 


tributary of the Bashkaus river, which, after its junction with 
the Tchulishman river, flows into the great Lake Teletskoi, a 
deep mountain-lake about 60 miles long, out of which flows the 
Bija river, the principal eastern source of the Obb. 

Before quitting the Tchuja Steppe, however, I must make 
some remarks upon the character of the country we saw from 
the tops of the mountains at the sources cf the Obb. These 
mountains are above 6500 feet, absolutely bare and treeless, 
though three or four species of willows are found along the 
streams up to about 7500 feet. They consist of steep, shaly 
mountains surrounded by great rolling downs of grass. From 
the tops of these mountains, at about 9000 feet, we could see the 
sourees of the Irtysch aud the Kemchik, which latter flows into 
the Yenesei, and of the Kobdo river, which loses itself in the 
Mongolian desert. Eighty or ninety miles to the southward we 
could see the high snow-peaks of the Southern or Mongolian Altai 
range, which have, according to the accounts of Russian travellers, 
dense forests on their sheltered slopes. I am informed that the 
Beaver occurs there, as it certainly does in the Sayansk moun- 
tains near the source of the Yenesel. 

To give an idea of the Alpine flora of the South-eastern Altai, 
T may mention a few of the plants which were most conspicuous 
for their beauty near our camp on the Darkoti, or Tachety river 
as Tehikatchetf spells it, 80 miles south-west of Kuch Agatch, 
at about 7000 feet. I have never, either in the Alps of Europe, 
in the Sikkim Himalaya, in Colorado, Califvrnia, or anywhere 
else, seen such a perfect natural garden of beaut ful alpime 
flowers as I saw here in the middle of July. Among the most 
couspicuous were the lovely Primula nivalis, Pall., which 
strongly resembles P. Parryi of Colorado ; Dracocephalum gran- 
diflorum, which grew in sheets of cerulean blue; Polemonium 
pulchellum ; Gentiana altaica; Pedicularis verticillata, P. 
foliosa, P. comosa; Allium sibiricum, or senescens, the most 
ornamental of its genus; Linum ceruleum ; Iris tigridia, Bunge ; 
Pyrethrum pulchellum ; a lovely blue Corydalis growing in wet 
places, which Mr. Baker cannot name, and which may be new ; 
a beautiful Agquélegia, named A. glandulosa at Kew, but much 
finer than that plant as we know it im our gardens; several 
pretty species of Astragalus, Lloydia serotina ; and many well- 
known Arctic and high Alpine plants, such as Papaver alpinum, 
Draba ochroleuca, and Saxifraga oppositifolia, which were found 


ZOOLOGY AND BOTANY OF THE ALTAI MOUNTAINS. 43 


as high up as 8500 feet, where the flora and scenery reminded one 
strongly of the high fjeld of Norway, and Dryas octopetala, which 
covered the curious dry gravelly ridges on the hill-sides in many 
places. 

Flying over these marshy alpine flower-gardens were some of 
the rarest and most beautiful butterflies of Siberia, the European 
Alps, and Lapland, many of then also found on the Alps of 
Colorado, such as Parnassius Hversmanni and P. delius, Erebia 
lappona, E. ero, E. tyndarus, aud EH. maurisius, Gneis bore and 
G. sculda, Argynnis Kefersteini, A. freya, and A. frigga, Colias 
mongola, Lycena orbitulus and L. pheretes. Whilst on the higher 
and more rugged mountain-tops were herds of the great Wild 
Sheep and Ibex, Marmots, and Alpine Hares. 

The Kirghiz, who were pasturing large herds of mares in this 
neighbourhood, on whose milk converted into kumiss they almost 
entirely live, had taken from the nest to train for faleonry two 
young falcons which I believe to be & sacer; but a rare species 
or variety known as Falco altaicus, allied to the Peregrine, also 
occurs in the country. 

Lhe only vegetables which we had during our stay at these 
altitudes were rhubarb (A. Rhaponticum) aud wild onions; but 
the Tartars were also fond of the young stem of a species of 
Heracleum, which was too strong for my taste. 

The larch, Larix sibirica, ascended here to a little over 7000 
feet. I saw young trees at this elevation about 14 diameter and 
7 feet high, which showed 25 years’ growth ; and a very remark- 
able stunted old larch, 3 feet in diameter and not more than 20 
feet high, which must have been many hundreds of years old. 
Away from these there was no fuel but willows and dry horse 
dung, the common fuel of Mongolia and Tibet. 

The change in the scenery and character of vegetation, fauna, 
and insects was most marked on crossing the watershed between 
the Tchuja and Bashkaus valleys. It seemed as though in one 
day we had passed trom Asia into Kurope, for a number of 
plants, such as Linnea borealis, Saxifraga umbrosa, and various 
Ericacee, familiar to me in the Alps, which I had not previously 
seen, were found there ; whilst ferns, which are conspicuous by 
their absence in the dry Tchuja valley, had also become abundant. 
Instead of thin larch forests, about which you could everywhere 
ride on horseback, we found dene forests of Pinus sylvestris, 


44 MR. H. J. ELWES ON THE 


Pinus Cembra, and spruce forests which exceed in their impassa- 
bility anything I have ever seen, even in British Columbia. 

When we reached the north end of Lake Teletskoi, we found the 
grass and herbaceous vegetation very high and rank, forming a 
most marked contrast to the dry stunted grasses of the Tchuja 
Steppe and the hills around it. Aconites, delphiniums, thistles, 
wild hops, and many other plants grew 6 or & feet high; but 
though many species of moths were seen, butterflies were much 
less numerous and interesting, most of them being common 
Huropean species. 

As there is no track passable for horses along the western shores 
of Lake Teletskoi, we had to traverse it in a boat, and it took two 
days of hard rowing. The western shore of this lake is so steep 
and rocky that in many places you cannot land for several 
miles, and we had great difficulty in hauling our boat ashore 
during a sudden storm which sprung up and threatened to 
swamp us. Along the whole western shore of this great Jake 
there is not a single vestige of man’s presence, and the forest is 
so dense, rocky, and impassable, that I do not think it would be 
possible for a man on foot to make more than four or five miles 
a day in summer, though these forests are no doubt more easily 
traversed in winter on snow-shoes by fur-hunters. The shores 
of Lake Teletskoi were formerly a favourite resort of the Elk, 
Deer, and Bear, but they are now much diminished since firearms 
have become common. There is something in the climate of this 
region which, as Helmersen remarks, must be very exceptional, 
for we were assured by the inhabitants, as he was, that ice rarely 
remains on the lake for more than a month at a time, whereas 
the Obb river is frozen up during three or four months every 
winter. 

The inhabitants of the country round Lake Teletskoi, and 
probably of the Bija valley, are of an apparently different race 
to the Altai Tartars, and are called Teleutes; in physique they 
appear to be much poorer, and nothing like so healthy and 
vigorous. They cultivate a little spring rye and oats in the 
valleys, and are now mostly Christianised to a certain extent by 
Russian missionaries, whereas the Altai Tartars and Kirghiz 
are either mussul nans or worshippers of spirits. All these 
natives are very much addicted to drink, and the chief, or Saisan, 
through whom we procured horses and men for our different 


ZOOLOGY AND BOTANY OF THE ALTAI MOUNTAINS. 45 


journeys, though nominally a Christian, was not free from this 
failing. 

As to the remarkable absence of glaciers in the Altai, a few 
words seem desirable. At the sources of the Katuna, where 
the mountains are higher and steeper than they are in the South- 
eastern Altai, I believe glaciers are larger and more abundant ; 
but the only place where we saw what could really be called a 
glacier was in a high mountain south of the Kurai Pass. Their 
scarcity is probably accounted for by the extremely dry climate 
and light winter snowfall of the higher mountains. Notwith- 
standing the extreme cold, the snowfall in the Upper Tchuja 
valley is so light that horses can procure food on the mountains 
all through the hardest winter, whereas in the low country around 
Lake Teletskoi and the Bija valley they have to be fed on hay. 

When we reached the north end of Lake Teletskoi, we had 
four days’ hard riding down the Bija valley before we reached a 
country over which carts could travel. This valley is remarkable 
for its magnificent forests of pine, Pinus sylvestris, which exceed 
anything I have ever seen before. In some places I counted as 
many as 20 or 30 irees to the acre, of an average girth near the 
ground of from 6 to 10 feet, and carrying their girth higher up 
than I have ever previously seen, so that at a height of 80 or 
100 feet the tree would still be over a foot in diameter. The 
Russians, however, who, as described by a well-known German 
forester, are ‘‘everywhere and at all times true wasters and 
destroyers of forests,’ are making rapid inroads upon this 
magnificent timber, which is felled and floated down the Bija and 
Obb for supplying the towns and villages in the Steppe country 
northwards. Fire also is rapidly wasting many of the hill-sides, 
and when the pine has been burnt off, its place is usually taken 
by poplar and birch. 

The climate of this Bija valley is evidently very much damper 
than that of the country to the soutiwards, and we had the 
greatest difficulty in getting our horses through some of the 
marshy forests. The horses in the Altai are, however, capable 
of going where even ponies in the Rocky Mountains could 
hardly scramble, and where the road is too steep and slippery 
to get a foothold, will clamber up through the thick brush- 
wood with dense undergrowth and herbaceous plants higher than 
their heads on an incline of at least 30 degrees. To give an 
idea of their endurance, I may state that one day we rode the 


46 DR. W. G. RIDEWOOD ON THE 


same horses from 9 a.m. till 9.30 p.m., without halting more than 
a few minutes, none of them being apparently the worse for it, 
and a sucking foal following its mother during the whole journey. 
The horse which I rode accomplished the last hour of the journey 
at a canter in the dark. 

Tn conclusion 1 would strongly urze upon any one who may 
think of visiting the country, that a knowledge of the Russian 
language is almost indispensable; and considering the large 
amount of valuable material which is practically buried in 
Russian scientific journals, 1 am surprised that so few English 
naturalists have hitherto thought it worth their while to do 
what many young Army Officers, for military purposes, now do 
every year. 


Some Observations on the Caudal Diplospondyly of Sharks. 
By W. G. Rrpewoop, D.Sc., F.L.S., Lecturer on Biology 
at St. Mary’s Hospital Medical School, London. 


{Read 19th January, 1899. ] 


Tr is a well-known fact in Ichthyology that in Selachian fishes 
the vertebrae of the tail are twice as numerous as the caudal 
segments, delimited by the spinal nerves and the intermuscular 
septa. 

The first clear reference to this remarkable phenomenon occurs, 
curiously enough, in Gétte’s memoir on the development of the 
Fire-bellied Toad (Bombinator), (6. p. 418, footnote). It has 
since been remarked upon by von Ihering (9. pp. 220-236), 
Hasse (7. p. 21), Balfour (1. p. 455), Schmidt (15. p. 756), Mayer 
(13. pp. 262-267), Gadow (4. pp. 194-196), and others. 

To each myomere and neuromere there occur two centra, two 
pairs of crural plates, two pairs of intercrural plates, and four 
neural spines. ‘The two centra are similar*, as also are the 
hemal arches and the neural spines, but the crural plates are 
alternately imperforate and perforated by the ventral nerve-root, 
‘while a similar relation exists between the intercrural plates and 


* Except in Galeus, in which they are alternately slightly longer and 
shorter. 


CAUDAL DIPLOSPONDYLY OF SHARKS. 47 


the dorsal nerve-roots. In some forms, such as Scylliwm, the 
nerve-roots pass out, not through the plates, but between them ; 
yet the alternation exists the same (see fig. 1, p. 50). 

In Hasse’s monograph on the vertebrae ot Elasmobranchs, to 
which one naturally turns for information on such a point, tie 
figures (7. pl. 34. fig. 14, Scyllium catulus, and fig. 22, Scyllium 
canicula) give an incorrect idea of the actual appearance of the 
vertebree, for the differences between the calcified and unealcified 
parts are exaggerated, while the margins of the cartilage plates, 
which are of far more morphological importance, are not shown 
at all. And, further, although Hasse was fully aware of the 
occurrence of diplospondyly in the tails of these animals (7. p. 21), 
he has indicated in these figures nerve-apertures on consecutive 
vertebre. The error cannot be excused on the ground that the 
part of the vertebral column figured is anterior to that where 
diplospondyly obtains, for the presence of large hemal arches 
proves the contrary. The figures given by Mayer (18. pl. 18. 
fig. 12, and pl. 19. fig. 1, Scyllium stellare) are considerably clearer 
than those of Hasse, but even they leave much to be desired in 
the delineation of the boundary lines between the neural plates 
and spines. 

The portion of the vertebral column of Scyllium catulus 
(=stellare) depicted in fig. 1 is taken immediately below the 
second dorsal fin, and shows the diplospondyious condition in its 
most typical form. The hemal arches (/) and the crural plates 
(cr) are fused on the centra (c), but the intererural plates (7c), 
alternating with the crural plates, remain distinct. The hemal 
spines (is) are not separate from the hemal arches, but the 
neural spines (zs) are small cartilages which fit with great recu- 
larity over the intervals between the crural and intererural plates. 
The dorsal roots of the spinal nerves issue through the foramina 
(d) on the posterior margin of every alternate intercrural plate, 
while the foramina for the ventral roots (v) lie at the lower end 
of the posterior edge of the crural plates. The dorsal foramina 
of the right and left sides lie in the same transverse plane; 
and similarly with the ventral foramina. 

This condition of diplospondyly obta‘ns not only in the tail of 
Scyllium catulus, but, as I can testify from personal observation, 
also occurs in Mustelus vulgaris, Galeus canis, Carcharias lati- 
cauda, Scyllium canicula, Cestracion Philippi, Acanthias vulgaris, 
Spinax niger, and Scymnus lichia. Considerable confusion has 


48 DR. W. G. RIDEWOOD ON THE 


been introduced into the literature of the subject by von [hering’s 
statement (9. p. 229 & p. 233) that Scymnus departs from the 
condition found in Acanthias and Scyllium by having no double 
vertebre in the tail, or only one or two vertebre with imperforate 
crural and intercrural plates ; for Gadow (4. p. 195) has repeated 
the statement in his memoir on the vertebral column of fishes, 
without having observed that Mayer (138. p. 265) had corrected 
von lhering on this poimt. The figure given by Mayer (13. pl. 18. 
fig. 9) shows that normal diplospondyly commences at the forty- 
third vertebra in Scymnus lichia. In a specimen of this species, 
which by the kindness of Mr. G. A. Boulenger I was allowed to 
examine at the Natural History Museum, I found that nine 
myomeres of the tail were diplospondylous. The first of these 
segments was situated immediately behind the cloaca, and the 
hemal arches commenced at the same place. The relations were 
thus exactly those which occur in other Selachians. 

The passage from the diplospondylous condition in the tail to 
the monospendylous condition in the trunk is not abrupt, but 
gradual; and the four or five vertebre involved in the transition 
offer an irregularity which deserves more notice than has hitherto 
been accorded to it. Although three theories at least have been 
propounded to explain the origin of the caudal diplospondyly, it 
does not appear to have occurred to anyone to inquire minutely 
into the detailed structure of these transitional vertebre ; yet 
they hold, as it were, the key to the situation. The transiticn 
is invariably in the proximity of the cloaca, as von Ihering and 
Maver (9. p. 228, and 18. p. 261) have already pointed out. It 
occurs behiud the last rib-bearing vertebra, and in front of the 
vertebra carrying the third or fourth complete hemal arch (see 
fiy. 2). As arule it does not extend through more than four or 
five myomeres, but according to Mayer (138. p. 266) six body- 
segments are involved in Mustelus. 

The transitional vertebre do not standin any constant relation 
with the dorsal fins, for they are anterior to the first dorsal fin 
in Rhina (18. pl. 18. fig. 1), below it in Scylliwm, between the two 
dorsals in Acanthias, and below the second dorsal in Scymnus. 
Since, however, the dorsal fins are variable in position with respect 
to the cloaca in different genera, and the transitional vertebre 
are definitely related to the position of the cloaca, it is but a 
logical conclusion that the vertebral transition shall not be related 
to the position of the dorsal fins. Since caudal diplospendyly 


CAUDAL DIPLOSPONDYLY OF SHARKS. 49 


occurs in forms like Acanthias, Scymnus, and Rhina which are 
destitute of the anal fin, the transitional region of the vertebral 
column cannot bear any relation to this appendage. 

It is evidently to this transition region that Miiller is referring, 
when he writes (14. p. 156):—“ Bei Zygena fand ich noch das 
merkwiirdige, dass an einigen Wirbeln des mittlern Theils der Wir- 
belsiule sogar 3 Bogenstiicke hinter einander auf einen Wirbel 
jederseits kommen, wiihrend die meisten Wirbel nur 2 Bogenpaare 
haben.” The regularity of the neural plates over the centra in 
the tail-region was such that the duplicity of vertebre passed 
without notice, but the irregularity in the transition region 
did not fail to catch the eye. Since Miiller did not notice the 
relations of the vertebra to the neuromeres and myomeres, the 
differences between the caudal and trunk vertebree escaped him- 
According to Mayer (18. p. 263), the much misunderstood state- 
ment of Kolliker’s with regard to Heptanchus (vide postea, 
p- 51 footnote) also refers to the few vertebre in the transition 
region. 

Gegenbaur (5. pl. 9. fig. 19) has figured a portion of the 
vertebral column of Cestracion, from the vertebra bearing the 
last rib to that with the fifth hemal arch. The figure, however, 
shows no nerve foramina nor introduction of new intercrural 
plates, but perfect regularity such as would occur in the trunk 
region. Although, therefore, this is obviously the region of 
transition, the irregularities which must have existed are not 
shown. 

The fullest information on the subject is that furnished by 
Mayer, whose illustrations (13. pls. 18 & 19) include the transi- 
tional vertebree of Scylliwm, Mustelus, Centrophorus, Heptanchus, 
Scymnus, Acanthias, and Rhina. He roughly describes (13. 
p- 266) the region in question in Scylliwm, but does not discuss 
the detailed relations of the neural arches. He merely states 
that the arches are irregular, and that the bodies may carry three 
_ pairs of arches. 

Tn the specimen of Scylliwm catulus depicted in fig. 2, the irre- 
gularities commence immediately behind the last rib-bearing 
vertebra. The vertebra marked 2 has the crural plate (er) ex- 
ceptionally broad, and an additional neural spine (ms) is super- 
posed upon it. This in itself is an exceptional occurrence, for 
the neural spines normally lie over the boundary-lines between 
the neural plates, and not directly over a plate. The third 

LINN. JOURN.—ZOOLOGY, VOL. XXVII. 4. 


50 DR. W. G. RIDEWOOD ON THE 


vertebra of the series differs from the second only in that its 
crural plate is still broader, and that the intercrural plate behind, 
which should lie over the hinder part of the centrum, has been 
pushed entirely off, and the third neural spine of this vertebra 
(ns') has suffered a similar backward displacement. The centra 
4 and 5 are the first to show the doubling. They are shorter 
than the preceding three, but are longer than the half of each of 
these. The second of the two centra (5) has over it a crural 
plate and the anterior half of the following intercrural, and 


Le ' ms. Gl. gp ie 


Ce 


Z V vr ul 
mi 


Fig. 1.—Sceyllium catulus ; caudal vertebree, left side, natural size. 
Fig. 2.—Sczlliwm catulus; transitional vertebrae, in the region of the cloaca, 
immediately below the first dorsal fin ; left side, natural size. 
ce, centrum; er, crural plate; zc, intercrural plate; ms, 2s', neural 
spines; #, hemal arch; hs, hemal spine; 7, last rib; d, foramen for 
dorsal root of spinal nerve ; v, foramen for ventral root. 
The dotted lines indicate the hypothetical limits of the perfect vertebra. 


above these are a half and a whole neural spine. The anterior 
centrum (4) has belonging to it a crural plate and the posterior 
half of the displaced intercrural in front, and a whole and a half 
neural spine. That only one half of the intercrural belongs to 
this vertebra is clear from the fact, that if in figure 2 the line be 
erased which separates the centra 4 and 5 and their corresponding 
crural plates, the now single vertebra will be a facsimile of those 
marked 2 and 3, except for the fact that there are two hemal 
processes instead of one, 


CAUDAL DIPLOSPONDYLY OF SHARKS. 51 


The next double vertebra (6° 7) is a repetition of (4°5) except 
that the three neural spines occupy the full length of the two 
centra, and that the hemal arch on centrum 7 is completed by a 
hemal spine (hs). The double vertebra (8°9) shows a further 
departure. Both hemal arches are complete; and between the 
two narrow crural plates an intercrural, unnotched on its anterior 
and posterior borders by nerve foramina, has been introduced. 
The middle of the three neural spines is thus again exceptionally 
placed, since it les immediately over a plate, and not over a 
boundary line as it did in (4°5) and (6°7). The double vertebra 
(10-11) is the first of the normal diplospondylous series, and 
differs from (8°9) only in the greater breadth of the crural 
plates. A feature of special interest in vertebra (8°9) is that 
the intercalated intercrural has only appeared on the left side. 
As seen from the right side, this double vertebra is an exact 
counterpart of (6° 7). 

To summarize the above description :—The transition is effected 
by steps taken in the following order—the broadening of the 
crural plate and the introduction of an additional neural spine ; 
the division of the centrum and crural plate, and the doubling 
of the hemal process; the mtercalation of an additional inter- 
crural between the two contiguous crurals. The great advantage 
attained by this gradual transformation is obviously the avoidance 
of excessively large or excessively small cartilages, while yet 
securing a diminution, on the whole, of the antero-posterior 
length of the elements. 

The most recent view on the subject of caudal diplospondyly 
in Selachians is that expressed by Dr. Gadow, who attributes 
(4. p. 194) the phenomenon to the “ chorda centra’”’ being inde- 
pendent of the “arcualia” and to the difference between the 
metamerism of the centra and that of the arches. He explains 
that in the middle of the trunk region of Heptanchus the centra 
are double their normal length, extend through two myomeres, 
and have four pairs of “ dorsalia”’*. After stating that it is the 


* It is not clear from Dr. Gadow’s text whether he is here describing obser- 
vations of his own, or is merely enlarging upon the unfortunate sentence of 
Kolliker’s (11. p. 199) “bei Heptanchus im hintern und vordern Theil der 
Wirbelsaule die Zahl der Wirbel um das doppelte grosser ist als in der Mitte.” 
But in either case, it should not escape notice that Hasse failed to confirm 
Kolliker’s observation, and stated (7. p. 46) that this anatomist was probably 
misled by the deceptive appearance of “ein dunkler, doppelt contourirter 


52 DR. W. G. RIDEWOOD ON THE 


variable length of the chorda centra which causes the discre- 
pancies, he proceeds :—“ These apparent irregularities reach 
their climax in the tail of many Elasmobranchs, where exactly 
the reverse takes place to what occurs in the trunk, in this way, 
that the chorda centra are so numerous, or so short, that two of 
them fall to the share of one true segment. The number of 
dorsal cartilaginous pieces varies extremely.” Now, as a matter 
of fact, the neural plates are arranged with the greatest regu- 
larity, as will be seen by a glance at fig. 1; namely, one plate 
united with each centrum and one plate intercalated, the median 
dorsal cartilages being regularly disposed over the intervals. In 
fact Mayer (whose paper is quoted by Gadow on p. 196) had 
already written (13. p. 266), “ Nur die an der Ubergangsstelle 
befindlichen Wirbel zeigen allerlei Unregelmassigkeiten in Lage 
und Anzahl jener Stiicke.’”” The only shark in which I have been 
able to detect any want of correspondence in the tail region 
between the neural plates and the centra is Galews. Since, as a 
rule, the elements of the caudal vertebre are as regularly dis- 
posed as are those of the trunk vertebre, the ‘“ explanation ” 
cannot be considered valid. 

Dr. Gadow further observes (4. p. 195) that the “‘ intercalation 
or wedging-in of these various cartilages can be followed from 
the simplest to the most complicated conditions in the Rajide.” 
It is to be regretted that he does not give illustrations of these, 
for he acknowledges that Hasse’s figures do not explain the facts. 
But it must here be pointed out that the Rays are less primitive 
than the Sharks; and that the fact of the phenomenon being 


Streifen, welcher der Mitte der Basen der Neur- und Haemapophysen entspricht 
und senkrecht verlaufend die tiberraschendste Aehnlichkeit mit einem Zwischen- 
wirbelgewebe hat.” And one year previously to this von thering (9. p. 222) 
had suggested that Kolliker’s error arose from his counting the neural spines 
as arches. The figure, moreover, which Mayer has given of Heptanchus (13. 
pl. 18. fig. 10) shows the usual Selachian diplospondyly, commencing on the 
fifty-ninth vertebra. The deceptive bands on the centra mentioned by Hasse 
are clearly shown, but there is nothing abnormal in the regularity of the neural 
arches. In the accompanying text (13. p. 263) Mayer shows that the statement 
of Kolliker’s is correct if read as referring only to the few transitional vertebree 
_ between the trunk and tail regions. But none of these explanations can apply 
to Dr. Gadow’s assertion (4. p. 194) that “each long centrum actually belongs to 
two true segments” —a statement which cannot refer to the alternation of the 
intercentral plates with those of the intermuscular septa, for that is normal in 
Sharks, and would not be worth mentioning. 


CAUDAL DIPLOSEONDYLY OF SHARKS. 53 
inexplicable without having recourse to the former is a sign of 
weakness in the argument. And besides, Dr. Gadow is here 
disregarding his own word of caution expressed on page 198 of 
his treatise, “Indeed, mischief enough has been done by the 
selection of the Rajide for the elucidation of fundamental 
morphological questions.” 

Much may be said in favour of the contention of Mayer and 
Gadow (13. p. 266, and 4. p. 195) to restrict the word “ vertebra C 
or “spondylus” to a complete scleromere, equal in value and 
antero-posterior extent to a neuromere and myomere ; although 
to agree in this is not necessarily to accept the conclusion of the 
latter writer that “diplospondylous” is a “term without any 
reasonable meaning.” For, after all, these “ vertebre”’ of the tail 
of Sharks are so regular and complete that we can scarcely deny 
them the title. Each consists of a centrum of cartilage, partially 
ealcified, with a conical depression in front and another behind, 
occupied by persistent notochord. Above each centrum, and united 
with it, is a pair of cartilaginous plates, and between every two 
consecutive “ vertebra ’’ a pair of intercalary plates, while located 
over the intervals between these alternating plates are median 
dorsal cartilages, twice as numerous as the centra. That is to 
say, the structure of each “ vertebra” of the tail is exactly the 
same (neglecting, of course, the distinctive features such as 
hemal arches and absence of ribs) as that of a trunk vertebra, 
except that every alternate one has no nerve foramina, while all 
the trunk vertebre are provided with them. 

If, therefore, we deny the caudal “vertebre” the right to 
rank as equal to those of the trunk region because of their failure 
to fall in with the metamerism of the muscular and nervous 
systems, we must yet admit for them a metamerism of their own, 
which is almost as perfect as that of the trunk vertebra. It is 
certain that there is no such “ discrepancy between chorda centra 
and arches,” or ‘‘ difference between the metamerism of the 
centra and that of the arches,” as Dr. Gadow would have. The 
only discrepancy occurs in the transition region, where it could 
~ hardly be avoided ; and even there it only affects four or five 
segments of the body. 

Von Ihering conceived the idea (9. p. 235) that in the primi- 
tive Selachians the whole vertebral column was diplospondylous, 
and that the monospondylous condition is secondary, and has 
been introduced by a fusion of parts proceeding regularly from 


54 DR. W. G. RIDEWOOD ON THE 


before backwards. According io this view, therefore, the fewer 
double vertebre in the tail in existing forms of Selachians, the 
more specialized the fish. But Mayer, in disposing of the case 
of Scymnus, upon which von Ihering placed so much reliance, 
has shown (18. p. 265) that the idea is altogether untenable. 
He holds that the diplospondylous condition is secondary, and is 
due to the halving of the normal vertebre. Hasse (7. p. 21), 
although lending active support to von Ihering’s theory, at the 
same time regards every alternate “vertebra” of the tail as 
intercalated, and therefore not homologous with the others. 

Embryology unfortunately throws no light upon the subject. 
Both Gotte and von Ihering (6. p. 418, and 9. pp. 222 & 235) 
were agreed that the ontogenetic segmentation of the caudal 
vertebree was precluded by the disposition of the crural and 
intercrural plates; and Balfour (1. p. 455) satisfied himself by 
actual examination of embryos that the duplicity of the caudal 
vertebre was not due to secondary segmentation, but was 
observable so soon as the vertebral column showed any signs of 
differentiation into vertebre. ‘The figure given by Klaatsch 
(10. p. 172, fig. 8) of the longitudinal section of the embryonic 
vertebral column of Mustelus fully confirms the conclusions of 
this embryologist. 

Baifour was inclined to explain the want of correspondence 
between the metamerism of the caudal vertebree and that of the 
nerves and muscles by the fact that the former are differentiated 
later than the latter. Since, however, he also showed (1. p. 458) 
that the segmentation of the continuous cartilaginous sheath of 
the notochord was determined by the muscle segments, and gave 
good reasons for the fact, the lateness of the differentiation of 
the vertebral segments cannot be taken as an eaplanation of 
diplospondyly. It merely leaves open the possibility of other 
influences coming into play and over-ruling the dominating 
metamerism of the muscles and nerves. 

Caudal diplospondyly being so widely spread among existing 
Sharks, and the fact that there are no traces of the actual 
process of doubling during ontogenetic development, point to the 
conclusion that the condition is a very ancient ove. With the 
object of ascertaining whether paleontology could assist in the 
solution of the problem, I availed myself of the kind assistance 
of Mr. A. 8S. Woodward in examining the specimens of fossil 
Sharks at the Natural History Museum. The results were 


CAUDAL DIPLOSPONDYILY OF SHARKS. 55 


disappointing; for, in the absence of the muscles, the only 
means of deciding the principal metamerism of the tail is by the 
nerve-foramina, and these could not be made out in any single 
instance. Yet, judging from the centra of the tail. being 
markedly shorter, in proportion to their height, than those in 
the trunk region, it is by no means improbable that the diplo- 
spondylous condition of the tail is of considerable geological 
antiquity. 

Embryology and paleontology both failing us im our efforts 
to divine the cause and origin of diplospondyly in Sharks, we are 
constrained to fall back upon the evidence afforded by the tran- 
sitional vertebre, and upon another important fact, that diplo- 
spondyly does not extend to the extreme posterior end of the 
vertebral column. ‘The only reference to this fact that I have 
been able to discover in the scattered literature of the subject is 
the remark by Mayer (18. p. 267), “Somit entspricht an der 
Schwanzspitze wenigstens jedem Myotom ein Sclerotom.” In 
the hinder three-fourths of the caudal fin of Acanthias the 
myomeres are as numerous as the centra. The change from the 
diplospondylous to the monospondylous condition occurs at 
about the twenty-fourth centrum from the end; but the rela- 
tions between the vertebre and the muscle-segments can only be 
made out for the anterior half of these; for in the hinder part 
there is scarcely any muscle at all between the skin and tke 
vertebre. The last ten or twelve vertebre are imperforate, as 
already shown to be the case in Seylliwm by von Thering and 
Mayer (9. p. 228, and 13. p. 269), and the little muscular tissue 
that is attached to these vertebre is innervated by a backward 
extension of the nerves supplying the preceding myomeres. 

Diplospondyly is thus confined to that part of the body lying 
between the cloaca and the greater part of the caudal fin; and 
the questions that most naturally present themselves are these— 
- What advantage does diplospondyly confer on this part of the 
body, and in what respect would this part suffer if the mono- 
spondylous condition prevailed? The answer, it seems to me, 
can be given in a single word—Flexibility. Those who have 
watched dogfish swimming in an aquarium will know how im- 
portant is the “tail,’? or post-cloacal region of the body, as an 
organ of locomotion, the paired fins playing but a small part in 
the actual progression of the body through the water. Yet, 
when the tail is lashed from side to side, the caudal fin at its 


56 DR. W. G. RIDEWOOD ON THE 


extremity is seen not to partake of the general lateral curvature, 
or only ina minor degree. The caudal fin is chiefly concerned 
with obtaining a purchase upon the water, so as to constitute a 
relatively fixed point, about which the rest of the body can be 
moved by appropriate muscular contraction. Flexibility is thus 
not required in the caudal fin itself, but is of great advantage in 
the part of the body immediately preceding. The need for this 
extreme flexibility ceases in front of the cloaca, for here the body 
is largely occupied by the alimentary and other viscera, and 
constitutes, with the head, the most important part of the body, 
compared with which the post-cloacal part is merely a sub- 
servient appendage. And, further, since the body is thicker in 
the trunk region, the proportion existing between the leneth and 
the breadth of a segment is much less than in the tail, and con- 
sequently one vertebra to each myomere gives the necessary 
amount of flexibility. 

There are not, in Sharks, synovial articulations between the 
centra as in Snakes, where the flexibility of the vertebral column 
is considerable; neither are there zygapophysial articulations 
between the neural arches as in most Vertebrates. The only 
movements possible are those due to the slight yielding power 
of the fibrous tissue around the margins of the centra, and 
between the various cartilages of the neural arches. To double 
the yielding power of this separating fibrous tissue would be to 
weaken the connection between the several vertebra, and to 
introduce the possibility of lateral displacement ; but by doubling 
the number of vertebre in any region, twice the amount of 
fibrous tissue is introduced, without the above-mentioned dis- 
advantage. 

This response by the skeletogenous tissue to the requirements 
of flexibility of the particular part of the body, is possibly 
referred to in the following sentence from Gadow (4. p. 192) :— 
“Tt is obvious that the chondrified chordal sheath is affected by 
the ‘ centra of motion,’ which establish themselves according to 
the way in which the fish ‘ wriggles’.”’ 

That the vertebree must be integral multiples of the segments 
of the body is evident from the relations which exist between 
the muscles and the vertebrae. Although a secondary feature 
(Balfour, 1. p. 458, and Gadow, 4. p. 192), it is a fact, that in 
the development of Elasmobranch fishes the chondrified sheath 
of the notochord is uniform and unsegmented at a time when 


CAUDAL DIPLOSPONDYLY OF SHARKS. 57 


the metamerism of the muscular and nervous system is pertect. 
As a rule the segmentation of this sheath is determined by the 
myomeres, in the manner explained by Balfour (1. p. 453), so 
that the vertebre are as numerous as the myomeres; but there 
is nothing to prevent the vertebral segments being twice, or even 
three times, as many as the primary segments. To have frac- 
tional parts, however, is obviously impossible. Even allowing 
that the transition between the trunk and tail regions is beauti- 
fully gradual, yet, as will be seen by a glance at figure 2, the 
last monospondylous skeletal segment (3) is followed imme- 
diately by one with two centra and two crural plates (4° 5). 

The most logical conclusions, therefore, from the facts at 
command, are, that the condition of diplospondyly in the tail of 
Sharks is secondary, but of ancient date; and, further, that it is 
purely adaptive, being calculated to maintain a due proportion 
between length of centrum and width of body, without dimin- 
ishing the length of the muscle-segments. In the region of the 
body from the cloaca to the caudal fin, the demand for increased 
flexibility is prepotent over the normal tendency of the chon- 
drified chordal sheath to segment in such a way that the centra 
are as numerous as the myotomes. 

This, of course, is not an explanation of diplospondyly, it is 
merely a suggestion for its raison d’étre. That the diminution 
in the length of the tail which would be entailed by a shortening 
of the myomeres, and consequent restoration by this means of 
the balance between the length of the centra and the width of 
the body, would be a disadvantage, is also a pure assumption, 
Indeed, the study of Teleostean fishes shows that a shortening of 
the tail by the abolition of the terminal vertebre may, and does 
occur; and this without any compensating increase in the 
skeletal parts, for the caudal segments of Teleosteans are mono- 
spondylous. But, in this connection, Ama comes to our assist- 
ance; for here, in spite of a homocereal tail and presumably 
abbreviated vertebral column, the segments of the caudal axis 
occur two to each myomere and neuromere (9. p. 231). And, as 
in Selachians, the last segments of the body, namely those in the 
hinder part of the caudal fin, are monospondylous (Franque, 3 ; 
Kolliker, 12; Shufeldt, 16; Hay, 8; and others). 

Whether, therefore, we accept the view of Franque (38. p. 10) 
that in Amza those vertebral bodies of the tail which are destitute 
of neural and hemal arches have been intercalated between the 

LINN. JOURN.—ZOOLOGY, VOL. XXVII. 5 


58 DR. W. G. RIDEWOOD ON THE 


true vertebrex ; that of von Ihering (9. p. 235), that the condition 
is due to the secondary segmentation of vertebre originally 
simple; that of Schmidt (15. p. 755), that two consegmental 
vertebre occurred primitively throughout the body, and now 
persist only in the caudal region; or that of Baur (2. p. 942), 
and Hay (8. p. 5), that the pleurocentrum and hypocentrum 
together constitute a single vertebra equivalent to those of the 
trunk region, where the fusion of pleurocentra and hypocentra 
is assumed to have occurred,—the same general conclusion 
will apply as that above specified for Selachians, namely, that 
the universal tendency to develop single vertebral bodies is, in the 
region between the anus and the caudal fin, over-ruled by the 
demand for increased flexibility. Indeed, the conclusion might 
even be further extended to the Stegocephali, in which the 
embolomerous type of vertebra prevails in the caudal region 
only. 


List oF REFERENCES. — 


1. Batrour, F. M.— Comparative Embryology. Vol. ii. 
London, 1881. 

2. Baur, G.—<On the Morphology of Ribs.” Amer. Nat. 
vol. xxi., Philadelphia, 1887 ; pp. 942-945. 

3. Franquz, H.—Nonnulla ad Amiam calvam accuratius co- 
gnoscendam. Berlin, 1847; pp. 14, one plate. 

4. Gavow, H., and Assorr, E. C.—< On the Evolution of the 
Vertebral Column of Fishes.” Phil. Trans. Roy. Soc., 
vol. 186. 1895, B (1896); pp. 163-221, numerous figures 
in the text. 

5. GeGenBaur, C.—“ Ueber die Entwickelung der Wirbelsiule 
des Lepidosteus, mit vergl.-anat. Bemerkungen.” Jena- 
ische Zeitschrift, vol. iii., Leipzig, 1867; pp. 359-420, 
three plates. | 

6. Gorrr, A.—Die Entwickelungsgeschichte der Unke. Leipzig, 
1875. 

7. Hassz, C.—Das natiirliche System der Elasmobranchier 
auf Grundlage d. Baues u. d. Entw. ihrer Wirbelsiiule. 
Jena, 1879. 

8. Hay, O. P.—“ On the Structure and Development of the 
Vertebral Column of Ama.” Field Columbian Museum 
Publications, Zool. Series, vol. i., no. 1. Chicago, 1895 ; 
pp. 54, 3 plates. 


10. 


11. 


12. 


13. 


14. 


15. 


16. 


CAUDAL DIPLOSPONDYLY OF SHARKS. 59 


. Inertne, H. von.—Das peripherische Nervensystem der 


Wirbelthiere. Leipzig, 1878. 

Kiaatscou, H.—“ Beitriige zur vergl. Anat. der Wirbel- 
saule. II. Ueber die Bildung knorpeliger Wirbelkorper bei 
Fischen.” Morph. Jahrb., Leipzig, 1898; pp. 143-186, 
one plate and six figures in the text. 

Kourrcer, A.—‘‘ Ueber die Beziehungen der Chorda dorsalis 
zur Bildung der Wirbel der Selachier urd einiger andern 
Fische.” Verhandl. d. phys.-med. Gesell. in Wiirzburg, 
vol. x., 1860; pp. 193-242, two plates. 

Korircer, A.—Ueber das Ende der Wirbelsiule der Gan 
oiden und einiger Teleostier. Leipzig, 1860; pp. 28, four 
plates. 

Mayer, P.—“ Die unpaaren Flossen der Selachier.” Mit- 
theil. Zool. Stat. Neapel, vol. vi., Berlin, 1885 (1886) ;. 
pp- 217-285, five plates. 

Mttter, J.—“ Vergl. Anat. der Myxinoiden. I. Osteologie 
und Myologie.” Abhandl. d. konig. Akad. der Wiss. zu 
Berlin, 1834 (1836) ; pp. 65-340, nine plates. 

Scumipt, L.—‘ Unters. zur Kenntnis des Wirbelbaues von 
Amia calva.” GZeitschr. f. wiss. Zoologie, vol. liv. Leipzig, 
1892; pp. 748-764, 1 plate and 5 figures in text. 

Sauretpt, R. W.—“ The Osteology of Amia calva.” US. 
Fisheries Comm. Report for 1883. Washington, 1885; 
pp. 747-878, 14 plates. 


LINN. JOURN.—ZOOLOGY, VOL. XXVII. 6 


60 MR. THOMAS SCOTT ON THE 


Report on the Marine and Freshwater Crustacea from Franz- 
Josef Land, collected by Mr. William 8S. Bruce, of the 
Jackson-Harmsworth Expedition. By Tuomas Scorr, 
E.L.S., Naturalist to the Fishery Board for Scotland. 


[Read 15th December, 1898.] 
(Puates 3-9.) 


TuroueH the kindness of Mr. W. 8S. Bruce I have had the 
privilege of examining the Crustacea which he collected in Franz- 
Josef Land during his sojourn there in 1896 and 1897. 

It was with some hesitation that I undertook the examination 
of this interesting collection. Fortunately, however, a large 
number of the organisms contained in it were already more 
or less familiar to me, either as recent or fossil species, and 
therefore the examination, though arduous, was less so than it 
would otherwise have been. Prof. G. 8. Brady and the Rev. T. 
KR. R. Stebbing have kindly assisted me with the identification of 
certain doubtful species ; while my son, Mr. Andrew Scott, gave 
me valuable help with the examination of the Copepoda, and 
by the preparation of a number of drawings necessary for the 
elucidation of some apparently new forms, and for the confirma- 
tion of others which, though already described, have not before 
been recorded from the Arctic seas. 

It was necessary, in describing the results of my examination, 
to adopt some recognized method in classifying them, and the 
general arrangement which I have followed is that proposed by 
the Rev. T. R. R. Stebbing in his ‘ History of Crustacea,’ * and 
exhibited in the synoptical table at page 49. In this table the 
Crustacea are divided into four Subclasses, viz.: the Manaco- 
STRACA, ENTOMOSTRACA, GIGANTOSTRACA, and THYROSTRACA (or 
Cirripedia). The collection of Crustacea made by Mr. Bruce 
has been found to contain representatives of the first, second, 
and last of these Subclasses; and I now proceed to indicate 
briefly the number of the species that belong to each of these 
three subdivisions. 


(1) The MatacosTraca. 


The Malacostraca, which comprise what are otherwise called 
the higher Crustacea, are still further subdivided into the two 


* Tnternat. Sci. Ser. vol. Ixxiv. (London, 1893). 


CRUSTACEA OF FRANZ-JOSEF LAND. 61 


‘Orders—PoporutTHatma (or Stalk-eyed Crustacea) and EprioPH- 
THALMA (or Sessile-eyed Crustacea). The first was represented 
in Mr. Bruce’s collection by seven species, five of which belong to 
the smaller Macrura and two to the Schizopoda: there were no 
representatives of the larger Brachyura in the collection. But 
if the Stalk-eyed Crustacea are few in number, the Sessile- 
eyed forms are fairly numerous: they comprise Cumacea, of which 
there are five species; Isopoda, represented by twelve; and 
Atnphipoda, of which there are forty-six species. 


(2) The EntomostTraca. 

The Entomostraca are divided into three Orders, but the first 
of these, Braghiopoda, is not represented in the collection ; there 
are, however, numerous examples of Ostracoda and Copepoda. 
Thirty-four species of the former and sixty-six of the latter, 
with one new variety, have been obtained, and are recorded in 
the sequel. 


(3) The Tayrostraca or CIRRIPEDIA. 


The only Cirripedia found in the collection are two species of 
Barnacles, both of which are widely distributed in the northern 


seas. 


Perhaps the preceding statements may be more clearly under- 
stood if put into tabular form, thus:— 

Taste showing the general classification and number of the 

species of Crustacea contained in Mr. Bruce’s Collection of 
Crustacea from Franz-Josef Land. 


Number 
Subclass. Order. Suborder. of species. 
(Podophthalma (“Stalk- f Macrura............... 5 
eyed” Crustacea) ...... Schizopoda ......... 2 
MALACOSTRACA. 4 Edriophthalma (‘‘ Sessile- Cumacea ....;....0+. 5 
[ eyed” Crustacea) ...... | Hisojgeae cenegoogas00005 12 
Amphipoda ......... 46 
Podocopa ......-.++ 32 
(Ostracoda .......--.....-« e | Myodocopa ......... 1 
| Cladocopa........- 1 
TOM OSE ACE: 1 Gnathostomata...... 65 
@Gpepod aliens easeerrnas te | Poisons Ne ae 1 
Siphonostoma ...... 1 
‘CIRRIPEDIA. MWOLTACICA .2..2-eess-008 «.-. Operculata ......... 2 


62 MR. THOMAS SCOTT ON THE 


‘It will be observed from this table and from the preceding 
remarks that the collection of Crustacea brought home by 
Mr. Bruce, though it does not contain any of the larger Brachyura,. 
is particularly rich in the smaller forms ; indeed, though twelve 
suborders are represented in the collection, by far the largest 
number of species belong to only three of these. The total 
number of species is 173, but 132 of them belong to the 
Amphipoda, Podocopa, and Gnathostomata. It may be also: 
remarked that this collection of Crustacea exceeds in importance 
as well asin number of species any other previously brought 
from Franz-Josef Land, except perhaps in the number of the 
larger forms. 

Of the species above enumerated, three, represented each by a 
single specimen, are reserved for further investigation—one being 
a very small Macruran of the family Hypolytide, and the other 
two minute Cumacea, probably new. 

In the following detailed list the locality where each species: 
was obtained in Franz-Josef Land is given, except in the 
case of a few that are comparatively common. Notes on the 
distribution of species are occasionally added, and especially of 
those that have been observed in the British seas. Descriptions 
of several apparently new species will also be found in the sequel. 
A list of works that have been specially consulted in the pre- 
paration of this report is added. 

The majority of the specimens were collected in the vicinity 
of Cape Flora and Cape Gertrude, Northbrook Island; Elm- 
wood, which is also frequently mentioned, is situated near to 
Cape Flora. Northbrook Island is somewhat V-shaped: one 
branch extends in a north-by-west direction, about 15 miles from 
Barents Cape; the other branch, which is nearly 40 miles in 
length, extends almost due west; and Cape Gertrude, Elmwood,. 
and Cape Flora are ali situated at the western extremity of this 
lower branch. Northbrook Island is one of the most southerly 
of the Franz-Josef Land group, and while the lower branch 
which terminates in Cape Flora is somewhat under the 80th 
degree of North latitude, the other reaches to some distance: 
beyond that. 


CRUSTACEA OF FRANZ-JOSEF LAND. 63 


MALACOSTRACA. 


MACRURA. 
(Tribe Caridea.) 


Genus Sprronrocaris, Spence Bate, 1888. 


SPIRONTOCARIS GAIMARDII (Milne-EHdwards). (P1. 3. figs. 1, 2.) 

1837. Hippolyte Gavmardit, Milne-Edwards (26), p. 378. 

The Bruce collection contained a single adult female with ova; 
it was taken about two-thirds of a mile south-west of Elmwood, 
in 18 fathoms water, on June 7th, 1897. The length of this 
Specimen was about 62 millim.; the armature of the rostrum 
consisted of seven teeth on the upper, and five on the lower edge, 
as shown in the figure (fig. 1). 


(?) SprrontocaRis Parppsit (Kroyer). (Pl. 3. figs. 3, 4.) 

1841. HMippolyte Phipps, Kroyer (42), p. 575. 

A single specimen of this species was dredged in 8 fathoms, 
off West Bay, Cape Flora, on July 24th, 1897. Kroyer states 
- that in this species there are four teeth on the front part of the 
lower margin of the rostrum, but in this specimen there are seven 
teeth ; the first three (counting from the apex of the rostrum) are 
each somewhat stouter than the next four and are moderately 
wide apart, the fourth, fifth, and sixth are close together, 
while between the sixth and seventh there is an interval about 
equal to that between the third and fourth as shown by the 
drawing (fig. 3). But though the Cape Flora specimen differs 
from the typical S. phippsii in this respect, it seems to agree 
with it otherwise; variation in the number of the teeth of the 
rostrum amongst the group of crustaceans to which this one 
belongs is not uncommon. 


SPIRONTOCARIS POLARIS (Sabine). 

1824, Alpheus polaris, Sabine (65), p. 238, pl. 2. figs. 5-8. 

There were several specimens of this species in the Bruce 
collection, some of them being females with ova. Considerable 
variation was observed in the number of teeth on the upper and 
lower margins of the rostrum; on the upper margin the number 
varied from four to seven, and on the lower margin from one to 
three. All the specimens were taken in the vicinity of Elmwood 
at the western extremity of Northbrook Island, and mostly 


64 . MR. THOMAS SCOTT ON THE 


during June and July 1897. One was taken on floe-ice, but the 
others were dredged in depths ranging from 5 to 19 or 20) 
fathoms. The variation in the armature of the rostrum may b 
thus indicated : 


OLOL Os Os On mie 


the number within parentheses indicates the number of specimens, 
while the number above and the number below the line indicates 
the number of teeth on the upper and lower margins of the 
rostrum. One of the specimens in which the arrangement of the 


teeth of the rostrum is represented by es measured 65 millim. 
in length. 


Genus ScLEROCRANGON, Sars, 1882. 


SCLEROCRANGON BOREAS (Phipps). 

1774. Cancer boreas, Phipps (61 a), p. 190, t. 12. fig. 1. 

This was represented by one specimen, which was taken in ten 
fathoms, off West Glacier in Giinther Sound, on September 9th,. 
1896. 


SCHIZOPODA. 
Genus Tuysanorssa, Brandt, 1851. 


THYSANOESSA (?) NEGLECTA (Ardyer). 

1842. Thysanopoda neglecta, Kroyer (44), pl. 7. figs. 3 a-d. 

Three specimens of Thysanoessa were included in the collection 
from Franz-Josef Land: one was obtained in West Bay, Cape 
Flora, on September 9th, 1896, and two near Hast Glacier, Cape 
Flora, on August Ist, 1897. The specimens appeared to belong 
to 7. neglecta, but were scarcely perfect enough to enable the 
species to be satisfactorily determined. 


Genus Mysis, Latrezile, 1803. 


Mysis ocunata (Fabricius). 

1780. (°) Cancer oculatus, O. Fabricius (28 a), n. 222, p. 245. 

A considerable number of specimens of this Mysis were 
included n the Bruce collection. Various stages of develop- 
ment were represented, but the majority of the specimens were 
adult. They appear to have been all captured at the west end. 
of Northbrook Island, in the vicinity of Cape Flora. Several 


CRUSTACEA OF FRANZ-JOSEF LAND. 65 


were taken in September 1896, but the largest portion were 
collected between the 4th and 23rd of June, 1897; they were 
dredged in from two to three fathoms water off West Point, 
Cape Flora. A specimen of Dajus mysidis was observed on two 
of the adult Mysis. 


CUMACHA, 
Genus Diasryuis, Say, 1818. 


Drastytis Ratuxi (Kroyer). 

1841. Cuma Rathkit, Kroyer (42), pp. 513 & 531, t. v-vi. figs. 17-30. 

This species is represented by a single specimen—a female— 
taken near Hast Glacier, Cape Flora, on August 1st, 1897. 


Genus Lamproes, Sars, 1863. 


Lamprops Fuscata, G. O. Sars. 

1865. Lamprops fuscata, G. O. Sars (69), p. 192. 

L. fuscata was represented by a number of specimens, most of 
which were dredged in West Bay in from two to ten fathoms. 
This has been recorded froma few places in Greenland by Hansen 
and G. O. Sars. 


Genus Perarosarsta, Stebbing, 1893. 


PETALOSARSIA ? DECLIVIS (G. O. Sars). 

1865. Petalopus declivis, G. O. Sars (69). 

A single specimen that seems undoubtedly to belong to this 
species was obtained in some material dredged in ten fathoms 
about two-thirds of a mile south-west of Elmwood, Cape Flora, 
in January 1897. Petalosarsia declivis, though somewhat rare, 
appears to have a wide distribution; it has been recorded from 
Norway, as well as from the British seas. Petalosarsia has also 
been recorded from Spitzbergen. __ 


ISOPODA. 
The names and arrangement of the Isopoda are as far as 
possible in conformity with the second volume of Prof. G. O. 
Sars’s new work on the Crustacea of Norway. 


Genus Typutotanats, G. O. Sars, 1880. 


TYPHLOTANAIS FINMARCHICUS, G. O. Sars. (PI. 3. figs. 5-7). 
1880. Typhlotanas finmarchicus, G. O. Sars (72), p. 36. 


66 MR. THOMAS SCOTT ON THE 


Several specimens of this Isopod were included in the Bruce 
collection ; they occurred chiefly amongst dredged material from 
West Bay, Cape Flora; a few were also taken amongst sand 
near East Glacier, in the vicinity of Cape Flora. Prof. Sars 
discovered the species “‘ many years ago, rather plentifully in the 
Harbour of Vads6 at a depth of 30 fathoms.” One of the largest 
of the specimens in the Bruce collection measured about 2:3 
millimetres. Besides the other characters that distinguish this 
_ species, the meral joints of the last three pereiopoda are furntshed 
with one (or two) minute but distinct spines near the end of the 
inside margin (see fig. 6). 


Genus Leproanarata, G. O. Sars, 1880. 


LEPTOGNATHIA LONGIREMIS (Lilljeborq). 

1865. Tanais longiremis, Lilljeborg (49), p. 23. 

A specimen of this species was obtained in a gathering of 
Crustacea from the vicinity of Cape Flora. The dactylus of the 
chele was not so distinctly serrate on the superior aspect as 
Scottish examples. This Isopod “ occurs alony the whole Norwe- 
gian coast from Vadso to Christiania ;” I also have it from various 
parts of the Scottish coast. 


Genus Pseuporanats, G. O. Sars, 1880. 


PSEUDOTANAIS FORCIPATUS (Lilljeborg). (Pl. 3. figs. 8, 9.) 

1865. Tanais forcipatus, Lilljeborg (49), p. 16. 

This species was represented in the collection by a single speci- 
men dredged in Giinther Sound at a depth of ten fathoms on 
September 9th, 1896. The form of the chele (fig. 8) readily dis- 
tinguishes this from other species of Pseudotanais. P. forcipatus 
is also represented in the British fauna; it was moderately fre- 
quent in a gathering of small Crustacea collected in the Moray 
Firth in 1895. 


Genus Gnatuta, Leach, 1814. 


GNATHIA ELONGATA (Ardyer). 

1842. Anceus elongatus, Kroyer (44), pl. 30. figs. 3 ag. 

A single specimen of a female Gnathia, apparently belonging 
to this species, was obtained in some dredged waterial collected in 


CRUSTACEA OF FRANZ-JOSEF LAND. 67 


lat. 77° 55' N., long. 53° 20' E., on July 18th, 1897. The distri- 
bution of G. elongata seems to be chiefly arctic. 


Genus Janrtra, Leach, 1814. 


JANIRA TRICORNIS (A7rdyer). 

1842. Henopomus tricornis, Kroyer (44), pl. 30. figs. 2 a—g. 

A somewhat imperfect specimen of Janira tricornis was taken 
in the vicinity of Cape Mary Harmsworth in from 53 to 938 
fathoms on August 7th, 1897 (Cape Mary Harmsworth lies between 
30 and 40 miles further north and nearly 100 miles further west 
than Cape Flora). 


Genus Munna, Kroyer, 1839. 


Munna Fasrici, Aroyer. (Pl. 3. figs. 10, 11.) 

1842, Munna Fabric, Kroyer (44), pl. 31. figs. a-q. 

A single specimen was dredged in 30 fathoms off East Glacier, 
Cape Flora, on July 21st, 1897. (Hig. 10 is a drawing of the 
superior antenna.) 


Munna Krovert, Goodsir. (Pl. 3. figs. 12-14.) 

1842. Munna Kréyert, Goodsir (33), p. 365, pl. 6. fig. 2. 

A few specimens of a Munna, apparently belonging to this 
species, occurred in gatherings of small Crustacea dredged in the 
vicinity of Cape Flora—One specimen was taken with the dredge 
in 15 fathoms one mile off Flora Cottage, on September 10th, 1896 ; 
others were dredged at West Bay and off Cape Gertrude in July 
1897, in from 5 to 30 fathoms. 


Genus PLevrogonivum, G. O. Sars, 1871. 


PLEUROGONIUM INERME, G. O. Sars. 

1883. Pleurogonium inerme, G. O. Sars (73), p. 67, pl. 2. fig. 5. 

Two female specimens of this species (with ova), and three 
others, probably males of the same species, were dredged off Hast 
Glacier, Cape Flora, in 30 fathoms. The three (?) males were 
narrower in general outline than the female, but resembled them 
otherwise. 


PLEUROGONIUM SPINOSISSIMUM, G. O. Sars. (P1.3. fig. 15.) 
1865. Pleurocantha spinosissimum, G. O. Sars (71), p. 30. 
Two specimens, undoubtedly belonging to this species, were 


68 MR. THOMAS SCOTT ON THE 


taken at the same place and time as P. inerme. The drawing 
(fig. 15) represents the more perfect of the two specimens. The 
late Dr. Robertson of Cumbrae has recorded this species from the 
Firth of Clyde. 


Genus Munnopsts, WZ. Sars, 1860. 


Munwnopsts tyrica, If. Sars. 

1860. Munnopsis typica, M. Sars (67), p. 84. 

The Bruce collection contained several specimens of Munnopsis 
typica ; they were collected chiefly in the vicinity of Cape Flora— 
such as, a quarter of a mile west of the flagstaff, on July 2nd, 
1897 (this specimen was taken in the surface tow-net) ; off Hast 
Glacier in 30 fathoms, on the 21st of July and again on August 
the Ist; and off West Point in 2 to 3 fathoms in July. A 
specimen was also taken at about two-thirds of a mile south-west 
of Elmwood on April 29th, 1897, in 18 fathoms. Some of the 
specimens were considerably damaged, but a few were very 


complete. Heller also records Munnopsis typica from Franz- 
Josef Land. 


Genus Dasus, Ardyer, 1846. 


Dasus Mysiv1s, Ardyer. 

1846. Dajus mysidis, Kroyer (47), pl. 28. fig. 1 A-B. 

Two specimens of this parasite were observed; they were 
adhering to specimens of MMysis oculata, being attached to the 
underside of the thorax between the posterior swimming-feet. 


Genus Popascon, Giard § Bonnier, 1895. 


Popascon STEBBINGI, Giard & Bonnier. 

1895. Podascon Stebbing?, Giard & Bounier (29), p. 462. 

A few specimens of an Epicaride occurred amongst gatherings 
of small Crustacea dredged in West Bay, off West Point, and off 
Hast Glacier (all in the vicinity of Cape Flora). This Hpicardie 
agrees very closely with a form recorded by Rev. Mr. Stebbing 
from the Arctic seas, and which Professors Giard and Bonnier 
describe under the above name in their memoir on Epicarides 
published in the ‘ Bulletin Scientifique de la France et de la 
Belgique,’ tome xxv. (1895). 


CRUSTACEA OF FRANZ-JOSEF LAND. 69° 


AMPHIPODA. 


This suborder of the Edriophthalma was represented in the 
Bruce collection by a considerable number of species, and some 
of the species, such as Anonyx nugaw, Onesimus Edwardsii, and. 
Gammarus locusta, by many individual specimens. Prof. G. O. 
Sars’s recent work on the Amphipoda of Norway is generally 
followed in the arrangement and names of the species. 


HYPERIIDEA. 
Genus Hyrvrrtia, Latreille, 1825. 


Hiyperia GALBA (Montagu). 

1815. Cancer gammarus galba, Montagu (53 a), p. 4, pl. 2. fig. 2. 

A single specimen of Hyperia galba was taken in West Bay, 
Cape Flora, on July 5th, 1897. According to Prof. Sars, the 
distribution of this species extends from the Arctic seas to the 
coasts of Britain and France. 


Genus Pararuemisto, Boeck, 1870. 


PARATHEMISTO OBLIVIA (A7réyer). 

1838. Hyperia oblivia, Kréyer (41), p. 70, pl. 4. fig. 19. 

A number of specimens of this Amphipod were obtained on the 
sand near Hast Glacier, Cape Flora, on August Ist, 1897. (No 
specimen of Huthemisto libellula (Mandt), so frequent and widely 
distributed in the Arctic sea, was contained in this collection.) 


GAMMARIDEA. 
Genus OrcHOMENELLA, G. O. Sars, 1890. 
ORCHOMENELLA MINUTA (Kroyer). 
1846. Anonyx minutus, Kroyer (47), 2 R. 2 B, p. 23. 
A single example of this species was captured in about one to 


two fathoms at West Bay, near Cape Flora, on August 20th, 
1896, and another near the same place on June 6th, 1897. 


Genus Anonyx, Kroyer, 1838. 


ANONYX NUGAX (Phipps). 
1774. Cancer nugax, Phipps (614), p. 192, pl. 12. fig. 3. 
A considerable number of specimens of all ages and sizes were 


70 MR. THOMAS SCOTT ON THE 


contained in the Bruce collection; most of the specimens 
were collected in the neighbourhood of Cape Flora, especially 
about two-thirds of a mile south-west of Elmwood. Some were 
collected in January 1897, and others during April, May, June, 
and July. Several of the specimens were of large size: Prof. 
Sars states that 40 millim. is about the maximum length of Arctic 
specimens; one or two of the largest in the Bruce collection 
measured from 40 to 42°5 millim. 

Anonyx nugax seems to be oue of the more commonly distri- 
buted species in the Arctic seas. Prof. Sars (loc. cit.) gives 
Franz-Josef Land among the places mentioned in his note on the 
distribution of the species. I have obtained on two separate 
occasions in the Firth of Forth what is certainly the same 
Species. 


Genus Hoprtonyx, G. 0. Sars, 1890. 


Hoptonyx stminis, G. O. Sars. (Pl. 9. figs. 11-18.) 

1890. Hoplonyx similis, G. O. Sars (75), p. 98, pl. 33. fig. 1. 

A single adult female specimen (with ova) of an Amphipod 
which I ascribe to this species was taken about two-thirds of a 
mile south-west of Elmwood, in 18 fathoms water, on May 22nd, 
1897. At first I thought it might be a specimen of Hoplonyx 
cicada (Fabricius), as that is said to be a widely distributed 
species in the Arctic Seas, but a more careful examination showed. 
that the form of the fourth coxal plates (fig. 11) and of the last 
epimeral plates (fig. 12) differed from those of that species, while 
they agreed very closely with those of H. sémilis, G. O. Sars. 
In this specimen the integument is ornamented with numerous 
circular depressions as exhibited by the figures. 


Genus Pseupatisrorus, Della Valle. 


* PSEUDALIBROTUS LITTORALIS (Avdyer). 
1845. Anonyx hittoralis, Kroyer (46), 2 R.1 B. p. 621. 
This species was obtained by Mr. Bruce on several occasions, 


* Prof. Sars, in ‘ Crustacea of Norway,’ vol. i. p. 102, expresses some doubt 
asto whether M.-Edwards’s generic name Alibrotus is correctly applied to this 
northern form, but he does not propose any substitute for it. More recently, 
however, Della Valle instituted a new genus, Psewdalibrotus, for Kroyer’s species, 
and I have adopted this name here. 


CRUSTACEA OF FRANZ-JOSEF LAND. vai 


and usually in comparatively shallow wateror on the shore; it 
appeared to be one of the more common of the Franz-Josef Land 
Amphipoda. The following are some of the localities where it 
was obtained:—West Bay, Cape Flora, in 2 to 3 fathoms, on 
the 22nd and 25th August, and off Flagstaff Point, Elmwood, 
on the 19th September, 1896. Fifty yards off West Point, Cape 
Flora, in 2 to 8 fathoms, on July 7th; and on sand near Hast 
Glacier, Cape Flora, on August Ist, 1897. A few days after- 
wards, viz. on the 7th of August, the same species was taken on 
the shore at Cape Mary Harmsworth. Prof. G. O. Sars (Joe. cit.) 
also records the species from Franz-Josef Land. 


Genus Onestuus, Boeck, 1870. 


Onestmvus Epwarpstt (Kroyer). 

1846. Anonyx Edwardsi, Kroyer (47), 2 R. 2B. p. 1. 

This species was also represented by numerous specimens in 
the Bruce collection ; it was taken at the surface of the water 
and at various depths down to 26 fathoms. In 1896 it was 
obtained at West Bay, Cape Flora, in 2 to 3 fathoms water, on 
the 22nd of August, and at about a mile off Flora Cottage in 15 
fathoms on the 10th September. Onesimus Hdwardsii was taken 
on several occasions during 1897 ; the first record of it for that 
year is on January 11th, when specimens were collected about 
two-thirds of a mile south-west of Elmwood, others were after- 
wards collected near the same locality during April, May, and 
June. The species also occurred in other places, but they 
were all in the neighbourhood of Cape Flora, near the western 
extremity of Northbrook Island. This is also one of the species: 
recorded by Prof. Sars from Franz-Josef Land. 


Genus Amputtocuts, Spence Bate, 1862. 


AMPHILOCHUS OcULATUS, Hansen. 

1887. Amphilochus oculatus, Hansen (86), p. 89, pl. iii. figs. 2-2 ¢. 

Only one or two specimens of this comparatively small species 
were observed in the Bruce collection ; they occurred in a gather- 
ing of material dredged in 2 to 10 fathoms in West Bay, Cape 
Flora, on July 2nd, and off Cape Gertrude in 30 fathoms on July 
21st, 1897, 


2, MR. THOMAS SCOTT ON THE 


Genus Grrana, Boeck, 1870. 


Grrana Sarstt, Boeck. 

1870. Gitana Sarsi, Boeck (6), p. 52. 

A single specimen of this species occurred in a gathering of 
Microcrustacea dredged off Hast Glacier, in 30 fathoms, July 21st, 
1897. 


Genus Meropa, Boeck, 1870. 


Meropa pusiuna, G. O. Sars. 

1890. Metopa pusilla, G. O. Sars (75), p. 256, pl. 90. fig. 1. 

One or two specimens of a Metopa apparently belonging to 
this species were obtained off Hast Glacier, at a depth of 30 fathoms, 
and off Cape Gertrude, on July 21st, 1897. 


Merora sinvuata, G. O. Sars. 

1890. Metopa sinuata, G. O. Sars (75), p. 263, pl. 92. fig. 2 

This species was represented by a single specimen which was 
dredged off Cape Mary Harmsworth, in 53 to 93 fathoms, on 
August 7th, 1897. 


Merora NEGLECTA, Hansen. 

1887. Metopa neglecta, Hansen (36), p. 96, pl. ili. figs. 9-9 e. 

This species was obtained in the same gatherings with J. 
pusilla and was represented by only one or two specimens. 

All these three species of Metopa have already been recorded 
from the Arctic seas, but not from Franz-Josef Land. 


Genus Par@piceros, G. O. Sars, 1890. 


PaR@DICEROS LYNCEUS (JZ. Sars). 

1858. Cidiceros lynceus, M. Sars (66 a), p. 143. 

This species, which is widely distributed in the Arctic seas, 
occurred very sparingly in the Bruce collection. It was first taken 
September 10th, 1896, about a mile off Flora Cottage, in 15 
fathoms, and it also occurred amongst some material dredged in 
‘West Bay, Cape Flora, in 2 to 10 fathoms, on July 2nd, 1897. 


Genus Monocutopzs, Stimpson, 1858. 


MonocuLopEs BOREALIS, Boeck. 
1870. Monoculodes borealis, Boeck (6), p. 88. 
Two specimens of I. borealis occurred in the Bee) collechone 


CRUSTACEA OF FRANZ-JOSEF LAND. 73 


they were obtained in West Bay, Cape Flora, in 2 to 3 fathoms, 
in July 1897. This species has been taken in the Firth of 
Clyde. 


MOoNOCULODES LATIMANUS (Goés). 

1866, Gidiceros latimanus, Goés (32), pl. ii. fig. 23. 

A single specimen of an Amphipod apparently belonging to 
this species was obtained amongst some dredged material from 
West Bay, Cape Flora; it seemed to differ slightly from J 
latimanus in the form of the rostrum, but it otherwise Tee 
very well with that species. 


MonocuLopes ScHNFIDERI, G. O. Sars. 

1895. Monoculodes Schnetderi, G. O. Sars (75), p. 692, pl. vi. (Suppl.) 
fie. 1. 

This pretty species was represented in the Bruce collection by 
a considerable number of specimens; they mostly occurred in the 
one gathering dredged in the vicinity of West Bay in 2 to 1C 
fathoms. 


Genus Monocutopsis, G. O. Sars, 1891. 


MonocuLopsis LONGICORNIS (Boeck). 

1870. Monoculodes longicornis, Boeck (6), p. 85. 

A single specimen of Monoculopsis was obtained in the 
gathering containing IZ. Schneideri from West Bay. This species 
has a superficial resemblance to Perioculodes longimanus, but the 
enathopods, and especially the first pair, differ considerably. 
I have this species also from Baflin’s Bay. 


Genus Barnymepon, G. O. Sars, 1891. 


BAaTHYMEDON OBTUSIFRONS (Hansen). 

1887. Halimedon obtusifrons, Hansen (36), p. 116, pl. v. fig. 1. 

A single specimen of an Amphipod whieh I identify as Bathy- 
medon obtusifrons (Hansen) was obtained in the same gathering 
as the two species last recorded. 


Genus AcERos, Boeck, 1860. 


AcERos PHYLLONYX (I. Sars). 
1858, Leucothoé phyllonyx, M. Sars (66a), p. 148. 
A specimen of this species was included in the Bruce collectior, 


74, MR. THOMAS SCOTT ON THE 


but was taken a considerable distance to the south of Franz- 
Josef Land. It was dredged in lat. 77° 53’ N., long. 53° 16’ E. 
(or almost midway between Novaya Zemlya and Northbrook 
Island), and at a depth of 130 fathoms. Heller records this 
species from Franz-Josef Land. 


Genus AcanTHOosTEPHEIA, Boeck, 1870. 


ACANTHOSTEPHEIA MatmGRENI (Goés). 

1865. Amphithonotus Malmgrent, Goés (32), p. 10, fig. 17. 

This species was represented by a single specimen dredged in 
the vicinity of West Bay, in from 2 to 10 fathoms, on July 2nd, 
1897. Mr. Stebbing records Acanthostepheia from lat. 75° 14° 
N., long. 44° 26’ E., as well as from other parts of Barents Sea ; 
it has also been recorded from Stor Fjord and other parts of the 
Spitzbergen coast by Goés, and from West Greenland by Hansen. 
The peculiar conformation of the eyes in this species gives it a 
somewhat owtré appearance. 


Genus ParampnuitHok, Bruzelius, 1859. 


PARAMPHITHOE PULCHELLA (Kroyer). 

1846. Amphithoé pulchella, Kroyer (47), pl. 10. fig. 2. 

Two specimens were captured with the dredge off West Bay, 
Cape Flora, at a depth of about 8 fathoms, on July 24th, 1897. 
Paramphithoé pulchella, though widely distributed, seems to be 
scarce in the Arctic seas; but appears to be more or less common 
on the west and north of Norway. 


PARAMPHITHOE BICUSPIS (Kroyer). 

1838. Amphithoé bicuspis, Kroyer (41), p. 273, pl. 2. fig. 10. 

This species was obtained amongst some small Crustacea 
dredged in West Bay, in from 2 to 10 fathoms, on July 2nd, 
1897. This species is widely distributed in the Arctic seas, and 
is also included in the British fauna. 


PARAMPHITHOE MoNocUusPIS, G. O. Sars. 

1892. Paramphithoé monocuspis, G. O. Sars (75), p. 351, pl. 123. 
fig. 2. 

The most prominent difference between this and P. bicuspis is 
that suggested by the name—this species having only one dorsal 


CRUSTACEA OF FRANZ-JOSEF LAND. 75 


cusp instead of two. The species otherwise are closely related ; 
but as the difference referred to appears to be fairly constant, 
and as there are one or two points in which a disagreement 
between the two forms is observed, it is more satisfactory to 
regard this as a distinct species than simply asa variety. More- 
over, both forms have an extensive distribution. P. monocuspis 
is represented in the collection of Franz-Josef Land Crustacea 
by two specimens, one of which was obtained off Bear Berg, in 
about 10 fathoms, on September 27th, 1896; and the other about 
one mile off Flora Cottage, in 15 fathoms, on the 10th of the 
same month. Prof. Sars records the species from Greenland 
and from Norway ; and it is also a member of the British fauna _ 


Genus Parapneustes, Buchholz, 1874. 


PARAPLEUSTES GLABER (Boeck). 

1860. Amphithopsis glaber, Boeck (4), p. 662. 

This species is represented by a single specimen captured off 
Bear Berg, in 10 fathoms, on September 27th, 1896. It has 
been recorded from various other Arctic localities, and south as 
far as Christiania Fjord and the Kattegat. 


Genus AcantHonorosoma, Boeck, 1876. 


ACANTHONOTOSOMA CRISTATUM (Owen). 

1835. Acanthonotus cristatus, Owen (61), p. 90, pl. B. figs. 8-12. 

This species is included in the collection of Crustacea from 
Franz-Josef Land, but was dredged somewhat to the south of 
that Archipelago in lat. 77° 53’ N., long. 53° 20’ E., at a depth 
of 130 fathoms, on July 11th, 1897. Goés records this from 
Spitzbergen, Hoek from Barents Sea, and Hansen from the Kara 
Sea; while Mr. Stebbing records the occurrence of a specimen 
from lat. 75° 14’ N., long. 44° 26’ E., from a depth of 130 
fathoms. Goés’s Spitzbergen Station ((Heenloopen Strat) is 
somewhat farther north than that of Mr. Bruce. 


Genus SyRRHOE, Gioés, 1865. 


SyRRHOE CRENULATA, Gloés. 

1865. Syrrhoé crenulata, Goés (32), p. 11, fig. 25. 

This species was dredged in 15 fathoms, about one mile off 
LINN. JOURN.—ZOOLOGY, VOL. XXVII. Uf 


76 MR. THOMAS SCOTT ON THE 


Flora Cottage, on September 10th, 1896, and off West Bay, Cape 
Flora, in 8 fathoms, on July 24th, 1897. Syrrhoé crenulata 
appears to be a somewhat scarce species in the Arctic seas, but 
seems to be more frequent off the coast of Norway. 


Genus Parpatisca, Kroyer, 1842. 


PARDALISCA CUSPIDATA, Kroyer. 

1842. Pardalisca cuspidata, Kroyer (43), p. 153. 

This species was also taken off Flora Cottage on September 
10th, 1896, along with the Syrrhoé. One specimen only was 
obtained. Pardalisca cuspidata, although apparently scarce in 
the Aret seas, is, according to Sars, not uncommon off the 
coast of ¥inmark. 


Genus EvsiRus, Kroyer, 1845. 


Eustrus cusprpatus, Kréyer. 

1845. Eusirus cuspidatus, Kroyer (46), p. 501. 

A single specimen of Husirus cuspidatus was captured about 
two-thirds of a mile south-west of Elmwood on May 21st, 1897. 
The distribution of the species seems to be almost restricted to 
the Arctic seas. 


Genus Ruacnorroris, 8. Smith, 1883. 


RHACHOTROPIS AOULEATA (Lepechin). | 

1778. Oniscus aculeatus, Lepechin (48), p. 247, pl. 8. fig. 1. 

This species was represented in the Bruce collection by 
several specimens, which were obtained as follows:—A few 
specimens, all more or less immature, were taken in about 10 
fathoms off West: Glacier, in Giinther Sound, on September 9th, 
1896; an adult species was captured two-thirds of a mile 
south-west of Elmwood on April 30th, 1897 ; other specimens, 
more or less immature, occurred off West Point, Cape Flora, in 
2 to 3 fathoms, on June 22nd; on June 26th three immature 
specimens were obtained south-west of Elmwood, in 18 fathoms; 
and on July 18th one or two more, also immature, were obtained 
in 4 fathoms off East Glacier, Cape Flora. Heller records this 
species also from Franz-Josef Land. 


CRUSTACEA OF FRANZ-JOSEF LAND. 77 


Genus Rozinante, 7. BR. R. Stebbing, 1894. 


ROZINANTE FRAGILIS (Goes). 

1866. Paramphithoé fragilis, Goés (32), p. 524, pl. 39. fig. 16. 

This Amphipod was described by Goés as a Paramphithoé, but 
it was afterwards referred by Boeck and others to Tritropis. 
As, however, this name was already in use, Prof. S. Smith, in 
1883, altered it to Rhachotropis. Along with these changes in 
the name, the characters also of the genus had been modified 
and restricted, with the result that Goés’s species was dis- 
inherited. Mr. Stebbimg came to the rescue of this unfortunate 
Amphipod and instituted a new genus (Rozinante) for its recep- 
tion, where it now seems to be at rest. The following are some 
of the localities where Rozinante was obtained by Mr. Bruce :— 
Off Flora Cottage, in 15 fathoms, September 10th, 1896; two- 
thirds of a mile south-west of Elmwood, in 20 fathoms, January 
11th, 1897; off West Point, Cape Flora, in 2 to 3 fathoms, 
June 22nd; off Cape Gertrude, in 30 fathoms, July 17th; off 
West Bay, Cape Flora, in 8 fathoms, July 23rd and 24th; and off 
Cape Mary Harmsworth, in 53 to 93 fathoms, August 7th, 1897. 
Goés records it from Wijde Bay, on the north, and Stor Fjord 
on the east side of West Spitzbergen, and also from Greenland. 
Hansen records it also from Greenland, and Mr. Stebbing from 
the Kara Sea (lat. 71° 19’ N., long. 63° 34’ E.). Its distribution 
seems to be limited to the Arctic seas. 

There seems to be considerable variation in the length of the 
cleft of the telson, in some instances it does not exceed one- 
sixth of the length, while in others it is as much as one-third. 


Genus Hattraces, Boeck, 1870. 


HatiraGEs FuLvocinetus (1. Sars). 

1854. Amphithoé fulvocincta, M. Sars (66), p. 141. 

This species was represented in the Bruce collection by a con- 
siderable number of specimens; they were collected at various 
localities during 1896 and 1897, but chiefly in the neighbourhood 
of Cape Flora, near the western extremity of Northbrook 
Island. The following is a brief summary of the places where 
Halirages fulvocinctus was obtained:—In West Bay, Cape 
Flora; south-west of Elmwood, Cape Flora; off Cape Flora 
itself; off Cape Gertrude ; near Hast Glacier ; off West Giacier, 

A 7% 


78 - MBE. THOMAS SCOTT ON THE 


off Bear Berg; and off Wilczek Island. H. fulvocinctus appears 
to be a common Arctic species; it also extends ‘‘ along the whole 
west and north coasts of Norway.” 


Genus CLEIPPIDES, Boeck, 1870. 


CLEIPPIDES QUADRICUSPIS, Heller. 

1878. Cleippides quadricuspis, Heller (37), pp. 25-40. 

This species was described by Prof. Camil Heller from spe- 
cimens captured during the Austrian North-Polar Expedition 
(1878). It was represented in the Bruce collection by a single 
adult specimen, which was dredged in 130 fathoms, in lat. 77° 53° 
N., long. 53° 16’ E., on July 13th, 1897. In this specimen the 
dorsal cusps were very prominent, even more so than is shown 
in Heller’s figure. 


Genus Caxtiorius, Lilljeborg, 1865. 


CALLIOPIUS LHVIUSCULUS (Kroyer). 

1838. Amphithoé leviusculus, Kroyer (41), p. 281, pl. 3. fig. 15. 

A single specimen of this widely distributed species occurred 
among some Crustacea collected near West Point, Cape Flora, 
on July 5th; while a second was obtained in a gathering 
collected about two-thirds of a mile south-west of Elmwood on. 
January 11th, 1897. 


Genus AmpuHitTHopsis, Boeck, 1870. 


AMPHITHOPSIS GLACIALIS, Hansen. 

1887. Amphithopsis glacialis, Hansen (36), p. 137, pl. 5. figs. 6-6 e. 

This species was taken off West Point, Cape Flora, in 2 to 3: 
fathoms, on the 20th of June; and algo inshore at Cape Mary 
Harmsworth, on August 7th, 1897. 


Genus Atyuus, Leach, 1817. 


ATYLUS CARINATUS (Fabricius). 

1798. Gammarus carinatus, Fabricius (28), t. ii. p. 515. 

The only localities where this species was obtained were off 
West Bay, Cape Flora, in 5 fathoms, on September 12th, 1896; 
and off West Point, Cape Flora, in 2 to 3 fathoms, on June 20th, 
1897— one specimen being obtained at each place. Atylus cari- 
natus has also been recorded for Franz-Josef Land by Miers. 


CRUSTACEA OF FRANZ-JOSEF LAND. 79 


Genus AmaTHILLA, Spence Bate, 1863. 


AMATHILLA HOMARI (Fabricius). 

1779. Astacus homari, Fabricius (27), p. 247. 
CNThis species was obtained off West Point, Cape Flora, in 2 to 
A fathoms, on two different occasions, viz., on June 22nd and 
July 5th, 1897. 


g 

AMATHILLA PINGUIS (Kroyer). (PI. figs. 14, 15.) 

1838. Gammarus pinguis, Kroyer (41), p. 24, pl. 1. fig. 5. 

This Amathilla was represented by several specimens captured 
in the neighbourhood of Cape Flora on different occasions during 
June and July, 1897.. Prof. G. O. Sars states that this species 
ought, in his opinion, to be removed as the type of a separate 
genus, as it differs considerably in several points “from the 
typical Amathilla.” In Amathilla pinguis the last epimeral 
plates of the metasome somewhat resemble those of Apherusa 
Jurinet. 


Genus Gammaracantuus, Spence Bate, 1862. 


GAMMARACANTHUS LoRICATUS (Sabine). 

1824. Gammarus loricatus, Sabine (65), p. 131, pl. 1. fig. 7. 

A single specimen of this fine species was captured about 50 
yards off West Point, Cape Flora, in 2 to 3 fathoms, on June 
22nd, 1897. G. loricatus appears, in its distribution, to be 
restricted to the Arctic seas, and is not even recorded from the 
coast of Finmark, a district that has furnished not a few Arctic 
forms to the fauna of Norway. 


‘Genus Gammarus, Fabricius. 


GAMMARUS Locusta (Linne). 

1767. Cancer locusta, Linné (50), p. 1055, 

Numerous examples of Gammarus locusta, comprising all 
' stages from embryos to adults, were included in the Bruce 
collection. Nearly all the specimens were from inshore, and 
formed part of every inshore gathering of invertebrates. Some 
of the specimens were of large size, while a considerable pro- 
portion of the adult females carried ova or embryos. Prof. Sars 
includes Franz-Josef Land in his notes on the Arctic distribution 
of Gammarus locusta. 


‘80 MR. THOMAS SCOTT ON THE 


Genus Puoris, Kroyer, 1842. 


Parorts renvscornts, G. O, Sars. (P1.M. figs. 16, 17.) 

1883. Photis tenuicornis, G. O. Sars (73), p. 110, pl. 6. fig. 4. 

This species was represented by a very few specimens collected 
chiefly in the neighbourhood of Cape Flora, as, for example, off 
Flora Cottage, in 15 fathoms, on September 10th, 1896, and off 
West Glacier, Giinther Sound, in 10 fathoms, on September 9th, 
1896. In the male of this species a row of minute spines extends 
in a diagonal manner across the basal joint of the posterior 
gnathopoda (fig. 17). 


Genus Iscuyrocerts, Kroyer, 1838. 
2 


Iscuyrocerus(?) anaurpes, Kroyer. (Pl. ¥. fig. 18.) 

1838. Ischyrocerus anguipes, Kroyer (41), p. 55, pl. 3. fig. 14. 

A few specimens of an Amphipod probably belonging to this 
species were among the Crustacea in the Bruce collection. In 
the male specimens the propodos (fig. 18) differed from those 
of the fully developed male of Ischyrocerus anguipes, but this 
difference may be due to the specimens being scarcely mature. 
The following are the localities where the specimens were col- 
lected :—Off West Glacier, Gunther Sound, in 10 fathoms, 
September 9th, 1896; and off West Point, Cape Flora, June 
20th, 21st, and 22nd, 1897. 


Genus Duticuta, Kroyer, 1845. 


DvuticHia sprnostssima, Kroyer. 

1845. Dulichia spinosissima, Kroyer (46), p. 512, pl. 6. fig. 1. 

This curious species was represented by a single specimen 
captured about two-thirds of a mile south-west of Elmwood, at a 
depth of 18 fathoms, on April 28th or 29th, 1897. It appears to 
be restricted to the Arctic seas. 


(O} N48 18) 1h 1h Te 1D) 1S NG 


Genus Alaina, Ardyer, 1843, 


AEGINA SPINOSISSIMA, Stimpson. 

1853. gina spinosissima, Stimpson (88), p. 44. 

Representatives of this species were obtained off Flora Cottage 
in 15 fathoms, September 10th, 1896; off West Bay, Cape 


CRUSTACEA OF FRANZ-JOSEF LAND. 81 


Flora, in 8 fathoms, on July 28rd or 24th, 1897; and off Cape 
Mary Harmsworth, in 53 to 93 fathoms, on August 7th, 1897. 
The largest specimen measured from rostrum to telson about 38 
millimetres, and the antennules 32 millimetres, or a total length 
of 24 inches. 


Genus Capre.LaA, Lamarck, 1818. 


CAPRELLA SEPTENTRIONALIS, Kroyer, forma e. parva, Mayer. 

1838. Capreila septentrionalis, Kroyer (41), p. 90. 

This species was dredged in 130 fathoms in lat. 77° 53’ N., | 
long. 53° 20’ E. One specimen only was obtained. 


CAPRELLA MICROTUBERCULATA, G. O. Sars. 

1865. Caprella microtuberculata, G. O. Sars (69). 

This species was taken in 10 fathoms, off Bear Berg, on 
September 28th, 1896; and off Cape Mary Harmsworth, in 53 to 
93 fathoms, August 7th, 1897. 


CaPRELLA DUBIA, Hansen. 

1887. Caprella dubia, Hansen (36), p. 217, pl. 4. figs. 8-8 d. 

A specimen of this Caprelia was dredged by Mr. Bruce, in 
10 fathoms off Bear Berg, on September 28th, 1896. Dr. Hansen 
recorded this species in his work on the Crustacea of Green- 
land, and he at first described it as Caprella microtuberculata, 
G. O. Sars, var. spinigera; but in a postscript to the same work 
(p. 217) he considered that the form he had so described should 
rank as a species, for which he proposed the new name of 
Caprella dubia. 


ENTOMOSTRACA. 


OSTRACODA. 


The Ostracoda contained in Mr. Bruce’s collection number 
thirty-four species; the first four are freshwater forms, all the 
others are marine ; the freshwater species are from ponds in the 
vicinity of Elmwood, at the western extremity of Northbrook 
Island. A pond near Cape Mary Harmsworth was also examined, 
but no Ostracoda were obtained in it. Three of the freshwater 
and one of the marine species appear to be undescribed ; but all 
the others belong to more or less well-known forms, and are all 
represented in the British fauna either as recent or fossil. As 


82 MR. THOMAS SCOTT ON THE 


to names and arrangement of the species recorded below, I have 
followed the Monograph of the marine and freshwater Ostracoda 
of the North Atlantic and North-western Europe by Prof. G. &. 
Brady and the Rev. Dr. A. M. Norman. 


PODOCOPA. 


(a. Freshwater.) 
& Genus Cyctocyrris, Brady ¢ Norman, 1889. 

CycLocypris aLoposa (G. O. Sars). (Pl. 4. fig. 1.) 

1863. Cypris globosa, G. O. Sars (68), p. 27. 

Single valves of an Ostracod apparently identical with C. glo- 
bosa were obtained in the gathering from Elmwood Pond. The 
valve figured measures about 1:1 millimetre in longest diameter. 
(This may be the Cypria lacustris of Lilljeborg, but the con- 
vexity of the shell is rather too great to fit that species.) 


Genus Hrerperocyrris, Brady & Norman, 1889. 


HERPETOCYPRIS (?) DUBIA, sp.n. (Pl. 4. figs. 7-11.) 

The shell seen from the side is subreniform ; the greatest height, 
which is equal to fully half the length, is situated about one- 
third of the entire length from the anterior end; the upper 
margin is considerably elevated in front of the middle; the 
posterior slope is gentle and slightly curved, but anteriorly the 
slope is more abrupt ; the anterior extremity is broadly rounded ; 
the posterior end is also evenly and gently rounded, but is 
rather narrower than the front end; lower margin slightly 
sinuated. Seen from above, ovate, widest in the middle, greatest 
width equal to about half the length; sides evenly curved ; 
extremities slightly acuminate. Shell surface smooth and 
yellowish in colour. Length 13 mm. 

This Ostracod, seen from above, somewhat resembles Cypris 
crassa, O. F'. Miiller, and the side view is also suggestive of the 
same species ; but when compared with the figures of that species 
in Brady and Norman’s Monograph, the present form, viewed 
laterally, is seen to be more boldly arched. It may also be noted 
that, when viewed laterally, H. dubia somewhat resembles 
Candona rostrata seen in the same position; but in that species 
the dorsal view is different. It is just possible that H. dubia 
may represent a somewhat immature stage of the next species. 


CRUSTACEA OF FRANZ-JOSEF LAND. 83 


HERPETOCYPRIS ARCTICA, sp. n. (PI. 4. figs. 2-6.) 

Shell, seen from the side, subreniform, highest in front of the 
middle; greatest height equal to fully half the length, extremities 
broadly rounded; dorsal margin moderately arched, ventral 
slightly sinuated. Seen from above, ovate, tumid, greatest width 
situated behind the middle and nearly equal to half the length: 
posteriorly the sides converge in a broadly rounded curve and, 
where they meet, form an obtuse angle; they taper more gradu- 
ally towards the anterior end, and the extremity there is slightly 
acuminate. Colour bluish green, ornamented with darker streaks 
and blotches. Length 1:9 mm. 

This Ostracod, which was moderately frequent in a freshwater 
pond near Elmwood, Cape Flora, was collected July 30th, 1897. 


Genus Canpona, Baird, 1845. 


Canpona Harmsworrtat, sp. n. (PI. 3. figs. 16, 17.) 

The shell, seen from the side, is somewhat subreniform ; the 
dorsal margin is considerably elevated near the posterior end, 
where the anterior and posterior slopes meet and form an obtuse 
angle; the greatest height is equal to rather more than half 
the length; the front slope curves gently downwards to the 
evenly rounded anterior extremity; the posterior end is sub- 
truncate and forms a slight curve from the obtuse dorsal angle 
downwards and backwards to where it meets the ventral margin ; 
the ventral margin is distinctly incurved in front of the middle. 
The shell, seen from above, is ovate; the greatest width, which 
is situated behind the middle, is equal to rather more than two- 
fifths of the length; extremities slightly acuminate. Length 
1 mm. 

I have named this quite distinct species after Mr. Harmsworth, 
of the Jackson-Harmsworth Expedition. 

Candona harmsworthi has a distant resemblance to C. candida, 
var. claviformis, when viewed laterally, but the dorsal view is 
dissimilar. 


(b. Marine Species.) 


Genus Pontocypris, G. O. Sars, 1865. 


Pontooypzis (?) HYPERBOREA, sp.n. (PI. 4. figs. 12-15.) | 
Shell, seen from the side, subreniform, dorsal margin boldly 
arched, being almost semicircular, height equal to half the length ; 


84 . MR. THOMAS SCOTT ON THE 


ventral margin nearly straight; anterior end narrowly rounded, 
posterior subangular. Seen from above ovate, the sides evenly 
curved; ends slightly acuminate, or forming an acute angle ; 
width scarcely equal to half the length. Seen from the end, 
somewhat triangular, with the underside (the base of the triangle) 
slightly oblique. Surface of the shell smooth, white, with scat- 
tered opaque circular markings. Length ‘9 mm. 

A single dead specimen was dredged in West Bay, Cape Flora, 
at a depth of 2 to 10 fathoms, on July 2nd, 1897. I only 
provisionally ascribe this Ostracod to Pontocypris, for without 
the animal it is hardly possible to determine the genus it may 
belong to. 


Genus CytHERE, Jfiiller, 1781. 


CYTHERE MARGINATA, WVorman. 

1862. Cythere marginata, Norman (56), p. 47, pl. 3. figs. 10-12. 

Dredged one mile off Cape Flora, in 15 fathoms, September LOth, _ 
1896. This has been recorded from Spitzbergen, Norway, and. 
from various British localities. 


CYTHERE LimicoLa, Norman. 

1865. Cythere imicola, Norman (58), p. 20, pl. 6. figs. 1-4. 

Dredged off Hast Glacier, Cape Flora, in 30 fathoms, July 21st, 
1897. This was a somewhat rare species in the Franz-Josef 
Land collection ; it is also one of the less common of the British 
species. It has been recorded from Baffin’s Bay by Dr. Brady, 
and from Norway by G. O. Sars. 


CYTHERE GLOBULIFERA, G. S. Brady. 

1868. Cythere globulifera, Brady (12), p. 406, pl. 31. fig. 42. 

This species was also rare in the collection, one or two speci- 
mens only having been observed. Dr. Norman records it from 
Norway, and Prof. Brady from Spitzbergen. Asa recent species 
it is rare in the British seas, but is less rare as a post-tertiary 
fossil.. It was taken off Hast Glacier with the last. 


CYTHERE CLUTH#, Brady, Crosskey, 5 Robertson. 


1874. Cythere cluthe, Brady, Crosskey, & Robertson (16), p. 153, oil 13. 
figs. 16, 17. 


This was also a rare species ; it was taken off East Glacier, in 
30 fathoms, on July 21st, 1897. C. cluthe was first described 
from fossil specimens, but has since been obtained as a recent 


CRUSTACEA OF FRANZ-JOSEF LAND. 85 


species in several British localities, as the Irish Sea (Malcolmson) ; 
Loch Fyne and Stromness Harbour (mihi). Prof. G. S. Brady 
records it from Cape Frazer, from specimens obtained in Capt. 
Feilden’s dredgings during Nares’s Arctic Expedition. 


CYTHERE SEPTENTRIONALIS, G. S. Brady. 

1866. Cythere septentrionalis, G.S. Brady (11), p. 875, pl. 60. figs. 4 af. 

This fine species was dredged one mile off Cape Flora, in 
15 fathoms, on September 10th, 1896, and off Hast Glacier, in 
30 fathoms, July 21st, 1897. Prof. Brady described the species 
from specimens obtained in Dr. P. E. Sutherland’s dredgings at 
Hunde Islands, Baflin’s Bay, in 60-70 fathoms. 


CYTHERE TUBERCULATA (G. O. Sarvs). 

1865. Cythereis tuberculata, G. O. Sars (71), p. 37. 

This was dredged in 15 fathoms about one mile off Cape 
Flora, September 10th, 1896; also off East Glacier and off Cape 
Gertrude, in 30 fathoms, July 21st, 1897. It is a common and 
widely distributed species. 


CYTHERE EwARGrINata (G. O. Sars). 

1865. Cythereis emarginata, G. O. Sars (71), p. 38. 

Dredged off Cape Flora, September 10th, 1896; also off West 
Point, July 5th, 1897, and on the 21st cf the same month oft 
Cape Gertrude. C. emarginata has been obtained in Loch Fyne; 
but the specimens had probably washed out from a submarine 
post-tertiary deposit ; it has been obtained recent at Shetland. 


CytHErE custata, G. S. Brady. 

1866. Cythere costata, G.S. Brady (11), p. 375, pl. 60. figs. 50a. 

This was dredged off East Glacier, Cape Flora, in 30 fathoms, 
July 21st, 1897; it did not appear to be very common. 


CYTHERE MIRABILIS, G. S. Brady. 

1868. Cythere mirabilis, G. S. Brady (12), p. 415, pl. 29. figs. 7, 8. 

This species was very ‘rare in the Franz-Josef Land collec- 
tions; it was dredged with the last off East Glacier. Prof. 
Brady records C. mirabilis from Spitzbergen. 


CYTHERE DUNELMENSIS (Worman). 

1865. Cythereis dunelmensis, Norman (58), p. 22, pl. 7. figs. 1-4. 

Dredged in 15 tathoms, one mile off Cape Flora, Septem- 
ber 10th, 1896, but very rare in the Franz-Josef Land collection. 


86 MR. THOMAS SCOTT ON THE 


Genus CyTHERIDEA, Bosquet, 1852. 

CYTHERIDEA PAPILLOSA, Bosquet. 

1852. Cytheridea papillosa, Bosquet (8), p. 42, pl. 2. figs. 5 a—d. 

This was one of the more common species of Ostracoda in the 
Collection, and was obtained at several places; it was dredged off 
East Glacier, September 10th, 1896 ; at West Bay, Cape Flora, 
in 2 to 10 fathoms, July 2nd, 1897, and off Cape Gertrude, in 
30 fathoms, on the 21st of the same month. It seems to bea 
common Arctic, as it isa common British species. 


CYTHERIDEA PUNCTILLATA, G. S. Brady. 

1865. Cytheridea punctillata, G. 8. Brady (9), p. 189, pl. 9. figs. 9-11. 

Several specimens of this Ostracod were obtained in a gathering 
dredged off Cape Flora, September 1896, and in another dredged 
off East Glacier, July 1897. 


CyTHERIDEA SORBYANA, Jones. 

1856. Cytheridea sorbyana, Jones (39), p. 44, pl. 4. figs. 6 a-e. 

A considerable number of examples of Cytheridea Sorbyana 
were obtained in the Franz-Josef Land collection ; they occurred 
mostly in some material dredged off Cape Flora, September 10th, 
1896. All these three Cytherideas have been recorded for Spitz- 
bergen by Prof. G. S. Brady. 


Genus Eucytuers, Brady, 1868. 


EucyTHERE DECLIVIS (Norman). 

1864. Cythere declivis, Norman (57), p. 192. 

This Ostracod occurred very sparingly in the Collection ; it was 
dredged off Cape Gertrude and off East Glacier in July 1897. 


Genus XEsTOLEBERIS, G. O. Sars, 1865. 


XESTOLEBERIS DEPRESSA,; G. O. Sars. 

1865. Xestoleberis depressa, G. O. Sars (71), p. 68. 

This was dredged off Cape Flora, in 15 fathoms, September 
1896; at West Bay, Cape Flora, in 2 to 10 fathoms, and off 
West Point, in 2 to 4 fathoms, on July 5th, 1897; very few 
specimens were observed. 


Genus Cyruerura, G. O. Sars, 1865. 


CYTHERURA UNDATA, G. O. Sars. 
1865. Cytherura undata, G. O. Sars (71), p. 75. 
Only a few specimens of this species were observed in the 


CRUSTACEA OF FRANZ-JOSEF LAND. 87 


Collection ; they occurred in two gathermgs—one dredged off 
West Point, the other off Hast Glacier. This has been recorded 
from Spitzbergen and other Arctic localities ; it is also frequent 
as a British species. 


CyTHERURA FULVA, Brady § Robertson. 

1874. Cytherura fulva, Brady & Robertson (21), p. 116, pl. 4. figs. 1-5. 

This was dredged off Hast Glacier, Cape Flora, at a depth of 
30 fathoms, July 21st, 1897, but very few specimens were ob- 
served. This is a widely distributed species in the British seas; 
but it does not appear to have been recorded from the Arctic 
seas. 


CYTHERURA CLATHRATA, G. O. Sars. 

1865. Cytherura clathrata, G. O. Sars (71), p. 77. 

This distinct and pretty species was dredged off East Glacier 
with the previous species, and, like it, was also rare, very few 
specimens being observed. 


Genus CyTHEROPTERON, G. O. Sars, 1865. 


CYTHEROPTERON LATISsstMuM (Norman). 

1865. Cythere latissima, Norman (58), p. 19, pl. 6. figs. 5-8. 

One or two specimens only of this species were observed in 
the Collection ; they occurred in some material dredged off West 
Point, Cape Flora, in 2 to 4 fathoms, in July 1897. 


CYTHEROPTERON PYRAMIDALE, G. S. Brady. 

1868. Cytheropteron pyramidale, G. S. Brady (12), p. 34, pl. 5. 
figs. 11-14. 

This was scarcely so rare as the last species; a few specimens 
were dredged off Cape Flora on September 9th, 1896, and off 
Cape Gertrude and East Glacier, in 30 fathoms, in J uly 1897. 


CYTHEROPTERON SUBCIRCINATUM, G. O. Sars. 

1868. Cytheropteron subcircinatum, G. O. Sars (71), p. 81. 

A considerable number of specimens of this Ostracod were 
dredged off Hast Glacier, Cape Flora, July 21st, 1897; this 
species does not appear to have previously been observed out of 
Norway. Prof. Brady agrees with my identification of the 
species. 


CYTHEROPTERON PuNCTATUM, G. S. Brady. 

1868. Cytheropteron punctatum, G. S. Brady (12), p. 449, pl. 34. 
figs. 45-48. 

This is a rare species in the Collection; it was obtained off 


88 MR. THOMAS SCOTT ON THE 


East Glacier, along with the last, and is also a new record for 
Arctic seas. 


CYTHEROPTERON ANGULATUM, Brady § Robertson. 

1872. Cytheropteron angulatum, Brady & Robertson (19), p. 62, pl. 2. 
figs. 7, 8. 

This also was very rare; it was taken off Hast Glacier along 
with C. punctatum and OC. subcircinatum. In view of these 
additions to the Arctic Ostracod fauna (for this also is now for 
the first time recorded for the arctic seas), it may be of interest 
to quote a remark made by the authors of the Monograph of 
the Marineand Freshwater Ostracoda of the North Atlantic and 
North-Western Europe concerning C. angulatwm, which is as 
follows :—‘ From its abundance in the glacial clays of Scotland 
it may be expected that this species (C. angulatum) will hereafter 
prove be a recent Arctic form.” It has been obtained as a 
recent species at quite a number of places around the Scottish 
coasts, a8 well as in Roundstone Bay, Ireland. 


Genus PsEUDOCYTHERE, G. O. Sars, 1865. 

PsEUDOCYTHERE CAUDATA, G. O. Sars. 

1865. Pseudocythere caudata, G. O. Sars (71), p. 88. 

Only one or two specimens of this curious species were 
obtained ; it was dredged off Hast Glacier, where so many other 
rare things were captured. It is of interest to note that Pseudo- 
cythere caudata is recorded by Prof. G. 8. Brady from the 
vicinity of Kerguelen Island, situated about 50° South latitude 
and 70° East longitude. 


Genus ScrERocHILUS, G. O. Sars, 1865. 

ScLEROCHILUS ConToRTUS (Norman). 

1862. Cythere contorta, Norman (56), p. 48, pl. 2. fig. 15. 

This was dredged off Cape Flora, September 10th, 1896, and 
off Cape Gertrude and East Glacier in July 1897. SS. contortus 
has already been recorded from the Arctic seas ; it is also one of 
the British species. It was moderately rare in the Franz-Josef 
Land collection. 


Genus ParapoxostoMa, Kischer, 1855. 
PARADOXOSTOMA VARIABILE (Baird). 
1835. Cythere variabilis, Baird (1), p. 98, pl. 5. figs. 7 a—b. 
Several specimens were obtained in some material dredged off 


CRUSTACEA OF FRANZ-JOSEF LAND. 89 


West Point and in West Bay, Cape Flora, in July 1897, at a 
depth of from 2 to 10 fathoms. This is recorded from Spitz- 
bergen and Greenland, and is also one of the more generally 
distributed species in Norway, as well as round the British 
Islands. 


PARADOXOSTOMA FLEXUOSUM, G. S. Brady. 

1866, Bythocythere ? fleawosa, G. S. Brady (11), p. 211. 

A few specimens of Paradoxostoma flecuosum were dredged 
off East Glacier in July 1897. It has been previously recorded 
from Davis Strait (lat. 69° 31’ N., long. 56° 1’ W.) by A. M. 
Norman, and is widely distributed southward to the Bay of 
Biscay. 


MYODOCOPA. 


Genus PuitomepsEs, Lilljeborg, 1853. 


PHILOMEDES BRENDA (Baird). 

1850. Cypridina brenda, Baird (3), p. 181, pl. 23. figs. 1 a-g. 

A number of specimens of this Ostracod were dredged in 
15 fathoms, one mile off Flora Cottage, on September 10th, 
1896, and off West Glacier, Giinther Sound, in 10 fathoms, on 
September 9th, 1897. Dr. Norman records Philomedes brenda 
from Holsteinbourg Harbour, Greenland; it has also been 
observed in different Norwegian localities as well as in Sweden. 
In Britain the only places where it has been obtained are off 
Noss, in Shetland, the Dogger Bank off the coast of Durham, 
and in the Clyde in the deep water to the east of Arran. 


CLADOCOPA. 
Genus Potycorr, G. O. Sars, 1865. 


PoLycoOPE ORBICULARIS, G. O. Sars. 

1865. Polycope orbicularis, G. O. Sars (71), p. 122. 

This species was dredged off Flora Cottage in September 1896, — 
and off East Glacier and Cape Gertrude in July 1897. Polycope 
orbicularis, though recorded from various parts of the British 
and Norwegian coasts, does not appear to have been previously 
observed so far within the Arctic circle; its occurrence there is, 
however, not surprising, seeing that it is moderately common as 


90 _ MR. THOMAS SCOTT ON THE 


a post-tertiary fossil, at least in the shell-bearing ‘Glacial clays” 


of Scotland. 


COPEPODA. 


Copepoda were fairly numerous in Mr. Bruce’s collection of 
Franz-Josef Land Crustacea. They represent a considerable 
number of species, and two of them are freshwater forms. 
Several of the species have been long known as denizens of the 
Arctic seas; but, on the other hand, this is the first time that a 
large proportion of them have been recorded from such high 
latitudes. Nearly all the Franz-Josef Land Copepoda belong to 
described species, only a very few being new to science. Another 
interesting point in regard to these Arctic Copepods is, that 
while the majority of the pelagic forms—that is such as are 
usually captured by tow-net—are of large size and belong to few 
species, the majority of those taken with the dredge are small 
and the species numerous. It may be remarked further that 
in recording the species of Copepoda obtained in the Franz- 
Josef Land collection, those belonging to the Calanide are 
placed first, then follow the Centropagide, the Mesophriade, the 
Cyclopide, the Harpacticide, andthe Onceade. The families 
Centropagide, Mesophriade, and Onceade are each represented 
by a single species, the Calanide and the Cyclopide by four 
species each, and the Harpacticide by 36 species, or 47 in all. 


Family CALANID&. 
Genus Catanus, Leach, 1819. 


CALANUS FINMARCHICUS (Gunner). 

1765. Monoculus finmarchicus, Gunner (35), p. 175, figs. 20-28. 

This species was fairly well represented in the Franz-Josef Land 
collection ; it occurred chiefly in tow-net collections from the 
vicinity of Cape Flora, Northbrook Island, as, for example, at 
West Bay; off east end of Cape Gertrude ; two-thirds of a mile 
south-west of Elmwood; off West Glacier, and also near Hast 
Glacier. 

CaALANUS HYPERBOREUS, Kroyer. 

1838. Calanus hyperboreus, Kroyer (41), p. 82, pl. 4. figs. 23 a-d. 

This is quite distinct from Oalanus finmarchicus, both in the 
junior and adult forms. Calanus hyperboreus was of rather more 


CRUSTACEA OF FRANZ-JOSEF LAND. 91 


frequent cccurrence than C. finmarchicus, and appeared to be more 
generally distributed ; but most of the specimens were collected 
in the neighbourhood of Cape Flora. Some of the adult females 
of this species were comparatively of large size; one that was 
measured was fully 8 millimetres (nearly one-third of an inch) 
in length from the forehead to the end of the caudal furca. 


Genus Psrvpocatanus, Boeck, 1872. 

PSEUDOCALANUS ELONGATUS (Boeck). 

1864. Clausva elongata, Boeck (5), p. 10. 

This species was observed in several tow-net gatherings and 
was moderately frequent in a few of them. Both male and 
female specimens were observed. ‘They occurred chiefly in 
gatherings from the neighbourhood of Cape Flora. 


Genus Evucumra, Philippi, 1843. 
KucH#@TA NoRVEGICA, Boeck. 
1872. Eucheta norvegica, Boeck (7), p. 40. 
A single, scarcely mature specimen of this Hucheta was 
captured with the tow-net about one and a half miles south-west 
of Elmwood on June 9th, 1897. 


Family CENTROPAGIDS. 
Genus Merripra, Boeck, 1865. 


Merripra tones (Lubbock). (Pl. 4. figs. 16, 17.) 

1854. Calanus longus, Lubbock (51), p. 127, pl. 5. fig. 10. 

This species was rather common in tow-gatherings collected in 
the neighbourhood of Cape Flora, as, for example, off Elmwood, 
near Hast Glacier, &c. A considerable proportion of the specimens 
were immature, Itis difficult to distinguish immature specimens 
of Metridia longa from those of Metridia hibernica; the adults 
of Wf. longa may be recognized by their larger size; but satis- 
factory identification can only be arrived at by the careful ex- 
amination of structural details in mature specimens. JJetridia 
longa has been recorded from the Faroe Channel. Some time 
ago I had the privilege of examining a series of tow-net gatherings 
collected between the Shetland Islands and Faroe, but the only 
Metridia obtained in these gatherings was IZ. hibernica (Brady 
& Robertson). [Metridia longa (Lubbock, 1854) is identified 
as MMetridia armata, Boeck (1865), but not Metridia armata, 

LINN. JOURN.—ZOOLOGY, VOL. XXVII. 8 


92 MR. THOMAS SCOTT ON THE 


Brady (1878): the latter is recognized as a distinct species by 
Giesbrecht under the name of Metridia hibernica (Brady & 
Robertson, 1873). ] 


Family MIisoPpHRIADa&. 


Genus Mrsopneta, Boeck, 1864. 


MisorHria PALLIDA, Boeck. 

1864. Misophria pallida, Boeck (5), p. 24. 

One or two specimens of this species were obtained in a 
gathering of micro-crustacea dredged at West Bay, Cape Flore, 
in 2 to 10 fathoms on July 2nd, 1897. Misophria pallida has 
been recorded from Norway, and it appears to be generally 
distributed around the British coast, but to be nowhere very 
common. In Scotland it has been obtained in the Firth of Forth, 
in Moray Firth, and in the Firth of Clyde. 


Family CycLopipS. 


Genus Orrnona, Baird, 1848. 


O1rHONA stmILis, Claus. (Pl. 4. figs. 18, 19.) - 

1866. Otthona similis, Claus (24). , 

This species was obtained in tow-net gatherings collected two- 
thirds of a mile south-west of Elmwood on May 22nd, 1897, but 
was comparatively scarce ; and it was taken with the tow-net in 
the same neighbourhood during the following month. This is 
certainly the Oithona similis, Claus, as described and figured by 
Dr. Giesbrecht in ‘ Pelagischen Copepoden des Golfes von 
Neapel.’ The same author refers doubtfully to this species the 
Oithona helgolandica, Claus (1863), and the Ozthona spinifrons, 
Boeck (1864); and with respect to Ovthona spinifrons, Boeck, 
Prof. Brady, in his ‘ Monograph of the Br.tish Copepoda,’ vol. i. 
published 20 years ago, states that he is “ not at all satisfied that 
this is not synonymous with O. helgolandica, Claus.” As there 
appears to be a consensus of opinion that all three species are 
synonymous, it would be better if the earlier name—O. helgo- 
landica—of Claus were adopted. Ovthona spinirostris, Claus 
(1863), is regarded by Dr. Giesbrecht as synonymous with 
O. plumifera, Baird (1843). 


CRUSTACEA OF FRANZ-JOSEF LAND. 93 


Genus Cyctoprina, Claus, 1863. 


CYCLOPINA GRACILIS, Claus. 

1863. Cyclopina gracilis, Claus (23), p. 104, pl. 10. figs. 9-15. 

A few specimens of this small but distinct species were 
obtained in some material dredged off Hast Glacier, Cape Flora, 
on July 21st, 1897. Cyclopina gracilis has a wide distribution 
in the British seas, but is not very common. 


Genus THoreLitA, Boeck, 1864. 


THORELLIA BRUNNEA, Boeck. 

1864. Thorellia brunnea, Boeck (5), p. 26. 

Thorellia brunnea was more or less frequent in several gather- 
ings of micro-crustacea collected by means of the dredge, as, for 
example, in gatherings from West Bay, off East Glacier, and 
south-west of Elmwood—this last was an inshore gathering in 
quite shallow water. 


Genus Crcroprs, O. &. Miller, 1776. 


Crctors Brucet, sp.n. (PI. 6. figs. 1-6.) 

This appears to be a new species, of which the following is a 
description :—Length of adult female 1-1 mm. (about Jy of an 
inch). Antennules (fig. 2) moderately short and stout, 12- 
jointed and sparingly setiferous ; the third and fifth joints are 
the shortest, the eighth, ninth, and last are rather longer than 
any of the others except the first. Both branches of the 
swimming-feet are short and stout and 3-jointed: in the first 
pair (fig. 3) the inner branches are armed with a strong terminal 
claw-like spine; the fourth pair (fig. 4) are less powerfully 
armed; the fifth pair (fig. 5) are small, the secondary joint is 
cylindrical in form, and the length rather more than twice the 
breadth, the truncate end bears interiorly a moderately short 
spine, and exteriorly a long seta that is at least four times the 
length of the joint from which it springs; the seta that springs 
_ from the exterior produced part of the basal joint is also elon- 
gate. The caudal furca (fig. 6) are nearly three times as long 
as the last abdominal segment; the small spiniform seta usually 
observed on the outer margins of the furca in Cyclops is situated 
at about a third of their length from the distal end; there is algo 


a minute notch near their base. 
ge 


94 MR. THOMAS SCOTT ON THE 


The species is named in compliment to Mr. Bruce, the Naturalist 
of the Jackson-Harmsworth Expedition. 

In some respects Cyclops Brucet resembles Cyclops bisetosus, 
Rehberg, but in that species the antennules are 17-jointed ; 
the armature of the swimming-feet also differs somewhat, and 
especially as regards the first pair. 

Hab. Pond near Elmwood, Cape Flora; not uncommon. 


Family HaARPACTICIDS. - 


Genus Brapya, Boeck, 1872. 


Brapya Typrca, Boeck. 

1872. Bradya typica, Boeck (7), p. 42. 

This species was somewhat rare in the Franz-Josef Land 
collections ; it was only observed in a gathering made off East 
Glacier, Cape Flora, in about 30 fathoms, on July 21st, 1897. 
Though Bradya typica appears to be widely distributed, it is not 
a common species anywhere. 


Brapya minor, 7. & A. Scott. 

1896. Bradya minor, T. & A. Scott (84), p. 425, pl. 35. figs. 5, 9, 18, 
21, 24, 31, 35, 42; pl. 36, figs. 5 & 9. 

This was one of several species described in the ‘ Transactions 
of the Linnean Society of London,’ vol. vi., 2nd ser., p. 425 ; 
it is distinguished from its near allies by its small size, brownish 
colour, and an “ eye-like dusky pigment-spot at the base of the 
antennules.” Bradya minor occurred along with B. typica in 
the gathering collected off Hast Glacier, Cape Flora, on July 21st, 
1897. Its distribution in the British Islands includes the Firths 
of Forth and Clyde and Liverpool Bay. 


Genus Ectinosoma, Boeck, 1864. 


EctinosoMa Sarst, Boeck. 

1872. Ectinosoma Sarst, Boeck (7), p. 45. 

This species was dredged off Hast Glacier in company with 
the two species of Bradya already referred to, and also at West 
Bay, Cape Flora, in from 2 to 10 fathoms on July 2nd, 1897. 
This is one of the larger and more common species of Kctinosoma 
in the British Copepod fauna. 


CRUSTACEA OF FRANZ-JOSEF LAND. 95 


Ecrinosoma PRopiInquuM, JZ. § A. Scott. 

1896. Ectinosoma propinguum, T. & A. Scott (84), p. 428, pl. 36. 
figs. 19, 27, 46, et seq. 

Two specimens of an Hetinosoma differmg little from the 
characters of H. propinquum were obtained in a gathering 
frum West Point, Cape Flora, at a depth of 2 to 4 fathoms, 
collected on 5th July, 1897. This species is one of the larger of 
the Hctinosomata, being not much less in size than Ketinosoma 
Sarsi; it possesses a distinctly hooked labium, the fifth pair of 
thoracic feet are of moderate size and resemble somewhat those 


of EF. Sarsi. 


EcrrnosoMa curticorne, Boeck. 

1864. Hetinosoma curticorne, Boeck (5). 

Dredged off Hast Glacier, Cape Flora, July 21st, 1897; rare. 
Like Bradya minor, this EHctinosoma is of a brownish colour. 
It has been observed in the stomachs of young flat-fish (Pleuro- 
nectes limanda) caught in Liverpool Bay. 


Ecrinosoma pyamaum, 7. & A. Scott. 

1896. Ectinosoma pygmeum, T. & A. Scott (84), p. 433, pl. 36. 
figs. 15, 41; pl. 37. figs. 5, 20, 39, 43; pl. 38. figs. 4, 26, 31, 55. 

This was also dredged off Hast Glacier, Cape Flora, in 30 
fathoms, July 21st, 1898. H. pygmeum is very small: it is even 
shorter, but somewhat stouter than H. atlanticum. Distribution: 
Firth of Forth, Scotland; vicinity of Port Erin, Isle of Man. 


EcriInosoMA MELANICEPS, Boeck. 

1864. Ectinosoma melaniceps, Boeck (5), p. 30. 

The colour of the head in this species is, as the name implies, 
distinctly different from the rest of the body, so that the species 
may be distinguished by this character alone. H. melaniceps 
occurred rather more frequently than others of the same genus. 
It was dredged at about 50 yards off West Point, Cape Flora, 
in 2 to 3 fathoms, June 18th, 1897; at West Bay, in 2 to 10 
fathoms, July 2nd, 1897; and off Hast Glacier, in 30 fathoms, 
July 21st, 1897. It is also of frequent occurrence around the 
shores of the British Islands. 


Eerrnosoma Normant, 7. f& A. Scott. 
1896. Eetinosoma Normant, T. & A. Scott (84), p. 435, pl. 36. figs. 21, 
29, &e. 


This was dredged off East Glacier, Cape Flora, July 21st, 


S6 MR. THOMAS SCOTT ON THE 


1897. EE. normani was one of the rarer species in the Franz- 
Josef Land collections. Its distribution includes the Firths of 
Forth and Clyde, Scotland ; and Barrow Channel, near Barrow- 
in-Furness, England. 


Ecrrnosoma attanticum (Brady 5 Robertson). 

1873. Microsetella atlantica, Brady & Robertson (20), p. 180, pl. 9. 
figs. 11-16. 

The small size and slender form of this species make it easily 
overlooked. It was very scarce in the Franz-Josef Land collec- 
tions, having only been observed in a gathering off Hast Glacier 
made in July 1897. This species kas a wide distribution, and is 
not unfrequent in the British seas. 


Genus Zostme, Boeck, 1872. 


ZOSIME TYPICA, Boeck. 

1872. Zosime typica, Boeck (7), p. 14. 

This was obtained at Cape Gertrude and also off Hast Glacier, 
Cape Flora, but was apparently not very common. It is also a 
British species. 


Genus Ropertsonta, Brady, 1880. 


Ropertsonia TENUIS (Brady & Robertson). 

1873. Ectinosoma tenue, Brady & Robertson (22), p. 196. 

This species was dredged at West Bay, Cape Flora, in from 
2 to 10 fathoms, July 2nd, 1897. Robertsonia somewhat re- 
sembles Kctinosoma, and was at first ascribed to that genus. 
It has been obtained at various places around the British coasts. 


Genus Amymone, Claus, 1863. 


AMYMONE SPHERICA, Claus. 

1863. Amymone spherica, Claus (23), p. 114, pl. 20. figs, 1-9. 

The Copepod so named is one of a very curious group of 
crustaceans, and quite unlike the usual Copepod forms. It 
appeared to be very rare in the Franz-Josef Land collections. 
The only gathering in which this species was obtained was dredged 
off Flagstaff Point, Elmwood, by D. W. Wilton, 20th September, 
1896. This is also a British species. 


CRUSTACEA OF FRANZ-JOSEF LAND. 97 


Genus STENHELIA, Boeck, 1864... 


STENHELIA REFLEXA, 7. Scott. 

1895. Stenhel:a refleva, T. Scott (80), p. 166, pl. 3. figs. 1-9. 

This was dredged off East Glacier, Cape Flora, on July 21st, 
1897. Stenhelia reflexa is described and figured in Part in. of 
the 18th Annual Report of the Fishery Board for Scotland 
(1895), from specimens obta:ned in. the Firth of Forth. 


Genus AmeEtra, Boeck, 1864. 


AMEIRA LONGIPES, Boeck. 

1864. Ameira lonyipes, Boeck (5), p. 49. 

This was obtained in a gathering dredged at West Bay, Cape 
Flora, on July 5th, 1897; it was also obtained off Hast Glacier 
on the 21st of the same month, but it appeared to be somewhat 
rare in these gatherings. It is not a very rare species in the 
British seas. 


Ametra Exraua, 7. Scott. 

1894. Ameira exigua, T. Scott (79), p. 243, pl. 6. figs. 15-23. 

This is a much smaller species than the last, and it was also of 
rare occurrence. It was obtained sparingly in the gathering 
dredged off East Glacier on July 21st, 1897. This species is 
described and figured in Part iii. of the 12th Annual Report of 
the Fishery Board for Scotland (1894). 


AMEIRA LONGIREMIS, 7. Scott. 

1894. Ametra longiremis, T. Scott (79), p. 241, pl. 5. figs. 29-32, 
pl. 6. figs. 1-5. 

This Ameira was also dredged off Hast Glacier; this was the 
only gathering in which it was observed. The species is described 
and figured in Part i. of the same Fishery Board’s Report ir. 
which Ameira exigua and Ameira reflexa are described. 

AMEIRA REFLEXA, 7. Scott. 

1894, Ameitra reflera, T. Scott (79), p. 240, pl. 5. figs. 20-28. 

Ameira refleca was obtained iu the same gathering with the 
last species, and appeared to be rare. 


Genus Jonestetia, Brady, 1890. 


JONESIELLA SPINULOSA (Brady & Robertson). 
1875. Zosima spinulosa, Brady & Robertson (22), p. 196. 
This Jonesiella was rather more frequent than some of the 


93a MR. THOMAS SCOTT ON THE 


other species ; it was dredged off Hast Glacier, off Cape Gertrude 
in 80 fathoms, and in 8 fathoms off Cape Flora, in July 1897. 
This is not an uncommon species in the British seas. 


Genus Detavatta, Brady, 1868. 


Detavatta roBusta, Brady & Robertson. 

1875. Delavalia robusta, Brady & Robertson (22), p. 196. 

This species was dredged off East Glacier in 30 fathoms and 
in 2 to 4 fathoms off West Point, Cape Flora, in July 1897. 
Several specimens were obtained. 


Detavatta mimica, 7. Scott. 

1897. Delavalia mimica, T. Scott (82), p. 150, pl. 1. figs. 1-9. 

A number of specimens of this distinct species were obtained 
in gatherings dredged off West Point, July 5th, and off Hast 
Glacier, July 21st, 1897. The species is described and figured 

Part ii. of the 15th Annual Report of the Fishery Board for 
Scotland (1897), from specimens obtained in the Firth of Clyde ; 
it has also been observed in the Firth of Forth. 


DELAVALIA REFLEXA, Brady & Robertson. 

1875. Delavalia refleca, Brady & Robertson (22), p. 196. 

A few specimens of Delavalia reflexa were obtained in a 
gathering collected off Hast Glacier. In this species the inner 
branches of the first .pair of swimming-feet want the stout 
spiniform terminal seta that distinguishes D. robusta; in 
D. reflexa the terminal sete are slender. 


DELAVALIA arctica, sp.n. (PI. 5. fig. 14; Pl. 6. figs. 7-11.) 

The female specimen represented by the figure (fig. 7, Pl. 6) 
measured fully 1 mm. (54 of an inch) in length from the end of 
the rostrum to the extremity of the caudal furea. The species 
somewhat resembles Delavalia palustris, Brady, in general outline. 
The antennules (anterior antenne) are 8-jointed (fig. 8, Pl. 6); 
the proportional lengths of the joints are indicated approxi- 
mately by the formula: 


Proportional lengths of the joints ............ 20). 12). 118 728.6 28 
INOUTMORTES OSE Was TOMAS” Goeconponnsooenontonenoeded Lea o aG as 


The mouth-organs are somewhat similar to those of Delavalia 
guesbrechti, T. Scott, except that the second foot-jaws are com- 
paratively robust; the first joint bears one slender and two stout 
spiniform set at the extremity of the inner margin; the terminal 


CRUSTACEA OF FRANZ-JOSEF LAND. 99 


claws are feeble (Pl. 5. fig. 14). The first pair of swimming-feet 
resemble those of Delavalia palustris, but both branches are more 
elongate and the lengths of the joints are proportionally slightiy 
different; they differ also in their armature, as shown by the 
figure (fig. 9, Pl. 6). The other swimming-feet are also moderately 
elongate and slender; the joints of the inner branches, as shown by 
the figure of the fourth pair (fig. 10, P1. 6), have their inner distal 
angles more or less produced downwards into spine-like processes 
as in Delavalia robusta, Brady & Robertson. The fifth pair also 
somewhat resemble those of that species, but the secondary 
joint is proportionally broader and of a somewhat different form ; 
and there is a peculiarity in the hinge arrangement of the joint 
by which it can be extended at nearly right angles to the body 
(fig. 11, Pl. 6). The caudal furca are about as long as the com- 
bined lengths of the last two abdominal segments. 

Hab. Cape Gertrude (Cape Flora), Northbrook Island; rather 
rare. 

Kemarks. This species partakes of the characters of Delavalia 
palustris on the one hand and of Delavalia robusta on the other ; 
but it differs from the first by the form of the fifth pair of 
thoracic feet and from the second in the structure of the first 
pair. A difference may also be observed in the proportional 
lengths of the joints of the antennules, as well as in the more 
robust form of the posterior foot-jaws. 


Genus Mararnosrotus, Mrazek, 1893. 


Mararnosiorus Vrespovsxyi, Mrazek. (PI. 6. figs. 12-17.) 

1893. Maraenobiotus Vejdouskyi, Mrazek (5%), p. 103, pl. 4. figs. 17-32, 
pl. 5, figs. 33-87. 

This is a freshwater species ; it was first discovered by Mrazek 
in Bohemia, and it has also occurred in one or two places in 
Scotland. It is a slender Copepod, and in this respect resembles 
certain species of Moraria—a genus with which it is closely 
related. One of the principal characters that distinguishes 
Maraenobiotus from Moraria is the very rudimentary form of the 
mandible-palp (fig. 14) ; whereas in Moraria the mandible-palp, 
though small, is normal in structure. The Franz-Josef Land 
specimens resemble those found in Scotland in almost every 
detail of structure: the chief difference observed is in the form 
of the secondary joints of the fifth thoracic feet ; in the Franz- 


100 MR. TITOMAS SCOTT ON THE 


Josef Land specimens this joint is subquadrangular, with straight 
margins, as shown in fig. 17. The species is very small—the 
average length of the specimens is about ‘56 mm. to “6 mm. 
qs to ,, of aninch). The species was moderately frequent in 
freshwater pools near Elmwood, Cape Flora. A few of the 
females carried ova, but a considerable proportion of the speci- 
mens were more or less immature. 

After the above remarks on Maraenobiotus had been prepared, 
I received from the author, Dr. Jules Richard, an interesting 
contribution to the literature of the Arctic freshwater fauna, being 
a report on the Entomostraca obtained in the freshwaters 
explored during the recent voyage to the Arctic seas of the 
steam yacht ‘ Princesse Alice. The places visited comprised 
Lofoten, Spitzbergen, Iles Beeren, Hope, de Barents, and 
Faroe. In this memoir Dr. Richard describes under the name 
of Mesochra Brucei an harpactid which appears to resemble 
the Franz-Josef Jand form referred to above; it may not, 
however, be the same species. 

With regard to the specimens from Franz-Josef Land, the 
rudimentary form of the mandible-palp, together with the 
structure of the first and fifth pairs of thoracic feet, identify 
them with Mrazek’s Maraenobiotus; there may be slight 
differences in the armature of the swimming-feet, but such 
differences are unimportant in view of the close similarity 
otherwise. 


Genus CantHocamptus, Westwood, 1835. 


(?) Canrnocamprus parvus, T. f A. Scott. (PI. 6. figs. 18-24.) 

1896. (?) Canthocamptus parvus, T.5-A. Scott (85), p. 6, pl. 2. figs. 14-22. 

This species, which is represented in the Franz-Josef Land 
collection by several specimens, was dredged off East Glacier, 
Cape Flora, in 80 fathoms, July 21st, 1897. (?) Canthocamptus 
parvus is a small species that was first described from specimens 
obtained in the Firth of Forth near Aberlady. The Franz-Josef 
Land specimens are somewhat larger than those from the Firth 
of Forth; there are also one or two other differences, but they 
are comparatively unimportant. The following is a brief descrip- 
tion of the Arctic specimens :— 

The antennules in the female are short and six-jointed (fig. 19). 
The antenne have the secondary branches small and two-jointed 
(fig. 20). The mandible-palp is moderately well developed and 


CRUSTACEA OF FRANZ-JOSEF LAND. 101 


bears a one-jointed branch which is subapical (fig. 21). A 
moderately long plumose seta springs from the basal joint of 
the palp a short distance below the one-jointed branch. The 
posterior foot-jaws are moderately stout, and are each furnished 
with an elongate and slender terminal claw. ‘The first pair of 
swimming-feet somewhat resemble those of Dactylopus longi- 
vostris, Claus, but are rather more slender (fig. 22); a small 
seta springs from about the middle of the inner margin of the 
first jomt of the inner branches, while the outer margin is 
fringed with minute hairs; the inner branches are also armed 
with a moderately stout terminal spine and an elongate slender 
seta; the outer branches, which are rather more than half the 
length of the inner, are composed of three subequal joints. The 
inner branches of the second, third, and fourth pairs are 
two-joited ; in the fourth pair (fig. 28) the two-jointed inner 
branches are short, but the outer, which are three-jointed, are 
elongate. The fifth pair (fig. 24.) are somewhat similar to those 
of Dactylopus minutus, Claus. The caudal furca are very 
short. 

Remarks. This species resembles very closely one of the smaller 
species of Dactylopus both in its general outline and in its six- 
Jomted antennules; but it is precluded from that genus by the 
structure of the mandible-palp and by the inner branches of 
the second, third, and fourth pairs of swimming-feet being only 
two-jointed, and in these respects it agrees more closely with 
Canthocamptus than with any other described genus. In the 
original description of the species, reference is made to one or 
two points in which the species does not agree with Ouantho- 
camptus, and which may by-and-by render its removal from that 
genus necessary. J am inclined, however, for the present to 
leave it as described. The length of the specimen figured 
(fig. 18) is 48 mm. (;'5 of aninch). The Firth of Forti is the 
only British habitat of the species known to me. 


Genus Laopnonte, Philippi, 1849, 


LAOPHONTE HORRIDA, Norman. 

1876. Laophonte horridu, Norman (60), p. 206. 

This well-marked form was dredged off East Glacier in 
30 fathoms, and also off West Point in 2 to 3 fathoms; both 
localities being in the vicinity of Cape Flora, Northbrook Island. 
lt was dredged at West Point on June 4th and on July 2nd and 


102 MR. THOMAS SCOTT ON THE 


21st,1897. Laophonte horrida is recorded from the Arctic seas by 
Buchholz, in his Report on the German North-Polar Expedition 
of 1869-70, but he, under some misapprehension, referred this 
Copepod to Miller’s Cyclops minuticornis. The species appears 
to be generally distributed round the British coasts. 


LAoPHONTE CURTICAUDA, Boeck. 

1864. Laophonte curticauda, Boeck (5), p. 55. 

This species was somewhat rare in the Franz-Josef Land 
collection; a few specimens were obtained in some material 
dredged off Cape Gertrude. L. eurticauda is also a British 
species. 


LaorHonte DEPRESSA, 7’. Scott. 

1894. Laophonte depressa, T. Scott (79), p. 245, pl. 6. figs. 24-31, pl. 7. 
figs. 1-3. 

This species was described and figured in Part iii. of the 
Twelfth Annual Report of the Fishery Board for Scotland 
(1894), from specimens found in the Firth of Forth. The Franz- 
Josef Land gathering in which it occurred was dredged off Hast 
Glacier, Cape Flora, July 21st, 1897. 


LAOPHONTE LONGICAUDATA, Boeck. 

1864. Laophonte longicaudata, Boeck (5), p. 55. 

This also was dredged off Hast Glacier, Cape Flora, and 
appeared to be somewhat rare. It has been long known as a 
British species. The outer branches of the first pair of 
swimming-feet in all the Franz-Josef Land specimens appear 
to be two-jointed, the first joint being short and the other about 
twice as long as the first. 


LaopHonre INTERMEDIA, J. Scott. 

1895. Laophonte intermedia, T. Scott (80), p. 168, pl. 3. figs. 10-20. 

This distinet species was obtained in West Bay on July 27th, 
1897, it occurred in a tow-net gathermg. L. intermedia was 
described and figured in Part i. of the Thirteenth Annual 
Report of the Fishery Board for Scotland (1895), from specimens 
found in the Firth of Forth. 


LAOPHONTE sIMILts, Claus. 

1866. Laophonte similis, Claus (24), p. 23, pl. 5. figs. 15, 14. 

One or two specimens of a Copepod which I have ascribed to 
Laophonte similis occurred in some dredged material from West 
Bay, Cape Flora, collected in July 1897. 


CRUSTACEA OF FRANZ-JOSEF LAND. 103 


LAOPHONTE PERPLEXA, sp.n. (PI. 7. figs. 1-7.) 

Several specimens of a Laophonte that appear to be undescribed 
were obtained in a gathering of small Crustacea collected off 
Hast Glacier, Cape Flora, in 30 fathoms, on July 21st, 1897. Its 
characters are somewhat intermediate between those of Laophonte 
curticauda and Laophonte similis. It is rather smaller than 
either of these species, being only about °63 mm. ({4 of an inch) 
in length. The body is in general appearance lke that of 
L. curticauda. The rostrum is short, and the antennules are 
seven-jointed; the fourth and fifth Joints of the antennules 
are shorter than the others, as shown by the formula, which 
indicates approximately the proportional lengths of all the 
joints :— 

Proportional lengths of the joints ............... Wee AD o UG 4D Be de ia 
gee Nuimibersioret hey OMMtsseeenereaesteeceeereeecrcse crc DS} bs NG, 7 


The posterior foot-jaws (fig. 3) resemble very closely those of 
L. similis except that the terminal claw is somewhat stronger. 
The first pair of swimming-feet (fig. 4) are intermediate in 
structure between those of L. similis and L. thoracica; the outer 
branches are short and two-jointed, the last joint being about 
twice the length of the first, the terminal claws of the inner 
branches are moderately stout. The fourth pair (fig. 5) have 
the outer branches elongate and slender, and the inner branches 
short and composed of two subequal joints. The fifth pair 
(fig. 6) are nearly like those of ZL. curticauda, but the secondary 
joints are proportionally rather smaller, and the armature of 
both joints is somewhat different. The caudal furea (fig. 7) 
resemble those of ZL. similis, beng rather longer than the last 
abdominal segment. 

Remarks. The Laophonte referred to above is one of those 
perplexing forms, met with now and again, which are somewhat 
difficult to dispose of satisfactorily. Its characters are such 
that it might be considered a variety of Laophonte curticauda 
as well as of L. similis ; in these circumstances it appeared to me 
_ to be better to give the form a distinct name. 


Genus LaopHontopgs, 7. Scott, 1894. 


Laoprnontopes trpicus, 7. Scott. 
1894. Laophontodes typicus, T. Scott (79), p. 249, pl. 8. figs. 2-8. 
This genus and species were described and figured in 1894 in 


104 MR. THOMAS SCOTT ON THE 


Part iii. of the Twelfth Annual Report of the Fishery Board 
for Scotland, from specimens obtained in the Firth of Forth. 
Its occurrence at Franz-Josef Land shows that it is widely 
distributed. It was obtained off Hast Glacier, Cape Flora, on 
July 21st, 1897. This is a very small Copepod, being only 
about ‘4 mm. (;'5 of an inch) in length. 


Genus CreroveEs, Brady, 1872. 


‘\LETODES stMintis, 7. Scott. 

1895. Cletodes similis, T. Scott (80), p. 168, pl. 3. figs. 22-26, pl. 4. 
figs. 1-3. 

This Copepod was dredged off Cape Gertrude in 30 fathoms ; 
it was also obtained off Hast Glacier. The species was described 
in 1895, in Part ii. of the Thirteenth Annual Report of the 
Fishery Board for Scotland, from specimens found in the Firth 
of Forth. 


CLETODES TENUIPES, 7. Scott. 

1897. Cletodes tenuipes, T. Scott (82), p. 170, pl. 1. figs. 19-27. 

C. tenuipes was also dredged off East Glacier, but very few 
specimens were obtained. It was described and figured from 
Clyde specimens in Part ill. of the Fifteenth Annual Report of 
the Fishery Board for Scotland (1897). 


CLETODES LonerIcauDATA, Brady § Robertson. 

1872. Cletodes longicaudata, Brady & Robertson (19), p. 196. 

This was also obtained off East Glacier, and appeared to be 
moderately rare. This species has long caudal furca. 


Genus Ennyprosoma, Boeck, 1872. 
EyHyDROSOMA cURVATUM (Brady § Robertson). (PI. 3. fig. 17.) 
1875. Rhizothrix curvata, Brady & Robertson (22), p. 197. 

A few specimens of what appears to be this species were 
obtained in some material dredged off Hast Glacier. There was 
a slight difference in the armature of the fifth pair of thoracic 
feet (as shown by fig. 17, Pl. 3), but otherwise the Franz- 
Josef Land specimens appeared to be identical with the species 
to which they are ascribed. 


Genus Dactytorus, Claus, 1863. 


DacTYLOPus TISBOIDES, Claus. 
1863. Dactylopus tisboides, Claus (23), p. 127, pl. 16. figs. 24-28. 
This, which is a common British species, was moderately 


CRUSTACEA OF FRANZ-JOSEF LAND. 105 


frequent in one or two of the Franz-Josef Land gatherings, 
7. e. in a gathering dredged about fifty yards off West Point, 
Cape Flora, in 2 to 3 fathoms, on June 18th, 1897; in another 
from West Bay, dredged in about 8 fathoms on July 5th, and 
in a third collected on the 23rd or 24th of the same month. In 
some of the specimens there is a tendency for both branches of 
the fifth thoracic feet to have pellucid markings along the edge 
of the outer margin. 


DacryLorus coronatus, 7. Scott. 

1894. Dactylopus coronatus, T. Scott (79), p. 255, pl. 9. figs 12-20, 

This species was obtained amongst sand near Hast Glacicr, 
Cape Flora, August 5th, 1896; it appeared to be rare. D. coro- 
natus was described and figured in Part iil. of the Twelfth 
Annual Report of the Fishery Board for Scotland (1894). 


DAcTYLOPUS TENUIREMIS, Brady § Robertson. (PI. 8. figs. 1-4.) 

1875. Dactylopus tenuwemis, Brady & Robertson (22), p. 197. 

Several specimens of what seems undoubtedly to be this 
species were obtained in a gathering of small Crustacea collected 
cif West Point, Cape Flora, in from 2 to 4 fathoms of water, on 
July 21st, 1897. D. tenuiremis is closely related to the next 
species, but differs from it in the form of the fifth thoracie feet, 
and in the structure of the first pair, as shown by the figures. 


DactyLopvus Lonairostris, Claus. (Pl. 3. figs. 5-S.) 

1863. Dactylopus longirostris, Claus (23), p. 127, pl. 17. figs. 4-6. 

This is one of the species described by Dr. Claus in his ‘ Die 
freilebenden Copepoden’ (pub. 1863), which were obtained in 
the vicinity of Heligoland. It resembles D. tenuwiremis in some 
respects, but the form of the fifth thoracic and especially of the 
secondary joints is distinctly different ; these outer (secondary) 
joints are in this species broadly oval and leaf-like (‘‘ das anssere 
ovale Blatt’), the proportion of the breadth to the length being 
nearly as 20 is to 29, whereas in D. tenuiremis it is nearly as 
15 is to 28; there is also a difference in the general outline as 
well as in the armature of both the basal and secondary joints 
of the fifth pair. It may be noted further that, besides the 
difference in the first swimming-feet already referred to, the 
structure of the posterior foot-jaws differs slightly in the two 
species, as indicated by the figures. D. longirostris occurred ina 
gathering from the vicinity of East Glacier, Cape Flora; only a 


106 MR. THOMAS SCOTT ON THE 


few specimens were obtained. This species, and probably also 
D. tenuiremis, should be regarded as belonging to the genus 
Diosaccus, as both appear to be furnished with two ovisacs. 


Dactrytorus Striémit (Baird), var. ARCTICUS, var. nov. (PI. 5. 
figs. 11-17.) 

1850. Canthocanptu Strémii, Baird (3), p. 208, pl. 27. fig. 3. 

A number of specimens of a Copepod, which can hardly be 
distinguished from Dactylopus Strémii (Baird), were collected 
about 50 yards off West Point, Cape Flora, during June and July, 
1897. The specimens are from comparatively shallow water— 
2 to 4 fathoms: and differ from British species of D. Strémiz 
chiefly in the following points :—(1) The antennules have nine 
instead of eight joints (fig. 12); this difference, however, appears 
to be immaterial, as the number of the joints of which the distal 
half of the antennules is composed seems to be liable to variation. 
(2) The posterior foot-jaws (fig. 18) are large and powerful, the 
second joint is elongate and subcylindrical instead of ovate, and 
they differ somewhat in their armature. (8) In the fifth pair 
of thoracic feet the outline of the secondary joint (fig. 15) and 
the arrangement of the sete with which it is furnished are 
somewhat dissimilar to the normal form of the species; and 
(4) the ova-bearing females carried two ovisacs instead of one. 
The first pair of swimming-feet (fig. 14), as well as the second, 
third, and fourth pairs, resemble very closely the same appendages 
in D. Stromit. The species bas a wide distribution in the North 
Sea, and seems to extend all round the British Islands. 


Genus THALEsTRIs, Claus, 1863. 


THALESTRIS HELGOLANDICA, Claus. 

1863. Thalestris helgolandica, Claus (23), p. 131, pl. 17. figs. 12-21. 

A few specimens of this Thalestris were obtained in a gathering 
frcm 30 fathoms, collected July 21st, 1897, off East Glacier, 
Cape Flora. Thalestris helgolandica is a well-marked species, 
and, though apparently not very common, it has evidently a wide 
distribution. It occurs sparingly at various places around the 
British Islands. 


THALESTRIS POLARIS, sp.n. (PI. 7. figs. 8-16.) 

A Thalestris, which I have named as above, occurred very 
sparingly in gatherings collected at the following places :—From 
sand near Hast Glacier on August 5th, 1896; off West Point, 


CRUSTACEA OF FRANZ-JOSEF LAND. 107 


Cape Flora, on June 4th, 21st, and July 5th, 1897; and also in 
July in West Bay, Cape Flora. The following is a description 
of the species :— 

Description of the female.—Body robust, especially the cephalo- 
thorax ; rostrum very short ; entire length from rostrum to caudal 
furca about °95 mm. (54 of an inch). Antennules (fig. 9) short, 
9-jointed ; the first two joints of moderate length and subequal, 
the next two shorter and also subequal; the second joint of the 
flagellum (the fourth from the end) is equal to twice the length 
of the preceding joint and to the combined lengths of the next 
two, the end joint is slightly longer than the penultimate one ; 
the formula shows approximately the proportional lengths of all 
the jeints :— 

Proportional lengths of the joints ...... 23..25.15.16.5.10.4. 

7 


4.6 
Numbers of joints ....0.....ccscesccsessesees IF) eo OO 8 9 


The posterior foot-jaws (fig. 10) resemble those of Thalestris 
hibernica, Brady & Robertson, both in their form and armature ; 
the other mouth-organs are somewhat similar to those of 
Thalestris mysis, Claus. First pair of swimming-feet moderately 
slender (fig. 11); terminal claws of both branches slender, and 
not much more than half the length of the branches from which 
they spring. Fourth pair (fig. 12) also slender; the inner 
branches reach to about the end of the second joint of the outer 
branches, and both are furnished on the inner margins with 
long plumose sete. The fifth pair somewhat resemble those of 
Thalestris hibernica, but the secondary joints are comparatively 
rather larger and extend as far as the end of the basal joints ;. 
the basal joint bears five apical sete, while the surface of both 
it and the secondary joint appears to be more covered with 
extremely fine cilia (fig. 18). The caudal furca (fig. 16) are 
elongate, the length being equal to fully twice the breadth. 
Description of the male—The male differs little from the 
female except that the antennules are modified for grasping. 
The inner branches of the second pair of swimming-feet (fig. 14) 
are two-jointed, and somewhat similar in their structure and 
armature to the inner branches of the same pair of feet in the 
male of Thalestris hibernica. The secondary joints of the fifth 
pair (fig. 15) are elongate-ovate; the inner margin is nearly 
straight and fringed with minute hairs ; the outer margin, which 
is slightly curved and tapers gradually towards the apex, bears. 
LINN. JOURN.—ZOOLOGY, VOL. XXVII. 9 


108 MR. THOMAS SCOTT ON THE 


three moderately short and stout sete, there is also a stout and 
moderately long apical seta and a smaller one at the termination 
of the inner margin; the basal joimt is scarcely produced 
interiorly, and is provided with three spiniform sete on the 
broadly rounded apex, the middle seta being considerably longer 
than the other two and plumose. 

Remarks.This species has a superficial resemblance to 
Thalestris hibernica, Brady & Robertson, and I was at first 
inclined to regard it as a northern variety of that form. I had 
recently, however, the opportunity to compare typical Scottish 
examples of Thalestris hibernica with the Franz-Josef Land 
specimens, and find that they are quite distinct, the difference 
in the length of the caudal furca being alone sufficient to 
distinguish the one from the other. 


THALESTRIS FORFICULA, Claus. 

1863. Thalestris forficula, Claus (23), p. 151, pl. 17. figs. 7-11. 

The Franz-Josef Land Copepod which I now record under 
this name is similar to a form described and figured in the 
‘Annals and Magazine of Natural History ’ for October 1893, 
by T. and A. Scott, under the name of Thalestris forficuloides. 
T am now inclined to consider that species as a ‘ form’ of Claus’s 
Th. forficula. Its occurrence at Franz-Josef Land extends con- 
siderably the distribution of the species. It was obtained in a 
eathering dredged off Cape Gertrude in 30 fathoms, July 21st, 
1897. In Scotland it has been obtained both in the Firths of 
Forth and Clyde. 


THALESTRIS FRIGIDA, sp.n. (Pl. 7. figs. 17-23; Pl. 8. figs. 
il 2) 

Description of the female.—Body robust, length 1°63 mm. 
zs of an inch). Rostrum prominent. Antennules short, 
moderately stout and 9-jointed; the sixth and last joints are 
each of them about equal to the combined lengths of the seventh 
and eighth (fig. 18); the proportional lengths of all the joints 
_ are shown approximately by the formula :— | 


Proportional lengths of the joints ... 48 .24.24.20.12.16.7.8.16 
Numbers of the joints .................. OPQ SAL TG eto 


The mouth-organs are somewhat similar to those of Thalestris 
mysis, Claus, except that there is a slight difference in the 


CRUSTACEA OF FRANZ=JOSEF LAND. 109 


armature of the posterior foot-jaws (fig. 19). The first pair of 
thoracic feet are moderately slender; the seta on the inner 
margin of the first jomt of the inner branches springs from near 
the middle of the joint; both branches are furnished with a 
strong terminal claw of moderate length, and the outer margins 
of the first and second joints of the outer branches, and of the 
first joint of the inner branches, are fringed with minute spines. 
The second, third, and fourth pairs are elongate and slender, 
and have both branches furnished with long plumose sete 
(fig. 21). The fifth pair are large and broadly foliaceous, and 
somewhat resemble those of Thalestris mysis, but the secondary 
joint is considerably smaller than the basal joint, and the 
armature of the secondary joint is also distinctly different ; 
moreover, both joints, besides being different in general outline, 
have their surface ornamented with what appears to be numerous 
minute papille (fig. 22). The caudal furca are short (Pl. 8. 
fig. 2) and about equal to the length of the last abdominal 
segment, and they are about as broad as long. 

Description of the male.—The male is very similar to the female 
except that the antennules have a modified and hinged structure, 
to permit of their use as grasping-organs. The inner branches 
of the second pair of thoracic feet are also modified as shown 
(Pl. 7. fig. 23). The fifth pair (Pl. 8. fig. 1) are much smaller 
than in the female; the inner portion of the basal joint, which is 
only slightly produced, is broadly rounded, and provided with 
three stout sete of unequal length—the middle one being the 
‘longest; the secondary joint is broadly ovate, the inner margin 
is furnished with only a few minute hairs, but several stout 
plumose setz spring from the outer margin and apex. 

Hab. Off Hast Glacier, Cape Flora, July 1897; only a few 
specimens were obtained in the Collection. 

Remarks. The large size and robust form of this Copepod, 
together with the large and broadly foliaceous fifth pair of feet 
-of the female, differentiated the species at once from all the 
others in the Collection. 


THALESTRIS JACKSONI, sp.u. (PI. 8. figs. 3-9.) 

Description of the female.—Body moderately stout. Length 
of the specimen figured 2°5 mm. (;/y of an inch). Rostrum very 
short. Antennules short, 9-jointed, the sixth joint is consider- 
ably longer than any of the other five end-joints (fig. 4); the 

9* 


110. MR. THOMAS SCOTT ON THE 


approximate proportional lengths of all the joints are shown by 
the formula :— 


Proportional lengths of the joints ... 17.21.15.13.6.10.5.4.6 
Numbers of the joints ...............+6 1 52) VS) 4 ib iG aS 


le) 


€ 


The posterior foot-jaws (fig. 5) are stout; the hand somewhat 
resembles that of Thalestris rufoviolascens, but the inner margin 
is more oblique and the marginal spinules are larger; the other 
mouth-organs are somewhat similar to those of Thalestris mysis. 
The first pair of thoracic feet are moderately stout, and both. 
branches are armed with strong terminal claws (fig. 6). In the 
fourth pair the inner branches only reach to about the end of 
the second joint of the outer branches; both branches bear long 
plumose sete (fig. 7). The fifth pair are large and foliaceous: 
the length of the basal joint is scarcely equal to twice the 
breadth ; this joint bears five sete on the broadly rounded apex, 
two of the intermediate (fig. 8) are considerably longer than the 
others : the secondary joint is oval in outline, its greatest breadth 
is equal to about half the length, and both the mner and outer 
margins are fringed with minute cilia; this joint is furnished 
with six sete, three on the lower half of the outer margin and 
three at the apex,—the two outer apical sete are close together 
and more slender than the others, which are moderately wide 
apart as shown in the figure. The caudal furca (fig. 9) are 
elongate, being fully twice the length of the last abdominal 
segment, and the two principal sete are as long as the whole 
length of the abdomen and furca combined. No males of this 
species were observed. 

Hab. Half mile off Cape Gertrude in 8 fathoms, June 6th, 
1897; about fifty yards off West Point, Cape Flora, in 2-8 
fathoms, June 20th; and off West Glacier, in 1-3 fathoms, 
July 6th, 1897. 

The species was of rare occurrence in these gatherings. 

Remarks. The large size of Thalestris Jacksoni, the peculiar 
form of the posterior foot-jaws, and the form and armature of the 
large foliaceous fifth pair of thoracic feet, combine to distinguish 
this from any other Thalestris known to me. ‘This fine species 
is named in compliment to Mr. Jackson, of the Jackson-Harms- 
worth Arctic Expedition. 


CRUSTACEA OF FRANZ-JOSEF LAND. lil 


Genus Westwoopta, Dana, 1855. 


WEStWwoopia NOBILIS (Baird). 

1845. Arpacticus nobilis, Baird (2), p. 155. 

This species was obtained in a gathering collected in West 
Bay, Cape Flora, July 27th, 1897; a single specimen only was 
observed. Westwoodia nobilis is found sparingly in many places 
around the British Islands, and appears to be otherwise widely 
distributed. It is a distinct species, and when living very 
prettily coloured. 


Genus Harpacticus, Wilne-Edwards, 18388. 


Harpacricus cHEeirer (Miller). (PI. 8. figs. 10-13.) 

1776. Cyclops chelifer, O. F. Muller (55), p. 2415. 

This Harpacticus, which is moderately common in the British 
geas, was of rare occurrence in the Franz-Josef Land collection ; 
the only gathering in which the species was observed was one 
from West Bay, Cape Flora, collected in July 1897. 

It will be observed from the figures on Plate 8, that the Franz- 
Josef Land specimens of Harpacticus chelifer differ somewhat 
from those from Heligoland and the British seas ; the antennules 
of these Arctic specimens (fig. 10) are 9-jointed, whereas in 
Dr. Claus’s description of the species they are stated to be 
8-jointed (achtgliedrig), and this agrees with what I have observed 
in Clyde specimens. The posterior foot-jaws (fig. 11) are 
scarcely so angular on the inner aspect as they are found to be 
in Clyde specimens. There does not appear to be much differ- 
ence in the structure of the first pair of swimming-feet (fig. 12), 
In the fifth pair the secondary joints are proportionally smaller 
than in Clyde specimens, and the inner produced part of the 
basal joints is more broadly rounded. Notwithstanding these 
differences, it seems better to regard this simply as a form of 
Harpacticus chelifer. 


HARPACTICUS CHELIFER, var. ARCTICUS. (PI. 8. figs. 14-17.) 


An Harpacticus, of a more robust form than the last, was of 
frequent occurrence in sowe of the gatherings. I was inclined 
at first to regard this as belonging to Harpacticus gracilis, 
Claus, which it in some respects resembles, but I now prefer to 
Jook on it as a robust variety of Harpacticus chelifer. The 


112 MR. THOMAS SCOTT ON THE 


antennules are 9-jointed ; the first four joints are proportionally 
rather shorter than those of H. gracilis. The posterior foot-jaws 
are larger than those of the form last described, but otherwise 
they resemble them very closely. The first pair of swimming- 
feet are also larger than those of the form referred to, and there 
is a slight difference in their structure: the inner branches 
have the two short end-joints distinct, but the short end-joint 
of the outer branches appears to be coalescent with the second, 
so that the outer branches are thus apparently only 2-jomted. 
The fifth pair (fig. 17) have the secondary joints broadly ovate, 
and somewhat resemble those of Harpacticus gracilis. 

This robust variety occurred chiefly in gatherings of Crustacea 
collected in 2-4 fathoms, about fifty yards off West Point, Cape 
Flora, during June and July, 1897. 


Genus Zavs, Goodsir, 1845. 


ZAUS SPINATUS, Goodsir. 

1845. Zaus spinatus, Goodsir (33 a), p. 326, pl. 11. figs. 1-8. 

This comparatively well-marked species was dredged at West 
Bay, Cape Flora, in 2-10 fathoms, July 2nd, 1897; only a few 
specimens were observed. Its British distribution extends from 
the Scilly Islands to Shetland. 


Genus Evretter, Claus, 1863. 


EvUPELTE PURPUROCINCTA (Worman). 

1868. Alteutha purpurocincta, Norman (59), p. 298. 

This species occurred in the same gathering with Zaus spinatus 
from West Bay, Cape Flora. It has sometimes been described 
as an Alteutha, but Prof. Claus and others consider it to be 
generically distinct. As a British species, it is to be found all 
round our shores, but seldom in large numbers. Ewupelte pur- 
purocincta appeared to be rare in the Franz-Josef Land 
gatherings. 


Genus Ipya, Philippi, 1848. 


Ipya FoRcaTa (Baird). 
1837, Cyclops furcatus, Baird (1 a), p. 330, pl. 9. fies. 26-28. 
A considerable number of specimens of Idya furcata were 


CRUSTACEA OF FRANZ-JOSEF LAND. 113: 


obtained in the Franz-Josef Land collections ; they could usually 
be distinguished at sight by the characteristic pale purpie bands 
across the dorsal aspect. They occurred in three gatherings 
collected off West Point, Cape Flora, about 50 yards, in 2-4 
fathoms: (1) on June 4th, (2) June 22nd, and (8) July 5th, 
1897. Some specimens were also obtained in a gathering 
dredged at West Bay, in 2-10 fathoms, July 2nd of the same 
year. Asa British species, Jdya furcata is moderately common. 


Ipya minor, 7. & A. Scott. 

1896. Idya minor, T. & A. Scott (84), p. 228, pl. 4. figs. 11-17. 

This is a distinctly smaller species than J. furcata, and appears 
to be a scarce form. The only gathering in which a few speci- 
mens were obtained was dredged in 2-3 fathoms, about fifty 
yards off West Point, Cape Flora, on June 18th, 1897. 


Genus ScureLtipium, Claus, 1866. 


ScUTELLIDIUM TIsBOIDES, Claus. 

1866. Scutellidium tisboides, Claus (24), p. 21, pl. 4. figs. 8-15. 

Several specimens of this species occurred in the same gather- 
ing with Idya minor, but in no other. It is a moderately large 
and easily recognized species. It is widely distributed, but 
apparently not very common. It is one of the rarer of the 
British species. © 


Family ASCOMYZONTID4. 


Genus Dermatomyzon, Claus, 1889. 


DERMATOMYZON NiGRIPES (Brady § Robertson). 

1875. Cyclopicera nigripes, Brady & Robertson (45), p. 197. 

This species was very rare in the Franz-Josef Land collections. 
The only gathering in which it was observed was one collected 
off East Glacier, Cape Flora, and which contained several other 
interesting species, such as Munna Fabricii and Pleurogoniwm. 
The British distribution of Dermatomyzon nigripes extends to 
the Shetland Islands. Dr. Giesbrecht refers doubtfully to the 
occurrence at Spitzbergen of this species. 


114 MR. THOMAS SCOTT ON THE 


Genus Myzoprontius, Giesbrecht, 1895. 


Myzorontius PUNGENS, Giesbrecht. (Pl. 9. figs. 1-10.) 

1895. Myzopontius pungens, Giesbrecht (53), p. 182. 

Description of the female.—Thorax broadly ovate, being about 
one-third longer than broad; the abdomen is narrow and fully 
half the length of the thorax; “thoracic segments scarcely 
produced into lateral processes, neither are the abdominal ” 
(fig. 1). Antennules short, 12-jointed; the terminal joint is 
elongate, being fully twice the length of the penultimate one 
and bearing an asthetask near the distal end (fig. 2); the 
formula shows approximately the proportional lengths of all the 


joints :— 
Proportional lengths of the joints 18.12.6.6.5.5.9.6.8.8.11.25 
Numbers of the joints............... LBs 42 Soy 8 Bud ml mH, 


Antenne (PI. 9. fig. 3) 4-jointed; third joint short, secondary 
branch very small. Mandibles (Pl. 9. fig. 4) in the form of 
long, slender stylets. Maxille (fig. 5) furnished with two 
lobes: inner short, oval, and bearmg one long and one short 
apical seta; outer lobe elongate and narrow, and provided with 
two apical sete of moderate length. The anterior foot-jaws 
(fig. 6) are armed with very long and curved terminal claws. 
The posterior foot-jaws are long and slender (fig. 7) ; the second 
joint is elongate, but the last three are shorter and narrower, 
and the terminal claw is moderately stout and about equal in 
length to the last three joints. The first four pairs of swimming- 
feet are somewhat similar in structure, and have both branches 
3-jointed; in the first pair (fig. 8) the end-jomts of the outer 
branches are armed with three spines and five plumose sete ; 
the end-joints of the inner branches have one plumose seta on 
the outer margin, three on the inner margin, and two at the 
apex, while the second joints are furnished with two setx, and 
the first joints with one on the inner margin. The end-joints of 
the outer branches of the fourth pair (fig. 9) are furnished with 
four spines and five sete; the number of sete on the inner 
branches is similar to that on the inner branches of the first pair, 
except that there are only two sete on the inner margin of the 
end-joint, and the inner one of the apical sete is replaced by a 
slender sabre-like spine. Fifth pair (fig. 10) small, 1-jointed, 


CRUSTACEA OF FRANZ-JOSEF LAND. 115 


cylindrical, the length being about equal to twice the breadth, 
and furnished with three terminal hairs. The first segment of 
the abdomen is only slightly enlarged anteriorly, the second and 
third joints are both shorter than the last joint ; the caudal furca 
are of moderate length, being about as long as the last two 
abdominal joints combined. 

Hab. Off East Glacier and near Cape Gertrude, Northbrook 
Island. A few specimens only were obtained. 

Remarks. There seems to be little doubt that this Franz- 
Josef Land species is identical with Dr. Giesbrecht’s Myzopontius 
pungens from the Bay of Naples, so far as can be made out from 
the description alone. The Arctic specimens appear to be 
somewhat larger than those from Naples; the specimen figured 
measured 1°6 mm. (;/5 of an inch), whereas Dr. Giesbrecht gives 
0°85 to 1:1 mm. as the size of the female. 


Family ONC HAD &. 


Genus Oncda, Philippi, 1843. 


OncAA MEDITERRANEA (Claus). 

1863. Antaria mediterranea, Claus (23), p. 159, pl. 30. figs. 1-7. 

A few specimens of Oncdéa mediterranea were obtained in a 
gathering collected about fifty yards off West Point, Cape Flora, 
in 2-3 fathoms, on June 4th, 1897. This Copepod appears to 
have a wide distribution; and it is also of interest to note that 
though Mr. Bruce obtained it in quite shallow water at Franz- 
Josef Land, it has, on the other hand, been found at considerable 
depths in the tropical seas. Dr. W. Giesbrecht records its 
occurrence at a depth of 4000 metres *, and I have obtained the 
same species in a gathering of micro-crustacea from the Gulf of 
Guinea collected at a depth of 360 fathomsy. What appears to 
be the same form was recorded from Spitzbergen by Dr. Lillje- 
borg in 1875 f. 


* * Pelagischen Copepoden des Golfes von Neapel,’ p. 591. 

+ Trans. Linnean Society, 2nd ser. (Zool.), vol. vi. p. 118. 

} “ De under Svenska vetenskapliga Expeditionen till Spetsbergen 1872-1873 
derstades samlade Hafs-Entomostraceer” (Ofvers. Akad. Férhandl. Stockholm, 
Aar 1875). 


116 MR. THOMAS SCOTT ON THE 


CIRRIPEDIA. 
Family BALANID &. 


Genus Bananus, Lister. 


BALANwus Porcatus, da Costa. 

1788. Balanus porcatus, Em. da Costa (25), p. 249. 

Several specimens of Balanus porcatus were included in the 
Franz-Josef Land collection; they were nearly all obtained in 
the vicinity of Cape Flora. They were obtained on floe-ice off 
Flora Cottage on 24th August, 1896, and others were collected 
near the same place in September. Other specimens were 
gathered off Cape Gertrude during June and July 1897, as well 
as some distance south-west of Elmwood, off Cape Flora, and in 
the vicinity of a glacier between Cape Flora and Cape Gertrude. 
It would thus appear that this species was more or less frequent 
all round the neighbourhood of Cape Flora. 


BaLANUS CRENATUS, Bruguiére. 

1789. Balanus crenatus, Bruguiére. 

This Balanus was very rare in the Collection ; the species was 
represented by only one specimen of the smooth variety, which 
was dredged in 8 fathoms off Cape Flora, July 1897. 

Both species occur in the Glacial clays of Scotland: B. por- 
catus is frequently observed, but B. crenatus is scarcer. 


InpDEx TO SPECIES. 


. Page Page 
Spirontocaris Gaimardii (M.-Edw.). 63 | Munna Fabricii, Ky. ...............68. 67 
(?) 5 Phippsii (Kr.) ......... 63 »,  Kroyeri, Goodsir ............ 67 

a polaris (Sab.) ......... 63 | Pleurogonium inerme, G. O. Sars... 67 
Sclerocrangon boreas (Phipps) ...... 64 p spinosissimum, G. O. 
Thysanoessa neglecta (Kr.) ......... 64 Sais’ sage foseseoagusce 67 
Mysis oculata (Fabr.) ................-. 64 | Munnopsis typica, M. Sars............ 68 
Diastylis Rathkii (Kr.) ............... Gomi Dajusimiysidisy KOsgeeneeeeceec steers 68 
Lamprops fuscata, G. O. Sars ...... 65 | Podascon Stebbingi, G. & B. ......... 68: 
Petalosarsia declivis (G. O. Sars) ... 65 | Hyperia galba (Mont.) ............06 69 
Typhlotanais finmarchicus,G.O.Sars, 65 | Parathemisto oblivia (Kv.)............ 69 
Leptognathia longiremis (Lillj.) ... 66 | Orchomenella minuta (Kr.) ........: 69 
Pseudotanais forcipatus (Lillj.)...... 66 | Anonyx similis (Phipps) ............... 69 
Gnathia elongata (Kr.) ..........6... 66 | Hoplonyx similis, G. O. Sars ......... 70 
Janira tricornis (Kv.) ...............06 67 | Pseudalibrotus littoralis (Kr.) ...... 70 


CRUSTACEA OF FRANZ-JOSEF LAND. 


Page 

Onesimus Edwardsii (Kr.) ............ 71 
Amphilochus oculatus (Hansen) ... 71 
Gitana Sarsii, Boeck .................. 42, 
Metopa pusilla, G. @#iSarst-cn.ccecess 72 
») sinuata,G. ©. Sars ......... 72 
peenieelectas) Elansenty.s.s---.52 72 
Parcediceros lynceus (M. Sars) ...... 72 
Monoculodes borealis, Boeck......... 72 
e latimanus (Goés) ...... 73 

2 Schneideri, G.O. Sars. 73 


Monoculopsis longicornis (Boeck) ... 73 
Bathymedon obtusifrons (Hansen)... 73 
Aceros phyllonyx (M. Sars) 
Acanthostepheia Malmgreni (Goés). 74 


Paramphithoé pulchella (Kr.) ....:. 74 
5 bicuspis (Kr. )......... 74 

“ny monocuspis, G.O.Sars 74 
Parapleustes glaber (Boeck) ......... 75 
Acanthonotosoma cristatum (Owen) 75 
Syrrhoé crenulata, Goés............... 75 
Pardalisca cuspidata, Kr. ............ 76 
Eusirus cuspidatus, Kr. ............... 76 
Rhachotropis aculeaia (Lep.) ...... 76 
Rozinante fragilis (Goés) ............ a, 
Halirages fulvocinctus (M. Sars) ... 77 
Cleippides quadricuspis, Heller...... 78 
Calliopius leviusculus (Kr.) ......... 78 
Amphithopsis glacialis, Hansen...... 78 
Atylus carinatus (Fabr.).............-- 78 
Amathilla homari (Fabr.) ............ 79 
3% jorbavesele) (UKGE,)) coscassoacconee 79 
Gammaracanthus loricatus (Sab.) ... 79 
Gammarus locusta (Linn.)............ 79 
Photis tenuicornis, G.O. Sars ...... 80 
Ischyrocerus anguipes, Kr............. 80 
Dulichia spinosissima, Kr............. 80 
Algina spinosissima, Stimps. ......... 80 
Caprella septentrionalis, Kr. ......... 81 
»  microtuberculata,G.O.Sars 81 
PeClultaseelanseniereeceeeeeeee 81 
Cyclocypris globosa, G. O. Sars...... 82 
Herpetocypris dubia, sp.n. ......... 82 
35 arctica, sp.n.......... 83 
Candona Harmsworthi, sp.n. ...... 83 
Pontocypris hyperborea, sp. n. ...... 83 
Cythere marginata, Norman ......... 84 
»,  limicola, Norman............ 84 


Page 
Oythere globulifera, G. S. Brady ... 84 
5 @lewtilnes,, 18% (Ob a5 1M couoauade 84. 


septentrionalis, G. S. Brady 85 
»,  tuberculata (G. O. Sars) ... 85 
»  emarginata (G. O. Sars) ... 85 


»  costata, G.S. Brady......... 85 
» mirabilis, G. 8. Brady ...... 85 
dunelmensis (Norman)...... 85 


Cyther idea papillosa, Bosq. ......... 86 
re punctillata, G. S. Brady 86 

55 Sorbyana, Jones ......... 86 
Eucvthere declivis (Norman)....... . 86 
Xestoleberis depressa, G. O. Sars .. 86 
Cytherura undata, G. O. Sars 
5 UK A, 18%, WS IR gosooand000006 87 

f clathrata, G. O. Sars...... 87 
Cytheropteron latissimum (Norm.) 87 
= pyramidale, Brady... 87 
subcircinatum, Sars 87 
punctatum, Brady... 87 
angulatum, B. & R. 88 


”? 


2) 


Pseudocythere caudata, Sars ......... 88 
Sclerochilus contortus (Norm.) ...... 88 
Paradoxostoma variabile (Baird) ... 88 
3 flexuosum, Brady ... 89 
Philomedes brenda (Baird) ......... 89 
Polycope orbicularis, G. O. Sars ... 89 
Calanus finmarchicus (Gun.) ......... 80 
my LENOSAOOEMIE, UTP, sonocoucoone 90 
Pseudocalanus elongatus (Boeck) ... 91 
Eucheta norvegica, Boeck ............ 91 
Metridia longa (Lubbock) ............ 91 
Misophria pallida, Boeck ............ 92 
Oithona similis, Claus.................. 92 
Cyclopina gracilis, Claus ............ 93, 
Thorellia brunnea, Boeck ............ 93 
Cyclops braceiispreneeeeeesceereacter 95 
Bradya typica, Boeck................... 94 
59 nwa, IN, a's Als SOO oscoca ce 94 
Ectinosoma Sarsi, Boeck............... 94 
* propinquum, T. & A. 8. 95 

- curticorne, Boeck ...... 95 

5 pygmeum, T. & A.S.... 95 

Fe melaniceps, Boeck ...... 95 


Normani, T. & A. Scott re 
5 atlanticum (B. & R.) .. 
Zosime typica, Boeck ......... ........ on 


118 MR. 


Robertsonia tenuis (B. & R.) 
Amymone spherica, Claus 
Stenhelia reflexa, T. Scott 
Ameira longipes, Boeck 
Gwalqne, AU, HEOwWs Gooacocdoone 
longiremis, T. Scott......... 
»  vreflexa, T. Scott 
Jonesiella spinulosa (B. & BR.) ...... 
Delavalia robusta, B. & R. 
reo, ID, SKEOUE osoosedes 
reflexa, B. & R..:.......... 
es artica, sp. n. 
Maraenopiotus Vejdovskyi, Mrk.... 
Canthocamptus parvus, T. & A. 
PME SCOGU sy “sie ciiutcurciiscctt aecueese rere oecleas 
Laophonte horrida, Norman 
curticauda, Boeck 
depressa, T. Scott ...... 
longicaudata, Boeck ... 
intermedia, I. Scott ... 
similis, Claus ............ 
pbexplexaysjssneleeseeeese 
Tmeebomtide: typicus, T. Scott . 
Cletodes similis, T. Scott 


eee 


3? 


39 


eer coccee 


a7 


39 


39 


. 103 


103 


THOMAS SCOTT ON THE 


Page 

Cletodes longicaudata, B. & R. ... 104 
Enkydrosoma curvatum, B. & R.... 104 
Dactylopus tisboides, Claus ......... 104 
aS coronatus, TT’. Seott...... 105 

Ss tenuiremis, B. & R. 105 

os longirostris, Claus ...... 105 

, Stromii (Baird), var. ... 106 
Thalestris helgolandica, Claus ...... 106 
7" TOOUETE, BO} 10, Soscccocerc0 106 

Fe forficula, Claus............ 108 

ie frigida, sp. n. 108 

Fe Jacksoni, sp. 0........0066- 109 
Westwoodia nobilis (Baird) ......... 111 
Harpacticus chelifer (Miller) ...... 111 
3453 oe Wats) ieacceeeaee 111 
Zaus spinatus, Goodsir ............... 112 
Eupelte purpurocincta (Norm.) ... 112 
lidyalturcatay (Baird) eereeeeceeeesees 112 
Hy emo, AD, A's ZN, ISICOWBocoocecoaco5 113 
Scutellidium tisboides, Claus ...... 113 
Dermatomyzon nigripes (B. & R.). 118 
Myzopontius pungens, Giesbrecht . 114 
Oncia mediterranea (Claus)......... 115 
Balanus porcatus, da Costa ......... 116 
55 OROMALDS, IBFAME,  cacconoaacae 116 


List of Works referred to in the preceding pages. 


le elss5: 
Gp USB a a is 
2 SAa a) 
5 elsON in _ 
4. 1860. Borck, A.—Forhandl. 
5. 1864. _,, bh 
(Gs NOVO Hb 
Th, LUSH i. ‘ 
8. 1852. 

Belgique. 
9, 1865. 
10. 1866. A 


bP) 


Mode. 

Oversigt Norges Copepoder. 

Crustacea Amphipoda borealia et arctica. 
Nye Slegter og Arter af Saltvands- 


Barrp, W.—Trans. Berwick. Nat. Club, vol. i. 
Mag. Zool. & Bot. vol. x. 

Trans. Berwick. Nat. Club, vol. u. 
Nat. Hist. Brit. Entomostraca. 


ved de Skand. Naturf. 8 de 


Copepoder. 


Bosquser.—Desceript. Entom. foss. terr. France et 


Bravy, G. S.—Ann. & Mag. Nat. Hist. vol. xxvi. 


Trans. Zool. Soc. vol. v. 


16. 


ihe 


CRUSTACEA OF FRANZ-JOSEF LAND. 119 


1866. Brapy, G. S.—Brit. Assoc. Report. 

1868. . x Monogr. Recent Brit. Ostracoda. 
(Trans. Linn. Soe. vol. xxvi.) 

1868. “ 5 Ann. & Mag. Nat. Hist. ser. 4, vol. ii. 

1878-80. ,, 8 Monogr. British Copepoda. (Ray 
Society.) 

1881. Brapy, H. B.—On some Arctic Foraminifera from 


Soundings obtained on the Austro-Hungarian 
North Polar Expedition of 1872-74. (Ann. & Mag. 
Nat. Hist. ser. 5, vol vii. pp. 393-418 ; contains 
notes on Ostracoda.) 

1874. Brapy, Crosskry, & Rogprertson.—Monoer. Post- 
Tertiary Entom. Scotland, including specimens 
from England and Ireland. (Palwontographical 
Society.) 

1889-96. Brapy & Norman.—Monogr. Marine and Fresh- 
water Ostracoda of North Atlantic and North- 
western Europe. (Trans. Roy. Irish Acad.) 

1872. Brapy & Rosertson.— Ann. & Mag. Nat. Hist. 

ser. 4, vol. ix. 


1872. a o Brit. Assoc. Report. 

1873. Be He Ann. & Mag. Nat. Hist. 
ser. 4, vol. xii. 

1874. i x Ann. & Mag. Nat. Hist. 
ser. 4, vol. xiii. 

1875. at Fe Brit. Assoc. Report. 

1863. Craus, C.—Die freilebenden Copepoden, mit beson- 


derer Berucksichtigung der Fauna Deutschlands, 
der Nordsee und des Mittelmeeres. 

1866. Cuaus, C.—Die Copepoden-Fauna von Nizza. 

1778. pa Costa, EManvurL.—Hist. Nat. Test Brit. 

1837. Epwarps, M.—Hist. Nat. des Crustacés, tom. i. & ii. 

1779. Faprictus, J. C.— Reise nach Norwegen, mit 
Bemerkungen aus der Naturgeschichte und Oeko- 
nomie. Hamburg, 1779. 

1793. Faxricius, J. C.—Systema Entomologiz, 2 vols. 


. 1780. Fasrictus, O.-—Fauna Greenlandieca. 


1895. Grarp & Bonnier. — Contrib. 4 ?Etude des Epe- 
carides (xx.). (Bull. Scientif. de la France et de 
la Belgique, vol]. xxv.) 


120 


33a. 


50. 


MR. THOMAS SCOTT ON THE 


. GIESBRECHT, W.—Pelagischen Copepoden des Golfes 


von Neapel. 
Ann. & Mag. Nat. Hist. ser. 6, 


vol. xvi. 


99 99 


. Gok&s, A. — Crustacea Amphipoda maris Spetsber- 


giani. (Ofvers. Kel. Sv. Vet.-Akad. Forhandl. 1865.) 


. Goopsir, H.—Edinb. New Philosophical Journal, 


vol. XXxXviil. 


Ann. & Mag. Nat. Hist. vol. xvi. 


99 29 


: Gus hk. — Notes on a Voyage to the Greenlaad 


Seas in 1888. (The Zoologist, January-March 
1889.) 


5. Gusner, J. E.—Nogle smaa rare og meestendelen 


nye Norske Soedyr. (Act. Hafn. x.) 


. Hansen, H. J. — Malacostraca marina Greenlandize 


occidentalis. 


. Heier, C.— Die Crust., Pyenog. u. Tunicaten d. 


kaiserl.-konig. Oesterr. Ungar. Nordpol.-Exped. 


. Hever, Tu. von.—Reisen nach dem Nordpolar- 


meer. 


. Jones, T. R.—Monogr. Tertiary HEntomostraca of 


England. (Paleontographical Society.) 


. Jones, T. R., & C. D. SuErBorn.—Supplementary 


Monogr. Tert. Entomostr. England. (Paleonto- 
eraphical Society.) 


. Kroyer, H.—Gronlands Amfipoder. 


Udsigt nord. Art. Hippol. (Naturh. 
Tidsskr. iii., xiii.) 

Mon. Fremst. af Slegt. Hippolyte’s 
nordiske Arter. 

Voy. en Scand., Crust. 

Nye nord. Slegt. og Art. af Amfip. 
(Nat. Tidsskr. 1 R., B. 4.) 

Karcin. Bidr. (Nat. Tidsskr. 2 R., 
B. 1.) 

Voy. en Scand., Zoology. 


Tamron aes Acad. Sci. Tivo, Petrop. 


. Littsesore, W.—Bidrag t. Kannedom. om de inom 
Sver. och Norr. forek. Crust. af Tan. fam. (Ups. 
Univ. Arsskr., Math. og Naturv. vol. i.) 


. Linnt.—Systema Nature, ed. 12. 


ol. 


52. 


53. 


61. 
61a. 
62. 


63. 


64. 


CRUSTACEA OF FRANZ-JOSEF LAND. 121 


1854. Lusgock, Sir J.— On some Arctic Species of 
Calanide. (Ann. & Mag. Nat. Hist. ser. 2, vol. xiv. 
p- 125.) 

1878. Mismrs, E. J.—Arctic Crustacea, with notes on the 
Copepoda by Rev. A. M. Norman, and on the 
Ostracoda by G. S. Brady. (Voyage to the Polar 
Sea in H.M. ships ‘ Alert’ and ‘ Discovery,’ 1875- 
76, under Captain Sir G.S. Nares, Vol. ii. Appendix 
no. Vii.) 

1880. Miers, E. J.—On a small collection of Crustacea 
made by Edward Whymper, chiefly in the North 
Greenland seas. (Journ. Linn. Soe., Zool. xv. 
pp: 59-73.) 

1815. Monracu.—Linn. Trans. vol. xi. pp. 1-26. 

1893. Mrazex.—Zoologische Jahrbiicher. 

1776. Miter, O. F.—Zool. Dan. Prodromus. 

1862. Norman, A. M.—Ann. & Mag. Nat. Hist. vol. ix. 

1864. 4s is Brit. Assoc. Report. 

1865. % de Nat. Hist. Trans. Northumberland 

and Durham, vol. 1. 
1868. Pe BY Brit. Assoc. Report. 
1876. Norman, Rev. A. M.—On some Crustacea and some 
other Groups of Invertebrates obtained by the 
‘Valorous’ Expedition to Greenland and Davis 
Straits in 1875. (In Report on the Zoology of the 
Expedition by Dr. J. Gwyn Jeffreys, F.R.S., and 
Dr. Carpenter, C.B., F.R.S., Proc. Roy. Soc. 
vol. xxv.) 
1835. OwEn.—Appendix to Ross’s Second Voyage in search 
of a North-west Passage. 
1774. Puipps.—Voyage au Pole boréal. 
1898. Ricuarp, Jurtms.—Sur la Faune des Eaux douces 
explorées en 1898 pendant la Campagne du yacht 
‘Princesse Alice.’ 
1888. Rozertrsoy, D.—On the Post-tertiary beds at Gar- 
vel Park, Greenock. (Trans. 
Geol. Soc. Glasgow, vol. vii. 
pt. 1.) 

1888-92. _,, » Amphipoda and Isopoda of the 


Firth of Clyde, and Supplement. 


122 
65. 


66. 
66 a. 
67. 
68. 
69. 
70. 
71. 
72. 


73. 
74. 


75. 


716. 
77. 


78. 
79. 


80. 
81. 


82. 


83. 


84. 


85. 


MR. THOMAS SCOTT ON THE 


1824. Sanine. — Suppl. to Appendix of Captain Parry’s 
Voyage. 
1854. Sars, M.—Christ. Vid.-Selsk. Forhandl. (1854). 


1858. 
1860. 


9? 9? 


9) 99 


99 99 99 99 (185 8). 
(1860). 


39 99 39 99 


1863. Sars, G. O.—Om en i Somm. 1862 foret. zool. 


1865. 


1865. 
1865. 
1880. 


1883. 
1885. 


99 2? 


39 29 


99 39 


99 3) 


obty) ” 


99 9? 


1890-95. ,, 


1896-96. ,, 
1892-98. Scumetn, Dr. Orro.—Deutschlands freilebende 


1898. 


29 


Reise Christiania. 

Om den aberr. Krebsdyrg. Cumacea, 
og d. nord. Art. (Forh. i Vid.-Selsk. 
i Christiania, 1864.) 

Oversigt af Norges marine Copepoder. 

Beretn. om en i Somm. 1865 foret. 
zool. Reise ved Kyst. af Christianias 
og Christiansands Stifter. 

Revision af Gruppen Isopoda Cheli- 
fera. 

Oversigt af Norges Crustaceer. 

Norweg. North-Atlantic Expedition— 
Crustacea. 

An account of the Crustacea of Nor- 
way: Vol. i., Amphipoda. 

Op. cit. Vol. u. parts i-vii, Isopoda. 


Siisswasser-Copepoden. 
Op. cit., Nachtrag. 


9 


1894. Scorn, T. Eneroleny ‘Aad Report, Fishery Board 


1895. 


1897. 


1897. 


29 99 


29 99 


rp) 


Loch Than, 


for Scotland (pt. 11.). 

Thirteenth Ann. Rept. Fish. Board for 

Scotl. (pt. 111.). 

Fourteenth Ann, Rept. Fish. Board for 

Scotl. (pt. 111.). 

The Marine Fishes and Invertebrates of 
(Fifteenth Ann. Rept. Fish. Board 


for Scotl., part 111.) 
1893. Scorr, T. & A.—On some new or rare Crustacea 


39 


29 


from Scotland. (Ann. & Mag. 
Nat. Hist. ser. 6, vol. xii.) 

A Revision of the genera Bradya 
and EHetinosoma. (Trans. Linn. 
Soe. ser. 2, vol. vi.) 

Annals of Scottish Natural History, 


CRUSTACEA OF FRANZ-JGSEF LAND. 123 


86. 1885. Scorr, T., & J. Sreer.—Notes on the Occurrence 
of Leda arctica (Gray), Lyonsia arenosa (Moller), 
and other organic remains in the Post-pliocene 
clays of Garvel Park, Greenock. (Trans. Geol. 
Soe. Glasgow, vol. vil. pt. il.) 

87. 1894. Sressine, T. R. R.—Amphipoda collected during 
the voyage of the ‘ Willem Barents’ in the Arctic 
Seas in the years 1880-84. (Bijdragen tot de Dier- 
kunde mitgegeven door het Koninklijk Zoologisch 
Genootschap “‘ Natura Artis Magistra” te Amster- 
dam, Afi. 17.) 

88. 1853. Srrwpson.—Marine Invert. Grand Manan. (Smiths. 
Contr. to Knowl. vol. vi.) 


EXPLANATION OF THE PLATES. 


Prats 3. 
Fig. 1. Spirontocaris Gaimardii, rostrum, enlarged. 
2. a a telson, enlarged. 
Bs (Oy oy Phipsii, rostrum, enlarged. 
4, ro = telson, enlarged. 
5. Typhlotanais finmarchicus, superior and inferior antenne, enlarged. 
6. i is one of the sixth pair of pereiopoda, 
enlarged. 
ie 60 3 one of the pair of uropoda, enlarged. 
8. Pseudotanais forcipatus, one of the chelipeds, enlarged. 
9. i es one of the pair of uropoda, enlarged. 
10. Munna Fabricit, one of the antennules, enlarged. 
Wk. i me one of the second pair of pereiopoda, enlarged. 
12. »  Kréoyeri, one of the antennules, enlarged. 
13. 4 Fe one of the antenne, enlarged. 
14. 5 ss one of the second pair of pereiopoda, enlarged. 


15. Pleurogonium spinosissimum, dorsal view, enlarged. 
16. Candona Harmsworthi, lateral view, enlarged. 
I Fr ss dorsal view, enlaryed. 


Puate 4. 


. (?) Cyclocypris globosa, lateral view, enlarged. 

. Herpetocypris arctica, lateral view, enlarged. 

dorsal view, enlarged, 

one of the antennules, enlarged. 
one of the antenne, enlarged. 
post-abdomen, enlarged. 


Fig. 


oh go po 


” ” 


LINN. JOURN.—ZOCLOGY, VOL. EXVII. 10 


124 MR. THOMAS SCOTT ON THE 


Fig. 7. Herpetocypris dubia, lateral view, enlarged. 


8. ie 5, dorsal view, enlarged. 

9. Bs » one of the antennules, enlarged. 
10. “ ,, one of the antenne, enlarged. 
11. a post-abdomen, enlarged. 


12. Pontocypris hyper Lanta, lateral view, X 40. 


13. ; si} dorsal view, x 40. 

14. : = ventral view, x 40. 

15. : end view, x 40. 

16. Metridia longa, fifth pair of thoracic feet (¢, right foot), enlarged. 

17. a 5 - 5 (3, left foot), enlarged. 

18. Oithona seal, one of third pair of thoracic feet (2 ), enlarged. 

19, », one of fourth pair of thoracic feet (2), enlarged. 

Prats 5. 
Fig. 1. Dactylopus tenwiremis, one of the antennules, x 253. 

2. 5 us posterior foot-jaw, X 253. 
3. 5 fb one of the first pair of swimming-feet, x 253. 
4 ys 3 one of the fifth pair, x 190. 
5 s longirostris, one of the antennules, X 253. 
6 " posterior foot-jaw, < 253. 
Ue 7 , one of the first pair of swimming-feet, X 253. 
8 A ne one of the fifth pair, x 190. 
9 i Strémii, var. arcticus, female, lateral view, X 27. 

10. A Pa is one of the antennules, x 127. 

ik x # e posterior foot-jaw, x 253. 

12. © 5 te one of the first pair of swimming- 

feet, x 127. 
1B, as o i one of the fifth pair, x 95. 


14. Delavalia arctica, posterior foot-jaw, x 380. 
15. Enhydrosoma curvatum, one of the fifth pair of feet, x 380. 


| Puatr 6. 


Fig. 1. Cyclops Brucei, female, dorsal view, x 27. 
a eh one of the antennules, x 95. 
i » one of the first pair of swimming-feet, enlarged. 
5 » one of the fourth pair, enlarged. 

one of the fifth pair, enlarged. 

abdomen and caudal furca, enlarged. 

Delavalia arctica, female, dorsal view, X 27. 

“ is one of the antennules, x 95. 

one of the first pair of swimming-feet, x 127. 

one of the fourth pair, x 63, 

be u one of the fifth pair, x 95. 


5 2 £2 © SN Sd Si ES Co 


Fig. 12. 


15. 
HES 
15. 
16. 
17. 


18. 
19. 
20. 
21. 
22, 
23. 
24, 


Fig. 


FS ORAR Ow wo 


ee 
SSP Cx ps OS) KS) 


18. 


oH 


CRUSTACEA OF FRANZ-JOSEF LAND. 125 


Maraenobiotus Vejdovskyt, female, lateral view, xX 86. 


one of the antennules, x 253. 

one of the mandibles, greatly enlarged. 
posterior foot-jaw, < 803. 

one of the first pair of swimming-feet, x 253. 
one of the fifth pair, x 760. 


(?) Gah nocanmntus parvus, female, lateral view, x 80. 


4 one of the antennules, x 570. 

one of the antennz, x 570. 

one of the mandibles, x 380. 

one of the first pair of swimming-feet, x 253. 
x one of the fourth pair, xX 253. 

3 one of the fifth pair, x 253. 


PuatEe 7. 


. Laophonte perplexa, female, lateral view, x 21. 


39 


one of the antennules, x 190. 

posterior foot-jaw, x 253. 

one of the first pair of swimming-feet, x 253, 
one of the fourth pair, x 190. 

one of the fifth pair, x 190. 

caudal fureca, x 99. 


Thalestris polaris, female, lateral view, X 32. 

one of the antennules, xX 95. 

posterior foot-jaw, x 95. 

one of the first pair of swimming-feet, x 84. 
one of the fourth pair, X 68. 

one of the fifth pair, x 84. 

inner branch of second pair (¢), x 126. 
one of the fifth pair (¢), x 126. 

caudal furea, x 40. 

frigida, female, lateral view, X 27. 


2) 


one of the antennules, x 95. 

one of the posterior foot-jaws, x 127. 

one of the first pair of swimming-feet, x 63. 
one of the fourth pair, x 63. 

one of the fifth pair, x 63. 

inner branch of second pair (¢), X 126. 


PuaTe 8. 


. Thalestris frigida, one of the fifth pair of thoracic feet (¢), x 126. 


caudal furea, x 80. 


Jacksont, female, lateral view, < 18. 


bY) 


EP) 


one of the antennules, x 84. 
one of the posterior foot-jaws, x 95. 
NO 


126 


Fig. 


CRUSTACHA OF FRANZ-JOSEF LAND. 


6. Thalestris Jacksoni, ove of the first pair of swimming-feet, x 63. 
lie rf: A one of the fourth pair, x 47. 
8. i re one of the fifth pair, x 63. 
9 i c caudal furea, x 40. 
10. Harpacticus chelifer, one of the female antennules, x 126. 
Ane 6 99 one of the posterior foot-jaws (2), x 126. 
12. 3 Ss one of the first pair of swimming-feet, x 95. 
13. m i one of the fifth pair (2), x 126. 
14. a . var. arcticus, one of the female antennules, x 126. 
By re be M, one of the posterior foot-jaws (@), 
x 126. 
16. # es i one of the first pair of swimming- 
feet, x 95. 
17. 0 Ns ie one of the fifth pair (2), x 126. 
Puate 9, 


Fig. 1. Myzopontius pungens, female, dorsal view, < 27. 


2 ae su one of the antennules (2), x 84. 
3 , a one of the antennez, x 126. 
4 a as mandible, x 95. 
5. a 45 one of the maxillz, x 126. 
6 os 7 one of the anterior foot-jaws, x 95. 
7 5 es one of the posterior foot-jaws, x 95. 
8 ms x one of the first pair of swimming feet, x 84. 
9 " - one of the fourth pair, x 84. 
10. “S AS one of the fifth pair of thoracic feet, x 126. 
11. Hoplonyx similis, one of the anterior coxal plates, enlarged. 
12: RS AS one of the last pair of epimeral plates of metasome, 
enlarged. 
He, Ps » telson, enlarged. 
14. Amathilla pinguis, one of the last pair of epimeral plates of metasome, 
enlarged. 
15. Ms es telson, x 85. 
16. Photis tenuicornis, one of the first pair of gnathopods (¢), enlarged. 
ile bp 3 one of the second pair of gnathopods, enlarged. 


18. Ischyroceras (?)anguipes, one of the second pair of gnathopods, 
enlarged. 


Linn. Soc. Journ. Lool. Vol. XXVII, PL 3. 


Scott. 


J.T,Rennie Reid, Lith. Edin® 


Scote, del, 


We 


LAND. 


JOSEF 


CRUSTACEA FROM FRANZ 


Scott. Linn. Soc. Journ, Zool. Vol. XXVII, PL. 4. 


a 


wien gg 


rer 


T.Scott, del. J.T, Rennie Reid, Lith. Edin” 


CRUSTAGEA FROM FRANZ JOSEF LAND. 


a 


Scots. Linn. Soc. Journ. Zool. Vol. XXVII, Pl 5. 


Se 
/ 


= 


IPR 
Iced a ee 
culpa 


ss 


Bae ieee rE 


aE ee et 


A.Scott, del. 


J.T,Rennie Reid, Lith. Edin? 


CRUSTACIBA IMSOM TIRVAINZZ  GiOS Ie WvANN}ID) 


Linn. Soc. Journ. Zool. Vol. XXVII, Pl. 6. 


Scott. 


arama ae, 
7 — TS ren ee 
Ty, 


J.T.Rennia Reid, Lith. Edin 


A.Scott, del, 


JOSE F 


FRANZ 


FROM 


NUANG E\es 


ND) 


GRU 


Linn. Soc. Journ. Zool. Vol. XXVIII. Pl. 7 


Scott . 


oe 


on Sa ESRI HN wer 


we <== oars 


oa 


s <= — 


~~ aoe le ce ras ee 


J.T. Rennie Reid, Lith. Edint 


A.Scott, del. 


CGHRUOSWNG EA WIR OWL IMRVAINIA GOS) iLyeNN|ID)> 


Linn. Soc. Journ. Zool. Vol. XXVII, Pl. 8. 


OcGobb. 


J.T. Rennie Reid, Lith. Edin? 


A-Scott, del, 


CRUSTACEA FROM FRANZ JOSEF LAND. 


Linn. Soc. Journ. Zool. Vol. XXVIL Pl. 9. 


Scott. 


J.T.Rennie Reid, Iath. Edin® 


Figs 1-10 A.S. del. 
others T.S. del. 


CRUSWAC IVA IIROIM URRVAINIZA GOS ILeNIN ID), 


MR. H. M. BERNARD ON RECENT PORITID#. D7 


Recent Poritide, and the Position of the Family in the Madre- 
porarian System. By Henry M. Bernarp, M.A., F.L.S. 


[| Read 2nd February, 1899.] 


In the last paper dealing with the Stony Corals which I had the 
honour of reading before this Society *, a phylogeny of the 
Madreporarian skeleton was sketched out, and suggestions were 
made as to the lines of development which had been taken by 
some of the better-known recent forms. I now propose to give 
an instalment of the morphological results of work done during 
the past eighteen months on one of the families mentioned in 
that paper—the Poritide. It is hoped that in the course of the 
next few years the remaining groups of the Madreporaria, both 
recent and fossil, will be dealt with, and the proposed phylo- 
genetic scheme completed and strengthened by additions and 
amendments suggested in the course of the work. 

In this paper I propose to confine myself to the systematic 
position of the Poritide, and to introduce only so much structural 
detail as is necessary. The variations of form assumed by Porites, 
a few of which are only incidentally referred to here, deserve 
separate description. The most important question raised by the 
work itself, however, was primarily the place of the Poritide 
among the Madreporaria. This point has therefore been worked 
out first, and the present paper will be shortly followed by a 
more detailed account of the structure and of the chief lines of 
differentiation found within the constituent genera. 

But, while reserving an account of the structure for fuller 
treatment in a second paper, a brief description of the corals 
known as Porites will doubtless add to the interest of the fol- 
lowing pages. Although both explanate and branching forms 
are known, the most familiar are smooth round masse-, some- 
times attaining a circumference of 60 feet. They are built up 
by a dense crowd of minute cylindrical polyps characterized by 
twelve short, thick, (?) rudimentary tentacles. ‘The polyps rise 
fairly high above their skeletal substratum, and when retracted 
are unable to withdraw entirely into their calicles, as is the usual 
habit in the Stony Corais. Examination of the dried skeleton 
shows that the pits are too shallow to take in the polyps; they 


* Journ. Linn. Soc., Zool. xxvi., read December 1897. 


128 MR. H. M. BERNARD ON RECENT PORITIDA. 


are, in fact, little more than surface depressions in a skeletal 
mass, Conspicuous on account of its bemg a fine porous 
reticulum. The shallow calicles may be rounded and separated 
by reticular walls of different thicknesses, or else so crowded 
together as to be polygonal in outline. The skeletal structure 
within the calicle is also remarkable. The radial symmetry, 
which is such a striking feature in the Stony Corals, is in Porites 
more or less obscured. Septa never stand out as radiating plates, 
but instead 5-6 or more red-like or granular pali rise up as 
a central ring from the reticulum which fills up the base of 
the calicle. Round this ring of pali the rudiments of septal 
formation can generally be traced. 

Such corals as these early attracted the attention of naturalists, 
and the significant name Porites was first associated with them 
by Pallas (1766), and was eventually accepted by Lamarck as 
the generic name. 

The position which Lamarck assigned to the new genus is 
given in his classification of the ‘‘ Lamelliferous ” Corals (Anim. 
s. Vert., ed. 1816, ii. p. 219). It is placed with Madrepora in 
the last section, viz., in that which contamed corals with cells 
circumscribed and covering the whole free surface of the stock, 
and it follows Astrea, which also has cells circumscribed but 
confined to the upper surface of the stock. 

In 1821 * Lamouroux followed Lamarck in placing the genus 
Porites after Astrea, but called the group in which the former 
occurred “ Les Madréporées,”’ and that in which Astrea occurred 
“Les Astrées.”” De Blainville ¢ divided the Stony Corals into 
“‘Madrephyllies’’ and “‘ Madrepores,” the Astreids being placed 
among the former and Porites near the end of the latter. 
Nevertheless, in his observations he admits the apparent kinship 
of the genus with both Astrea and Madrepora as suggested by 
Lamarck, but thought it was much closer with the latter than 
with the former. The recently established genus G'oniopora, 
Q. & G., was rightly placed by De Blainville next to, but in front 
of. Porites. 

In 1834 Ehrenberg ¢ placed Porites in the “ Madreporina,”’ 
which were quite distinct from the “ Astreina.’’ The Madre- 
porina were divided mainly according to their methods of 

* ¢Hxposition Méthodique,’ pp. 56, 60. 
t ‘Manuel,’ pp. 334, 395. 
{ ‘Corallenthiere des Rothen Meeres,’ pp. 91-115. 


MR. H. M. BERNARD ON RECENT PORITIDE. 129 


budding into two genera, Heteropora (= the modern genus 
Madrepora) and Madrepora, Ehr. This latter consisted of two 
subgenera, Madrepora-phyllopora (=Astreopora, de Bly.) and 
Madrepora-porites, which was a heterogeneous group consisting 
of Montipores, Stylophores, Alveopores, with a few true Poritids. 

Dana* was the first to found the family Poritide, of equal 
value with and closely allied to the family Madreporide. It 
contained two genera, Porites and Goniopora. 

This arrangement of Dana’s was, in the main, accepted by 
Milne-Edwards and Haime in 1851. The Poritide, enlarged by 
the addition of many more genera, constituted, together with a 
very large family the Madreporidw, the great section Madre- 
poraria Perforata, as opposed to the greater bulk of the remaining 
Stony Corals, which were grouped as Madreporaria Aporosa. This 
recognition of the structure of the coral-skeleton as a feature of 
fundamental taxonomic importance is the chief merit of the work 
of these authors, which is the last comprehensive attempt to 
classify the whole coral system. It was, however, hardly to be 
expected that this first attempt to solve the difficult morphological 
problems presented by the coral-skeleton would be successful. 
It is not, therefore, surprising to find that every advance in our 
knowledge of corals has led to some sweeping revision of Milne- 
Edwards and Haime’s system. At the present day only two of 
its five original sections can be said to have held their own, viz., 
the two most important, the Madreporaria Aporosa and M. Per- 
foratat. That these two are now in their turn on the eve of 
modification, the extent of which cannot yet be predicted, because 
the researches which render revision necessary are still too recent, 
will, it is hoped, be made clear in the following pages. Criticisms 
of details have not been wanting, but they have mainly referred 
to the relative positions of families or genera. 

No change has, so far, been made affecting the position of the 
Poritide, which is the matter we have especially in hand in this 
paper. The only expressed doubt as to their affinity with the 
Madreporide with which I am acquainted is in the recent work 
of Miss Ogilvie ¢, who found it impossible to decide whether the: 
two families were or were not related. 


* * Zoophytes,’ 1848. 

t See Martin Duncan’s revision‘of the system in the Journ. Linn. Soc., Zool. 
Xvili. 1884, p. 3. 

¢ Phil. Trans. vol. 187, 1896, p. 327. 


130 MR. H. M. BERNARD ON RECENT PORITIDA. 


This brief historical sketch shows that almost all who have 
studied the Madreporaria have come to the conclusion that 
Porites is in some way related to the Madreporids. The reasons 
for this conclusion may be briefly arranged under the following 
heads :— 

1. The general similarity of their polyps, with twelve tentacles 
in a single ring. 

2. The fact that the septa are mostly in some low multiple of six. 
In Porites there are almost invariably twelve. 

3. The skeletal walls are porous, and in both tend to form reti- 
cular coenenchymas *. 

Although the real value of this last pot, viz., the pessession 
of porous or reticular walls, has never till recently been under- 
stood, it was nevertheless a common character in striking contrast 
with the solid mural structures found in the Astreeide. 

As opposed to these ccmmon characters uniting the two 
families, we have certain differences keeping them apart. ‘These 
were described by Dana, who limited the family to two genera, 
Porites and Goniopora, as follows:—(1) Extrao:dinary porosity 
of the Poritid skeleton as compared with the more regularly 
lamellate skeleton of the Madreporide ; (2) the fact that, in the 
Poritide, the skeleton in its relation to the polyp is purely basal 
and never rises to a deep cup; (8) that, as it grows, the small 
central depression of the calicle fills up, so that the stars are 
hardly or not at all traceable thrcugh the substance of the 
corallum, as they always are in the Madreporide. 

On the other hand, Milne-Edwards and Haime, carried along 
by their theory of the origin of the Madreporarian skeleton, 
believed that the “ trabecular” character of the septa in Porctes 
was the fundamental distinction, the septa in the Madreporide 
being lamellate. The remaining differences above quoted from 
Dana were no longer applicable, because several other genera 
with the so-called “ trabecular’ septa were now iucluded. 

As it is useless to attempt to discuss these resemblances and 
differences until we understand clearly what is meant by the 
terms used, it is necessary to sketch the fundamental theory 

* Milne-Fdwards and Haime also added that the families agreed in having 
no tabula, which are so ccmmon in the Astreids. This distinction is incorrect. 


I have already described tabule in Astreopora and Turbinaria, and find them 
also in Goniopora and specially numerous in Porites. 


MR. H. M. BERNARD ON RECENT PORITIDA. 131 


upon which the system of Milne-Hdwards and Haime was based. 
It must be remembered that this system, though 50 years old, 
has never yet been superseded so far as the two chief divisions of 
the Stony Corals are concerned, the perforate and imperforate. 
Though worker after worker finds it obsolete, no comprehensive 
criticism of it has yet been attempted in connection with syste- 
matic work*. I make no excuse, therefore, for reviewing the 
situation which recent researches have brought about. If I 
needed any excuse I should find it in the great trouble and 
perplexity which the word “trabecular”? has caused me during 
the past few years. The reason of the confusion can best be 
explained by showing with what total absence of precision the 
term was originally used. I hope to make this quite clear by 
means of concrete examples. 

According to Milne-Edwards and Haime the Madreporarian 
skeleton was built up by the fusion of vast numbers of spicules 
like those found isolated in the Alecyonaria. Fusion by terminal 
erowth of isolated spicules would naturally result in a reticulate 
corallum. Direct evidence of this theory was found in the 
fact that Madreporaria still existed i which the skeleton 
was reticular, the septa being a lattice-work. Porites showed 
these primitive skeletal conditions best, but others, e. g. Glonio- 
pora, Alveopora, and Montipora, were all sufficiently reticular, 
or, as it was called, “ trabecular’”’ T, to be united with Porites in 
one family. Corals showing a further degree of fusion were the 
Madreporide, in which the septaare for the most part lamellate, 
and only the walls are reticular. Thus the Poritide and the 
Madreporide were classed as Perforata in contrast to those 
corals, such as the Astreids, which showed a still higher degree 
of specialization, both septa and walls being solid throughout. 

It is needless to criticize the details of this scheme, since it has 
been recently proved to rest upon an entirely erroneous con- 
ception of the origin of the Madreporarian skeleton. We now 
know, primarily through the researches of Dr. von Koch, that 
the Madreporarian skeleton is a purely ectodermal secretion, 
and that the septa which appear to be internal are always clothed 


* The bearing of Miss Ogilvie’s work will be.alluded to in the course of what 
follows. 
| For criticisms of this term see pp. 137 & 145. 


132 MR. H. M. BERNARD ON RECENT PORITID. 


with a layer of ectoderm-cells. The whole skeleton is thus 
outside the polyp, and could never have been built up by a fusion 
of spicules developed inside the body. 

With regard to the general bearing of Dr. von Koch’s dis- 
covery on the origin and classification of the Madreporaria, I 
venture to believe that the phylogenetic scheme which was pre- 
sented in my last paper to this Society supplies us with a solid 
foundation on which to build up a natural system. That this 
scheme was only partially seen by Dr. von Koch himself is not 
to be wondered at; the clue to it lay hid entirely in the epi- 
theca, the great importance of which seems everywhere to have 
escaped attention. Dr.v. Koch’s conclusion was that the “ basal 
plate ’’ with the “ epitheca” (that term being commonly limited 
to the continuation of the basal plate a short way up the sides 
of the polyp) together formed the primitive skeletal cup of the 
Madreporaria *. 

This description, though correct in fact, fails to recognize the 
fundamental morphological importance of the epitheca. My own 
systematic studies had, on the other hand, led me, along quite 
independent lines, to the conclusion that the epitheca, from which 
it is impossible to separate the basal plate as a distinct morpho- 
logical unit, had been at one time the most important element in 
the skeleton, and that, though it is now very generally vestigial, 
it was the original cup-like exoskeleton of the Stony Corals from 
which all the later internal (septal) skeletons had been developed 
by infoldings. This view is fully supported by the facts :— 
(1) That the epitheca forms such cup-like exoskeletons in the 
earliest stages of many (? all) Stony Corals; (2) that transitional 
forms such as Alveopora occur, in which the primitive importance 
of the epitheca is much longer retained; (8) that many 
Paleozoic corals are almost purely epithecate; and (4) that 
published drawings of sections of Flabellum show the septa as if 
they were still formed as simple infoldings of an external 
wall t. 

This was summed up in my previous paper (/. c. p. 514) in the 
following words :—“The Madreporarian skeleton may be de- 
scribed as the rigid secretion of the basal portion of the columni- 
form body of a polyp into which the flexible upper portion may 


* Gegenbaur’s Festschrift, ii. 1896, p. 272. 
t Cf. p. 134. 


MR. H. M. BERNARD ON RECENT PORITID®. 133 


be invaginated. In its earliest development a simple cup, it has 
become complicated in various ways; primarily, by the develop- 
ment of radial infoldings of the stiff external wall, comparable 
with the infoldings of the chitinous cuticle of Arthropods: 
secondarily, (1) by further complications of these infoldings so 
as to form an intricate ‘internal’ skeleton, which may render 
the primitive external cup unnecessary, and hence lead to its 
becoming vestigial ; (2) by a process of repeated sheddings of 
the external hard secretions, and the formation of new ones 
(dissepiments and tabule) across and among the existing ‘ in- 
ternal’ skeletal structures.” 

Further work with Madreporarian skeletons has only con- 
firmed this generalization. One or two points, however, require 
attention. In my former paper these septal infoldings were 
likened to the apodemes of Arthropods, formed by the infoldings 
of the chitinous skeleton which sometimes, e.g. in the cephalo- 
thorax of the Spiders, form together an elaborate internal frame- 
work. While this resemblance is structurally accurate, the 
comparison must be received with caution. The apodematous 
system of the Arthropods can be shown, even in detail *, to be 
due, at least in their earliest stages, to muscular action, either 
directly drawing in the chitin to which it is attached, or causing 
deep wrinkles or folds across the line of the muscles. But it 1s 
difficult to see how the infoldings of the calcareous exoskeleton 
of the early Madrepores to form septa could have been due to 
muscular action. Dr. von Koch (/.c.) thinks that the septa 
might have arisen in connection with certain endodermal ridges 
found in some larve. But we shall be probably nearer the truth 
if we can find a cause for them in the ectoderm itself. Until 
recently I thought that they were due to increased local activity 
in the secretion of calcareous matter, which would therefore push 
in the body-wall. From this point of view I found fault (Geol. 
Mag. 1897) with Miss Ogilvie’s description of the process as an 
“invagination which became filled up” with skeletal matter. 
But this terminology, though not felicitous, need not be alto- 
gether wrong. It seems to me not unlikely that the puckering 
which gave rise to the septa was caused by the growth of the 
basal, and probably best nourished, wall of the polyp, and that 
this wall, cramped by the primitive exoskeletal epitheca, could 


* Ann. & Mag. Nat. Hist. (6) x. p. 67 (1892). 


134 MR. H. M. BERNARD ON RECENT PORITIDA. 


ouly increase by the formation of folds. We can well under- 
stand how such puckering might be radial in the bases of the 
mesenterial chambers, but quite irregular in the base of the 
central cavity, where the radial puckers would meet and fuse 
together with twists and curves. Under the radial puckers, the 
ectoderm would secrete the septa; under the central, the 
columella. This view finds some support in the fact that fresh 
septa are added by puckering of the ectoderm just below the rim 
of the epitheca (or subsequently of the theca) of the growing 
coral, 7. e. just where the polyp is trying to expand *. 

Again, in endeavouring to establish my argument that the 
septa arose from infoldings of the epitheca, I appealed to sections 
of Flabellum. While this appeal is, I think, perfectly justifiable, 
the sections demonstrating in a remarkable way the point it was 
desired to establish, yet I confess that, at the time, I did not see 
that this case itself (#labellum) required explanation. For such 
d:rect infoldings of the epitheca from the external surface cannot 
be considered ag primitive. As far as we can see, the epitheca 
must primitively have formed a continuous calcareous layer, and, 
when infoldings began, they must have risen from its inner 
surface without the possibility of there being any external scar 
such as necessarily exists in the case of chitinous infoldings in the 
Arthropods, at least until secondarily obliterated. The direct 
infoldings of the epitheca of Flabellum with external scars are 
therefore somewhat startling. Dr. Ortmann’s sections of 
Flabellum, it is true, show an external layer but with a cir- 
cumferential dark linet, indicating that this layer itself was 
formed under a fold. I have already suggested that this discre- 
pancy between Dr. Ortmann’s sections and those of Dr. Fowler, 
Mr. G. C. Bourne, and Dr. von Koch can be explained by sup- 
posing that, in the case of his specimens, there had been a bagging 
of the soft parts over the rim of the epitheca which would cause 
it to grow as a fold. That some specimens do thus bag over we 
know from Moseley’s account of Flabeliwm (Chall. Rep. u. p. 162, 
1881). But this folded rim is not exactly what is wanted. We 
should have expected a simple rim of epitheca without any dark 


* In my former paper I described other results of this effort to grow, viz. 
the bagging of the polyp or even its overflowing over the rim of the epi- 
thecal cup. 

+ Zool. Jahrb. iv. (Syst.), pl. xviii. fig. 9 (1889). 


MR. H. M. BERNARD ON RECENT PORITIDA. 135 


line outside the first beginnings of the septa. The fact that 
this is not found—cf,, e. g., Dr. Fowler’s sections (Q. J. M.S. 
XXvill.)—required explanation; and that probably lies in the 
fact that the puckering of the expanding ectoderm in the early 
stages of the epithecal cup extends beyond its rim, and that 
consequently, when this puckered skin secretes its exoskeleton, 
the latter is puckered or folded from the first. 

The last matter to which I wish to refer is still more important, 
and, moreover, it brings us back to the main subject of our paper, 
viz., the affinities of Porites. One of the special difficulties im 
dealing with the morphology of the Madreporaria lies in the 
fact that, for precision’s sake, we have to idealize the parent 
polyp and picture to ourselves the possible transformations of its 
skeleton, as if it remained stationary. But, as a matter of fact, 
buddimg and colony-formation come in to complicate matters 
greatly. This, therefore, we must face as a difficulty in the way 


A B Cc D 


A and B. Hpithecate stages; the septa developed in A, become exsert in B. 
C. Thecate stage; the exsert septa replace the epitheca, which becomes 
vestigial. D. Diagrammatic section of Porites. 


of our line of argument. We have, for instance, assumed that 
the epithecal cup became vestigial because it was rendered 
unnecessary by the rising up of the septa (fig. 1) above the edge 
of the epithecal cup to form an internal theca, which supplied in 
every way a stronger and better-defended retreat for the polyp 
than the epithecal cup itself with its edges tending to be filmy 
and friable. Now, while a comparative study of the different 
forms of calicle leads me to believe that this is actually what took 
place, yet, when the habit of budding and colony-formation is 
taken into account, we are forced to ask whether a reason for 
the degeneration of the epitheca might not also be found some- 


136 MR. H. M. BERNARD ON RECENT PORITIDA. 


where in this latter. That such may indeed have been the case we 
know from the fact that species of Alveopora occur in which the 
lateral expansion of the colony is so pronounced that the usually 
conspicuous epitheca becomes little more than a film protecting 
the coral from the substratum, although there are here no exsert 
septa out of which to form an internal theca in the manner 
shown in the diagram. The same can also be shown in the 
genus Goniastrea, which multiplies by what is called fissiparity. 
Two prominent septa mark off the skeleton of the bud*. The 
skeleton of the colony is here again septate, and the epitheca is 
flattened out by colony-formation, that is, not in the way shown 
in the diagram. 

This point was not evident in my former paper, even though 
T left it undecided whether Porites was to be regarded as related 
to Madreporide or to Alveopora. It was quite clear that the 
epitheca of Porites was flattened out, and that the theca was 
therefore internal (fig. 1, D); but I saw only two ways in which 
this could have occurred, and in both the epitheca was slowiy 
replaced in an essentially similar way, viz., by the rising up of an 
internal theca, formed by the septa becoming more and more 
exsert. The theca of Porites might, I thought, be either a 
secondary modification of that of the Madreporide by the per- 
foration of the lamellate septa, or an independent development 
from a form like Alveopora with horizontal spine-like septa. In 
this latter case, as the epitheca flattened out, the spines would 
become vertical and form the vertical “ trabecule ” of Porites. I 
now see, however, that the epitheca might be flattened out in the 
process of colony-formation, when the skeleton of the bud is 
marked off by the meeting of septa which cut off a portion of the 
parent calicle. 

We have, then, three apparently possible origins of Porites. 
Of these we may, I think, safely dismiss this last supposition, 
viz., that the flattening out of the epitheca was due to the rapid 
lateral budding of some fissiparous coral. Such an origin would 
give us no explanation of the radial series of “trabecule”’ or of 
the thick intervening walls. 

Returning, then, to the main alternatives, we have to decide 

* This method of budding may be compared with that described in a 


former paper (Journ. Linn. Soce., Zool. yol. xxvi. p. 495, pl. 33. fig. 10) as that 
of an Astreeid ! 


MR. H. M. BERNARD ON RECENT PORITIDA. 137 


whether the internal theca of Porites has been developed out of 
the exsert septal spines of a Favositid, or is a secondary modifi- 
cation of that of the Madreporide, with its ring of lamellated 
septa. 

Recent work with both Goniopora and Porites has led me to 
the conclusion that this latter view is the correct one. The 
septa in both these two genera were once purely lamellate. Proof 
of this can be seen in the fact that, both in Porites and in 
Goniopora, every transition can be tound between the forms with 
almost purely lamellate septa and others with purely “trabecular” 
septa. Further, in a great many forms, the cost round the 
growing edges run out as vertical lamelle to the rim of the 
epitheca, that is, just where primitive conditions might be 
expected. 

This conclusion is not only of permanent importance for the 
solution of the problem we have in hand, but it deserves the 
special attention of all students of Stony Corals, because it relieves 
them for ever of the “ trabecula ” as a unit of morphological value. 
The “ trabecula,” which is in reality merely so much formative 
tissue, was brought into the system by the theoretical scheme 
of Madreporarian tectonics put forward by Milne-Edwards and 
Haime, a scheme which Miss Ogilvie * has recently endeavoured, 
by considerable emendation and amplification, to place on 
the surer basis of extended histological research. How great a 
snare it has been I have already shown f in the case of Montipora, 
which, on account of its “trabecule,” was ranked by Milne- 
Edwards and Haime among the Poritide. The “trabecule” of 
Montipora, that is, if what I called the trabecule of that genus 
are what Milne-Edwards meant, turned out on examination to 
be very different from those of Pordtes (see further on this point 
below). Again, if the trabecule in Porites and Goniopora, 
in which genera of all others they appear to play the most 
important part, can yet be shown to have no real morphological 
value, their case finally breaks down. 

My own experience is as follows. I began work first with 
Goniopora, its larger calicles admitting of easier examination. 
At the outset the “trabecula” was accepted as a morphological 


* Phil. Trans. vol. 187, 1896. 
ft Ann. & Mag. Nat. Hist. xx. 1897. 


138 MR. H. M. BERNARD ON RECENT PORITIDA. 


unit. The following reasons seemed to justify this acceptance :— 
(1) Well-developed “ trabecule ” occur in the walls of many 
species. (2) The pali appear to be the tips of others. (3) A 
vertical section through a corallum frequently shows it to have 
been built up of long nodnlated threads (trabecule) running in 
the line of growth and joined together at intervals by cross- 
pieces arranged parallel with the surface: this, however, is truer 
of Porites than of Goniopora. (4) In some forms there appeared 
to be a regularity in position and arrangement of the trabecule 
which suggested their having real value. 

After examining a great number of specimens, I reconstructed 
on the simplest possible plan an ideal primitive skeleton of a 
Goniopore built up of trabecule (see fig. 2). But the longer 
the actual specimens were studied with this hypotketical ancestral 


Fig. 2° 


Ideal arrangement of the “ trabecule,” if regarded as morphological units, 
necessary to explain the skeleton of a Goniopore, the columellar tangle 
being omitted, A, in ground plan: B, in vertical section ; ~, central pali. 


form, the more impossible it became. The meshes of the lattice- 
work were always pores, often very irregular in size and arrange- 
ment, in otherwise lamellate septa. Surely some forms would 
have retained the rectangular lattice-work with the trabeculz per- 
sisting in their primitive importance. But no such condition was 
found. Then, again, the pali failed as tips of growing trabecule. 
They were plates when the septa were but slightly perforated, 
and were only tips to the narrow divisions between the large 
perforations in other cases. Lastly, the finding of the growing 


MR. H. M. BERNARD ON RECENT PORITIDS. 139 


edges, already mentioned, in which lamellate cost ran out to 
the rim of the epitheca, finally convinced me that the so-called 
“trabecular” septa are merely perforate lamellate septa. 

We have thus reached an important stage in our enquiry as 
to the position of the Poritide: their so-called “ trabecular ”’ 
structure belongs to the terminology of the past. Their thecz 
were originally built up of lamellate plates like those of the 
Madreporids, and the perforation of these plates has to be con- 
sidered as a secondary characteristic. To this difference between 
the Madreporid and the Poritid septum we shall return when we 
have discussed the next most striking contrast between the two 
families, which may be stated as follows :— 

In the Madreporide, except in JZontipora with its immense 
development of the cenenchyma, the thecz are tall and conical. 
In the Poritide, on the contrary, the thec are low and shallow. 
The septa in the latter are therefore not only perforated, z. e. 
poor in quality, but also poor in quantity, that is in size. 

This contrast is shown diagrammatically in fig. 1, p. 135, in 
which C and D are intended to represent individual calicles 
(ideal parent calicles) of a Madreporid and of a Porites respec- 
tively. In the former, the septa rise above the flattened epitheca 
to form a new theca, being mutually supported by synapticule 
which would project from the plane of the figure in the dotted 
areas. In the latter we have the low basal skeleton of Porites ; 
the septa with their synapticule being together reduced to a 
reticulum. Can any explanation be given of these differences ? 
I think so. The diagrams of themselves seem to suggest that 
the conditions found in Porites are due to arrested development. 
The suggestion is therefore made that these swarms of minute 
polyps, which are so ubiquitous and appear in such vast numbers 
that they are reckoned among the principal builders of the coral- 
reefs, may be regarded as Madreporids arrested at an early stage 
in their developwent. ‘This is, in fact, the position I have found 
myself compelled to assign to the Poritide in the Catalogue of 
the British Museum Madreporaria *. 


* Vol. I. (by the late George Brook) deals with Madrepora. Vols. II. & III. 
contain the Madreporid genera Turbinaria, Astreopora, Montipora, and Ana- 
eropora. Vol. IV., which is nearing completion, contains the two Poritid 
genera Porites and Goniopora, somewhat extended (see below, pp. 148-148). 

LINN. JOURN.—ZOOLOGY, VOL. XXVilI. rl 


140 MR. H. M. BERNARD ON RECENT PORITIDA. 


Let us then see what are the arguments in favour of this 
suggestion. For the moment, dealing only with Porites, we find 
the polyps, like their calicles, small and degenerate, 7. e. little 
grown and with only twelve tentacles. Their skeletal secretions 
are purely basal, and the animals retreat down upon them rather 
than into them*. Thus, in relation to the animal, the skeleton 
is but feebly developed, so feebly indeed that the coralla rarely 
have any elasticity or beauty of form. They are, for the most 
part, merely rounded massive concretions such as could be built 
up of small flat dises. 

It may perhaps be objected that such a poorly developed and 
rudimentary skeleton might also be primitive, but this is certainly 
not the case here; for a glance at fig. 1 shows that the skeleton 
of Porites belongs to the highest known type, viz., that in which 
an internal theca has replaced the primitive epithecal cup. This, 
as above stated, I believe it could only have done by the internal 
thecabeing pronounced enough to replace the epithecal cup as 
a more eflicient refuge into which the polyp could contract. 
Hence we can only account for the internal theca of Porites by 
assuming that it was at one time tall and deep, forming with its 
jagged septal edges a stronger and better guarded receptacle for 
the polyp than the primitive epitheca. In other words, the 
theca of Porites must at one time have been tall and composed 
of lamellate septa, and the fossa, now shallow and quite incapable 
of containing the polyp, must at one time have been large enough 
to have allowed the whole polyp to smk down into its recesses 
(fossa, interseptal and intercostal spaces). The theca, from being 
a true calicle, has become, in Porites, a mere basal pedestal for 
the comparatively speaking tall polyp which secretes it, it bemg 
one of the peculiarities of Porites (and of Goniopora) for the 
polyps to rise high above their skeletons. 

The internal theca of Porites can therefore only be regarded 
as rudimentary. It is not a vanishing structure, but it belongs to 
the most specialized type of Madreporarian skeleton, secondarily 
arrested in its development. This interpretation is further 
confirmed by comparing the skeleton of Porites with that of 
almost any minute young single coral, such as is frequently found 

* Thurston describes the polyps of Porites which can no longer retreat into 


their calicles as protecting themselves when exposed by a layer of slime. 
Bulletin of Madras Government Museum, No. 3 (1895), p. 93. 


MR. H. M. BERNARD ON RECENT PORITIDS. 141 


on the corroded bases of large stocks. The septa of such young 
forms are seen to be irregular and granular, and, as a rule, to rise 
but little above the columella-tangle. The fossa is consequently 
shallow. Further, the skeleton has always naturally to pass 
through a stage when it is small and incomplete, as compared 
with its secreting polyp, which rises in a column above it. 
Arrest at such a stage would account for the polyp in Porites 
rising high above its shallow calicle. 

Lastly, this argument is quite in keeping with the tendency 
to bud very early, which I have already noted as characteristic 
of recent Madreporide*. The conditions in Porites are simply 
explained if we assume them to have acquired the habit of 
budding still earlier, ¢.e. when the skeleton is quite immature. 

So far, then, as the genus Porites, with its minute polyps and 
feebly-developed skeletons, is concerned, the above arguments 
appear to me to be fairly conclusive as to their relationship with 
some primitive Madreporide as fixed young forms. 

The chief qualification of this conclusion would tend towards 
suggesting a polyphyletic origin to Porites. There is no reason 
to suppose that this arrest of development happened only once. 
If it is possible at all, it is likely to have taken place more than 
once and at different stages in the phylogenetic development of 
the Madreporide. Indeed, we might ask whether it is absolutely 
necessary to assume an exclusively Madreporid origin. ‘These 
points must be left for future discussion. They require a much 
wider survey of forms than we now possess, and a more profound 
insight imto the essential morphology of the Madreporarian 
groups. 

The genus Goniopora, Q. & G.—The first known forms of this 
genus led to their being placed near, and even among, the 
Astreide (Milne-Edwards & Haime). These last-named authors 
kept the name Goniopora for forms with thick-walled, rather 
shallow calicles, but gave the name Porastrea to those with thin 
walls. This latter name explains itself. Dana first placed the 
genus with Pordtes, with which it agrees in almost every respect 
except in size of calicles. The only difference I have myself 
been able to discover can be referred simply to increased growth. 
A third cycle of septa appears, which may be merely rudimentary, 


* Brit. Mus. Madrep. vol. iii. p. 12. 
IU 


142 MR. H. M. BERNARD ON RECENT PORITID®. 


but is most often well developed and with a fourth cycle indicated. 
The lamellate character of the septa is more evident in the larger 
septa of Goniopora than in the smaller septa of Porites, the 
perforations being about the same size. Thus the perforations 
as such are of less account in Goniopora than in Porites, and 
the vertical section is more of an irregular reticulum than a 
regular lattice-work, as it frequently is in Porites. The close 
relationship between Porites and Goniopora suggested by Dana 
has been universally accepted, Milne-Edwards and Haime aban- 
doning their former position as soon as possible after Dana’s 
work appeared. There has, however, been a tendency to limit 
the genus too much to forms which have tall thin walls and 
consequently deep calicles. As a matter of fact, the range of 
variation is very great; and the collection in the Natural History 
Museum contains many new and beautiful forms. 

Admitting this genus, then, as a near ally of Porites, the much 
greater size of its calicles raises an objection to our conclusions. 
Porites, by the small size of its calicles, might easily be accounted 
for in the way above suggested as fixed young forms. But 
how shall we explain the much larger size of the calicles of 
many Goniopores ? mits 3 

It seems to me that these need abt present any great difficulty. 
Passing over the possibility above suggested, that in these 
Poritidea we may have a group made up of fixed young forms 
of several different corals, whose separate ancestries it would 
now be extremely difficult to unravel *, there need be no 
difficulty in deducing the Goniopores from Porvtes directly ; and 
this seems, for the practical purposes of classification, the simplest 
course to pursue, provided, however, we do not lose sight of the 
above-mentioned possible polyphyletic origin. 

I propose, then, to regard the Goniopores as merely enlarged 
Porites, a kind of giant race which retains the skeletal habit of 
Porites. If once that habit became fixed, there is no reason 
why further growth should not simply enlarge it without 
necessarily running it into ancestral Madreporidan lines. 

In the present state of our knowledge, I regard anything like 
certainty in these relationships as unattamable. What I have 


* Here it is of great interest to note that Dana himself suggested that 
Goniopora might occupy a position in the Caryophyllacea corresponding to that 
which Porites oecupies in the Madreporacea (Zoophytes, p. 407). 


MR. H. M. BERNARD ON RECENT PORITIDS. 143 


here sketched out is intended to serve merely as a working 
hypothesis. It may be that a closer study of fossil forms will 
reveal to us new possibilities. In the meantime, however, we 
have to analyse the structures of the forms which we have at our 
disposal, and to arrange them as best we cau io a natural order. 


Several other genera, recent and fossil, were boldly classed 
among tne Poritide by Milne-Edwards and Haime. Any corals 
showing the “trabecular” structure were placed in the family, 
which was divided into two subfamilies, Portting and Monti- 
porine. 

The Poritine contained the genera Porites, Rhodarea, Gonio- 
pora, Litharea (foss.), Protarea (foss.), Alveopora, Microsolena 
(toss.), Aeandrarea (foss.), Coscinarea. In addition to these, 
Porites was divided by Verrill into Porites and Synarea; by 
Duchassaing and Michelotti into Porites, Neoporites, with anew 
genus Cosmoporites; while Quelch added another, Mapopora, and 
described a new genus, Tvchopora, as closely allied to Rhodarea 
and Goniopora. 

Any adequate discussion of these genera should be preceded 
by a detailed anatomical account of Porites and Goniopora, 
showing their ranges of variation. Such an account is in course 
of preparation. But in the meantime enough has already been 
said to make the following short notes on the claims of the various 
genera to a place in the family intelligible. Further, of these 
genera | propose only to refer to those which I know at first 
hand. I am not sultficiently acquainted with the fossil forms 
(which require a much closer study than I have yet been able to 
give to them) to desire to offer any opinion as to their claims 
to a place in the family. 

Synarea, Verrill.—This genus way separated from Porites by 
Dr. Verrill *, on the suggestion of Milne-Edwards and Haime, to 
contain certain forms in which the ealicles are quite filled up by the 
intercalicular skeleton, 2. e. which show a mere variation in the 
depth of the calicle. My own study of the variations in Porites 
makes it doubtful whether this is always even a specific, much 
less a generic distinction. 

Napopora, Quelch.—In the genus Porites there exist species 


* Bull. Mus. Comp. Zool. vol. i. (1854) p. 42. 


144 MR. H. M. BERNARD ON RECENT PORITID A. 


in which the thickened walls show tendencies to form extra 
ridges and hillocks closely resembling those of Montipora ; 
indeed, but for the calicles, such specimens would certainly be 
classed in that genus. These were not known when Quelch 
made his new genus*. There are, however, a good mary in the 
British Museum collection. It seems to me as impossible to 
separate them from Porites because of this rising of the wall, as 
it is to separate Synarea on account of the sinking of the wall. 
Tf the calicles are built on the same plan, variations in height 
of the wall can hardly be considered ag generic distinctions. 

Both these genera therefore, Synarea and Napopora, are 
merged in the genus Porites. 

Rhodarea.—This genus was established by Milne-Edwards and 
Haime t, and was thought to differ from Goniopora in that the 
latter had tall thin walls and spongy columella, while Rhodarea 
had thick low walls with a rosette of pali rising off the columella. 
These differences are only slight variations on the same essential 
structure. Even in individual stocks, the development of the 
pali is always the inverse of that of the walls; where walls are 
low, the pali are high and conspicuous. In any extended survey, 
it is found absolutely impossible to separate the specimens on 
these lines. I propose therefore to merge this genus into 
Gontopora. 

Tichopora, Quelch {.—The union of Gonzopora and Rhodarea 
forms a group which absorbs this proposed genus, in that it came 
somewhere between them, differing but slightly from either. 

Alveopora.—This genus was the subject of my former paper 
(1. ¢.), so that I need only repeat the conclusion at which I have 
arrived, that, in spite of its occasional resemblance to individual 
forms of Goniopora, as a primitive type of coral it is yet very 
far removed from the Poritide, which must rank among the most 
specialized of the Madreporaria. 

Coscinarea, M.-E. & H.—Very little is known of this genus. 
Only one species seems to be known. It was first figured in 
Savigny’s ‘Descr. de Egypte,’ pl. v. fig. 4, 1809, and named 
Meandrina. These are very puzzling figures, and hardly suggest 


* Ann. & Mag. Nat. Hist. xiii. (1883) p. 296. 
t C. R. xxix. p. 259 (1849). 
+ Chall. Rep. xvi. (1886) p. 188. 


MR. H. M. BERNARD ON RECENT PORITID®. 145 


any affinity with Porites. A second figure, however, is given, 
evidently of another specimen, by Milne-Edwards and Haime 
(Ann. d. Sci. Nat. 3 ser. ix. pl. v. fig. 2, 1848). This was first 
named Coscinarea Botte, M.-E. & H., but afterwards, being 
identified with Savigny’s figure, became Coscinarea meandrina. 
Dr. Klunzinger has fortunately re-discovered and photographed 
it as Coscinarea monile of Forskal, and regards it as a Fungid*. 

The remaining subfamily of the Poritide, M.-E. & H.—the 
Montiporine—consisted of two genera, Montipora and Psammo- 
cora, Dana. 

Montipora.—This genus is one of those which Milne-Edwards 
and Haime forced among the Poritide solely on account of its 
“trabecular” septa. I have already analysed the skeleton of 
Montipora and compared it with that of Porites. I was, how- 
ever, all the while conscious of some misunderstanding; the 
confusion lay in the word “ trabecula.” I endeavoured to show 
that the trabecule of Porites were not the same as the vertical 
rods which form such a conspicuous element in most sections of 
Montipora, the secondary development of which could be traced 
within the genus. I am now, however, not satisfied that Milne- 
Edwards and Haime meant these vertical rods at all. The word 
‘“‘trabecula’’? must have meant for them both vertical and 
horizontal rod-like skeletal processes; and the trabecule of 
Montipora were, for them partly at least, the short blunt septal 
teeth, and not exclusively the long nodulated rods which, in the 
sections of some forms, so closely resemble the vertical rods in 
sections of Porites. Their express words, ip discussing the claims 
of Montipora and Psammocora to be classed among the Poritide, 
are: “ La structure trabiculaire de leur polypier et principale- 
ment de leurs cloisons ne peut laisser aucun doute sur leurs 
véritables affinités”’z. Further, their description of the septal 
apparatus of Alveopora as “trabecular” leaves httle doubt that 
in their use of the word they meant either vertical or horizontal 
rod-like skeletal matter. 

It is not surprising, therefore, if the word “ trabecula” (“ pou- 
trelle”) has caused confusion, for this indefinite application of 


* ‘Corallenthiere,’ iii. 1879, p. 78. 
+ Ann. & Mag. Nat. Hist. xx. 1897; cf. also Introd. vol. iii. Brit. Mus. Mad. 
{ Ann. Sci. Nat. 3 ser. xvi. p. 54 (1851). 


146 MR. H. M. BERNARD ON RECENT PORITIDE. 


the term is quite inconsistent with the original description, where 
the trabecule are said to have “l’aspect de petites tiges noueuses, 
étranglées d’espace en espace” *. The septal teeth of Aontipora 
do not conform to this description, while the vertical rods in that 
same genus and in Porites, as a rule, do. - 

For the future, however, the word “ trabecula’ 
merely so much formative tissue, and, if it and its adjectives are 
retained at all, they can only have descriptive significance. Their 
intrusion into the morphology of the Madreporaria has so far 
only led to confusion. I must again, therefore, express my con- 
viction that the more exact description of the trabecula given us 
by Miss Ogilvie is solely of histological importance, and cannot 
again give it any weight as a morphological unit. 

Returning, however, to the genus Montipora, ry researches have 
led me fully to endorse the widespread opinion that it belonged 
to the Madreporide, and not to the Poritide. A comparative 
study of their skeletons showed them, as I thought, to be quite 
distinct. It is therefore not without interest to note that the 
conclusion we have now arrived at as to the origin of Porites has 
ence more brought them in a manner together. Both may be 
called Madreporids in which the pelyps are fixed at a very low 
stage of development, but the processes in the two cases are in 
strange contrast. In Porztes the whole organization, polyp and 
skeleton, never gets beyond the early stage at which their 
development is arrested. But, in Jontipora, the dwarfing of the 
polyps seems to have been due to the excessive development of 
the skeleton as such. ‘The Madreporarian skeleton, in fact, 
reaches its highest level of specialization in this genus, though 
at the expense of the polyps. The dwarfing of the polyps in the 
two genera gave some colour to the classification which placed 
them in the same family. Indeed, forms occur in both genera 
which it is not easy at first sight to assign to the one or to the 
other. Milne-Edwards’s distinction, that one has an interstitial 
ceenenchyma and the other has not, does not hold good, for there 
are many Porites with such thick walls that no difference in 
this respect can be recognized. The real distinction is found in 
the calicles. The septa of Montipora, composed of six vertical 
rows of small horizontal teeth round a deep fossa, are unknown 


” represents 


* ‘Les Coralliaires,’ vol. i. (1857) p. 32. 


t 


MR. H. M. BERNARD ON RECENT PORITID®. 147 


in Porites; while, on the other hand, the columella-tangle with 
the paliform granules or rods rising from it, which are character- 
istic of Porites, are never seen in Montipora. 

Psammocora, Dana, is the last genus which Milne-Edwards 
and Haime placed in their Montiporine subfamily of the Poritide. 
Dana, from a study of the living coral and on account of its 
skeletal structure, placed it among the Fungide. Its “ trabe- 
cular ” structure, however, compelled Milne-Edwards and Haime 
to transplant it. So far as I can see, beyond the granular inter- 
rupted edges of the septa, which thus appear to be built up of 
“trabecule,” this genus has no claim whatever to be classed 
anywhere near Porites, and I agree with Dr. Klunzinger, who 
replaces it among the Fungide, in removing it from the 
Poritide. 

Neoporites and Oosmoporites, Duchassaing and Michelotti *.— 
We can discuss these suggested genera together: the differences 
between them are slight, and the real question is whether they 
should be separated from Porites at all. The type of the sug- 
gested Neoporites may be taken to be the West-Indian Porttes 
astreotdes, Lamarck, which, with a few other West-Indian forms, 
differs from all the recorded Porites in having deeper calicles 
and either no pali or else mere traces of them (‘“ pallulis nullis 
vel subevanidis”’). This absence of pali and greater depth of 
fossa are certainly remarkable characteristics. But I find myself 
compelled to agree with Dr. Briiggemann in claiming them to 
be true Porites. On the one hand, it may be urged that 
the pali are an essential characteristic of Porites and Gonio- 
pora; and here we have forms in which the pali have been 
secondarily obscured or even suppressed; hence the need for 
establishing a new genus, and if so the name Neoporites is 
most felicitous, because it betokens an advance on the main 
genus; further, all the forms which might be grouped as 
Neoporites, and which have been so far described, occur in the 
West Indies, 7. ¢. they have a certain geographical unity which 
greatly supports the structural evidence in favour of their 
being a new geueric development. On the other hand, I would 
suggest that, if these specimens are removed from Porites 
on account of the absence of pali, they should for the same 


* *Coralliaires des Antilles,’ Suppl. 1864. 


148 MR. H. M. BERNARD ON RECENT PORITIDS. 


reason be removed from the family. Further, a review of 
the structural variations not only within the genus, but even often 
on one and the same specimen, reveais a correlation between the 
wall and the pali, so that when the one is specially well developed 
the other is correspondingly aborted. In Goniopora we have 
the species G. Stokesi, in which the walls are high and the 
pali are either absent or else only hinted at. And lastly, forms 
occur, and will shortly be described, in the Indo-Pacific area 
which show this same variation, viz., absence of pali. There 
seems to me, then, no special advantage in separating a few 
specimens of Porites because the deepening calicles have led to 
the partial or complete suppression of the pali. This variation 
seems to me not too great to be comprised within the range 
embraced by the genus. 

In connection with what has been said above about the relation 
of Porites to the Madreporide, the resemblance between these 
‘“ Neoporites” of Duchassaing and Michelotti, in which the pali 
are absent from the deep central calicles, and Jlontipora is very 
interesting: it shows how along two different lines almost the 
same structure may be reached. That these forms are not 
Montiporids may be gathered—(1) from their habit, which is 
more like that of Porites than of Montipora; (2) from the walls 
being more boldly reticular than in the majority of Montipores ; 
(3) in the presence of a columella-tangle slowly filling up the 
fossa, this being characteristic of the Poritide but not of the 
Madreporide (excl. Turbinaria) ; (4) in the traces of pali in the 
shallower young calicles; (5) in the twelve septa nearly equal in 
size, whereas in Montipora six, with a rudimentary second cycle, 
is the usual septal formula. 


SUMMARY. 


The foregoing pages contain a preliminary instalment to a 
revision of the classification of the Madreporaria by Milne- 
Edwards and Haime, which has been rendered necessary by 
recent advances in our knowledge of the morphology of the 
coral-skeleton. 

The object of the paper is to record the results, obtained 
during my work of cataloguing the specimens in the Natural 
History Museum, as to the position of Porites among the 
Madreporaria. A brief sketch of the history of the question 


MR. H. M. BERNARD ON RECENT PORITIDA. 149 


led to a review of the present situation, in which the funda- 
mental theory on which the existing system of classification 
which we owe to Milne-Edwards and Haime rests was criticised in 
the light of recent research. This criticism entailed a re-state- 
ment, with slight amplification, of the author’s phylogenetic 
scheme, along the lines of which it is maintained the classifica- 
tion of the Stony Corals will have, for the future, to proceed, 
2. é. until it is again superseded by further advances in morpho- 
logical science. The chief new points of interest with regard 
to this scheme related, (1) to the origin of septa; (2) to the 
various possible methods in which the primitive external epi- 
thecal cup may have been flattened and become replaced by an 
internal skeleton. 

This last discussion brought us naturally to the object of the 
paper, viz., to enquire along what lines of development Porites 
obtained its peculiar internal andso-called “trabecular”’ skeleton. 

The conclusions arrived at were :— 

(1) The “trabecular” septum is only a misleading name for 
perforated lamellate septum. 

(2) The Poritid skeleton can be explained as an immature 
Madreporid skeleton, arrested in its growth by very early 
budding. 

(8) As this may have happened more than once, Porites may 
be polyphyletic in origin. 

The paper concludes with a brief discussion of the various 
genera which have been from time to time united with Porites ia 
the same family or else separated from it as generically distinct. 
The revision suggested leaves the family Poritide as Dana left it, 
with only two genera, Porites and Goniopora,—Porites enlarged 
by the absorption of Synarea, Napopora, Neoporites, and OCosmo- 
porites, and Goniopora also enlarged by the merging with it of 
Rhodarea and Tichopora. 


My best thanks are due to my friend Prof. F. Jeffrey Bell for 
much kindly assistance, and for the warm interest he has 
_ taken in these investigations, not only as the Officer in charge 
of the Collections on the study of which they are based, but also 
in the interest of zoological science. 


150 MESSRS. J. C. MELVILL AND R. STANDEN ON 


Report on the Marine Mollusca obtained during the First Expe- 
dition of Prof. A. ©. Haddon to the Torres Straits, in 1888—89. 
By James Cosmo Metvi11, M.A., F.L.S., F.Z.S., and Ropert 
SranpEN, Agsistant-Keeper, Manchester Museum. 


[Read 16th February, 1899. ] 
(Puates 10 & 11.) 


Earty in 1898 Prof. Alfred Cort Haddon, before starting upon 
a fresh journey of exploration to New Guinea, the coasts of 
tropical North Australia, and Queensland, favoured us with the 
request that we would take charge of all the Marine Mollusca 
collected at low tides, er dredged, during his first expedition to 
the same region, ten years previously, on the understanding that 
we would catalogue them and decribe any new forms. This 
large mass of material had been for a long period lying at 
Cambridge, almost untouched, only a very few species having 
been identified by the Rev. A. H. Cooke. 

The general condition of the specimens contained in the col- 
lection is, as might be expected, variable; but, though many 
of them are only in a fragmentary state, and the larger pro- 
portion of the Pelecypoda are mainly represented by single valves, 
they are in only rare instances past recognition, and, with the 
exception of an exceedingly small residuum, we have succeeded 
to our satisfaction in the work of identification. 

Some few mdeed, mainly Polyplacophora of three or four 
kinds, also Haliotide and various Cypree, are preserved in 
spirit, but all these Mollusca are well known, both anatomically 
and systematically. 

The collection is, we consider, of more than usual interest, 
since its component parts differ in several notable particulars 
from the gatherings previously made in the same neighbourhood, 
thus tending to prove the extraordinary richness of molluscan 
life to be found there. And the area is by uo means large, geo- 
graphically speaking. Situate Long. 142° to 144° E., and Lat. 9° 
to 11°S., it is hardly more than 75 miles from the mainland of 
New Guinea, or at all events Saibai Island, to Cape York, 
N. Australia ; this narrowing of the passage between the Arafura 
and the Coral Seas, through which the South Equatorial Current 
pours its waters, being some 90 miles in width, and universally 
known as the Torres Straits. 


THE MOLLUSCA OF TORRES STRAITS. 1155) 


The whole of this area is contained within the great Indo-Pacific 
Marine Province, as proposed by S. P. Woodward, 1856, though 
the merging of that Province with the Australian is, speaking from 
the molluscan point of view, hazily defined only and, naturally, 
gradual. This Province is the largest by far, and likewise the 
most prolific in marine life of all, and though almost unwieldy, 
it should nevertheless, in our opinion, be still further extended 
so as to include Southern Japan, treated by Woodward and all 
who have followed him, as well as by Agassiz, as part of a 
separate region. The Indo-Pacific Province would then extend 
from the Hast coast of Africa, north of the Tropic of Capricorn, 
to the Red Sea, Persian Gulf, and Arabian Sea, round the whole 
coasts of India and its adjacent islands, southwards so as to 
include Madagascar, Mauritius, Bourbon, to Malaya, the East 
India Islands, and China coasts as far as, and inelusive of, 
Southern Japan, also taking in tropical Australia, and finally the 
Pacific Islands with Hawau. Though so vast, we cannot see how 
with advantage this tract can be lessened or modified; and one 
is strengthened in this view when the distribution of many of 
the tropical Marine Mollusca is considered. The range, for 
instance, of the most abundant, e. g., Cyprea helvola, L., Nassa 
arcularia, Li., or Terebellum subulatum, L., being that of the 
Province, even extending beyond its limits into the subtropical 
waters of Natal, or of Queensland and New South Wales. 

Many of the species in the following Catalogue have this 
wide distribution ; and with reference to this fascinating subject, 
it may be not out of place to refer briefly to the instructive 
remarks made by Prof. HE. von Martens, when enumerating 
the Mollusca of the Mergui Archipelago*, a few years since. 
He mentions that out of nearly four hundred species, only 
one (Natica unifasciata, Lam.) was known to have occurred 
in the New World, besides three, also found on Atlantic shores, 
and even these were species liable to spread by means of drift- 
wood and other agencies. 

There can be little doubt that an unusually large number 
of endemic foims occur in the region bounded north-westward 
and noithward by the Philippines and Ladrone Islands, west- 
ward and to the east by the Arafura Sea and New Caledonian 
archipelago respectively, the Torres Straits forming part of its 
southernmost boundary. 


* Journ. Linn, Soc. Zool. vol. xxi. p. 157 (1889). 


152 MESSRS. J. C. MELVILL AND R. STANDEN ON 


From a scientific point of view, the following four Expeditions 
are the most iaportant that have been made in past years to 
this region. 


1. THE VoyaceE or H.MS. ‘ Fy.’ 


Dr. J. B. Jukes*, in 1842-46, exploring part of North 
Australia, with the South Papuan coasts, assiduously collected 
Mollusca. These were mostly described by Arthur Adams and 
Reeve, while Dr. J. E. Gray, in an appendix to Jukes’s narrative, 
gave diagnoses of several important discoveries, e. g. Voluta 
Sophia, Cyprea Compton, &c. 


2. Tur VoraceE or H.M.S. ‘ CuaLLENGER.’ 


The ‘ Challenger’ Expedition 7, 1873-76, dredged as follows :— 

Station 184, August 29, 1874. E. of Cape York, N.E. Aus- 
tralia, 1400 fathoms; Globigerina-ooze. 

Stations 185 & 185°, Aug. 31, in the same locality, from 128 
to 135 fatboms. 

Sept. 7 & 8. Torres Straits and Flinders’ Passage, 3-11 
fathoms. 

Station 186, Sept. 8, 1874. Wednesday Island, Cape York, and 
Albany Island, 83-12 fathoms ; coral-mud. 

Station 187, Sept. 9, 1874. W. of Cape York, 6 fathoms. 

Station 188, Sept. 10. Off the 8.W. of Papua, 28 fathoms ; 
ereen mud. 

Most of these soundings were successful, and a profusion of 
new Mollusca of great interest the result. 

We are surprised that so few of these are in the collection now 
before us, as in some instances almost the same localities would 
seem to have been searched and traversed. 


8. Tur VoYAGE oF THE ‘ CHEVERT. 


This expedition started from Sydney early in 1876, Mr. John 
Brazier being the malacologist attached to the staff. He has 
published the results in a series of articles, dealing thoroughly 


* Narrative of the Surveying Voyage of H.M.S. ‘Fly, commanded by 
Captain F. R. Blackwood, R.N., in Torres Straits, New Guinea, and other 
Islands of the Eastern Archipelago, during the years 1842-46. By J. Beete 
Jukes. 

+ Report on the Voyage of H.M.S. ‘ Challenger’ during the years 1873-76. 
Zoology, vol. xiii. pp. 16-18; vol. xv. pp. 710-714. 


THE MOLLUSCA OF TORRES STRAITS. 153 


with the Gastropoda, but the Pelecypoda, so far as we are aware, 
have not yet been treated. Many new forms, especially amongst 
the minutiora, are described, but unfortunately not figured, the 
types all remaining in Australia, we believe mostly in the Aus- 
tralian Museum, Sydney. For our own part, we have been as 
careful as possible in the comparison of all our undetected forms 
with these descriptions. 

It is to be deplored that Mr. Brazier has not always used 
language the reverse of vague; still more unfortunate is it that 
as the Mollusca of the ‘Chevert’ Expedition were published 
before those of the ‘ Challenger,’ we are given to understand that 
they claim priority in not a few mstances over Dr. Boog Watson’s 
concise, clear, and admirably illustrated diagnoses. This increase 
of synonymy is much to he deprecated, and we would press for an 
insistence of the rule that no mere verbal description should 
suffice, but that a figure must be published simultaneously, before 
a new species be recognized. 


4. Tue Voyace or H.M.S. ‘ Aner’ *. 


The full and very interesting account of the Mollusca obtained 
during this expedition has been of the utmost service to us 
while preparing this paper. 

Indeed we are now, as upon many previous occasions, under 
a deep sense of obligation to Mr. Smith, who has compared with 
us all doubtful forms and given us every assistance at the British 
Museum (Natural History). 


The specialities of this collection will be dealt with in their 
proper place. Suffice to state here that of the twenty-four species 
we consider new to science, one, a Neritoid with some superficial 
resemblance to Vanikoro, must stand as the type of a new genus; 
and a remarkable Pholadomya, with almost equilateral valves, is 
also noteworthy. 

The arrangement adopted in the sequence of the following 
Catalogue is that of Paul Fischer, as given in the ‘Manuel de 
Conchyliologie,’ and we have also followed the same author to a 
ereat extent in the nomenclature. 


* Report on the Zoological Collections made in the Indo-Pacific Ocean 
during the Voyage of H.M.S. ‘ Alert,’ 1881-82. London (Brit. Mus.), printed 
by order of the Trustees, 1884. Mollusca, by Edgar A. Smith (pp. 34-116). 


154 MESSRS. J. C. MELVILLE AND R. STANDEN ON 


List oF Coitectine Stations *, Torres Straits, 1888. 

1. Fringing reef and shore, Thursday Island. 

2. 20 miles N.N.W. of Warrior Is]., 53 fathoms; broken shells 
and sand. Aug. 15, 1888. 

3. Channel between Saibai and New Guinea, 10-17 fathoms ; 
mud and rolled stones, dead shells; very few live animals came 
up in dredge, mainly sponges; of 3 dredge hauls 1 came up clean 
with Aleyonarians. Aug. 17, 1888. 

4. Between Ormans Reef and the “ Brothers Island,” 6-7 
fathoms. Aug. 18, 1888. 

5. Boydong Cays shore. Aug. 26, 1888. (N. Queensiand.) 

6. 2 miles west of Boydong Cays, 14 fathoms; coral-mud and 
dead shells. Aug. 27, 1888. 

7. Cockburn group, shore of small islet near the south of the 
three high islands. Aug. 27,1888. (N. Queensland.) 

8. Albany Pass, 10 fathoms; large quantity of sponges. 
Aug. 29, 1888. 

9. Prince of Wales Channel, 8 fathoms; clear rocky bottom 
and red seaweed. Sept. 15, 1888. 

10. Channel between Hammond Isl. and Wednesday spit, 
5 fathoms; sand and small rounded pebbles, dead shells. Sept. 15, 
1888. 

11. Fringing reef, Mabuiag. 

12. Channels between reefs, Mabuiag. 

13. Fringing reef, Mér (Murray Island). 

14. Channels between reefs, Mer. 


CATALOGUE OF THE SPECIES. 
PTEROPODA. 
CAVOLINIIDE. 


1. CAVOLINIA LONGIROSTRIS, Lesson. 
Station 5, Boydong Cays. 


GASTEROPODA. 
SIPHONARIID A. 


2. SIPHONARIA STPHO, Sow. 

Station 5, Boydong Cays ; Station 13, Murray Island. 

* Where no precise locality is referred to, it must be understood that the 
labels attached to those species contained nothing more definite than ‘‘ Torres 
Straits.’ 


THE MOLLUSCA OF TORRES STRAITS. 155 


ACTXONID 2. 


3. ACTON soLiIpuLus, L. 
Station 5, Boydong Cays. 


TORNATINID &. 


4, TORNATINA GRACILIS, A. Ad. 
Station 14, Mer. 


SCcAPHANDRIDA. 


5. ATYS DEBILIS, Pease. 
Station 2, Warrior Island; Station 5, Boydong Cays. 


6. Atys (Aticuna) cyniInpRica, Helbling. 
Station 13, Murray Island. 


7. CyLIcHNA ARAcHTS, Q. g G. 


BULLIDs. 
8. Burna Apamst, Menke. 
Station 13, Meér; Station 5, Boydong Cays. 


9. Buia PuncTuLATA, A. Ad. 
A West-American species (Panama, &c.), also New Caledonia 
(Hadfield), and various Australian localities (Cox, Angas, &c.). 


10. HamInea BREVIS, Q. & G. 
Station 2, Warrior Island. 


11. Hamrinea crocarta, Pease. 
Station 2, Warrior Island ; Station 5, Boydong Cays. 
APLUSTRID&. 


12. APLUSTRUM AMPLUSTRE, ZL. 
Station 18, Murray Island, 


TEREBRIDA. 


13. TexEBRA (SUBULA) MuUscARIA, Lam. 


14, TEREBRA (ABRETIA) AFFINIS, Cray. 
Station 18, Mér ; Station 2, Warrior Island. 
15. TereBra (MyvuRELLA) suBuLaATA, L. 
Station 13, Mer. 
LINN. JOURN.—ZOOLOGY, VOL. XXVII. 12 


156 MESSRS. J. C. MELVILL AND R. STANDEN ON 


ConID az. 
16. Conus mMarmorevus, L. 
Station 13, Meér. 
17. Conus (Coronaxis) minimus, L. 
Station 18, Mer (Murray Island). 


18. Conus (NuBEcuLA) stRIaTUS, L. 
19. Conus (DENDROCONUS) FIGULINUS, L. 


20. Conus (Lirnoconus) LitrERATUS, JZ. var. MILLEPUNC- 
TATUS, Lam. 


21. Conus (RHIzOcONOS) MUSTELINUS, Hwass. 
22. Conus (RuIzoconvs) viruLinus, Hwass. 
23. Conus (CHELYCONUS) RADIATUS, Gimel. 
24. Conus (TExTILE) canontcus, Hwass. 

25. Conus (HERMES) NUSSATELLA, L. 


26. Drittia ToRRESIANA, Sinith. 
Station 8, Albany Pass. 


27. Maneiita (GLyPHOsTOMA) RUGOsA, MWigh. 
Station 13, Mer. 


28. Manerria (CyTHara) GRaciis, Rve. 
Station 2, Warrior Island. 


29. Manernra (CYTHARA) PULCHELLA, Rve. 


30. Manernia (Cyruara) cHIONEA*, sp.n. (PI. 10. fig. 4.) 

M. testa ovato-fusiformi, delicatula, nivea, levigata ; anfrac- 
tibus, inclusis duobus apicalibus levibus, septem, oblique et 
fortiter costatis, costis nitidus perlevibus, interstitiis levigatis, 
parum nitentibus ; apertura oblonga, labro paullum effuso, levi; 
columella obscure denticulata. Long. 8, lat. 3°50 mm. 

Station 18, Mer (Murray Island). 

A white, fusiform, delicate and very smooth species, seven- 
whorled (inclusive of the two apical), the lower whorls being 
strongly obliquely ribbed, the ribs not crowded, the interstices 
smooth, but not shining; aperture oblong, outer lip slightly 
effuse, smooth ; columella in fine examples minutely denticulate, 
but worn in most of the examples seen. 


* yiwv, ShOW. 


THE MOLLUSCA OF TORRES STRAITS. 157 


We also have this delicate Mangilia from Thursday Island, 
collected by Mr. Arnold Umfreville Henn, in fine condition. 


OLIvID#. 
31. Oxtva (STREPHONA) EPISCOPALIS, Lam. 
An albino form only; slightly deformed in the centre of the 
last whorl. 
Station 5, Boydong Cays. 
32. OLIVANCILLARIA (AGARONIA) NEBULOSA, Lam. 
Station 18, Mer. 
MARGINELLID2. 


33. MARGINELLA (PERSICULA) ovuLUM, Sow. 
Station 18, Murray Island. 


VoLuTID&. 


34, MeEto piaDEMA, Lam. 


35. Votura (Autica) Ruckert, Crosse. 

Perhaps too near V. piperita, Sow., to be more than a variety ; 
but the red coloration is uniform in all examples we have 
inspected. 

Station 2, Warrior Island. 


86. Vortura (Avtica) rutTILA, Brod. 
Station 2, Warrior Island. 


37. Votuta (Avxtica) Sopuim, Gray. 

With V. nivosa, L., and V. Norrisi, Gray, this constitutes a 
peculiar group—V. Sophie being quite constant in its markings 
of rows of transverse blackish-brown spots—extending in range 
only from the west coast of Australia to the Torres Straits. 

Station 2, Warrior Island. 


38. Votura (ScAPHELLA) TURNERI, Gray. 
Station 5, Boydong Cays. 


Mirrip». 
39. Mrrra EPIScoPALis, L. 


40. Mirra (CHRYSAME) PEREGRA, Reve. 
12* 


158 MESSRS. J. C. MELVILLE AND R. STANDEN ON 


41. Mirra (CHRYSAME) RUBRITINCTA, five. 

Considered by some authors only a stunted variety of JL. ferru- 
ginea, Lam., but in our opinion entitled to good specific rank. 

Station 2, Warrior Island. 


42, Mirra (STRIGATELLA) DECURTATA, Five. 
Station 4, between Ormans Reef and the Brothers Island. 


43. Mirra (Turricura) corrvue@ata, Lam. 
Station 13, Murray Island. 


44, Mirra (VULPECULA) INTERMEDIA, Kiener. 
Station 13, Murray Island. 


45. Mirra (CASTELLARIA) MODESTA, Ave. 
Station 2, Warrior Island. 


46. Mirra (Pusta) prcHroa, Ad. & Rve. 

This species, with J. tricolor, Montr. I. Greffei, Cr., 
M. levizonata, Sow., and others, is merged by Tryon in Pusia 
luculenta, Rve., but, we think, erroneously. 


FascloLARIID&. 
47, FAScIOLARIA FILAMENTOSA, Lam. 


48. Larrrus poLyGonus, Gmel., var. TESSELLATUS, Kobelt. 
Characteristic examples of a form only hitherto reported from 
Mauritius. 


49, Lavirus (PERISTERNIA) AUSTRALIENSIS, ve. 
Station 5, Boydong Cays. 


TURBINELLIDA. 
50. CYNODONTA CORNIGERA, Lam. 


51. Semrrusus (MEGALATRACTUS) PROBOSCIDIFERUS, Lam. 

No full-grown specimens were in the collection, but a large 
operculum, and a mass of nidamental capsules, containing, 
roundly speaking, fully one hundred and twenty embryonal 
young. These are all, without exception, decollate, seven- to 
eight-whorled, averaging 80 mm. longitudinally. The canal is 
fairly long, the last whorl showing a little of the adult sculpture, 
and slightly tumid. 


THE MOLLUSCA OF TORRES STRAITS. 159 


This species is classed by Fischer (Man. de Oonch. p. 623) as 
the type of his subgenus MMegalatractus (1884). It is confined 
to the shores of Australia. 


BUCCINID2. 


52. CANTHARUS MOLLIS, Gild. 

Fine individuals, which we have compared with and cannot 
separate from the Japanese type. 

Station 8, Albany Pass. 


53. ENGIna concrnna, Rve. 
Station 2, Warrior Island. 


54. Enaina Reever, Tryon. 
Hab. North America and Mexico, also Australia (J. Brazier). 
Station 5, Boydong Cays. 


55. PHos SCALAROIDES, A. Ad. 

Station 5, Boydong Cays. 

56. Nassaria suTuRALIS, A. Ad. 

This well-marked form is sparingly distributed from the 
Arabian Sea eastward, being particularly frequent at Bombay 
(Abercrombie) and Karachi (Townsend). 

Station 8, Albany Pass. 


NaASsID&. 


57. Nassa (ALECTRYON) suTURALIS, Lam. 
Station 8, Albany Pass. 


58. Nassa (Nrorma) anBescens, Dkr. 
Station 13, Murray Island. 


59. Nassa (Nrorma) cremata, Hinds. 
Station 8, Albany Pass.. 


60. Nassa (ALECTRYON) FRETORUM *, sp. n. (PI. 10. fig. 3.) 

N. testa breviter oblonga, spira acuminata, albo-cinerea, 
nitida; anfractibus octo, inclusis apicalibus tribus, levibus, 
vitreis, ceteris regulariter longitudinaliter plano-costatis, costis 
nitidis, levibus, apud suturas impressis, et infra, juxta suturas, 
transversim uwnisulcatis, et sparsim brunneo-maculatis, liris 
transversis aliter absentibus, ultimo anfractu infra medium 


* Fretorum, i. e. Torresianorum, inhabiting the Torres Straits. 


160 MESSRS. J. C. MELVILL AND R. STANDEN ON 


usque ad basim transversim multiplicato ; apertura ovata, labro 
extus paullum effuso, cirea columellam nitide calloso, perlevi, 
intus striato, ad marginem denticulato, canali brevi. Long. 12, 
lat. 6 mm. 

Station 2, Warrior Island. 

We have in vain tried to match this small species. It is 
undoubtedly an Alectryon, its congeners being N. monile, Kien., 
and allies, although, from the description, WV. (Miotha) multi- 
costata, A. Ad., a species included in the section remarkable for 
the deep transverse sulcations, but which are obsolete in it, 
comes very near IV. fretorwm. Our species would seem to differ 
in the conspicuous spiral sulcation just below the sutures, in the 
broader contour in proportion to the length, and one or two 
minor details. We are especially indebted to Mr. Ernest R. 
Sykes for carefully comparing this Massa with the incomparable 
series in the National Collection, South Kensington. 


61. Nassa (NrorHa) roTunpa, Melv. § Stand. (Journ. of 
Conch. vol. vil. p. 273, pl. ix. fig. 2, 1896.) 

One of the few molluses, lately described by us from Lifu, 
that has occurred in this collection. Its nearest ally is 
NV. plebecula, Gld. 

Station 2, Warrior Island. 


62. Nassa (Hima) PLEBECULA, Gild. 
Station 5, Boydong Cays. 


CoLUMBELLID&. 


63. COLUMBELLA VARIANS, Sow. 
Station 13, Mér. 


64, CoLUMBELLA (Prema) FULGURANS, Lam.—a. eufulgurans; 
B. punctata, Lam. 

A series embracing the type with zigzag lines on a black 
ground, merging into the condition in which the white streaks 
are broken up into rounded or triangular white spots, and 
evidently occurring together. 

Station 2, Warrior Island ; Station 18, Mér (Murray Island) ; 
Station 5, Boydong Cays. 


THE MOLLUSCA OF TORRES STRAITS. 161 


65. CotumBELLA (Pyemma) TYLERI, Gray. 
Station 5, Albany Pass. 


66. CoLUMBELLA (Pya@m mA) VERSICOLOR, Sow. 
Station 12, Mabuiag, 4 fathoms (Oct. 1888). 


67. Conumpe ta (ArIntA) GaLaxtias, Rve. 
Station 5, Boydong Cays. 


| 68. CotumBEnia (Mirren) SEMICONVEXA, Lam. 
Station 5, Boydong Cays. 


69. CotumMBELLA (ConorpEa) FLAVA, Brug. 
Station 18, Murray Island ; Station 8, Albany Pass. 


Mouricip2. 


70. Murex BREVISPINA, Lam. 
Station 18, Murray Island. 


71. Murex (Carcorsus) apustus, Lam. 
Station 13, Mer. 


Var. Fuscus, Dkr. 

Excepting in the reddish-brown colour of the varices, we 
cannot trace any specific distinction between this and the typical 
MM. adustus, Lam. 

Station 8, Albany Pass. 


72. Murex (CHICOREUS) AXICORNIS, Lam. 
Station 18, Mer. 


73. Murex (CHiIcorEus) capucinus, Lam. 

The vague locality “ Philippines,” that has been given in our 
text-books for this species, has been considered unauthentic, as 
the west coast of South America (Chili) seems to be its head- 
quarters. Its occurrence therefore in the Torres Straits is very 
interesting. 


74, Murex (PreRonotts) SAIBAIENSIS, sp. n. (Pl. 10. fig. 1.) 

M. testa trigona, tenui, calcarea, albido-cinerea; anfractibus 
5-6, apicali mamillari, vitreo, ceteris trivaricosis, varicibus 
foliatis, uni-angulatis, expansis, spiraliter liratulis, apud medium 
anfractiis ultimi simul ac penultimi, inter varices noduliferis ; 
apertura ovata, peristomate foliaceo, delicate squamulato, canali 
producta, lata. Long. 18, lat. 13 mm. 

Between Saibai I. and New Guinea, 10-17 fathoms (Station 3). 


162 MESSRS. J. C. MELVILL AND R. STANDEN ON 


A delicate Pteronotus which, though apparently full-grown, 
may yet add two or three varices to its stature before attaining 
perfection, without in any way altering the characteristic contour 
of its whorls. It is impossible to match this form exactly with 
any described species, though its affinities lie with triformis, 
Reeve, acanthopterus, Lam., and eurypteron, Reeve. It is 
trigonous, light in texture, of a chalky consistency, cinereous 
white, five- or six-whorled, these each possessing three varices, 
and towards the centre of the penultimate whorl, as well as the 
lowest, nodulous squame are noticeable between each varix. 
The surface is spirally, finely but rather unevenly lirate, the 
varices being once-angled on the last whorl, immediately below 
the sutures. Aperture oval, outer lip beautifully fimbriate- 
squamose, canal prolonged and somewhat broad. Operculum 
not present. 


75. Murex (OcINEBRA) SALMONEUS™, sp.n. (PI. 10. fig. 2.) 

M. testa parva, breviter fusiformi, carnea, purpurea vel 
pallide punicea, multum corrugata; anfractibus septem, arcte 
variciferis, varicibus pulchre fimbriatulis, costis fistulosis trans- 
versim connexis ; apertura rotunda, labro extus paullum expanso, 
fimbriato, incrassato ; canali brevi, recurva. Long. 12°50, lat. 
6 inm. 

This little Ocinebra is shortly fusiform, brightly coloured, 
either pale flesh-colour, pink, or purple, seven-whorled, many- 
variced, the varices beautifully fringed or squamose, crossed 
spirally by fistulose ribs; aperture round; canal short, slightly 
recurved; outer lip expanded, thickened, fringed. 


76. Murex (Cuicornvs) CERVICORNIS, Lam. 


77. Murex (Curcorevs) ramosvs, Z. 
Station 18, Murray Island. 


78. PURPURA ALVEOLATA, ve. 
Station 18, Murray Island. 


79. Purpura PEeRsIcA, L. 
Station 138, Murray Island. 


80. Purpura (THALESsa) HIPPOCASTANUM, Lam. 
Station 2, Warrior Island. 


* Salmoneus, from the prevailing carneous hue. 


THE MOLLUSCA OF TORRES STRAITS. 163 


81. Purpura (STRAMONITA) RUSTICA, Lam. 
Station 13, Murray Island. 


82. Purpura (CRONIA) AMYGDALA, Kzener. 
Station 13, Murray Island; Station 8, Albany Pass. 
83. SISTRUM ARACHNOIDES, Lam. 

Station 2, Warrior Island. 

84. SISTRUM CAVERNOSUM, Lve. 

Station 13, Mer; Station 5, Boydong Cays. 
85. SIStTRUM CHRYSOSTOMUM, Desh. 
Station 2, Warrior Island. 

86. SISTRUM CONCATENATUM, Lam. 

Station 13, Murray Island. 

87. SISTRUM ELATUM, Blainv. 

Station 5, Boydong Cays. 

88. SISTRUM FISCELLUM, Chemn. 

Station 5, Boydong Cays. 

89. SISTRUM HEPTAGONALE, ve. 

Station 5, Boydong Cays. 


90. SISTRUM MARGARITICOLUM, Brod. 

Originally described by Broderip as a Murer from Lord 
Hlood’s Island, this has been found, in its typical form, generally 
distributed throughout the eastern tropical area, occurring as 
far north as the coasts of Baluchistan (Townsend). 


91. SisrRuM ocHROSTOMA, Blainv. 
Station 2, Warrior Island. 
TRITONID A. 


92. AQUILLUS (SIMPULUM) GEMMATUS, Rve. 
Station 18, Murray Island. 


93. AQUILLUS (SIMPULUM) PILEARIS, L. 
94, EprIpRoMUS ANTIQUATUS, Hinds. 


95. Eprpromus Bepnawu, Brazier. 
Varying in size, but not in sculpture, our largest specimen 
being long. 11, lat. 5 mm. 


96. GyRINnEUM (LAMPAS) RANELLOIDES, Reve. 
Station 18, Murray Island. 


164 MESSRS. J. C. MELVILL AND R. STANDEN ON 


97. GyrinEumM (EupPLevRA) sucunpDuM, A. Ad. 

In almost every particular, save that of size, this is the exact 
counterpart of the well-known Hupleura perca (Perry), pulchra 
(Gray). 

Station 13, Murray Island. 


98. Gyrineum (APOLLON) PUSILLUM, Brod. 
Mostly of the white variety with violaceous aperture, the 
largest example of all being uniformly whitish-ochreous. 
Station 13, Murray Island. 
CaSsIDIDA. 


99. Cassis (CasMARIA) VIBEX, L. 


100. Cassts (Semicassts) TORQUATA, Eve. 


Do.ipaz. 
101. Dotium PErprx, L. 
102. Dotium variecatum, Lam. 
103. Dotium (Marea) pomum, LZ. 
Station 5, Boydong Cays. 
CYPREIDA. 


104. Ovuna (Raprius) Anaast, A. Ad. 
Station 11, Mabuiag. 


105. Cyprma FELINA, Cray. 
Station 5, Boydong Cays. 


106. Cyprma (Artcra) annuus, L. 
107. Cyprma (Artcta) arastoa, L. 


108. Cyprma (Luponta) ERRoNEs, L. 

Some of the nidamental capsules are also present, consisting 
of coagulated masses of one hundred or more pellucid oval eggs. 

Station 9, Prince of Wales Channel. 


109. Cyprma (Luponta) FLAVEOLA, L. 
Station 18, Murray Island. 


110. Cypraa (Luponta) tynx, L. 


111. Cyprma (Luronta) ziczac, D. 
Station 138, Murray Island ; Station 2, Warrior Island. 


THE MOLLUSCA OF TORRES STRAITS. 165 


112. Trivia oryza, Lam. 
Station 18, Murray Island. 


113. Trivia staPHYLmMA, L. 
Station 13, Murray Island. 


STROMBIDS. 
114. SrromBus (GaLtinuta) CAMPBELLI, Gray. 
Station 10, Channel between Hammond Island and Wednesday 
Spit. 
115. StromBvs (GALLINULA) CANARIUM, L. 
Station 18, Murray Island. 


116. StrompBus (GALLINULA) ISABELLA, Lam. 

In our opinion a mere variety of S. canarium, L., but some 
authors place specific reliance on the thinner substance and 
absence of freckled marking. The distribution of the two species 
is the same. 

Station 18, Murray Island. 


117. Stromevs (Gatirnvuta) Srpparpr, Sow. 

Hitherto considered endemic in Ceylon, aremarkable extension 
of range is now presented. 

Station 2, Warrior Island. 


118. StrompBus (GALLINULA) URCEUS, ZL. 
Station 13, Murray Island. 


119. Strompus (GALLINULA) VARIABILIS, Swazns. 


120. StromsBus (GALLINULA) viTTATUS, L. 

Young, but finely coloured, being tesselated with brown. 
There is also in the collection, from Mer, a very remarkable but 
juvenile form, slightly variced, white, fusiform, and closely 
striate throughout. 

Station 4, between Ormans Reef and the Brothers Island; 
Station 18, Mer ; Station 5, Boydong Cays. 


121. SrromBus (Canarium) pENTATUS, L. 

Only, we think, a sub-species of S. wreeus, L., but at all events 
it is the prevailing form. 

Station 13, Mer (Murray Island). 


122. StrromBus (Monopactytvus) Lamarckt, Gray. 


123. StromBus (MonopactyLvs) MELANOSTOMUS, Swains. 


166 MESSRS. J. C. MELVILL AND R. STANDEN ON 


124, Srrompus (CoNOMUREX) LUHUANUS, L. 
Station 13, Murray Island. 


125. Prerocera (HEPTADACTYLUS) LAMBIS, L. 
Station 13, Mér (Murray Island). 


126. SERAPHS TEREBELLUM, L. 
Station 13, Mer. 


CERITHIID 2. 


127. Trrroris aiaas, Hinds. 


The largest example measures 42 mm. in length. 
Station 13, Mer. 


128. Triroris (INO) EXCELSIOR, sp.n. (PI. 10. fig. 5.) 

T. testa producta, multum attenuata, angusta, brunnea, hie 
illic rufo-maculata; anfractibus quinque- vel sex-et-viginti, 
tornatis, apud suturas elevatis, transversim arcte tricarinatis, 
levibus; carina infra, juxta suturas, minore, duabus alteris magis 
conspicuis, interstitié interveniente planata, ultimo anfractu 
quadricarinato, carina bina ad peripheriam addita; apertura 
rotunda, parva, canali breviter reeurvirostri, in uno specimine 
pone aperturam ipsam clausa. Long. 30, lat. 5 (sp. maj.) mm. 

Four or five, mostly imperfect, examples. The form is much 
acuminate and attenuate, narrow; colour light brown, here and 
there indistinctly flecked with rufous spotting; whorls 25 or 26, 
tornate, smooth, elevated at the sutures, closely thrice-keeled 
transversely, the keel just below the sutures is smaller and less 
conspicuous than the two lower, the last whorl is four-keeled, 
there being two on the periphery ; the aperture is roundish, small; 
in one (the most perfect) specimen the recurved and beaked 
canal is closed with shelly matter behind the aperture. 

The only species to which, in size, this very conspicuous 
Triforis could be referred is 7. gigas, Hinds, also occurring in 
the same localities. The sculpture, as seen by the above 
description, is however totally different, being smooth, with no 
interstitial pitting or gemmuled ribs of any kind. Owing to 
no one specimen being in a state of absolute perfection, we 
have been compelled to estimate the number of whorls and the 
dimensions generally with the aid of two or more examples, each 
complete in some one particular. 


THE MOLLUSCA OF TORRES STRAITS. 167 


129. CERITHIUM CITRINUM, Sow. 
Station 4, between Ormans Reef and the Brothers Island. 


130. CERITHIUM ECHINATUM, Lam. 
Station 5, Boydong Cays. 


131. CeritHiuM Morus, Lam. 

Very abundant. <A reddish variety also occurs at the Mér 
Station. 

Station 2, Warrior Island; Station 13, Mer; Murray Island ; 
Station 4, between Ormans Reef and the Brothers Island. 


132. CERITHIUM NOVH-HIBERNIA, A. Ad. 
Station 13, Mer. 


133. CrerITHIUM PETROSUM, Wood. 

The name “ tuwberculatum,” having been twice used by Linnzeus, 
must be abandoned in favour of that of Wood. 

Station 2, Warrior Island; Station 4, between Ormans Reef 
and the Brothers Island. 


134, CERITHIUM PIPERITUM, Sow. 
Station 13, Murray Island. 


135. CERITHIUM SALEBROSUM, Sow. 

Only broken individuals occurred, referred with some little 
doubt as above. The finest examples we have seen of C. sale- 
brosum come from Lifu (Hadfield). It is rare, and found 
generally, though sparingly, in the Philippines, with south- 
eastward extension of range. 

Station 5, Boydong Cays. 


136. CERITHIUM VARIEGATUM, Quoy. 

This, as identified by Mr. E. A. Smith (Voy. of H.M.S. ‘Alert,’ 
1881-2, p. 64), is a variety of the protean C. morus, L., from 
which it differs in its more attenuate fusiform shape, and the 
eolour-banding, with a smaller series of apical tubercles. 


137. CERITHIUM ZEBRUM, A7vener. 


138. CeritHium (VERTAGUS) aLUco, L. 
Station 5, Boydong Cays. 


139. CertrHium (VERTAGUS) LINEATUM, Lam. 
We do not agree with Tryon (Man. Conch. ix. p. 148) that 
this is a variety of C. asper, Lam. The shell is uniformly 


168 MESSRS. J. C. MELVILL AND R. STANDEN ON 


coarser, besides being, in proportion to its length, larger in all 
its parts, and often spirally banded. 


140. CeriTHiuM (VERTAGUS) PULCHRUM, A. Ad. 
Station 2, Warrior Island. 


141. Crritatum (VeRtTAGUS) SOWERBYI, Kiener. 
Station 5, Boydong Cays. 


142. CreriratumM (VERTAGUS) VERTAGUS, L. 
Station 5, Boydong Cays. 


143. Certrnium (LAMPANTIA) ZONALE, Brug. 
Station 5, Boydong Cays. 


144. CEertrHium (Colina) conrraotum, Sow. 

We consider this distinct from the type of Colina, C. pinguis, 
Ad., being attenuate, contracted, nine- or ten-whorled, the three 
last whorls almost straight and of uniform thickness. The 
species are now being frequently found in the North Indian 
Ocean, and in other places, C. pinguis having been formerly 
considered peculiar to South Africa. 

Station 13, Mér. 


145. Certtaium (Corina) TantatuM, Sow. 
Station 13, Murray Island. 


146. BirTIuM TORRESIENSE, sp. n. (PI. 10. fig. 6.) 

B. testa parva, elongato-cylindracea, pallide ochracea vel 
fulva et albo-maculata; anfractibus undecim, apicalibus inclusis 
tribus, albidis (vel, in altero specimine, fulvis), uniformibus, per- 
parvis, leevibus, tribus his proximis pulchre gradatis, ceteris apud 
suturas impressis, longitudinaliter recte gemmato-costulatis, 
quinque ordinibus spiraliter dispositis, in specimine typico 
gemmis vel tuberculis hic ochraceis, illic albidis, ultimo anfractu 
ad basim liris quinque concentrice predito, apertura ovata, 
canali brevissimo, peristomate tenui. Long. 5, lat. 2 mm. 

Station 14, channels between reefs, Mer. 

We have given much attention to the deep-sea forms of 
Bittium, especially those described in the Reports of H.M.S. 
‘Challenger’ from N.E. Australia and the Torres Straits. 
The only one which could challenge comparison with the shells 
before us is B. wanthum*, Watson, but in the following 


* Cf. Voyage H.M.S. ‘ Challenger,’ Gastropoda, vol. xv., Zoology, p. 537. 


THE MOLLUSCA OF TORRES STRAITS. 169 


particulars B. torresiense seems to dilfer :—first, the texture 
is more delicate in Watson’s species; secondly, the tubercu- 
lated gemme, forming longitudinal ribs, are closer, and the in- 
terstices simple, with no intervening lire whatsoever. The whorls 
are eleven in our species, as against eight, and the apical whorls 
are similar, both in number and form, but the painting 1s different, 
being flecked in one type with white. The dimensions are similar. 
Three examples. 


147. Potamriprs (TyMPANOTONUS) PALUSTRIS, L. 


148. PoramripEs (TYMPANOTONUS) RETIFERUS, Sow. 
Station 1, Thursday Island. 


149. PoramrpeEs (TEREBRALIA) SULCATUS, Born. 
Station 13, Mér. 


Mopvutip#. 


150. Mopvutus optusatis, Phil. 


PLANAXID&. 


151. PLANAxis sutcatus, Born. 

“Tn interstices of coral shore-rock, the smaller being from 
extreme high water-line, the larger a very little lower down ”’ 
(A. C. H.). 

Station 5, Boydong Cays. 


152, PLanaxis (QUOYIA) DECOLLATUS, Quoy. 
All of a dwarf form. 


TRICHOTROPIDID&. 


153. SEPARATISTA BLAINVILLEANA, Petit. 

A very large and perfect example, the dimensions of which 
are: long. 11, diam. 17 mm., thus much exceeding that of the 
others recorded. It is a rare inhabitant of the Pacific islands, 
and has not before been known to occur in the region embraced 
by our paper. 

Flinders’ Entrance, near Mer, 20 fathoms, Dec. 21st, 1888. 


VERMETID. 


154, VERMETUS (SIPHONIUM) Maximts, Sow. 
Station 12, Mabuiag. 


170 MESSRS. J. C. MELVILL AND R. STANDEN ON 


155. Vermetus (Brvonta) Quoyi, H. & A. Ad. 
Three varieties—cinereous, pule purple, and flesh-coloured. 
Station 13, Mer. 


156. VERMETUS (THYLACODES) NOVH-HOLLANDIZ, Rouss. 
An obliquely concentrically rugose shell, attached to branches 
of madrepore. Endemic, so far as Australian shores are con- 


cerned. 
Station 13, Mer. 


157. Sinsquarta Cuminait, Worch. 

Station 5, Boydong Cays. 

158. SILTQUARIA PONDEROSA, MWorch. 

A fine series, showing much variation in torsion. 

Station 4, between Ormans Reef and the Brothers Island ; 
Station 1, Thursday Island; Station 13, Mér. 


TURRITELLIDE. 
159. TURRITELLA MULTILIRATA, Ad. JF Reve. 
Station 5, Boydong Cays. 


160. Marninpa EuRyTIMA, Melv. & Stand. 
(Journ. of Conch. vol. viii. 1896, p. 310, pl. xi. fig. 73.) 
Precisely agrees, in sculpture and detail, with our Loyalty 
Island type. 
LitToRINIDs. 


161. Lirrorina FiLosa, Sow. 

Station 13, Murray Island. 

162. Lirrorina (MELARAPHE) MAURITIANA, Lam. 
Station 5, Boydong Cays. 

163. TEcTARIUS MALAccANUS, Phil. 

Station 5, Boydong Cays. 


Lirroprp &. 
164. Diana aLBuGo, Wats. 
Station 5, Boydong Cays. 
165. ALABA PYRRHACME, Melv. § Stand. 
(Journ. of Conch. vol. vii. 1896, p. 310, pl. ix. fig. 70, as 
Rissoa.) 


THE MOLLUSCA OF TORRES STRAITS. 171 


Agreeing with the Loyalty Island types (Hadfield), and also 
with examples from Thursday Island (A. U. Henn). 
Station 5, Boydong Cays *. 


RISSomIDz&. 


166. Rissorna sconopax, Sowvb. 
Identical with Loyalty Island Saale (Hadfield Coll.). 
Station 5, Boydong Cays. 


167. Rissorva THAUMASIA, Melv. & Stand. (Journ. of Conch. 
vol. ix. 1898, p. 31, pl. 1. fig. 3.) 

This agrees exactly with the Madras type (J. R. Henderson 
and H. Thurston, in Mus. Brit.). 


168. Rissorna TRIANGULARIS, Wats. 
We have this from Thursday Island (A. U. Henn). 
Station 5, Boydong Cays. 


169. Rissorna (PHOSINELLA) CLATHRATA, A. Ad. 
Station 5, Boydong Cays. i 


170. Rissorna (MOERCHIELLA) SPIRATA, Sow. 

Two varieties, one of which approaches var. deformis, Sow., 
the other var. artensis, Montr., a New Caledonian form. 

Station 2, Warrior Island; Station 13, Mer. 


CapuLIDz. 


171. Catyprrma TORTILIS, Rve. (=C. equestris, L., var. P). 
Station 5, Boydong Cays. 


VANIKORID&. 


172. VANIKORO CANCELLATA, Lam. 

A smooth variety, exhibiting hardly any dorsal cancellation. 
It is the type of the genus, named after the Island Vanikoro 
in the Pacific Ocean. The name, though barbarous, should, in 
our opinion, be retained, as the genus was definitely described 
by Quoy and Gaimard in 1832, thus having precedence over 
Merria, Gray, 1839; and a yet longer period over Leucotis, 
Swainson, 1840, and Narica, Récluz, 1841. 


* Mr, C. Hedley, in his Mollusca of Funafuti (Mem. Austral. Mus. iii. p. 43, 
1899), makes our 4. pyrrhaeme the type of his new genus Odtortio. 


LINN. JOURN.—ZOOLOGY, VOL. XXVII. 13 


172 MESSRS. J. C. MELVILL AND R. STANDEN ON 


NATICID&. 


173. Navica AREOLATA, Récluz. 
Station 5, Boydong Cays. 


174. Natica GavLtiertana, Petit. 
Station 5, Boydong Cays. 


175. Natica (NEVERITA) BICOLOR, Phil. 


176. Nattca (Mamma) avrantia, Lam. 
Station 6, Mer. 


177. Navica (Mamma) Freminerana, Récluz. 
Station 5, Murray Island; Station 6, Mer; Station 5, Boy- 
dong Cays. 


TANTHINIDA. 
178. Janruina Suirura, Rve. 
An Australian form hardly differmg from the common 
I. fragilis, L. 
Station 5, Boydong Cays. 


ScALARIIDA. 


179. ScALARIA LYRA, Sow. 
Station 5, Boydong Cays. 


180. ScaLARIA OBLIQUA, Sow. 
Station 5, Boydong Cays. 


181. ScALARIA SUBAURICULATA, Souvb. 
Station 5, Boydong Cays. 


182. Eexista TRICARINATA, Ad. f Rve., Voy. ‘Samarang,’ p. 49. 


The single example is very imperfect, but possesses two 
anfractual carine only, on the upper whorls. Turritella lepto- 
mita, Melv. & Sykes (Proc. Mal. Soe. ii. p. 176, pl. xiii. figs. 12, 
12 a), represents a very nearly allied species from the Andamans 
the whorls possessing three to four keels on the upper, four y, 
least on the lowest whorl. So rarely found are these nearly 
allied Hgliste that the anatomy is practically unknown, but the 
operculum being scalaroid they have on this account been 
transferred from the Turritellide to the Scalaride. 

Station 5, Boydong Cays. 


THE MOLLUSCA OF TORRES STRAITS. 173 


183. Scattona ELATA, Semper. 
Station 5, Boydong Cays. 


EULIMID 2. 


184. HULIMA AUSTRALASIACA, sp.n. (PI. 10. fig. 7.) 

E. testa fusiformi, politissima, crassiuscula, paullum incurva ; 
anfractibus 10-11, condensatis, lacteis, apud suturas paullulum 
impressis, ultimo anfractu crasso, varicibus obsoletis; apertura 
ovata, peristomate preesertim apud basim, incrassato, levissimo. 
Long. 7, lat. 3 mm. 

A very shining, polished, milky-white, opaque Hulima, fusi- 
form, slightly incurved, ten- or eleven-whorled, apex small ; 
whorls very slightly impressed at the sutures, the last whorl 
thickened, especially towards the base; aperture oval; peristome 
thick, exceedingly smooth. 

This species slightly resembles EH. latipes, Watson, coilected 
at Flinders’ Passage, Torres Straits, in 7 fathoms, during the 
‘ Challenger ’ Expedition, but is much larger in every part, and 
the spire does not so rapidly contract. The aperture, too, is large 
proportionately. The lateral varices are quite inconspicuous or 
obsolete. 


185. Eurima (Lerosrraca) acrcuna, Gould. 
Station 14, Mer. 


PYRAMIDELLID 2. 


186. OBELISCUS TEREBELLOIDES, A. Ad. 

Possibly a variety, but, if so, a very remarkable one, of 
O. dolabrata, L., being much narrower in proportion, more 
numerous in the whorls, with the columellar plice but two m 
number. 


Station 5, Boydong Cays. 
187. Cineurina spina, Cr. & Fischer. 


Station 5, Boydong Cays. 


NERITIDA. 


188. Nerrra ponrra, Z.; and var. aurora, Dkr. 
Station 18, Murray Island; Station 5, Boydong Cays. 


13* 


174 MESSRS. J. C. MELVILL AND R. STANDEN ON 


189. Nerrra stenata, Macleay. 
Station 5, Boydong Cays. 


190. Nerira unpata, L. 
Station 4, Ormans Reef; Station 18, Murray Island; Station 5, 
Boydong Cays. 


191. Nuerira (THELICOSTYLA) ALBICILLA, L. 
192. Neriva (PELERONTA) FUNICULATA, ve. 


193. Nerrra (PELERONTA) PLicaTA, L. 
Station 18, Mér (Murray Island). 


Maganis *, genus novum. 


Testa imperforata, depressa, orbicularis, albo-calcarea, longi- 
tudinaliter arcte lamellata, lamellis irregularibus; anfractus 8, 
quorum apicalis obtusus, globularis, interstitie inter lamellas 
arcte et pulcherrime sub lente spiraliter striate ; apertura lunaris ; 
peristoma rotundum, fere continuum; labri intus margo colu- 
mellaris latus, planatus, dentibus minutissimis preditus, extus 
crassiusculus, planatus, levis. Operculum..... P 


194. MaGapis EUMERINTHA T, sp.n. (PI. 10. figs. 8, 8 a.) 

M. testa ut supra. Alt 2°25, diam. 4°50 mm. 

Station 14, Mer, channels between reefs. 

A truly remarkable mollusc, and one for the reception a 
which it is imperative to create a new genus. It is undoubtedly 
a neritoid, but the close longitudinal lamelle suggest affinity 
with Vanikoro, Quoy, e.g. V. Gueriniana, Récluz. 

It is unlike any shell known, either recent or fossil, being 
small, depressed, orbicular, imperforate, of a chalky yellowish- 
white consistency, longitudinally closely lamellate, three-whorled 
(inclusive of the obtuse and globular apical whorl); the inter- 
stices between the lamelle are everywhere closely and very 
beautifully striate; aperture lunar and wholly neritoid; colu- 
mellar area straight, wide, and furnished with numerous very 
fine and minute teeth; peristome almost continuous; outer lip 
lunar, smooth, somewhat thickened. We have not seen the 
operculum. 

* payadois, a harp. 
t+ Eumerintha, cd prowys, well furnished with strings or lamellx. 


THE MOLLUSCA OF TORRES STRAITS. 175 


TURBINIDA. 


195. PHAsIaNELLA (OrTHOMESUS) NIvosA, Rve. 
Station 8, Albany Pass; Station 5, Boydong Cays. 


196. TURBO MaRMoratUs, L. 
Station 13, Murray Island. 


197. TurBo PerHotatus, L. 
Station 13, Mér. 


198. Turzo (SENEcTUs) ar@yrostomus, L. 
Station 2, Warrior Island ; Station 13, Mér. 


199. TurBo (SENEcCTUS) cHRYsostomus, L. 
Station 2, Warrior Island; Station 13, Mér. 


200. TurzBo (SENEcTUs) FoLIAcEUs, Phil. 
201. TuRBo (SENECTUS) SPARVERIUS, G'mel. 


202. ASTRALIUM PETROSUM, Wart. 
Station 2, Warrior Island; Station 13, Mér (Murray Island). 


TROCHIDE. 
203. Trocuus (TECTUS) FENESTRATUS, Gel. 
204. TRocuus (LaMPROSTOMA) MACULATUS, L. 


205. CLANCULUS ATROPURPUREUS, Gd. 

Station 10, Channel between Hammond Island and Wednesday 
Spit. 

206. CLANCULUS UNEDO, A. Ad. 

Station 13, Mer. 


907. CANTHARIDUS TORRESI, H. A. Smith. 
Station 13, Mer. 


208. MonoponTa CANALIFERUS, Lam. 
Station 13, Mer. 


209. CHRYSOSTOMA PARADOXUM, Born. 
Station 13, Murray Island; Station 5, Boydong Cays. 


210. Umsonium vEstTiarivum, L. 
Station 13, Mér; Station 5, Boydong Cays. 


176 MESSRS. J. C. MELVILL AND R. STANDEN ON 


211. Minonia GuaPHYRELLA, Velv. § Stand. (Jour. of Conch. 
vol. viii. 1895, p. 125, pl. ii. fig. 18.) 

Agreeing with the type from Lifu (Hadfield Coll.). The 
peculiar style of apical liration, and the presence of a distinct 
peripherial angle, together with the difference in coloration— 
glaphyrella being almost invariably straw-coloured and blotched 
below the suture with equidistant brown spots—distinguish this 
from Jf. pudibunda, Fischer. 

Station 5, Boydong Cays. 


212. MInoLia PuDIBUNDA, Mischer. 
Station 5, Boydong Cays. 


218. Minoxta vitinieingEa, lke. 
Station 14, Mer. 


214. CaLLiostoma (EUTROCHTS) SEPTENARIUM™, sp. n. (PI.10. 
fig. 9.) 

O. testa conica, profunde sed anguste umbilicata, solida; 
anfractibus septem, duobus apicalibus applanatis, vitreis, ceteris 
arctissime et pulchre transversim septem-costulatis, costis gem- 
muliferis, interstitiis obliqui-striatis, ultimo anfractu apud 
peripheriam acutangulo, ad basim novem lirarum concentricis 
ordinibus preedito, circa umbilicum ipsum paullum calloso; aper- 
tura quadratula, labro crassiusculo, marginem apud columellarem 
unidentato, apud umbilicum parum reflexo. Alt. 10°50, diam. 
9 mm. 

Station 8, Albany Pass, 10 fathoms, Aug. 29, 1888. 

A pyramidally conical shell, deeply but narrowly umbilicate, 
solid; whorls seven, all, with the exception of the two apical 
which are smooth and glossy, being closely spirally seven-ribbed, 
these ribs thickly and regularly formed of gemme, contiguous 
and crowded, the interstices between these beaded riblets are 
indistinctly obliquely striated; the last whorl is sharply angled 
at the periphery, and at the base there are eight or nine con- 
centric rows of the same kind of beaded ribs as under the whorls, 
the interstices very similarly obliquely striate; the aperture 
square, outer lip slightly thickened, whilst at the columellar 


* Septenarius, “ consisting of seven,” in allusion to the spiral beaded riblets 
uniformly seven in each whorl. if 


THE MOLLUSCA OF TORRES STRAITS. 177 


margin a small central tooth is observable. A resemblance, but 
merely superficial, may be traced to C. fragum, Phil. 


215. CALLIOSTOMA SPECIOSUM, A. Ad. 
216. EucHELUS ANGULATUS, Pease. 


217. HUcHELUS AaTRATUS, Gmel. 
Station 13, Mér; Station 2, Warrior Island. 


DELPHINULID2. 


218. DELPHINULA FoRMOSA, Ave. 
Station 138, Mer. 


219. Liorra vARIcosA, Rve. 
Station 1, Thursday Island. 


CYCLOSTREMATID &. 


220. Microrneca Acrpatta, sp.n. (Pl. 10. figs. 10, 10a.) 

M. testa candida, depressc-globosa, sculpturata, profunde um- 
bilicata ; anfractibus quatuor, quorum apicali simplici, nitido, 
ceteris transversim pulcherrime carinatis, antepenultimo tribus, 
penultimo, cum ultimo, quatuor carinis preditis, arcte noduloso- 
gemmatis, gemmis nitidissimis, interstitiis transversim arcte 
liratis, ultimis anfractu effuso ; regionem circa umbilicarem in- 
crassato, nitido, plus minus crenulato, fere levi ; apertura rotunda, 
peristomate crassiusculo, continuo; operculo....? Alt. 3°50, 
lat. 5 mm. 

Though small, one of the most exquisite shells yet discovered. 
It is pure white, depressedly globular, four-whorled, inclusive of 
the small and shining apical whorl, the remaining three being 
in the uppermost or antepenultimate whorl thrice transversely or 
spirally keeled, in the two lower whorls four times ; these keels 
or spiral ribs are closely studded with, or mdeed composed of, 
white shining gemme which are round, crowded, and brilliant ; the 
rest of the shell is finely and delicately spirally lirate or sculptured. 
Aperture round, white within; peristome thin, continuous. The 

basal portion of the surface is thickened round the umbilical 
region and more or less crenulate. 

MM. crenellifera, Ad., the type of the genus, which we have 
received from Bombay (Abercrombie), though originally reported 
from Japan only, bas the base and thickened crenulate umbilica 


178 MESSRS. J. C. MELVILL AND BR. STANDEN ON 


ornamentation very similar, but differs in form, being more 
elongate and quite smooth and plain to the apex. Pilsbry * 
queries WMicrotheca as a probable subgenus of Tecnostoma, Ad., 
but our strong opinion is that it should be separated generically. 
In Teinostoma the callosity entirely covers the umbilicus. We 
should, indeed, consider MWicrotheca nearer Cyclostrema, Marryat. 

The type, I. crenellifera, to which allusion has just been made, 
was originally described by Adams as a species of Lsanda, Ad., 
with which it possesses a certain amount of conchological affinity ; 
but until the anatomy of these minute molluscs be more dili- 
gently studied, the compilation of special monographs is useless, 
and no arrangement can be other than purely tentative. 

The species before us is, however, much nearer I. crenellifera, 
Ad., than Zsanda coronata, Ad., which last we have received from 
the Papuan region, though it does not occur in Dr. Haddon’s 
present collection. We have also obtained it from shell-sand 
collected a few years ago by Mr. Arnold Umfreville Henn, at 
Magnetic Island, Queensland, where it is very ‘abundant. 

Microtheca crenellifera occurring, as just pointed out, both in 
Japan and Bombay, will no doubt be found in intermediate 
stations, but it has hitherto occurred almost singly, and is very 
searce in European collections. It is five-whorled, globose, 
white, shining, smooth save for one spiral costulate crenulation 
below the sutures, which are much impressed, and for the 
crenate callosity round the umbilicus, which has its counterpart 
in WM. Acidalia. 

Of this new species two examples, quite perfect and precisely 
similar, occurred, which we think worthy of one of the epithets 
formerly bestowed on Aphrodite. 


STOMATIIDA. 
221. SromaTeLta Martetr, Cr. 
Station 18, Mer; Station 5, Boydong Cays. 


222. STOMATELLA SULCIFERA, Lam. 
Station 18, Mer. 


228. GENA STRIATULA, A. Ad. 
Station 2, Warrior Island ; Station 8, Albany Pass. 


* Trvon, Man. Conch. x. pp. 16, 106. 


THE MOLLUSCA OF TORRES STRAITS. 179 


HALIoripz&. 


224. Hattoris varia, L. 
Station 13, Mer (Murray Island). 


225. Haxtoris (PapoLius) ovina, Chemn. 
Station 18, Mer. 


226. Haxtotis (TErnorts) asintna, L. 
Station 5, Boydong Cays. 


FISSURELLIDZ. 

227. FissuURELLA (GLYPHIS) CORBICULA, Sow. ? 

Not quite normal, as the perforation is more rotund than in 
either typical F. corbicula or F. lanceolata, Sow. The solitary 
example was so covered with extraneous growths and somewhat 
broken, that we cannot with absolute certainty pronounce upon it. 

Station 8, Albany Pass. 


998, FIssuRELLA (Guypuis) JuKEst, Rve. 
Station 2, Warrior Island. 


229. Scutus uneuis, L. 
Station 10, Channel between Hammond Island and Wednesday 
Spit. 


230. RimvuLa Exguisita, A. Ad. 

Station 10, Channel between Hammond Island and Wednesday 
Spit. 

231. PHENACOLEPAS LINGUA-VIVERR& *, sp.n. (Pl. 10. fig. 11.) 

S. testa ovata, candida, parum nitida, subpellucida, rotunde 
antice convexa, apice incurvo, acuminato, marginem posticum 
superimpendente, undique longitudinaliter arcte radiatim cos- 
tulata, costulis arctissime et minute papilliferis, intus albida, 
margine purum incrassato. Alt. 5, lat. 11, diam. 8 mm. 

Two specimens, precisely alike, of a beautiful, thickly longi- 
tudinally radiate shell, the radiations being closely studded with 
white papille; the form is ovate, apex incurved, much over- 
hanging the posterior margin, front side elevated and convex; 
surface of the interior white, the margin hardly thickened. 

Phenacolepas, Pilsbry 1891, must take the place of Scutellina, 
Gray 1847, non Agassiz 1841 (Hehinodermata). 


* Lingua-viverre, civet-cat’s tongue, from the small papille. 


180 MESSRS. J. C. MELVILL AND RB. STANDEN ON 


POLY PLACOPHORA. 


- CHITONIDA. 


232. Curron pictus, fve. 

We are indebted to Mr. EH. R. Sykes for the following and 
other notes on the Polyplacophora of the collection :—“ Having 
examined Reeve’s types, I am unable to separate his species 
from that of Gould, originally described from Fiji. Two sets of 
figures in the Moll. U.S. Explor. Exped. bear the number 4384: 
the present species is represented by the right-hand set; the 
others being apparently a slip for 431. The slits in the largest 
specimen are: head-valve 8, tail-valve 12” (#. #. S.). This 
species was also found very sparingly by Mr. Brazier during the 
‘Chevert’ expedition, at Darnley Island. Endemic in the 
Torres Straits. 

Station 13, Mer. 


233. IscHNnocHIToN (HAPLOPLAX) sp. 

One very young specimen of a species of Haploplax; when 
alive it measured about 4 to 5 mm. “It is closely allied to 
I. (Haploplax) purus, Sykes; but the girdle-scales are large and 
striate. Too young to describe” (#. #. S.). 

Station 10, Channel between Hammond Island and Wednes- 
day Spit. 

234. TONICIA FORTILIRATA, five. ? 

“One immature specimen, which may belong to this species ; 
it does not show the granulose lateral areas, but this may be due 
to its youth” (#. R. 8.). 

The localities given for 7. fortilirata by Mr. Edgar Smith 
(Voy. ‘ Alert,’ 1881-82, p. 84) are Port Darwin, 8-10 fathoms 
(Coppinger) ; and Raine’s Island (Reeve). Endemic. 

Station 14, Channel between Mer and Davar. 


235. Tonicra conrossa, Gould. 
Station 13, Mer. 

236. ACANTHOPLEURA SPINIGERA, Sow. 
Station 13, Mer. 


237. ACANTHOCHITON. ‘Two species. 

i.) “One, immature, striate at the beaks and granulose on 
the lateral areas” (H. R. S.). 

Station 13, Mér, on reef. 


THE MOLLUSCA OF TORRES STRAITS. 181k 


(u.) “Two young specimens of an interesting form, chiefly 
noteworthy for the sculpture, which consists of radiating riblets 
from the beaks of the valves” (H. BR. S.). 

Station 10, Channel between Hammond Island and Wednesday 
Spit. 


238. Scuizocurton inctsus, Sow. 
Station 18, Mer. 


239. CRYPTOPLAX STRIATUS, Lam. 
Station 13, Mer. 


240, CRYPTOPLAX OCULATUS, Q. & G. 
In many stages of growth, some being unusually fine. 
Station 13, Mer. 


SCAPHOPODA. 


DENTALIID&. 
241. DENTALIUM PSEUDO-SEXAGONUM, Desh. 
Station 1, Thursday Island. 


242. DENTALIUM JAVANUM, Sow. 
Station 8, Albany Pass. 


PELECYPODA. 


OsTREIDA. 


243. OsTREA TUBERCULARIS, Lam. 
Station 8, Albany Pass. 


ANOMIIDS. 
244. Puacunanomia (Monta) tonr, Gray. 
Station 18, Mer. 


245. PuacuNna LoBATA, Sow. 
Station 2, 20 miles N.N.W. Warrior Island, 52 fathoms, 
Aug. 15, 1888 (A. C. H.). 


SPONDYLID &. 


246. PLicaTULA AUSTRALIS, Lam. 
Station 18, Meér. 


182 MESSRS. J. C. MELVILL AND R. STANDEN ON 


247. PuicaTuLA imBRicata, ke. 
Station 5, Boydong Cays. 


248. SPONDYLUS BARBATUS, Ltve. 
Station 18, Mer. 


249. SPONDYLUS FOLIACEUS, Chemn. 
250. SPONDYLUS NICOBARICUS, Chemn. 
251. SPONDYLUs OcCELLATUS, Rve. 


252. SPONDYLUS PACIFICUS, Ave. 


Limip#. 

253. Lima sguamosa, Lam. 

The Torres Straits form is indistinguishable from that occurring 
in the Mediterranean. We have traced it across the Isthmus of 
Suez to Red Sea waters, then along the coasts of India and 
Ceylon, eastward, uninterruptedly. 

Station 138, Murray Island; Station 14, Mer, channel between 
reefs ; Station 2, Warrior Island. 


254. Lima (LimaTuLa) BULLATA, Born. 
Station 5, Boydong Cays. 


255. Lima (Limatuna) Torrestana, HE. A. Smith. (‘ Chal- 
lenger’ Rep. vol. xiii. p. 291, pl. xxiv. figs.5,5 a.) (Pl. 11. 
figs. 19, 19 a.) 

Many examples, but for the most part only disassociated 
valves, of this rare endemic species. These are less convex 
than the type, and we therefore consider a figure necessary. 
Measurements as follows :—Alt. 12, lat. 10, diam. 8 mm. 

Station 13, Mér (Murray Island); Station 8, Albany Pass ; 
Station 5, Boydong Cays. 


256. Lima (CTENOIDES) FRAGILIS, Chemn. 
Station 1, Thursday Island (Sept. 1888); Station 2, Warrior 
Island; Station 12, Mabuiag (Oct. 1888). 


257. Lima (CTENOIDES) TENERA, Chemn. 
Station 2, Warrior Island. 


258. Lima (Mantettum) arcuara, Sow. 
Station 10, between Hammond Island and Wednesday Spit. 


259. Lima (ManrEetium) inriata, Lam. 
Station 13, Mer. 


THE MOLLUSCA OF TORRES STRAITS. 183 


PECTINID#. 
260. AMUSSIUM PLEURONECTES, L. 
Station 5, Boydong Cays. 
261. PectEeNn (PsSEUDAMUSSIUM) ARGENTEUS, Rve. 
Station 2, Warrior Island. 
262. PEcTEN BLANDUS, Jive. 
Station 14, Mér, channels between reefs. 


263. PEcTEN cRasstcostaTus, Sow. 
Station 14, Mer. 


264. Precten Crovucuti, Smith. 
Station 10; Station 8, Albany Pass. Mauritius (Smith), where 
it has hitherto been considered endemie. 


265. PECTEN LEMNISOCATUS, Lve. 
Station 8, Albany Pass. 


266. PECcTEN LENTIGINOSUS, Rve. 
Station 2, Warrior Island ; Station 5, Boydong Cays. 


267. PECTEN LiMaTULA, Rve. 
Station 5, Boydong Cays. 

268. PECTEN MADREPORARUM, Pett. 
Station 5, Boydong Cays. 

269. PrcTEN paLiium, LD. 

Station 5, Boydong Cays. 

270. PECTEN sENATORIUS, Gimel. 
Station 8, Albany Pass. 


271. Pecren (CHLAMYS) CUNEATUS, Ltve. 
Station 2, Warrior Island. 
AVICULIDS. 


272. AVICULA AQUATILIS, Rve. 
Station 2, Warrior Island. 


273. AvicuULA ALA-coRvi, Chemn. 
- Station 14, Mér. 
274, AVICULA cROCEA, Chemn. 


275. AVICULA MALLEOIDEs, Rve. 
Station 14, Meér. 


276, MELEAGRINA ANOMIOIDES, Jve. 


184 MESSRS. J. C. MELVILL AND R. STANDEN ON 


277. MELEAGRINA MARGARITIFERA, L. 
278. MELEAGRINA TEGULATA, Rve. 


279. VULSELLA LINGuLATA, Lam. (Mya vulsella, L.) 
Station 8, Albany Pass. 


980. CRENATULA FLAMMEA, five. 
Station 2, Warrior Island. 


981. PERNA ATTENUATA, Ltve. 
Station 5, Boydong Cays. 


282. PERNA AUSTRALICA, five. 


983. PERNA LENTIGINOSA, ve. 
Station 10, between Hammond Island and Wednesday Spit. 


284. Pinna FuMaTA, Hanley. 
Station 14, Channel below Mer. 


285. Pinna (Arrina) NiaRA, Chemn. 


Mytinip az. 
286. Myrinus HoRRIDUS, Dkr. 
287. Myrinus (AuLAcoMYaA) HIRSUTUS, Lam. 


288. SEPTIFER NICOBARICUS, Chemn. 
Station 18, Murray Island; Station 5, Boydong Cays. 


289. MopioLA ARBORESCENS, Chemn. 
Station 5, Boydong Cays. 


290. MopioLa AUSTRALIS, Gray. 
Station 10, Hammond Island and Wednesday Spit. 
291. Mopiona ciInNAMOMEA, Chemn. 


292. Moptona (ApuLA) LaNrGERA, Dkr. 
Station 2, Warrior Island. 


298. Mopioia LIGNEA, Fve. 


294. Moptota PHILIPPINARUM, Hanley. 
Station 5, Boydong Cays. 


295. Mopronarra Cuminerana, Dkr. 
Station 2, Warrior Island; Station 5, Boydong Cays. 


296. LiInHOPHAGUS CANALIFERUS, Hanley. 
Station 5, Boydong Cays; Station 11, Mabuiag. 


THE MOLLUSCA OF TORRES STRAITS. 185 


297. LITHOPHAGUS @RACILIS, Phil. 
Station 2, Warrior Island. 


298. LirHopHacus Hantryanus, Dkr. 


299. LirHorHacus TERES, Phil. 
Station 2, Warrior Island ; Station 18, Mér. 


ARCID&. 


300. ARCA NAVICULARIS, Brug. 
Station 2, Warrior Island ; Station 5, Boydong Cays. 


~ 


301. Arca ZEBRA, Swains. 

Station 5, Boydong Cays. 

302. ARCA ZEBUENSIS, ve. 

Station 5, Boydong Cays. 

303. BarBATIA DECUSSATA, Sow. 

Station 14, Mer ; Station 8, Albany Pass, 10 fathoms. 


304. BaRBaTIA voLtucris, fve. 
Station 2, Warrior Island; Station 11, Mabuiag. 


305. BarBATIA FUSCA, Brug. 
Station 18, Murray Island; Station 5, Boydong Cays. 


306. BarRBATIA IMBRICATA, Brug. 

Station 2, Warrior Island; Station 8, Albany Pass; Station 5, 
Boydong Cays. 

307. BaARBATIA LIMA, Eve. 

Station 10, Hammond Island; Station 5, Boydong Cays. 


£308. BarBaTIA TENELLA, Eve. 
‘Station 5, Boydong Cays. 


309. BaRBATIA TRAPEZINA, Lam. 
Station 5, Boydong Cays. 


310. Barpatia (VENUSTA) LACTEA, L. 

This species ranges throughout the whole of the Eastern 
hemisphere, being a common British shell, and its headquarters 
may be considered the Mediterranean. It is one of the few 
European marine Mollusca that occur the other side of the 
Isthmus of Suez. 

Station 8, Albany Pass. 


186 MESSRS. J. C. MELVILL AND R. STANDEN ON 


311. Barpatia (ACAR) ACERHA*, sp.n. (Pl. 10. fig. 15.) 

B. testa delicata, candida, pulchre cancellata, subquadratim 
oblonga, convexiuscula, umbonibus incurvis, prominulis, nequa- 
quam approximatis, margine dorsali utrinque recto, antice trun- 
cato, demde breviter marginem apud ventralem rotundato, latere 
postico oblique producto, subcarinato, superficie alba, delicatis- 
sime liratim costulata, liris ad latera crassioribus, sub lente 
nodosis; pagina intus alba, concava, cardine dentibus plurimis 
parum rectis predito, ligamento externo, sinu palliali inconspicuo. 
Alt. 7, lat. 15, diam. 6 mm. (spec. maj.). 

Four examples, all quite perfect, of an exceedingly highly 
sculptured little Acar, white, elegantly chased, and furnished 
with many close longitudinal radiating lire; these, when viewed 
with the aid of a lens, are nodose, and are thicker, both posteriorly 
and anteriorly, than in the centre of the shell-surface. In form 
it is quadrately oblong, very convex, the umbones not approxi- 
mate, but incurved and conspicuous ; dorsal margin straight, pro- 
duced on both sides of the umbones, anteriorly truncate, and 
finally merging in the rounding off of the ventral margin, this 
being exactly parallel to the dorsal edge. Posteriorly, the shell is 
obliquely produced, subcarinate. Within, the surfaceis white, very 
concave; pallial sinus only obscure; hinge with very many oblique 
teeth in either valve, regularly arranged. 


312. Barpatra (Acar) prvaricata, Sow. 

Station 5, Boydong Cays. 

313. Barpatra (AcaR) DOMINGENSIS, Lamarck. 

A West Indian species that seems to be making more head- 
way, so far as distribution is concerned, in the Eastern tropics 
than almost any other Pelecypod. It has also recently come 
across our notice from the Arabian Sea and North-Indian Ocean 
(Townsend). 

Station 5, Boydong Cays. 


314. Barpatta (AcAR) SCULPTILIS, Rve. 
315. ANADARA ANTIQUATA, L. 

Station 138, Mér (Murray Island). 

316. ANADARA CLATHRATA, ve. 

Station 5, Boydong Cays. 

317. PARALLELIPIPEDUM SEMITORTUM, Lam. 
Station 5, Boydong Cays. 


* dxépatos, pure. 


THE MOLLUSCA OF TORRES STRAITS. 187 


318. Pectuncutus Hover, sp.n. (PI. 11. fig. 24.) . 

P. testa orbiculari, equilaterali. equivalvi, solidiuscula, um- 
bonibus prominulis, approximatis, dorsaliter recta, ad latera 
utringue bis rufomaculata, radiatim fortiter costata, costis ad 
octo et viginti, uniformibus, pulchre nodoso-squamatis, nodulis 
interdum superficiem apud mediam evanidis, interstitiis trans- 
versim squamato-liratis; cardine dentibus ad viginti in valva 
utraque instructo, albis, simplicibus ; pagina intus albescente, ad 
latera interdum roseo-tincta. Alt. 24, lat. 25, diam. 15 mm. 
(spec. max.). 

Station 2, Warrior Island; Station 5, Boydong Cays. 

Superficially this interesting new form approaches P. nodosus, 
Reeve, a rare Ceylonese species, in which the ribs are of the 
same character, but bolder, wider in proportion, and consequently 
much fewer numerically, being only eighteen as against twenty- 
eight in P. Hoylet. The coloration also is more varied in the 
Ceylonese shell. Our species is almost colourless, being whitish, 
tinged here and there with faint red streaks, and conspicuously 
twice-blotched with red maculations both posteriorly and an- 
teriorly. The surface is thickly ribbed, the ribs being beautifully 
scaly-nodulous, the interstices between them being also squamoso- 
lirate. The teeth of the hinge in either valve are some twenty 
in number, simple. Interior whitish, tinged in most specimens 
laterally with faint rose-colour. 

We dedicate this Pectunculus to Mr. William Evans Hoyle, 
Director of the Manchester Museum, as a small recognition of 
many kindnesses experienced at his hands. 


819. PecruNncULUS vITREUS, Lam. 

This, the flattest and most placunoid of all Pectunculi, was, at 
the time of its discovery, one of the rarest of Mollusca, and 
for many years was only known by one valve, brought home by 
the voyageur M. Péron, and deposited in the Musée Royale, 
Paris. Prof. Haddon dredged about a dozen specimens, one of 
them being alive and pertect, though young. This shows 
orange-brown flecking and pure white interior. It has occurred 
at Mauritius and one or two other places in the Eastern tropics, 
and this is an interesting extension of its range. 

Station 5, Boydong Cays; Station 10, between Hammond 
Island and Wednesday Spit. 


LINN. JOURN.—ZOOLOGY, VOL. XXVII. 14 


188 MESSRS. J. C. MELVILL AND R. STANDEN ON 


320. LIMOPSIS CANCELLATA, [ve. 
Station 5, Boydong Cays; Station 10, between Hammond 
Island and Wednesday Spit. 


321. Limops1s Woopwarpt, A. Ad. 

This may possibly be only a variety of L. cancellata, Rve. 

Station 5, Boydong Cays; Station 10, between ‘Hammond 
Island and Wednesday Spit. 


322. CUCULLHA CONCAMERATA, Chemn. 
Station 10, Hammond Island. 


NvucuLipa#. 
323. NucuLa opiigua, Lam. 
324. Nucuna SIMPLEX, A. Ad. 


325. Lepa Darwint, Smith. 
Station 5, Boydong Cays. 


TRIGONIIDA. 


326. TRIGONIA UNIOPHORA, Gray. 
Station 14, Meér. 
CARDITIDA. 


327. CARDITA CARDIOIDES, Rve. 
Station 5, Boydong Cays. 


328. CARDITA MARMOREA, Ave. 
Station 14, Mér. 


329. MyTILICARDIA cRASsIcosTaTA, Lam. 
Station 2, Warrior Island; Station 8, Albany Pass; Station 
14, Mér. 


330. MytTiticarDIA MURICATA, Sow. 
Station 14, Meér. 
331. MYTILICARDIA VARIEGATA, Brug. 
Station 14, Mer. 

CRASSATELLIDA. 


332. CRASSATELLA ZIczac, Rve. 
Station 8, Albany Pass. 


THE MOLLUSCA OF TORRES STRAITS. 189 


ERYCINID 2%. 


333. KELLIA PHYSEMA*, sp.n. (PI. 10. fig. 14.) 

K. testa pellucida, vitrea, globulari, bullacea, convexa, nitida, 
equivalvi, umbonibus haud conspicuis, contiguis, sub lente deli- 
catissime irregulariter concentrice lirata, margine dorsali rapide 
declivi, postice, simul ac antice, leniter rotundata ; intus pellucida, 
ad margines lactea, ligamento interno, cardine dentibus duobus 
in utraque valva predito. Alt. 8, lat. 7°50, diam. 4mm. 

A most fragile and delicate globular pellucid Keldia, round 
the margins somewhat milky, otherwise vitreous, equivalve, with 
inconspicuous umbones, smooth, very microscopically concentri- 
cally lirate, the margins rounded, orbicular ; ligament internal ; 
inner surface of shell almost pellucid; the hinge provided with 
two teeth, one central one lateral, in each valve. 


334. TELLIMYA EPHIPPIOLUM 7, sp. un. (PI. 10. figs. 13, 18 a.) 

T. testa tenui, albescente; valvis cymbeformibus, profunde 
convexis, umbonibus parvis, antice obliquis, contiguis, superficie 
in medio depressa, utrinque lobata, precipue marginem apud 
ventralem, antice subproducta, rotundata, postice curta; intus 
alba, subpellucida, sub lente cirea margines minutissime longi- 
tudinaliter striata. Alt. 4, lat. 5, diam. 5°50 mm. 

Station 5, Boydong Cays. 

The ventrally depressed surface of the valves of this little 
species are very peculiar, and they are also extremely convex, the 
umbones contiguous, inclining anteriorly; within, the margins 
are microscopically longitudinally striate. The nearest ally 
would seem to be 7, triangularis, Gould. 


GALEOMMIIDA. 


300. SCINTILLA ALBERTI, Smith. 
Station 2, Warrior Island. 


336. SCINTILLA HYALINA, Desh. 
Station 8, Albany Pass. 


TRIDACNID A. 


337. TRIDACNA ELONGATA, Lam. 
Station 13, Murray Island. | 
* gvonpa, a bubble. 
t Ephippiolum, dim, of ephippium, a saddle. 
14* 


190 MESSRS. J. C. MELVILL AND R. STANDEN ON 


838. TRIDACNA SERRIFERA, Lam. 
Station 14, Mer. 


339. Tripacna squamosa, Lam. 
Station 14, Mer. 


CARDIID®. 


340. CarDIUM TENUICOSTATUM, Lam. 
Station 5, Boydong Cays. 


341. Carpium (TRACHYCARDIUM) DIANTHINUM™, sp. n. (PI. 11. 
figs. 25, 25 a.) 

C. testa oblique rotunda, tenui, pallide straminea, sepe ad 
medium carneo vel puniceo-suffusa, sepe unicolore, vel straminea 
vel undique punicea, longitudinaliter costata, costis quinque et 
triginta, planatis, postice serrulatis, et antice minute squamatis, 
umbonibus contiguis, parvis, ligamento externo; intus variante, 
interdum straminea, interdum carnea, marginibus postice dentatis, 
superficie interna delicate longitudinaliter lirata. Alt. 14, lat. 
12, diam. 8 mm. (spec. maj.). 

Station 10, Channel between Hammond Island and Wednesday 
Spit. 

A variably coloured, delicate, and bright little shell, its chief 
peculiarities being the obliquely rounded outline (in one variety 
somewhat squarely set towards the ventral margin), and the 
smooth ribs, fine and close, some 35 in number, only ser- 
rulate posteriorly and in front minutely scaly. Within, the 
surface varies as do the outer valves in coloration, the delicate 
ribs showing through, so thin is the substance of the shell. 

Several specimens. 


342. Carpium (TRACHYCARDIUM) ELONGATUM, Brug. 

One magnificent example, of narrower contour than the type 
fizured by Reeve, and which does not possess so many ribs, these 
being only 38 in number, as against 43 or 44 in normal specimens. 
The colour too is less vivid, and shows no trace of rufous, being 
of a pale cinereous-ochre. 

Station 14, Mer. 


343. Carpium (TRACHYCARDIUM) LACUNOSUM, Fve. 
A variety only, the ribs being but 30 in number, the super- 


* Dianthinus, resembling a carnation. 


THE MOLLUSCA OF TORRES STRAITS. 191 


ficies highly coloured, and somewhat stunted and incrassate. 
Reddish ochraceous without, pure milky-white within; marginal 
teeth acute and red-tipped. 

Station 8, Albany Pass. 


344. Carpium (TRACHYCARDIUM) MACULOSUM, Wood. 
Station 2, Warrior Island. 


345, CaRpiuM (TRACHYCARDIUM) RUBICUNDUM, Jive. 
Station 5, Boydong Cays. 


346. Carprium (TRacHycarpIuM) RuUgosUM, Lam. 
Station 5, Boydong Cays. 


346a@. Carpium (T'RACHYCARDIUM) SERRICOSTATUM, sp. 0. 
(Pl. 11. fig. 20.) 

C. testa ovata, crassiuscula, cinereo-alba, fere equilaterali, 
umbonibus incurvis, contiguis; valvis longitudinaliter costatis, 
costis ad quadraginta, prominulis, delicate serrulatis, antice et 
postice squamulosis, interstitiis excavatis, margine dorsali utrin- 
que leniter declivi, regulari, lateribus posticis, simul ac anticis 
marginem ad ventralem rotundatis ; intus cinereo-alba, circa mar- 
eines striato-dentata; dentibus cardinalibus fortibus, lateralibus 
prominulis. Alt. 20, lat. 18, diam. 12 mm. 

Station 4, Ormans Reef; Station 13, Murray Island. 

Several valves of a distinct species we have been quite unable 
to match. The peculiarly serrated, squarely-set ribs, some forty 
in number, are characteristic. The interstices are deeply ex- 
cavate, the umbones not very prominent, contiguous. Shell 
almost equilateral, the dorsal margin on each side of the umbones 
gently rounded off, both posteriorly and anteriorly, towards the 
ventral margin. Within, the shell is ashy-white, towards the 
margins striato-dentate, hinge teeth strong, laterals prominent. 

Three or four examples. 


347. Carpium (TRACHYCARDIUM) TRANSCENDENS, sp. 2. 
(Pl. 11. fig. 21.) 

C. testa ovata, delicata, arctissime et pulcherrime longitudina- 
liter tenuicostata, costis circa octo et quinquaginta, umbonibus 
levibus, nitidis, contiguis, paullulum antice inversis; valvis con- 
vexis, margine dorsali utrinque rapide declivi, postice et antice 
leniter marginem apud ventralem rotundatis, superficie pallide 


192 MESSRS. J. C. MELVILLE AND R. STANDEN ON 


straminea, ad latera puniceo-maculata, et undique carneo- et 
puniceo-suffusa, ad margines flavescente, ligamento externo ;. 
intus superficie pallide carnea, ad medium ochraceo-punicea,. 
nitida, delicatissime striata, apud margines serrata. Alt. 25, 
lat. 21, diam. 15 mm. 

A remarkably elegant Trachycardiwm, its nearest ally being 
C. maculosum, Wood, which it resembles in coloration, and to 
some extent in form, it being, however, more equally oval, and 
possessing about 58 as against 38 ribs. C. obovale, Sow., and 
C. mauritianum, Desh., likewise C. arenicolum, Reeve, are akin, 
but each possesses salient and marked individual characteristics 
of its own. The species now before us differs from all others of 
its section in its exceedingly fine longitudinal costulations and 
very delicate pink and pale yellow colour. It is roundly oval, 
thin, with the umbones red, shining, closely approximate; valves 
extremely convex, dorsally speedily sloping both ways, and then 
gently rounding off into the ventral margin. Within, the 
surface is delicately longitudinally striate, pale flesh-colour, 
yellowish red in the contre, shining, the margin of the valves 
serrated. 


348. Carprum (TRACHYCARDIUM) VARIEGATUM, Sow. 


345. Carpium (PapyRIDHA) PAPYRACEUM, Ohemn. 


350. Carpium (Lavicarpium) Brcuet, Ad. & Rve. 

One young valve only. A species of extreme rarity, and 
almost unknown in collections. Two magnificent specimens, 
formerly in the Colonial Exhibition of 1884, are now in the 
British Museum (Nat. Hist.). 

Station 2, Warrior Island. 


351. CarpiuM (L&VICARDIUM) BIRADIATUM, Brug. 
Station 13, Mér; Station 8, Albany Pass. 


352. Carpium (LAVICARDIUM) LyRATUM, Sow. 


353. HEMICARDIUM SUBRETUSUM, Sow. 
Station 10, Hammond Island. 


354, Hemicarpium (Fracum) unevo, L. 
Station 13, Mér; Station 5, Boydong Cays. 


THE MOLLUSCA OF TORRES STRAITS. 193 


355. HemicarpDiIuM (CTENOCARDIUM) FORNICATUM, Sow. 

For many years unique, this exquisite shell has been lately 
found, though always rarely, in the Arabian Sea (Ff. W. Towns- 
end), as well as now in the Torres Straits. 


356. HemMIcarDIUM (CTENOCARDIUM) FrAGUM, ZL. 
Station 13, Murray Island; Station 5, Boydong Cays. 


357. Hemicarpium (CTENOCARDIUM) IMBRICATUM, Sow. 
Station 18, Murray Island. 


CHAMIDA, 


358. CHAMA DIVARICATA, Rve. 
Station 10, between Hammond Island and Wednesday Spit- 


359. CHAMA FIBULA, ve. 


360. CHAMA FIMBRIATA, [tve. 
Station 5, Boydong Cays. 


361. CHAMA PELLIS-PHOCA, Rve. 
Station 14, Mér. 


362. CHaMA PULCHELLA, ve. 
Station 14, Mer. 


363. CHAMA REFLEXA, Fve. 
Station 5, Boydong Cays. 


364. Caama (ARCINELLA) SPINOSA, Brod. 
Station 5, Boydong Cays. 


CYPRINIDZ. 


365. Lipitina anauLata, Lam. 
Station 8, Albany Pass. 


VENERID&. 


3866. DIoNnE INFLATA, Sow. 
Station 5, Boydong Cays. 


367. LioconcHa HEBR&A, Lam. 
The smaller Lioconche are somewhat difficult to differentiate. 
Though they may not be considered particularly variable, yet so 


194 MESSRS. J. C. MELVILL AND R. STANDEN ON 


many so-called “species? were described in the earlier years of 
this century by Lamarck, Deshayes, and others, that much con- 
fusion naturally exists, and we should not be surprised if some 
future monographer were not only to merge this with ZL. tigrina, 
Lam., and Z. Sowerbyi, Desh., but also make more sweeping 
reforms still. 

Station 5, Boydong Cays. 


368. LroconcHa Prcta, Lam. 
Station 14, Mér. 


369. Prrar rEe@vLaris, Smith (as Caryatis). (‘ Challenger’ 
Rep. vol. xiv. pl. 1. figs. 38-3 b.) 

Pitar, Romer, has priority of five years over Caryatis of the 
same author. 


370. CIRcE CASTRENSIS, Z. 
Station 2, Warrior Island; Station 5, Boydong Cays; Station 
4, Ormans Reef. 


371. Circe PecTINATA, L. 

We believe that it will be found that the Circe (or Crista) 
pectinata, L., and C. gibbia, Lam., are the extreme forms of one 
variable species. Gibbia, as its name would imply, is the large 
swollen-valved shell, coarsely longitudinally nodulous-costate ; 
whilst normal forms of pectinata are flatter, simply oblong, and 
the nodules, though present, hardly developed. Gibbia, again, is 
as a rule colourless, with the exception of a dark bluish-brown 
variegation posteriorly ; while pectinata is, in all examples we 
have seen, more or less flecked throughout with zigzag brown 
markings. The distribution of both species is much the same, 
and embraces the whole of the tropical East Indies. 

Station 14, Mér ; Station 5, Boydong Cays. 


372. CIRCE RIVULARIS, Born. 
Station 5, Boydong Cays. 


373. DosIntA cmHRULEA, Rve. 


374. DOSINIA HISTRIO, Gmel. 

It is common and variable in the Arabian Sea and Persian 
Gulf (Zownsend); Andamans (Booley); Madras and S. Indian 
Ocean (Henderson) ; Ceylon (Layard), &c. <A variety from 
Murray Island is the D. lyrata, Sow. 


THE MOLLUSCA OF TORRES STRAITS. 195 


Station 2, Warrior Island; Station 18, Murray Island; 
Station 5, Boydong Cays ; Station 4, Ormans Reef ; Station 10, 
Hammond Island. 


375. CHIONE (OMPHALOCLATHRUM) CuEmNirzi1, Hanley. 
Station 8, Albany Pass. 


376. CHIONE (OMPHALOCLATHRUM) COSTELLIFERA, Ad. 
Station 13, Murray Island. 


377. CHIONE (OMPHALOCLATHRUM) EMBRITHES*, sp.u. (PI. 11. 
figs. 23, 23a.) 

C. testa percrassa, multum convexa, ovato-rotunda, equivalvi, 
ineequilaterali, umbonibus inversis, approximatis, superficie con- 
centrice costulata, costis tornatis, interstitils arctissime sculptis, 
deinde calcareo-tessellatis, longitudinaliter quadriradiata, in- 
terdum sparsim brunneo-maculata, interdum unicolore, lunula 
perexcavata, latere antico multum compresso, deinde marginem 
apud ventralem rotundato, postico abbreviato ; pagina intus alba, 
nitida, circiter impressionem muscularem posticam pulchre 
rubescente, linea palliali conspicua, sinuosa ; cardine incrassato, 
dentibus cardinalibus simul ac laterali postico conspicuis, antico 
perminimo, marginibus intus serrulatis. Alt. 28, lat. 30, diam. 
22 mm. (spec. maj.). 

Station 8, Albany Pass ; Station 4, Ormans Reef. 

Hitherto mixed up and confounded with C. torewna, Gould, 
also inhabiting the same seas, this species can be with ease 
differentiated by the extremely excavate lunule and less frequent 
concentric laminz. Besides these distinctions, the interstitial 
spaces between the lamine are more delicately tornate, and 
assume a lateritial pattern, resembling layers of minute bricks, 
laid regularly, some being of chalky-white consistency. The 
substance of the shell, below the umbones especially, is peculiarly 
massive. Hach valve is ornamented with three or four con- 
spicuous longitudinal rays. Within, the surface is white, 
shining, beautifully stained with red round the posterior mus- 
cular impression, the pallial line being conspicuous and 
sinuous; the hinge is thickened; the cardinal teeth and the 
posterior lateral large, the anterior exceedingly small; the 
margins of the valves being serrate within. 

* éuBorO1)s, heavy. 


196 MESSRS. J. C. MELVILL AND R. STANDEN ON 


378. CHIONE (OMPHALOCLATHRUM) LAMARCKII, Gray. 
Station 2, Warrior Island. 


379. CHIONE (OMPHALOCLATHRUM) MARICA, L. 
Station 5, Boydong Cays; Station 13, Murray Island. 


880. CHIonE (OMPHALOCLATHRUM) ListERI, Gray. 
Station 13, Murray Island. 


381. CHIONE (OMPHALOCLATHRUM) SUBNODULOSA, Hanley. 
Station 10, Hammond Island. 


3882. CHIONE (OMPHALOCLATHRUM) TOREUMA, Gould. 
Station 2, Warrior Island. 


383. ANAITIS CALOPHYLLA, Hanley. 
Station 5, Boydong Cays. 


384. ANAITIS THIARA, Dillwyn. 
Station 4, Ormans Reef ; Station 10, Hammond Island. 


385. PAPHIA GLABRATA, Desh. 
Our specimens are more rostrate than the type. 
Station 10, Hammond Island. 


386. Papua mitis, Desh. 


387. Tapes DusHayvesn, Hanley. 


388. TAPES MALABARICA, Sow. 
Station 4, Ormans Keef. 


389. Tapes (PAREMBOLA) LITTERATA, L. 
Station 2, Warrior Island. 


390. Tapes (PAREMBOLA) RADIATA, Chemn. 


391. Tapes (TExTRIx) suLcosa, Phil. 
Station 2, Warrior Island; Station 4, Mer. 


392. Tapes (TExTrIx) TEXTRIX, Chemn. 


393. KATELYSIA SCALARINA, Lam. 
Station 8, Albany Pass. 


THE MOLLUSCA OF TORRES STRAITS. 197 


UNGULINID 


394. DipLoponTa ETHIMa*, sp.n. (PI. 11. figs. 17, 17a.) 

D. testa oblique globulari, tenui, profunde convexa, umbonibus 
contiguis, incurvis, margine dorsali antice rapide declivi, rotun- 
dato, postice primum fere recto, deinde abrupte marginem circa 
ventralem rotundato, superficie extus alba, parum nitente, con- 
centrice rudi-lirata; pagina intus alba, cardine duobus dentibus 
parvis in valva utraque predita, linea palliali simplici, inconspicua. 
Alt. 11, lat. 11, diam. 8 mm. 

A thin, globular, somewhat oblique Diplodonta ; the umbones, 
owing to the great convexity of form, rather conspicuous and 
incurved anteriorly. Though this shell possesses no very marked 
characteristics, we have been unable to match it with any of the 
numerous described species. 


395. DipLoponra suBcrassa, Smith. (Rep. Zool. Coll. Voy. 
H.M.S. ‘ Alert,’ 1881-82, p. 104, pl. 7. figs. L, LI.) 

We have been confirmed in our naming by the author, with 
whom we compared our examples with the type, which came 
from West Island, Prince of Wales Channel (Station 9). 


396. Diptoponta suBaLoBosa, Smith. (‘ Challenger’ Rep., 
Lamellibranchs, p. 197, pl. 14. figs. 10, 10 a.) 

Two valves, on which Mr. H. A. Smith has also kindly pro- 
nounced an opinion, remarking that these are more adult than the 
types, and have a slight difference in shape. 


PSAMMOBIIDZ. 


397. PSAMMOBIA RASILIS T, sp. nu. (Pl. 11. fig. 18.) 

P. testa oblonga, tenui, fere levi, umbonibus inconspicuis, 
roseo-suffusis, superficie carnea, longitudinaliter septem-radiata, 
concentrice rudi-striata, margine dorsali antice recta, prolongata, 
ad juncturam marginis ventralis uniangulata, postice abbreviata, 
longitudinaliter corrugata, ligamento externo; intus pallide 
punicea, subpellucente, cardine in altera valva dente bifido, in 
altera indiviso, preedita, linea palliali sinuata. Alt. 16, lat. 27, 
diam. 3°50 mm. (spec. maj.). 

An elegantly rayed oblong species, almost smooth, tinted with 
rose at the inconspicuous umbones, flesh-coloured as to its 


* é0.uos, usual, commonplace. t+ Rasilis, smoothish. 


198 MESSRS. J. C. MELVILL AND R. STANDEN ON 


surface, and longitudinally seven-rayed; its dorsal margin is 
prolonged, and almost straight till its sudden angle on merging 
into the ventral margin, anteriorly, while the hinder margin is 
abbreviated, the surface being longitudinally wrinkled; the 
ligament is external; within, the surface is pale pink; the hinge 
has in one valve a bifid, in the other an undivided single tooth ; 
the pallial line is sinuate. 


398. PsamMoBia (GARI) ANOMALA, Desh. 

Our specimens do not show the usual radiations of this 
delicate shell. 

Station 5, Boydong Cays. 


399. PsamMMoBia (GARI) MARMOREA, Desh. 
Station 18, Mér. 


400. Psammosta (Garr) ornata, Desh. 
A yellow, prettily rayed variety. 
Station 13, Mer. 


401. PsamMosia (Gant) PRmSTANS, Desh. 
402. Psammopia (GARI) PULCHERRIMA, Desh. 


403. SOLETELLINA VIRESCENS, Desh. 

A curious shining pale ochraceous variety, with two longitudinal 
broad orange flames, one on each side of the umbones, very 
evanescent towards the ventral margin. It is a rare form, and 
seldom met with. 

Station 138, Murray Island. 


404. ASAPHIS DEFLORATA, DL. 

It is possible that all the Asaphis of the Western hemisphere, 
e. g. A. dichotoma, Auton, A. coccinea, Mart., &c., are but varieties 
of one variable form, equally abundant in the East. 

Station 2, Warrior Island; Station 5, Boydong Cays. 


SOLENIDA. 
405. SOLENOCURTUS (AZAR) COARCTATUS, Gimel. 
Station 18, Murray Island. 
MESODESMATIDA. 


406. MrsopESMA PR&#HCISA, Desh. 
Station 8, Albany Pass. 


THE MOLLUSCA OF TORRES STRAITS. 199 


Macrrip2#. 


407. Macrra aprcina, Desh. 
Station 13, Murray Island. 


408. Macrra acHatina, Chemn. 
409. Macrra (HemMImMactRA) aspersa, Sow. 


410. Macrra (Oxyprras) Copprncert, Smith. 


This very interesting form occurred but sparingly, and much 
encrusted with nullipores. 
Station 4, Ormans Reef. 


411. Razta Grayi, H. Adams. 
Hab. Borneo. 


412. Lurrartra arcuata, Desh. 
413. LUTRARIA RHYNCHANA, Jonas. 
Myipa. 


414. CorBuLA CRASSA, Hinds. 
Station 10, Hammond Island ; Station 5, Boydong Cays. 


415. Corputa Maceriiivrayi, Smith. 
416. CoRBULA TAHEITENSIS, Lam. 


417. CorspuLa TRUNCATA, Hinds. 


G-ASTROCH ANID. 
418. GASTROCHHENA CUNEIFORMIS, Lam. 


419. GASTROCHEHNA PLICATILIS, Desh. 
TEREDINID ®. 
420. TEREDO NUCIVORA, Speng. 
Station 14, Mer. 
DIBRANCHIA. 
LUCINACEA. 
Lucrnipa. 


421. Luctna (Divaricetta) Macannrem, A. Ad. 
We have seen this only previously from the Red Sea (Capt. 
Shopland). Slightly coarser in sculpture than L. ornata, Rve., 


200 MESSRS. J. C. MELVILL AND R. STANDEN ON 


but very similar. Indeed all the Divaricelle need a more 
critical differentiation. 
Station 5, Boydong Cays. 


422. Luctna (DIVARICELLA) ORNATA, Pve. 


493. Lucina (CoDAKIA) EXASPERATA, Ave. 
Station 5, Boydong Cays. 


424. Lucrina (CoDAKIA) FIBULA, Mve. 
Station 5, Boydong Cays. 

425. LucIna (CODAKIA) INTERRUPTA, Lam. 
Station 138, Murray Island. 


426. LORIPES ICTERICA, Rve. 
Station 2, Warrior Island. 


427. Lorrers Happont, sp. un. (PI. 10. fig. 12.) 

L. testa alba, orbiculari, equivalvi, fere xquilaterali, tenui, 
umbonibus rostratis, haud prominulis, contiguis, inflexis, lunula 
excavata, angusta, superficie concentrice striata, ligamento 
interno; cardine valve sinistre dentibus primariis duobus, dextree 
uno dente majore centrali predito ; intus marginem circa ven- 
tralem longitudinaliter striata, linea pallial conspicua, haud 
multum sinuosa. Alt. 13, lat. 18, diam. 6 mm. 

Station 2, Warrior Island. 

A typical Loripes, white, rather thin, orbicular, finely con- 
centrically striate, the umbones not very prominent, rostrate, 
inflexed; lunule narrow, excavate; the hinge of the left valve 
with two small primary teeth, of the right with one large tooth. 
Within, the surface is white, rudely longitudinally striate ; pallial 
line conspicuous, broadly sinuous. 

Two perfect examples and one valve. 


428. CorBIS ELEGANS, Desh. 
Station 13, Mer. 
TELLINID A. 
429, TeLiina (TELLINELLA) ASPERRIMA, Hanley. 


430. Te~uina (TELLINELLA) IRIDESCENS, Bens. 
Station 2, Warrior Island. 


431. Tenuina (TELLINELLA) STAURELLA, Lam. 


432. TELLINA (TELLINELLA) VIR@ATA, L. 
Station 5, Boydong Cays. 


THE MOLLUSCA OF TORRES STRAITS. 201 


433. TELLINA (TELLINELLA) VULSELLA, Chemn. 
Station 5, Boydong Cays. 

434, TELLina (DoNACILLA) RHOMBOIDES, G'mel. 
Station 5, Boydong Cays. 


435. TELLINA (DONACILLA) SEMITORTA, Sow. 

436. TELLINA (Donactiia) vireuLatTa, Hanley. 
437, TELLINA (TELLINIDES) EMARGINATA, Sow. 
438. TELLINA (ANGULUS) PHILIPPINARUM, Hanley. 


439. TELLINA (ANGULUS) PROcRITA™, sp. n. (PI. 10. fig. 16.) 

T. testa nitida, lata, ovata, tenui, pallide straminea, delicate 
radiis puniceis decorata, planata, umbonibus inconspicuis, planis- 
simis, antice vix declivi, deinde abrupte rotundata, latere postico 
breviter rotundato, margine ventrali leniter utrinque declivi ; 
intus pellucente, puniceo-suffusa, radiis conspicuis, cardine parvo, 
normali. Alt. 18, lat. 21, diam. 3°50 mm. 

A shining, very flattened, broadly ovate Tellina, pellucid, with 
pale straw ground-colour, radiately ornamented with pale pink; 
the umbones are very flattened and inconspicuous; anteriorly the 
sides hardly siope until they suddenly merge into the rounded 
ventral margin, the posterior side is roundly contracted. Within, 
the radiation shows through, the hinge is small and normal, 
pallial sinus very obscurely seen. 


440, TrLtina (ANGULUS) VERNALIS, Hanley. 
Station 5, Boydong Cays. 
441. Tecuina (ARcopaAata) prneuis, Hanley. 
Station 5, Boydong Cays. 
442, Tentina (Arcopacta) Savienyt, A. Ad. 
Station 5, Boydong Cays. 


443, TELLINA (ARCOPAGIA) TESSELLATA, Desh, 


SCROBICULARIID A. 
444, SEMELE DUPLICATA, Sow. 


445, SEMELE JuKsEsI, A. Ad. 

Station 10, Channel between Hammond Island and Wednesday 
Spit, Sept. 15, 1888. 

446. SEMELE LAMELLOSA, Sow. 


* spdxpiros, select, choice. 


202 MESSRS. J. C. MELVILLE AND BR. STANDEN ON 


CUSPIDARIID A. 


447. CUSPIDARIA LATISULCATA, Ten.-Woods. 
Station 18, Mér; Station 5, Boydong Cays. 


PANDORID®. 


448. PANDORA sp. 

It is unfortunate that the only specimen in the collection is 
in such unsatisfactory condition as to preclude a more detailed 
description, for we are convinced it is new. Valve acinaciform, 
trapezoid, inequilateral, arcuate and produced posteriorly, 
ultimately suddenly truncated and merged into the rounded 
ventral margin; anterior margin much abbreviated ; umbones 
pointed, not conspicuous; surface radiately unequally wavy- 
costulate, which show through on the interior surface. 

Station 138, Mer (Murray Island). 


PHOLADOMYID&. 


PHotapomya, Sowerby, 1823. 
§ Parinmmya, subgenus novum. 


Valve fere equilaterales, antice solum paullum hiulce, cost 
longitudinales valde obscure, umbones parvi. 


449, P. (Parttrmya) Happont, sp. nov. (PI. 11. figs. 22, 
22 a, 22 b.) 

P. testa cinereo-albida, delicatula, tenui, equivalvi, fere xequi- 
laterali, oblonga, postice clausa, antice paullum hiante; umboni- 
bus parvis, antice incurvis, approximatis, latere antico declivi, 
breviter truncatulo, marginem versus ventralem subrotundato, 
postice paullum magis producto, denique leniter rotundato, 
superficie fere levi, concentrice inequiliter rudi-striatula, radiatim 
costulis arctis, pellucidis, quasi-internis pulchre ornata; ligamento 
externo ; pagina intus cardine utriusque valve fere simplici, dente 
marginali obscuro, parvo, nitido, quasi detrito, fossa triangulari 
valvam apud dextram ; linea palliali fere obsoleta. 

Alt. 17, lat. 22, diam. 9 mm. (spec. maj.). 

Alt. 9, lat. 13, diam. 6 mm. (spec. min.). 

“Station 2, Warrior Island, at 53 fathoms, amongst broken 
shells and sand. August 15, 1888.”—A. C. H. 

Shell thin, equivalve, and almost equilateral, very delicate, 


THE MOLLUSCA OF TORRES STRAITS. 203 


smoothish, oblong, and slightly gaping anteriorly, the posterior 
portion of the valves closed in repose. The umbones are hardly 
prominent, and incline forwards, being closely approximate; the 
anterior margin is at first sloping, then somewhat truncately 
rounded off, and merging into the ventral margin, while 
posteriorly it is slightly produced and gently rounded. The 
surface of both valves is alike, smoothish, milky- or ashy-white, 
semipellucid, indistinctly but rudely concentrically striate, and 
closely beset with radiating quasi-internal riblets, hardly standing 
out beyond the superficies; indeed in young examples these 
radiations appear as pellucid lines imbedded in the internal 
layers of the shell. The ligament is external. Within, the 
hinge of either valve is almost destitute of teeth or any processes 
whatsoever, save that a small, worn-looking, centrally situated 
lamellar tooth is present, most conspicuous in the left valve, 
though there is a corresponding tooth and an elongately 
triangular pit in the right valve also. 


A few remarks on this peculiarly interesting genus may not 
be out of place here. 

Pholadomya, instituted by the elder Sowerby in 1823, is one 
in which but very few recent species are embraced, since certain 
forms described by Agassiz as nearly akin to the type (for example, 
P. caspica) have been, and rightly, relegated to the toothless 
section of the Cardiidex, e. g. Adacna, Hichwald. 

P. candida, Sow., the original type, is still of extremely in- 
frequent occurrence, nearly all the specimens hitherto obtained 
having been cast ashore from considerable depths after storms 
and hurricanes off the island of Tortola, in the Antilles. The 
anatomy has been worked out by Sir Richard Owen, who de- 
scribes the mantle-margins as being united, save where space is 
left: for the foot and outlets occur for the siphonal and anal 
orifices ; the gills on either side single, the outer lamina produced 
dorsally, and there is also an accessory, bifureate foot. A fine 
example of the shell in the private collection of one of us 
exhibits a large, papyraceous, fragile, white, extremely inezqui- 
lateral surface, equivalve, very convex centrally and towards the 
umbones, widely gaping both anteriorly and posteriorly. In 
form obliquely trapezoid, anteriorly roundly truncate, concen- 
trically rudely striate, and, centrally only, rayed from the 

LINN. JOURN.—ZOOLOGY, VOL. XXVII. 15 


204 MESSRS. J. C. MELVILL AND R. STANDEN ON 


umbones to the ventral margin ; rays much elevated, about eleven 
in number, broader towards the ventral margin, nodulous where 
the rude concentric strie cross them. Within, slightly nacreous, 
showing, owing to its extreme tenuity, the precise obverse of 
the external sculpture; hinge with an elongate trigonal “ fossa,” 
and a marginal “ lamina” in either valve. 

The great thinness of the substance causes the shell often to 
be worn away at the umbones; this is owing to incessant friction 
in opening and shutting the valves. Here we have the typical 
Pholadomya, a more appropriate cognomen than which could 
not have been possibly coined, for the resemblance to a large 
Pholas (e.g. P. costata, lu.) is very striking. Many extinct forms 
approach the recent shell both in size, facies, and ornamentation, 
and several even exceed it in beauty and magnitude. These are 
mainly Secondary and Tertiary fossils, especially abounding in 
the Liassic or Jurassic rocks. In the majority of instances, owing 
to the shell-substance having been originally so thin, they are 
mere casts, but are exact replicas of their former analogues in 
all salient particulars. 

In 1842* Agassiz subdivided the genus into various sections, 
considering the presence or absence of a posterior area (‘aire 
cardinale circonscrite”’) of primary consequence. Thirty-four 
years later, Meek, in 18767, considered that only two divisions 
were worthy of establishment, viz.: Pholadomya (type, as before, 
P. candida, Sow.), distinguished by an elongate inequilateral 
form, gaping at both extremities, and roundly truncate in front ; 
and Procardia, to contain the short, gibbous, subtrigonous, ex- 
tremely truncate, and often elevated forms, to which the greater 
proportion of the fossil species belong. These now exceed two 
_ hundred in number. 

Turning again to the recent forms, the second, discovered only 
afew years ago, is P. Loven, Jefir., of the typical section of 
the genus, and we consider it here worth while reproducing 
Dr. Gwyn Jeffreys’s original description, so as to place it side by 
side with that of our new species, to which, however, it does not 
bear much near resembiance. This is a remarkably interesting 


* IL. Agassiz, Etudes critiques sur les Mollusques fossiles: Neuchatel, 
1842-45. 


+ A Report on the Invertebrate Cretaceous and Tertiary Fossils of the 
Upper Missouri country: U.S. Geol. Survey of the Territories (Meek), 1876. 


THE MOLLUSCA OF TORRES STRAITS. 205 


molluse, evidently of wide distribution, ranging from the mid- 
Atlantic Ocean to Sicilian localities in the Mediterranean. 

“Pholadomya Lovent, Jeffreys, Proc. Zool. Soc. 1881 
(‘lightning’ and ‘Porcupine’ Exped., 1868-70), p. 934, 
pleplxexu ties (7, 

“Shell inequilateral, wedge-shaped, gaping at the posterior 
end, convex, of a pearly nature, partly semi-transparent, lustre- 
less; sculpture, 10-12 longitudinal ribs, besides some intermediate 
striz ; these are more or less interrupted by strong periodical 
marks of growth, so as to give the ribs a nodulous appearance ; 
the sides are ribless; the whole surface is covered with minute 
prickly tubercles ; colowr white; margins rounded on the anterior 
side, inclining upwards towards the other side, which is also 
rounded but slightly truncate, sloping at the back from each 
side of the umbo; beaks bluntly triangular, turned inwards ; 
umbones prominent; ligamental pit in the right valve obtuse- 
angled, placed outside underneath the beak, and defined outwards 
by a thin plate ; hinge-line sloping towards the posterior side ; 
hinge-plate thin, sinuous, reflected ; teeth none; inside highly 
glossy and nacreous; scars conspicuous. L. 0°4, B. 0°5. 

“« Porcupine’ Exped. 1870; Atl. St. 22, 28a; Med. 55. 

“None of the specimens are quite perfect. One of them 
indicates twice the size given in the description. That figured is 
from the ‘ Josephine’ Expedition. 

“ Dist. Palermo, frequent (Monterosato), 162 fathoms. Off 
Marseilles (‘Travailleur’ Exped. 1881); Villa Franca, Azores 
(‘ Josephine ’ Exped.), 320-600 fathoms.”—J. G. J. 

Into the two groups proposed by Meek we are unable precisely 
to allocate the P. Haddont. 'o this, indeed, we can find nothing 
near akin, unless an obscure species described by Mr. Bullen 
Newton in 1892*, as occurring in the Upper Keuper Sandstone, 
Shrewley, Warwickshire, prove its ally. Certainly from figure 
and description it appears analogous, particularly as to being 
nearly equilateral, with radiately ribbed surface, these ribs, how- 
ever, being far more pronounced. It was provisionally named 
Pholadomya(?) Richardsit, after Mr. H. P. Richards, the dis- 
coverer. 

From the differentiation we have given above, and which need 
not be recapitulated, we plead justification for the creation of a 


* J. of Conch. vii. p. 411, fig. 2 (1894). 


206 THE MOLLUSCA OF TORRES STRAITS. 


new subgeneric division. It only remains to say that several 
examples (mostly, however, single valves, though one at least 
occurred quite perfect) were found of P. Haddoni, dredged at 
about 5 to 6 fathoms in the localities above cited. It is not, 


- therefore, a deep-sea form. 
In dedicating this molluse to its discoverer, we would once 


again tender him our best thanks for having given us every 
facility and latitude in the preparation of this Report; and, we 
may add, it is by his express desire that all the types have been 
permanently deposited in our National Collection at the British 


Museum (Natural History), South Kensington. 


EXPLANATION OF THE PLATES. 


Puare 10. 

Fig. 1. Murex (Ptercnotus) saibaiensis, sp. n. 
2. Murex (Ocinebra) salmoneus, sp. n. 
3. Nassa (Alectryon) fretorum, sp. n. 

4. Mangilia chionea, sp. n. 

5. Triforis (Ino) excelsior, sp. n. 

6. Bittium torresiense, sp. n. 

7. Hulima australasiaca, sp. n. 

8, 8a. Magadis ewmerintha, sp. u. 

9. Calliostoma (Hutrochus) septenarium, sp. n. 
10, 10a. Microtheca Acidalia, sp. n. 
11. Phenacolepas lingua-viverre, sp. n. 
12. Loripes Haddoni, sp. u. 

13. Tellimya ephippiolum, sp. n. 

14. Kellia physema, sp. n. 

15, Barbatia (Acar) acerea, sp. n. 
16. Zellina (Angulus) procrita, sp. n. 


Prats 11. 
Fig. 17, 17 a. Diplodonta ethima, sp. n. 
18. Psammobia rasilis, sp. n. 
19, 19a. Lima (Limatula) torresiana, B. A. Smith, var. 
20. Cardium (Trachycardium) serricostatum, sp. n. 
21. Cardium (Trachycardium) transcendens, sp. n. 
22, 22a, 226. Pholadomya Haddoni, sp. n. 
23, 23a. Chione (Omphaloclathrum) embrithes, sp. n. 
24, Pectunculus Hoyle, sp. n. 
25, 25a. Cardium (Trachycardium) dianthinum, sp. n. 


Linn. Soc. Journ. Zoot. Vou. XXVII.Pu 10. 


Melvill &Standen, 


7 


Mantern Bros.imp. 


J.Greendel.et lth. 


MOLLUSCA FROM TORRES STRAITS. 


Lousn.Soc.Journ. Zoot. Von. XXVII. Pri. : 


Melvill & Standen. 


Mintern Bros .imp. 


J.Green del.et lth. 


MORE C SCATE ROM TORINE S (SaiRAIGE Sy. 


ON THE EXTERNAL NARES OF THE CORMORANT. 207 


Note on the External Nares of the Cormorant. 
By W. P. Pycrart, A.L.S. 


[Read 2nd March, 1899.] 


Tue following short note, with two figures, is intended as a 
supplement to a similar contribution to the pages of this Journal 
made by Prof. J. C. Ewart *. 

In that communication Prof. Ewart described the external 
nostril as a “mere slit situated at the end of a shallow superficial 
groove, which runs backwards along the beak parallel with its 
lower edge, and lying between its lower and middle third.” 
And concerning it he wrote: ‘‘ Whena bristle is introduced into the 
slit, ij never succeeds in forcing a passage into the nasal cavity. 
If the skin which forms the outer boundary of the slit is carefully 


Fig. 1. 


Left side view of the head of a Cormorant (Phalacrocorax carbo) in which the 
rhamphotheca has been displaced to show the horny plug which has 
been withdrawn from the aperture of the external nares.—, bristle; g, 
gnathotheca ; g', posterior maxillary portion of the gnathotheca; .a., 

- external narial aperture. 


reflexed, a groove is exposed which runs from the external slit-like 
nostril to a narrow canal lined apparently by modified mucous 
membrane ......it is possible to pass through this canal, 


* Vol. xv. 1881, p. 455. 


208 MR. W. Py, PYCRAFT ON THE 


without forming a false passage, a bristle about the size of an 
ordinary horse-hair..... 

The external nostril in every species of adult Cormorant which 
I have examined appears to lie without and below the rhino- 
thecal groove (fig. 1, v.a.), and not at its end as just described. 
As seen in the figure, it is represented by a shallow groove 


Fig. 2. 


Left side view of the head of an embryo Cormorant to show the relatively large 
narial aperture lying within the rhinothecal groove.—rh.g., rhinothecal 
groove. 


pointing downwards and forwards to the tomium. Its upper end 
joins the rhinothecal groove at its base, where it passes into the 
naked skin of the lores. 

I have not been able to find any trace of this narial groove, or 
aperture, in the Gannets. 

I failed entirely to pass even the finest bristle up this groove 
into the nasal cavity, though I tried in many species of Cormorant— 
one a fresh, and the rest spirit-specimens. 

My next step in this investigation was to force the rhampho- 
theca from the jaw. The rhinotheca was first raised and then 
the gnathotheca. This last brought away with it a short rod-like 
plug (fig. 1, p. 207)—apparently an inward and backward con- 
tinuation of that part of the rhamphotheca surrounding the 
external narial aperture, indicated by the groove just described 
(fig. 1, n.a.). Thus, instead of surrounding the aperture and 
giving place in this region to mucous membrane, the sheath 
seems to have grown inwards so as to form a horny tubular 


EXTERNAL NARES OF THE CORMORANT. 209 


lining. Microscopical examination—made in the first instance 
by my friend Mr. H. M. Bernard, and afterwards confirmed 
by myself—showed that the lumen of the “plug” was com- 
pletely blocked up by delamination of fragments of the horny 
layers from its inner surface. A bristle passed down the narial 
aperture, now thrown open by the removal of the plug, made 
its exit as usual at the posterior nares. 

The nasal cavity is a small chamber devoid of any trace of 
turbinal folds, and more or less imperfectly divided into anterior 
and posterior moities by a vertical transverse partition depend- 
ing from the roof into the chamber, the ventral border of the 
partition being free. 

In the embryo (fig. 2) the narial aperture lies within the 
rhinothecal groove, much as described by Ewart in the adult, 
and is still open. It is interesting to note that the oblique 
groove for this aperture in the adult is not yet indicated, neither 
is the segmented portion of the posterior end of the gnathotheca, 
so marked in the Gannets though comparatively slightly developed 
in the Cormorants (fig. 1, 9’). 

Mr. F. A. Lucas in ‘The Auk’ (vol. xiv. p. 87) has an interest- 
ing note on the external nares of the embryo and nestling 
Cormorant. He finds the nostrils still open in the oldest of 
the nestlings in his collection; the age of these he estimates at 
28 days. He considers “ that the external nostrils close about 
the time the young Cormorants take to the water and begin to 
feed themselves.’ 


LINN. JOUBN.—ZOOLOGY, VOL. XXVII. 16 


210 MR. P. CHALMERS MITCHELL ON SO-CALLED 


On so-called “ Quintocubitalism”’ in the Wing of Birds; with 
special refererce to the Columbe, and Notes on Anatomy. 
By P. Cuatmers Mrrcuett, M.A., F.L.S.* 


[Read 16th March, 1899.] 
(Puates 12 & 13.) 


Since the results of Wray’s investigations were published (1), 
the occurrence of two well-marked modes of disposition of the 
quill-feathers on the upper part of the wing of birds has been 
well known, and the explanation of the existence of the two 
conditions has been sought by many zoologists. In one mode of 
disposition, that known as “ quintocubitalism,” the quill-feathers 
which abut on the ulna are arranged in a regular and even series, 
each feather with its upper and lower covert being of approxi- 
mately the same size and lying at the same approximate distance 
from its neighbours on the distal and proximal sides. In the 
disposition termed “‘ aquintocubital” the first four quills, counting 
from the distal towards the proximal end of the ulna, are arranged 
precisely as in the quintocubital wing, but, after the fourth, 
there is a gap in which there is an upper and lower covert pre- 
cisely as in the regular arrangement but no quill between them. 
Thereafter the quills follow in regular series. It appears as if 
the fifth quill had been lost without any other disturbance of the 
series, and the condition was called “‘ aquintocubital,” 2. e. with- 
out the fifth cubital, on account of this ready interpretation. 
It appears to me that it would be more convenient to state the 
facts in another way. Immediately distad of the cubital quills 


* [This Memoir is complementary to that by Mr. W. P. Pycraft, which 
follows (infra, pp. 286-254). During the autumn of 1898, Mr. Pycraft inti- 
mated his intention of early presenting to the Society a memoir on the so-called 
Aquintocubitalism in the Bird’s wing, and in subsequent conversation with 
Mr. Mitchell he discovered that the latter had already arrived at the same 
main conclusion as himself, and that he had lodged a preliminary statement 
‘concerning it with the Editor of a scientific journal. When these facts became 
known to the Officers of the Linnean Society, they approached the two 
gentlemen with a proposal that their memoirs might be presented at one of 
the Society’s meetings and published together in its Journal, and to this they 
willingly agreed, Mr. Chalmers Mitchell very generously withdrawing the 
afore-mentioned press notice. 

Except that the authors agree on the main issue, to which they came 
‘‘independently and unknown to one another,” their papers will be found to 
supplement each other—one author having approached the subject through 
the study of development, the other through that of adult anatomy.—Eb. | 


“ QUINTOCUBITALISM ” IN THE WING OF BIRDS. 211 


there is a variable remex, which when normally developed, as in 
Nothura,.is in obvious series with the cubital remiges, and which 
is frequently smaller in size but connected to the cubital series by 
a special plica of membrane as in most pigeons, but which may be 
absent; although in the case of the Columbe, to which this memoir 
has special reference, I have not noticed a case of absence. 
I believe the simplest way to state the facts is to adhere strictly 
to the division of the remiges or wing-quills into primaries and 
secondaries, to consider the carpal remex as the first secondary, 
and then to say that, after the fifth secondary, there may be a gap 
more or less equivalent to the space which would be occupied by 
a sixth secondary in even series, after which the secondaries 
continue in normal series; or that the secondaries may all lie in 
normal series without the occurrence of a gap. For the first 
condition, that hitherto known as “ aquintocubital,” I propose 
the term “ diastataxic”; for the second condition, that known 
as “quintocubital,” I propose the term “ ewtawic.” These new 
terms are simply descriptive ; they convey no implication as to the 
‘way in which the two conditions arose, and they appear to me to 
‘be equally applicable, whether we accept the current view that 
tthe diastataxie condition has come about by the disappearance 
of a fifth secondary from an eutaxic series, or if, as I believe, 
there is no lost feather. If there be no missing feather, it is 
obvious that the diastataxic condition might have arisen from 
the eutaxic condition by elongation of the wing im the region 
of the gap without the addition of a quill to the series; or that 
the eutaxic condition might have come from a diastataxic con- 
dition by the closing-up of the quills without consequent oblite- 
ration of the gap. I hope in this memoir to show reasons for the 
latter view—for, in fact, the view that the diastataxic condition 
is architaxic. 

For some time I have been engaged in a special study of the 
Columb. These, like most of the larger groups of birds, have 
been described as diastataxic, and, without question, the wing 
in the majority of them presents a well-marked gap. ‘This is 
well seen in the wing of a common pigeon, where the gap is as 
large as in a duck orin am eagle, and is occupied by an apparently 
normal covert. In Turtur chinensis (P1. 18. fig. 7) there are ten 
primaries with their major coverts placed (as I find invariably 
among the Columbide) distally to the corresponding quills; then 
comes a moderately-sized carpal remex bound down by a special 

16* 


212, MR. P- CHALMERS MITCHELL ON SO-CALLED 


fold of the wing-membrane to the next secondary ; the carpal 
covert is, like all the secondary major coverts, proximad of the 
corresponding quill in insertion. After the carpal remex there 
follow four secondaries, then a gap which is slightly exaggerated 
in the drawing, then six other secondaries i even series. The 
major covert in the gap is bound to the major covert proximal to 
it by a thin slip of membrane. In Geotrygon montana (PI. 13. 
fig. 5) the carpal covert is very small, and under it lies a small 
remex bound down to the next secondary in normal fashion. 
Then follow twelve secondaries placed at almost equal distances, 
so that at first sight the arrangement appears to be eutaxic.. 
Examination of the interspaces, however, shows that there is a 
small additional covert clearly belonging to the major series but 
with no corresponding quill. The same condition is very plaim 
in Gna capensis; it is only the presence of the covert that makes 
it possible to regard the wing as diastataxic. The odd covert is 
plainly crowded, a condition which is still more plain in the 
specimen than in the diagram, in which for convenience of drawing 
the relative size of the feathers as compared with the interspaces 
has been minimized. I have found in a certain number of the 
Columb that the wing is practically the same as in Geotrygow 
and (na, with the most important difference that there appears 
to be no extra covert, and that, in consequence, the wing must be 
regarded as eutaxic, and the group Columbide added to those 
among which both eutaxic and diastataxic conditions occur. 
Nearly two years ago, when I found this eutaxie condition in 
Columbula picut, 1 showed thespecimens to my friend Mr. Beddard, 
in whose laboratory in the Zoological Gardens I was prosecuting 
my work, and with my consent he noticed the observation in his. 
recent work on Birds (2. p. 305). With that exception the 
observation is new to literature, and since then I have found 
the same condition in a number of other pigeons. In Geopelia 
cuneata (Pl. 12. fig. 1) and in G. tranguilla there are ten pri- 
maries, each with a distal major covert ; then comes the small 
carpal covert and remex, the latter with its usual slip binding it 
to the adjacent secondary ; then follow in even series ten fully- 
developed secondaries, each with normal proximally-placed major 
coverts, and in the diastataxic interspace there is neither any 
trace of gap nor an extra covert. In Lewcosarcia picata (PI. 13. 
fig. 2) the conditions are identical, except that there are eleven 
fully-developed secondaries. In Geophaps plumifera (PI. 13. 
fig. 3) the same condition exists, except that there are twelve 


““ QUINTOCUBITALISM ’’ IN THE WING OF BIRDS. 213 


fully-developed secondary quills. In Columbula picui there are 
ten primaries and also twelve fully-developed secondaries. In 
Starnenas cyanocephala (P1. 13. fig. 6) there are ten primaries, 
then an unusually iarge carpal remex with the normal binding 
fold and covered by a small carpal covert; then follow twelve 
fully-developed secondaries without any trace of the diastataxic 
gap. In Phiogenas cruentata there are nine primaries, then a 
normal covert and remex, then twelve fully-formed secondaries, 
with no trace of the gap. 

Tt appears, then, that the Columbe form an interesting addition 
to Kingfishers and Swifts, some members of which groups exhibit 
the one condition, others the other; and it is among such groups 
that we may hope to find an explanation of the divergence in 
structure *. The mere statement of the facts as they occur among 
the Columbz seems to me at once to suggest an extremely prob- 
able explanation. In most pigeons the wing is diastataxic, 
with a large gap occupied by a “covert. In some pigeons 
{e. g. Gina and Greotrygon) there is practically no gap but a 
covert crowded into the interspace which forms the gap in most 
pigeons ; in Geopelia cuneata and G. tranquilla, in Columbula, 
Leucosarcia, Starnenas, Geophaps, and Phlogenas there is no 
gap, and there is apparently no extra covert. 

It is now necessary to consider a third row of feathers, for 
simplicity not represented in the figures in the Plates, but shown 
in fig. 1 in the text. Lying apparently in between the quills’ 
and coverts, but really belonging to a more dorsal series, there 
is a small feather marked 3 in the figure; except in the 
gap marked x there is plenty of room for these third-series 
feathers. In the gap, even in a diastataxic bird (fig. 1, I), there 
is a certain crowding of these feathers, which are rather smaller 
than the others of their series. In a diastataxic bird where 
there is no actual gap, as in Gna or Greotrygon (fig. 1, 11), the 
two feathers on either side of the actual covert are exceedingly 
crowded, and are markedly degenerate, relatively much more so 
than in the diagram, while the covert itself is smaller in size. 
Jn the eutaxic birds (fig. 1, III) there is only one feather in this 
interspace ; as a result the interspace does not differ from the 
other interspaces. Which of the three crowded feathers has 
remained I cannot be certain, but I think it probable tbat it is 

* The late Mr. Seebohm made the suggestion that eutaxie species may 


possibly have arisen from diastataxic ancestors by suppression of the coverts 
(Classification of Birds, 1895, Suppl. p. 8). “4 


214 MR. P. CHALMERS MITCHELL ON SO-CALLED 


the so-called diastataxic covert in a form so much reduced as to 
appear to be one of the ordinary third series. My explanation 
is, then, that pigeons were originally diastataxic with a full gap: 
occupied by one feather of the upper-covert series and two small 
feathers of the third series. In some pigeons there has been a 
closing-up of the ranks of quills so that there is no actual gap: 
in the diastataxic space, while into this reduced space there are 


Willi 


iff 
44d 


biti phpper 
att 


2- 


Lb 
IC 


I. Diagram of secondaries (1), coverts (2), third-series feathers (3), im 
Columbx ; x, diastataxic gap. Complete diastataxic condition. 
II. Geotrygon: same lettering, showing reduced diastataxic gap (x) with 
three crowded feathers in it. 
IIT. Leucosarcia: same lettering. Hutaxic condition. No gap in the inter- 
space (x), and only one feather, which may be one of series 3, or 2. 
crowded three feathers, the two smaller markedly reduced and 
the covert itself not seldom noticeably smaller. Finally, a certain 
number of pigeons have become absolutely eutaxic by the sup- 
pression of the two smaller feathers, leaving the covert in a 
reduced form, or at least by the suppression of two of the three 
feathers. The eutaxic condition is a modification of the more 
primitive diastataxie condition, and intermediate conditions. 
oceur. 
Systematic Position of the Eutaxic Pigeons. 


The nomenclature I have followed is that used in the ‘ List of 
Animals’ published by the Zoological Society, as I obtained my 


“ QUINTOCUBITALISM ”’ IN THE WING OF BIRDS. 215 


specimens from the Zoological Society’s Gardens. These names 
are identical with those in the British Museum Catalogue, except 
in the cases of the bird I name Phloganas eruentata, which is 
there ealled P. Juzonica, and the bird I name Geophaps plumifera, 
which is there named Lophophaps plumifera. 

Garrod (8) considered the classification of pigeons from the point 
of view of anatomy. He does not place Geophaps, but of the 
other eutaxic forms, Lewcosarcia is placed in his family Phapide, 
and the others each in a different division of his family Treronide. 
According to the British Museum system, all the eutaxie forms 
belong to the family Peristeride but are distributed among four 
different subfamilies. It is clear that these eutasic forms cannot 
be regarded as forming a separate group by themselves. Here 
and there, almost at random, among the great mass of diastataxic 
forms occur a few eutaxic forms. If, as I have attempted to 
show in the earlier part of this paper, the eutaxic condition be 
a simple derivative of the diastataxic condition, then it is not 
surprising to find that quite different genera are convergent in 
this respect. I shall now show that there is considerable reason, 
based on anatomy, for regarding these eutaxic pigeons as con- 
vergent in other respects. 


Anatomical Reasons for the supposition that Hutaxie Pigeons 
are not Primitive. 


To avoid constant repetition of the names of the seven pigeons 
the wing of which displays the eutaxic condition, I shall refer 
to them according to the following list :— 


CrAO Dalit) COMGHED oooc00cec005 dhe 
Geopelia tranquilla. > ee a. 
Phloqanas cruentatu 
Phlogenas ? luzonica pieces. Re 
Starnanas cyanocephala ...... OC. 
Geophaps plumifera 
ee lagi. D. 
? Lophophaps ,, 
Givn DUG WUCUE. sonra boosece E. 
View COSareia PUCatm 2.12 4 ire 


When the anatomical differences presented by a large number 
of closely-related individuals are considered, it is generally 
possible to come to a conclusion as to what conditions are more 


216 MR. P. CHALMERS MITCHELL ON SO-CALLED 


primitive and what are secondary. In different groups certain 
structures are on the wane, others are in process of development, 
and such general changes appear to characterize whole groups ; 
the complete assemblage of related animals appears to be moving 
in the same direction; the same structures appear to be in 
process of advancement or of degeneration, so that the special 
characters of the whole group tend to become more and more 
accentuated. In this general progress, individual species or 
genera may advance specially rapidly or may lag behind; and 
those which are most or least advanced in the direction of the 
whole group are by no means necessarily most closely related to 
one another, although in the structures most affected by the 
general course of the changes, strong convergent resemblances 
result. In the case of pigeons for instance, there are anatomical 
considerations by which one may consider particular individuals 
or species more or less “ pigeony ” pigeons (to coin a convenient 
melegancy), and the species which are most “ pigeony”’ are not 
specially related one to the other. For some time I have been 
engaged on the anatomy of this group with the special view of 
tracing such progressive advances and degenerations. The work 
involves dissection of many hundreds of individuals, and it will 
be long before I am in a position to publish final results. There 
are many sources of error, some of which can be eliminated only 
by comparison of the anatomy of many individuals of the same 
species and the same variety. I have for the present purpose 
drawn on my notes only for such points as appear to me to be 
unusually clear; and in this way, although I leave out many 
features which I have no personal doubt will prove of interest, 
I gain in immediate certainty and brevity. The general con- 
clusion to which I come is that the eutaxic forms display a 
number of anatomical features which show them to be well to 
the front among pigeons generaily in the progressive changes for 
which evidence is to be found among pigeons—that, in fact, they 
are more “ pigeony ”’ than their diastataxic allies. I do not mean 
that every eutaxic pigeon exhibits every progressive advance or 
degeneration more notably than every diastataxic form, but that 
on the whole they do exhibit such changes in an unusual degree. 

In the argument which 1 am attempting to develop there is 
the apparent flaw that advance in one direction is not necessarily 
associated with advance in other anatomical structures. It is 
a familiar condition to find extreme specialization in certain 
directions associated with extremely primitive conditions in other 


“ QUINTOCUBITALISM ”’ IN THE WING OF BIRDS. 217 


directions. From this point of view, it might be said that even 
were the eutaxic condition primitive it would not be surprising 
to find that birds primitive in that respect were much specialized 
in other directions. I am not yet prepared to meet this argument 
fully in its application to pigeons; I can only say that I find 
that where the lines of progressive change are clear, there appears 
to me to be a high average of association among the changes. 
Taking only the changes characteristic in a group, individuals with 
one of the changes well marked have a high average of the other 
changes. Groupsare, in fact, characterized by a tendency to par- 
ticular variations in particular structures; these variations are 
individual and in a state of flux in the more primitive species, but 
tend to become fixed as specific or generic characters in more 
advanced types. 


Muscular Anatomy. 

M. rhomboideus superficialis—In the majority of Columbide 
according to Fiirbringer (4), and I am able to corroborate him, 
the origin of this muscle is fleshy. In A, a, B, C, and FE it has 
become tendinous, while in E part of the anterior end of the 
muscle is a degenerate fibrous sheet. In the vast majority of 
birds this muscle has passed into what is certainly the secondary 
condition of being tendinous in origin. The Columbe are peculiar 
in that most of them present the more primitive condition, but 
in five out of the seven eutaxic forms this primitive condition 
has been lost. 

M. supracoracoideus— The great development of this muscle 
is one of the special features of the anatomy of the Columbe. 
Tn its highest development, it extends to the extreme tip of the 
sternum and invades the keel to a considerable extent. A special 
feature, which will be noticed with the osteology, is the tendency 
to formation of a strong smooth ridge of insertion which carries 
the line of the coracoid across the anterior edge of the keel. In 
all the seven eutaxic pigeons the muscle and its ridges has 
reached the extreme development found in the group. Asso- 
ciated with this extreme development is a markedly bipinnate 
arrangement of the muscle-fibres on their central tendon; the 
extent to which this occurs varies among pigeons, but is strongly 
marked in all the eutaxial forms. 

M. coracobrachialis externus.—This muscle from the coracoid 
to the planum bicipitalis of the humerus is a large muscle in 
Ratites, but in Carinates is on the wane, entirely disappearing 
in some of the Passeres. In the Columbe generally it is very 


218 MR. P. CHALMERS MITCHELL ON SO-CALLED 


small and partly covered by the biceps tendon, but frequently 
possesses a fair proportion of muscular fibres. In all the 
cutaxic pigeons it is extremely small, in some of them being 
practically reduced to a tendon. 

MM, latissimus dorsi anterior et posterior.—These muscles are 
extremely variable among birds, and pigeons show a considerable 
range of difference. The general tendency among them is for 
the anterior muscle, originally narrow and strap-like, to extend 
its origin and to broaden out; the posterior, on the other hand, 
is becoming reduced. In all the eutaxic pigeons, the anterior 
muscle has a fairly broad origin ranging from about the third 
last cervical, or just in front of that, to the first or second dorsal. 
In this respect these pigeons cannot be said to be markedly in 
front of the diastataxic forms, but they occupy an advanced 
place among the progressive forms. On the other hand, they 
are all markedly advanced in the degeneration of the posterior 
muscle. In E there is a slight fibrous representative of it, in 
the others it is completely absent. In diastataxic forms it is 
frequently present, with distinct origin and insertion ; in Columba 
it is variable individually and specifically. 

Group of Alar Muscles.—In the angle between the humerus 
and forearm there are a number of muscular structures to which 
great attention has been paid by Garrod, Gadow, Firbringer, 
Beddard, and others ; and those structures have been shown to 
possess considerable systematic value. In pigeons generally 
there is great variety in the component parts of this group of 
structures, and the differences both in their general aspect and 
in their individual details have great significance. The general 
tendency undoubtedly is to increase the musculature attached 
nearer the proximal end of the forearm and to decrease that 
more distally inserted. The mechanical effect of these changes, 
which obviously are progressive in the group, is to strengthen 
the muscular pull, which from its insertion nearer the fulerum 
increases the rapidity of the upward movement of the forearm 
on the humerus. In this general tendency towards change, a 
more Passerine-like condition is being attained by many pigeons. 
The eutaxic pigeons show the changes, some of them in an 
extreme form, all of them notably; and in this respect it 1s In~ 
teresting to remember that the Passerines are a eutaxic group. 

Biceps patagialis—This importaat patagial muscle is in origin 
a slip froin the biceps brachialis which runs to the long tendon 


Z “ QUINTOCUBILALISM ”’ IN THE WING OF BIRDS. 219 


of the deltoides patagialis. In fig. 2 B, I show it in a condition 
which is usually primitive among pigeons, and which I found 
as an individual abnormality in Columba livia. The whole slip 


Alar muscles of Columba livia. 


PAT. Deltoides patagialis. Pec. Pectoralis propatagialis. B. Biceps pata- 
gialis slip, an unusually primitive condition not common in Columha 
livia. L. Longustendon. A. Anconzus scapularis. D. Deltoides major. 
BH. Extensor metacarpi radialis. a. Firbringer’s ‘alpha’ slip. 


from its origin to insertion is muscular. In fig. 3 I the normal 
pigeon-condition is shown. The origin from the biceps is 
reduced to a rounded tendon or even a thin aponeurosis arising 
partly from the wing-membrane. Jn fig. 3 I! the most advanced 
pigeon-development is represented. The muscle is now fully 
specialized, and has a long tendon at each end with a consider- 
able muscular belly. The insertion is now very low down on 
the longus tendon, the tendon of the biceps slip running parallel 
with that for a considerable distance. The muscle is in fact 
losing its primitive function and becoming really more of am 
accessory to the radial extensors. It appears to me to be a 
probable suggestion that this extreme development may be 
tending really to the final extinction of the muscle, and therefore 
to a Passerine-like condition. My immediate point, however, is 


220 | MR. P. CHALMERS MITCHELL ON SO-CALLED ° 


that in the eutaxic pigeons the biceps slip tends to be highly 
specialized. In D the normal pigeon-condition is present ; in A, 
EH, and F, the normal insertion with a considerable origin by 


Fig. 3. 


I. Geophaps plumifera. Il. Geopelia tranquilla. III. Phlogenas cruentata, 
showing differentiation of brevis tendon into a, B, and y slips of Fiir- 
bringer. 


aponeurosis from the wing-membrane; in a, B, and C, the 
extreme development with a long tendon at either end. 
Deltoides patagialis.—In tig. 2, Pat., the normal pigeon-condi- 
tion is shown. The belly of the muscle divides into two peaks, 
that to the longus being certainly smaller but quite well marked. 
The distal end of the brevis is divided only into a distal slip 
running to the extensor, and made known by Firbringer as 
‘alpha’; the two slips ‘beta’ and ‘gamma’ are not differen- 
tiated, but are represented by a single diffuse sheet. In A, a, 
D, E, and F the longus slip is proportionately rather smaller ; in 
Band C it is exceedingly small, almost absent, and the small 


“ QUINTOCUBITALISM ” IN THE WING OF BIRDS. 221 


longus tendon arises almost directly from the general muscle. 
As for the differentiation of the brevis tendon of insertion into 
‘ alpha,’ ‘beta,’ and ‘gamma, it occurs in A, a, B, E, and F, the 
pectoralis slip forming a considerable portion of ‘beta.’ In the 
others the normal condition is attained, not passed. 

Pectoralis slip.—tin the simple pigeon-condition, as in fig. 2, 
Pec., the pectoralis muscle gives off two slips partly muscular 
and continuous at origin, and these run in almost equal propor- 
tions to the longus and brevis tendon. The general tendency is 
for the longus slip to weaken or disappear, and for the brevis 
slip to become an exceedingly sharply-marked round tendon 
passing under the deltoides patagialis, and ultimately forming 
part of the ‘alpha’ or ‘beta’ tendon. This ultimate condition 
recalls that common in Passerines, where there is only a single 
tendon running to the brevis. The longus part of the patagial 
is absent in B, C, D, and very short and slight in the others. 

Deltoides major.—The series of progressive changes found 
among pigeons in this muscle are of very great significance and 
interest. They concern its insertion, size, and subdivision. 
Typically, the muscle is moderately well developed and is inserted 
for a varying distance down the humerus. In pigeons there are 
two grades in its development. In the first grade, as Firbringer 
showed, the muscle tends to increase the length of its insertion 
until ultimately it reaches almost to the distal end of the 
humerus, where it may be pierced by the large nerve which 
supplies the forearm. This condition, represented in fig. 4 I, 1s 
shown at its maximum in most of the specimens of Columba livia 
which I have dissected, but in some specimens it has not been 
reached. Among the eutaxic pigeons it occurs in B. In the 
others, a second series of changes begin. In D the muscle is 
completely divided into two portions. The upper and smaller 
portion is inserted to the humerus near the distal end of the 
supracoracoideus muscle-tendon. lia AX, @ (C5 Jd, arnel ID ong 
first part is similarly separated off. The second part in D reaches 
down the humerus in the usual fashion, but is not actually 
divided by the radial nerve. In A and F the second part has 
reached right down the humerus and is pierced by the radial 
nerve. It is, however, very thin and attenuated, while the 
deltoides patagialis and anconzeus are both unusually large. In 
FE (fig. 4, I1) the conditions are similar, but that part of the 
attenuated long division of the muscle which lies next the 


222, MR. P. CHALMERS MITCHELL ON SO-CALLED 


anconzus (D. 3) is partly fused with it, although it has a separate 
insertion. In C and a (fig. 4, Il) the final stage has been 
reached. The distal parts of the long division of the muscle have 


I 


— 


Deltoides muscle. 
PAT. Delt. patagialis. 8. Tendon of Supracoracoideus. D. Deltoides major, 


A. Anconzeus scapularis. 
I. Columba livia. I. Columbula picui. III. Geopelia tranquilla. 


apparently quite disappeared, but are represented actually by 
small slips which leave respectively the distal ends of the deltoides 
patagialis and the anconzus to be inserted on the distal end of 
the humerus on either side of the radial nerve. Thus all the 
eutaxic pigeons show extreme development of this muscle. In 
six of them it has passed beyond the development into stages of 
degeneration. It is interesting to notice that im the Passerines 
there is usually a similar complete division of the deltoid, but I 
do not know of the existence of stages of degeneration of the 
long division. 

M. metapatagialis or Expansor secuncariorum.—This curious 
slip, upon which so much stress was laid by Gairod, is obviously 
decaying among pigeons. In Columba lita it is extremely 


“¢ QUINTOCUBITALISM’’ IN THE WING OF BIRDS. 223 


variable, nearly every condition from full formation to complete 
absence being present. Traces of it are more common than not 
among pigeons generally. In the eutaxic forms it is never well 
developed. Traces of it occur in a, B, C, and D; in the others, 
as in Passeres, it is completely absent. 

At this point, I may sum up the evidence from the muscular 
structure of the wing in the eutaxic pigeons. They all exhibit 
high stages in the progressive changes to be found among pigeons, 
and im many of these changes they recall conditions to be found 
among the Passeres. Ido not for a moment wish to suggest that 
there is any genetic relation between Pigeons and Passeres ; but 
it appears to me that in many points the specialized pigeon-wing 
shows convergent resemblances to the specialized wing of Passeres. 
These convergences are peculiarly well marked among the 
eutaxic forms. 

M. ambiens.—This muscle is one of the most interesting and 
variable structures among birds, and it is only natural to find 
that the conditions it presents among pigeons are variable and 
significant. It is impossible to doubt that the pigeons are 
among that great group of birds characterized by Garrod as 
Homalogonate, from the normal presence in them of an ambiens 
muscle. Among some Homalogonatous birds (as, for instance, 
Parrots, Herons, and Storks)the ambiens may be present orabsent. 

In a former paper (5) I showed that there may be found 
among Homalogonatous birds, apparently devoid of an ambiens, 
distinct vestiges of the former existence of that muscle. In 
another memoir (6) I was able to show that there occurred as 
individual variations in Opisthocomus almost every stage in the 
degeneration of the muscle, from complete presence to such 
reduced vestiges as I had described in Parrots and Herons. I 
do not think it open to doubt that the ambiens muscle is a 
normal and ancestral constituent of the musculature of pigeons, 
and that when it is degenerate or rudimentary or absent, such 
conditions are secondary. In the great majority of pigeons 1 
have dissected it is present in the ordinary form. On the other 
hand, among the eutaxic pigeons it shows marked traces of 
reduction. In F alone is it present in the complete normal 
condition. In a and C it is present above the knee in an 
extremely reduced form; the usual channel through the fascie 
over the knee is absent and the slender tendon disappears. 
Below the knee,and quite unconnected with the upper part, 


224, MR. P. CHALMERS MITCHELL ON SO-CALLED 


there is an origin from the head of the fibula representing the 
usual accessory tendinous head, and from this slips go to the 
flexors of the toes. In D it is present, but very slender, above 
the knee, and does not cross the knee-joint, being completely 
absent below. In B and E it is quite absent above the knee, 
but below is represented by the fibrous rudiment from the fibula 
which I made known in Parrots and Herons. In A it is com- 
pletely absent above and below the knee. 

Thus the eutaxic pigeons show a strong tendency to de- 
generation in this muscle, which is generally present in a fairly 
well-developed form among diastataxic forms of pigeons. It is 
interesting to notice that in the eutaxic Opisthocomus, which in 
many respects resembles pigeons, the ambiens is degenerate. 

MM. peroneus profundus.—This muscle is very variable in birds, 
and is present in all pigeons; but in the eutaxie forms it is 
markedly degenerate. 


Visceral Anatomy. 


I have found no indications of very great importance from 
the point of view now under consideration. The oil-gland 
is absent in only one of them (C), but it is small in B and 
F. It is almost certain that the presence of large ceca 
is a characteristic of the more primitive forms of the gut in 
birds (7). In pigeons, generally, the cxca are on the wane, but in 
the great majority of forms they are definitely present as a pair 
of nipple-like structures. Among the eutaxic forms they are 
present in B, but extremely small and showing patches of dark 
pigment as are to be found frequently on degenerate organs. 
In C, according to Garrod, they are present; in the only speci- 
men I have seen they were absent, so that it is probable that: 
individual variation occurs. In the other five eutaxic forms 
they are totally absent. The gall-bladder is absent in all the 
eutaxic forms, and as it occurs in a comparatively large number 
of diastataxic forms, its absence may have some significance. 
In all the eutaxic forms, the gut shows the arrangement of coils 
and loops which I have deseribed as typical for pigeons (7) in a 
strongly-marked form. In A, a, B, and F the middle loop, so 
characteristic of pigeons, is particularly long and shows the 
peculiar spiral twisting in of the parallel distal and proxin.al 
limbs in a strongly-marked fashion. So far as the viscera are 
concerned, it may be said with confidence that the eutaxic 
pigeons exhibit the ty pical pigeon-variations in a high degree. 


Q 


“ QUINTOCUBITALISM ’ IN THE WING OF BIRDS. 22% 


Osteological Characters. 


I have not yet proceeded so far with the collection and colla- 
tion of osteological facts in pigeons as I have done in the case 
of the muscular anatomy, and I cannot speak with the same 
conviction as to primitive and ancestral characters, but there 
are a few definite points upon which the evidence seems to me 
more clear. 

Anastomosis of Dorsal Vertebre.—In pigeons, as in some other 
groups of birds, there is a marked tendency to anastomosis of 
some of the dorsal and posterior cervical vertebra, with the 
result of giving a greater rigidity to the vertebral column. This 
anastomosis affects the vertebral centra, the neural arch, and the 
articular processes, and is the result of ossification of the liga- 
ments, such as occasionally may be found in old specimens of 
almost any group. ‘Typically, in pigeons three vertebra, usually 
the three anterior dorsals, are anchylosed in this way; and I have 
not yet found in any diastataxic form a greater number of 
vertebre affected. But among the eutaxic forms the fusion has 
proceeded further. In A, a, C, and F it affects four dorsal 
vertebre, while in B, D, and E five are united. Similarly, the 
extent to which the ilia are increased by secondary ossification 
of the adjoining ligaments and membranes is on the whole more 
extensive in the eutaxic forms. 

Another point of some little importance in pigeons, as in 
many other groups of birds, is the degeneration of the fibula. 
I do not think it can be doubted that a fully formed fibula 
is a primitive character, while degeneration implies secondary 
modification. This degeneration is usually accompanied by re- 
duction of the deep or second peroneal muscle which springs 
from a varying area of the distal end of the fibula, and which 
shows special sigus of degeneration amoug eutaxic pigeons. 
Beddard and Parsons have shown that the absence of this muscle 
among Parrots is a feature of systematic importance (8). In 
Pigeons generally the second peroneal is a strong well-developed 
muscle, and the ossification of the fibula extends almost to the 
extreme distal end of the tibia, where it is continued in a strong 
fibrous band. Among the eutaxic forms, the ossification is never 
so much as seven-eighths of the length of the tibia, and seldom 
so much. In A,a, B, C,and D it does not reach more than 
three quarters of the length of the tibia, and the whole bone is 
ZOOLOGY, VOL. XXVII. 17 


LINN. JOURN. 


226 MR. P. CHALMERS MITCHELL ON SO-CALLED 


very degenerate, the characteristic fusion with the tibia towards 
the proximal end being specially well marked. 

Supracoracotdeal crest.—The supravoracoideus muscle (pecto- 
ralis secundus) is one which tends to become very highly 
developed among pigeons. It is well known that its insertion 
on a special ridge of the humerus is a peculiar feature of pigeon 
osteology. This ridge is specially well developed among the 
eutaxic forms, but 1t would be difficult to say more than that 
this special development of pigeon anatomy was well marked in 
them. On the other hand, there is a crest of origin for this 
muscle to which I cannot find that attention has been directed. 
In its fullest development it is a T-shaped crest, the stem of the 
T carrying out the line of the coracoid across the carina of 
the sternum at right angles to the long axis of that bony pro- 
tuberance, and the cross-piece of the T lying parallel to the 
anterior edge of the carina, the lower limb running along the 
line on the carina which separates the insertion of the two 
pectoral muscles. This crest is extremely well developed 
in C, the ventral limb of the cross-piece extending nearly half- 
way down the carina (fig. 5). In all the eutaxic forms it is 


Fig. 5. 


Sie tir 


Starnenas cyanocephala.—Carina of sternum, showing supracoracoideus crest. 


very well formed, while the extent of its development among 
diastataxic forms varies. Itisjust visible in Gowra and Oalenas ; 
it is well marked, so far as the stem of the T is concerned, in 
Columba and Turtur, and in pigeons generally it is noticeable, 
as in most Passeres. But in the eutaxic forms the stem is 
always very strong and the lower limb of the cross-piece well 
marked. 

Coraco-sternal articulation.—A primitive feature, familiar in 


“ QUINTOCUBITALISM ’ IN THE WING OF BIRDS. 227 


reptiles, is that the coracoids should overlap at their junction 
with the sternum. This is well marked in some birds, and 
occurs occasionally among species of Columba and in individuals 
of C. livia. In birds generally the coracoids tend to be more 
and more separated at their sternal articulation. In most 
pigeons they either actually meet or are very close together 
indeed. Among the eutaxic forms they just meet in C and E; 
in the others they either do not touch at all or are well 
separated. 

Angle of Scapula and Coracoid.—This angle varies with the 
development of certain wing-muscles, and on the whole varies 
inversely as the power of flight. While it is impossible to lay 
much stress upon it from the ordinary systematic point of view, 
it may at least be said that there is a close connection between 
wide angle and degenerate wings. The angle among pigeons 
varies considerably ; among the eutaxic forms it is never more 
than 50 angular degrees, and generally is considerably less. 


Fig. 6. 


1 sxe 


I. Columba. I. Phlogenas. III. Leucosarcia. IV. Corvus.—Articulation 
at shoulder-girdle. The coracoid is dotted. Cl. Clavicle. P.C. Pro- 
coracoid. Sc. Scapula. 


Procoracoid Process.—Firbringer has investigated the pro- 
coracoid process and the relations of the scapula, coracoid, and 
clavicle with the greatest care, and has drawn a number of im- 
portant conclusions from their conditions. It may be said 
generally that a large procoracoid articulating with the clavicle 
aud a separation of clavicle and scapula are comparatively simple; 


iy 


228 MR. P. CHALMERS MITCHELL ON SO-CALLED 


while in more specialized forms the procoracoid process becomes 
smaller, loses its connection with the clavicle, the latter deve- 
loping an epicleideal plane which meets and may overlap the end 
of the scapula. In fig. 6 J,a comparatively primitive condition, 
as seen in Columba, and with minor details in most diastataxic 
pigeons, is represented. The procoracoid is large and articulates 
prominently with the clavicle, while the latter is not expanded 
at its end and is well separated from the scapula. In fig. 6 IV, 
the more advanced condition, as seen in a typical Passerine, 1s 
represented. The procoracoid is much reduced, and does not 
meet the clavicle. The clavicle is expanded at its end, and 
meets and overlaps the scapula. Among the eutaxic forms, the 
procoracoid is generally small and never meets the clavicle by a 
broad articulation, but is either free from it, as in B (fig. 6, 11), 
a, C, and E; or just meets it above, as in A, D, and F 
(fig. 6, 11), the junction being in a different place and due 
rather to the growth of the epicleideal plane than to size of the 
procoracoid. The clavicle among the eutaxic forms is either 
quite close to the scapula, on account of growth of the epicleideal 
plane, as in B (fig. 6, IJ), or actually meets it, as in the other 
eutaxic forms (fig. 6, III), while in all the diastataxie forms 
I have seen it remains separate. 


Summary of Anatomical Argument. 


When the anatomical differences between eutaxie furms and 
their diastataxic allies are compared, it appears that the eutaxic 
forms are on the average distinctly more specialized. If there 
1s any general progress along special lines among pigeons, the 
eutaxic forms are well advanced. Their anatomical features 
app: ar to show that, if eutaxy be a derivative of diastataxy, it is 
at least a striking coincidence that in other respects eutaxic 
pigeons are more specialized than diastataxic forms. For my 
own part I am personally so much impressed by the evidence for 
the gradual modification of a whole group in the same direction, 
that I cannot avoid regarding the eutaxy of these pigeons as 
being simply a part of their general specialization. 

Size and Kutaxy.—The eutaxic pigeons, like eutaxic birds 
generally, are small compared with their allies. A, a, D, and E 
are very small pigeons, B and C are moderately small, and F is 
the only fair-sized bird. They are certainly smaller than the 
sand-grouse, which would appear to be the nearest diastataxic 


“ QUINTOCUBITALISM ”’ IN THE WING OF BIRDS. 229 


allies of pigeons. Bnt, unless one knew the size of the ancestral 
pigeon, it would be impossible to draw any strong argument 
from size; in a general way, it may be said that a large number 
of very primitive birds are large, while many of the most 
extremely specialized forms are minute, as among the Passeres. 


Theory of the Origin of the Diastataxic Condition. 


As my general argument involves the supposition that eutaxy 
is a derivative of diastataxy, it may be worth while to advance a 
speculation as to the reason why there should have occurred a 
gap in the wing of birds, and why that gap should be in a fixed 
position. 

The first point upon which I wish to insist is that the quills 
are not different in kind but only in degree of development, 
as compared with other feathers. From a uniform covering 
certain individual feathers have become enlarged to serve the 
purpose of flight, instead of the more primitive purpose of pro- 
tection. Precisely in the same way in the case of Elasmobranch 
fishes, from the general protective covering of toothed scales all 
over the body, certain individual scales or rows of scales become 
specially enlarged, and form series of exaggerated teeth, either 
at special points, as in the case of the dorsal spines of the spiny 
dogtish, or in rows along the dorsal surface of the tail, as in 
some skates, or in one or more rows along the edges of the jaws 
where the external skin folds in to form the stomatodeal lining 
of the mouth, when such rows are developed. It may fairly be 
taken for granted that birds had feathers or feather-like scales 
before they had quills, as quills are simply exaggerated feathers, 
and that the quills are simply rows of extremely developed 
feathers. 

When rows of large structures follow body contours, as in the 
ease of true teeth, caudal spines, or quills, the series appear to 
possess a longitudinal coherence and integrity which may be 
fallacious. If pins be placed vertically on diagonal lines crossing 
the surface of one of the irregular stuffed cushions familiar ou 
old fashioned toilet-tables, the pins which happen to lie along 
the contour edge appear to have a special symmetry with regard 
to these contours, and could we imagine these pins to increase 
in length because of their position, it would be difficult to avoid 
supposing that the long contour pins were planted specially in a 


230 MR. P. CHALMERS MITCHELL ON SO-CALLED 


contour row. In the feathering of a wing two series of rows 
are evident, the intersections of the rows forming a diamond 
pattern, most conspicuous on the surface of a plucked wing. 
One series is horizontal, or at least more or less parallel to the 
contour of the ulnar edge. Of these horizontal series, the rows 
of quills and of majorand second and third coverts are most obvious, 
partly because of the large size of the individual feathers, but 
these horizontal rows are much more difficult to trace outside 
above the larger feathered rows. The other series run rather 
diagonally to the ulnar edge and are starting upwards and with a 
forward slope from the quills. They are very beautifully seen 
in some of Mr. Pycraft’s figures (9. plate xxiv.), but are obvious 
enough in most wings. These diagonal rows run round the sur- 
face of the wing posteriorly almost at right angles to its long 
axis, but anteriorly with an increasingly forward inclination, to 
which I shall presently refer. They resemble the general dis- 
position of colour-markings or scales on a cylindrical surface 
which usually occur as hoops running round it; and it appears 
1o me that the apparently longitudinal rows are composed of 
members of the transverse rows at different levels, being in fact 
simply the enlarged individuals of the transverse rows which 
come to lie on the ulnar edge. 

In actual development in the wing of the chick, it is true 
that the two great horizontal rows which are to form the quills 
and the major coverts appear first as longitudinal rows. It is 
only when these become obvious, and when one or two other 
longitudinal rows appear from before backwards, that the 
diagonal rows begin to be marked. Later on these latter 
acquire increasing coherence until the adult stage is reached. 
J am not, however, prepared to attach great importance to 
this early ontogenetic appearance of the longitudinal rows as 
such. In the first place, considering what we know of the 
extraordinary accelerations and retardations that occur during 
larval development, it would appear to be pushing the recapitu- 
Jation theory to a ridiculous point, to attempt to found a theory 
of the ancestral nature of the rows from the order of their 
ontogenetic appearance. Secondly, there is a very obvious 
reason for the early appearance of the guill-rows. These 
aud the coverts are much larger than the other feathers; they 
take longer to grow and must begin first; they are more 


“ QUINTOCUBITALISM ” IN THE WING OF BIRDS. Jail 


important to the future bird, and early provision must be made 
for them. 

I suggest that the quills, although appearing to be a horizontal 
row, are really different members of diagonal transverse series. 

There now remains to suggest an explanation of the origin of 
the gap. In a general way, morphologists have been inclined to 
regard simple series, as in the case of metameric repetitions, as 
more primitive than discrete and incoherent series; but in a 
number of cases, as for instance in the nephridia of earthworms, 
it would appear that the diffuse condition is simpler and more 
primitive than the orderly repetition found in the common 
earthworm. I think we have learned to be on our guard against 
taking for granted that an apparently simpler condition is in 
reality more ancestral, and that we should approach the problem 
of the diastataxic gap without any prejudice in favour of its 
being secondary. Whether it be secondary or primitive, we have 
to account not only for its existence but for its appearance in a 
definite place after five quills. If the primitive condition were 
eutaxic and the gap produced in a whole series of different birds 
by secondary lengthening of the wing, we should have to explain 
why this lengthening always occurred after five quills. If we 
think of the gap as being primitive, there is no difficulty what- 
ever in supposing that convergent closing should have occurred 
independently in auy number of groups, or even of species and 
genera; and [| think it is not difficult to form some idea as to 
how a gap in that position might have come into existence in 
an ancestral wing. Consider an ancestral pentadactyle wing 
provided with scales or scale-like feathers. The most common 
arrangement of these, as may be seen by looking at the scales 
on any lizard or crocodile or on the feet of birds, is that longi- 
tudinal rows should run along the digits and diagonal transverse 
rows should surround the arm. Now these two series have 
to meet somewhere, and when different series of scales or 
markings meet, there must be a transition of some kind from 
one to the other. Sometimes one set of series gives way to the 
other; sometimes there is intercalated between the two a wedge- 
shaped set of rows, as Professor D’Arcy Thompson pointed out 
to me in the case of the markings on the zebra. In the foot of 
a bird (fig. 7, A) the transverse rows are represented by large 
single scutes in many cases, while a line of enlarged scutes may 


232 MR. P. CHALHERS MITCHELL ON SO-CALLED 


run down each digit. Where the digit-scutes come in contact 
with the other series, they may run down parallel with them and 
appear on the distal edge of the tarsus. In the arm of a lizard 
(fig. 7, B) the interference is more strikingly apparent, there 


Fig. 7. 


2 3 4 Cc 


A. Foot of a Passerine bird. B. Hand ofa Lizard. OC. Diagram of scales or 
feathers on a hypothetical ancestral bird’s wing. The quills of modern 
birds are represented by darker spots. 


being along the distal edge five scales which are in series with 
the digital rows, and there is an abrupt transition from these 
transverse rows to the modified digital rows. In fig. 7, C 
represents diagrammatically a simple mode of feather-distribution 
on a pertadactyle wing which would result in a gap. Rows run 
along the digits, and where these meet the transverse rows a 
wedge-shaped piece is intercalated, forming the transition. I 


“ Q@UINTOCUBITALISM ’’ IN THE WING OF BIRDS. 233 


have put larger dots to represent the actual quill-series of modern 
birds. The primaries are the enlarged series of the index digit ; 
the carpal remex belongs to the pollex series; the next four 
secondaries belong to the digital series of the four digits; theu 
comes a diastataxic gap due to the alteration of curvature (as in 
the transition from the neck to the body series in a Burchell’s 
zebra) being bridged over by a wedge; then follow the other 
secondaries, which are members at different levels of the suc- 
ceeding diagonal rows. Naturally this supposition is entirely 
theoretical, but it is enough to show that the occurrence of a 
gap after five quills is a phenomenon which might have arisen in 
a very simple fashion. 

So far as the underlying bones are concerned, the gap in my 
diagram occurs much more distally than in the adult ordinary 
bird. I do not think, however, that this presents the slightest 
difficulty. We know that ontogenetically and phylogenetically 
the wing of birds elongates. Actually in development the feather- 
papille shift along the wing. Superficial skiu-areas generally are 
exceedingly primitive in their character, but may shift almost in- 
definitely in their topographical relation to underlying structures. 
In the human body, these changes of skin-area in topographical 
position have been worked out in relation to the phenomena of 
referred pain. 


Eutaxy and Diastataxy in Aves generally. 


I am not at present prepared to extend the argument from the 
Columbe to birds in general at length. But I may point out 
that there isa general parallel to be found between the relations of 
eutaxy among Columbine forms and the relations of eutaxy 
among birds generally. In Columbine forms the majority are 
diastataxic, and the few eutaxic forms are isolated among their 
diastataxic congeners. Among the great groups of birds most 
are diastataxic. We may leave out of consideration the Struthious 
birds and the Spheniscide, as in these the wing is so greatly 
modified as to make any comparisons misleading, especially as we 
cannot be certain whether the modifications are from a high, or 
from a very low type of wing. As for the other eutaxic forms, 
the great majority of anatomists would agree that the Passeres 
were extremely specialized birds, and therefore forms in which, 
if diastataxy be primitive, one would not expect to find it. 
Similarly the Pici, the Cuculide, and Opisthocomus are birds 


234 MR. P. CHALMERS MITCHELL ON SO-CALLED 


in which one would not readily look for primitive features. 
Opisthocomus is an exceedingly peculiar and specialized form. 
Among the Ralline birds generally, the three eutaxic forms 
Psophia, Dicholophus, and hinochetus present many striking 
peculiarities which are not generally regarded as primitive, and 
anatomists generally have regarded them as modified forms of the 
diastataxic Gruide. There remains the great group of Gallina- 
ceous birds, which in many respects, such as the alimentary canal 
and the completeness of the muscles, present primitive features 
and are yet eutaxic. On the other hand, they are specialized in the 
sternum and in the vertebral column. The Anseriformes, which 
are diastataxic, are quite as primitive in their muscular structure 
and less specialized in the sternum and vertebral column, while 
among them are to be found the Palamedeide, certainly archaic 
forms, and sometimes regarded as forming a link between the 
typical Anseriformes and the Galli. These are diastataxic, as 
primitive as the Galli in muscular anatomy, more primitive 
osteologically, and with alimentary canals displaying what 
appears to be an exceedingly primitive disposition. So far as a 
general survey goes, there is nothing against and a good deal 
in favour of supposing that diastataxy is architaxic in Aves 
generally. 

The Alcedinide and the Cypselide are groups in which, as in the 
Columb, some members are eutaxic, while the others are diasta- 
taxic. I have not had an opportunity of dissecting a sufficient 
number of these forms to obtain an idea as to which condition 
of the wing is associated with greater specialization. 


Summary. 

The Columbe, which have been regarded hitherto as a diastataxic 
group, have several members with the eutaxic condition. Com- 
parison of the feathering in these forms makes it probable, or at 
least plausible, that the eutaxic condition has been attained by 
closing up of the gap, with first crowding, and then disappear- 
ance, of two of the three feathers occupying the primitive gap. 
Intermediate stages between true wide-gapped diastataxic forms 
and true eutaxic forms occur. Comparison of the anatomy of 
the eutaxic forms with that of the diastataxic forms shows that 
the former are on the whole more advanced in the general pro- 
gressive modification of the whole group. It is easy to show 
that in a hypothetical pentadactyle wing a gap in specialized rows 


Mitchell, 


P.cM. del. 


Lrxw Soc. Journ. Zoon. Vou. XXVIL.Pr Ja. 


Geo, West & Sons imp. 


Parker & Peneyrlith. 


QUIN DOC UB MPA SMa TING Richy SWAN CS TO ss eA) Se 


Y 


—— 


&.C.M. del. 


Luyw Soc. Journ. Zoot, Vou. XXVIT.Pr.13. 


Geo, West & Sons imp. 


Parker & Perey lith. 


QUINTOCUBITALISM IN THER WING OF BIRDS. 


‘¢ QUINTOCUBITALISM ’’ IN THE WING OF BIRDS. 235 


of incipient quills might arise in the position required for the 
diastataxic gap. Among Aves there is a general correspondence 
with the conditions among Columbe. The diastataxic condition 
of the wing is primitive among birds; it is the architaxic 
condition. By closing up of the ranks any architaxic wing may 
become eutaxic, and this change has been made by some whole 
groups and by individuals of other groups. 


Bibliography. 


(1) Wray.—“‘ On some Points in the Morphology of the Wing 
of Birds.” P.Z. S. 1887, p. 343. 

(2) Brepparp.—Structure and Classification of Birds. London, 
1898. 

(3) Garrop.—“ On some Points in the Anatomy of the Co- 
lumbe.” P.Z.S. 1874, p. 249. 

(4) Férprinerr.—Untersuchungen zur Morphologie und 
Systematik der Vovel (1888). 

(5) CHatmers Mrronurnn.—“ On the Perforated Flexor Muscles 
in some Birds.” P.Z.S. 1894, p. 495. 

(6) Cuatmers Mrtrcuetn.—“ A Contribution to the Anatomy of 
the Hoatzin.” P.Z.S. 1896, p. 618. 

(7) Coatmers Mrtcuetit.— On the Intestinal Tract of Birds.” 
P. Z. 8. 1896, p. 136. , 

(8) Bepparp & Parsons.—“ On certain Points in the Anatomy 
of the Parrots.” P.Z.S. 1893, p. 507. 

(9) Prcrart.—“ A Contribution towards our Knowledge of the 
Morphology of the Owls.’ Trans. Linn. Soc., Zool. vol. vii. 
(1898) p. 223, pls. xxiv.—xxix. 


EXPLANATION OF THE PLATES. 


Puats 12. 


Fig. 1. Geopelia cuneata, left wing: showing primaries and coverts, carpal 
remex and covert, and even series of secondaries and coverts; no gap, 
EHutaxic condition. 

ig. 2. Leucosarcia picata, left wing: showing primaries and coverts, carpal 
remex and covert, and even secondary series without gap. LHutaxic 
condition. 

Fig. 3. Geophaps plumifera, left wing: showing the primaries and coverts, the 

carpal remex and covert, the secondaries and coyerts without a gap, 
Hutaxic condition. 


eA 
Q 


236 MR. W. P. PYCRAFT ON 80-CALLED 


Prater 13. 


Fig. 4. Columbula picur, left wing ; carpal covert and remex; series of coverts 
and secondaries: showing the eutaxic condition. 

Fig. 5. Geotrygon montana, left wing ; 1st primary; carpal remex and covert ; 
series of secondaries and coverts: showing reduced diastataxic condition. 

Fig. 6. Starnenas cyanocephala, left wing; carpal remex and covert; six 
secondaries and coverts in even series. Hutaxic arrangement. 

Fig. 7. Turtur chinensis, left wing; carpal covert and remex; four secondaries 
and coverts; diastataxic gap with covert ; two secondaries with 
coverts. 


Some Facts concerning the so-called “ Aquintocubitalism ” in 
the Bird’s Wing. By W. P. Pycrarr, A.L.S.* 


[Read 16th March, 1899.] 
(Prates 14-16.) 


ConTENTS. 

Page 
Introductory Remarks ................-.2+2s-ce0ee aiaisiccicaes ese Neseeee 236 
Hutaxie and Diastataxic)\Wanes\essecenseesesnteeensecce eee seereee 237 
Dhe'Hmbry MWA gaan seecaeee aeeceacee nels sestctlsiseettoe eet seisteltects 238 
Summary of the foregoing Remarks ..................ceceeceeeees 241 
Evidence in support of the above Conclusions .................. 241 
AGpossible Objectioninecscmanceseewerecesesn see ace eee eeeeeeeee 244 
MherCarpaly@ oventiand PRemexg pene.) see kerceeeepeececee te teat 245 
ExplanabionstoteDiastabaxygeceneeresceesccscerecccressetechesse 246 
1DECGe) BLO} AIS) pe goo apeeceatios odsucboaReBeobbS oss cose so nd uoT}OIeDSseDeOI a0 ae 247 
Facts correlated with Diastataxy .............-.secsess-sersseeoe- 249 
Some) Degenerate Wangs tcc assneraccssccn ce ereaseeereeseeecreeties 250 
The Probable Origin of the Diastataxic Wing ...............0+ 252 
Diastataxy as a Factor in Classification.............cccceeseeeeees 254 


Introductory Remarks. 


Tur feathers in the typical bird’s wing, e.g., the Common 
Fowl, are divisible into two groups—tectrices or coverts, and 
remiges or flight-feathers, commonly known as “ quills.” 

The remiges form a single row of feathers running along the 
post-axial border of the wing from the tip of the index-digit 
inwards to the elbow-joint. Those of the hand constitute the 
primaries, those of the forearm the secondaries. With the 
primaries we have little or nothing to do in this connection ; 
suffice it to say that they never, in the Carinate, exceed 12 in 


* Cf. Editorial footnote on p. 21 


“ AQUINTOCUBITALISM”’ IN THE BIRD’S WING. 237 


number. The number of the secondaries varies greatly from 
9 to 387. Though the primaries are packed closely together at 
their bases, the secondaries are more or less widely spaced 
(Pl. 14. fig. 5). The exigencies of flight demand this. 

The tectrices are separable into several distinct series, furming 
the major, median, minor, and marginal coverts, to which may 
be added the ala spuria, the hypopteron, and parapteron. With 
these last we have nothing to do now. The tectrices clothe the 
dorsal and ventral surfaces of the wing. ‘The major coverts are 
the most post-axial, and are seated in pairs on the bases of the 
remiges—two to each remex, one dorsal and one veutral—to 
which they are firmly attached. The median form the row next 
in front of the major coverts, both on the dorsal and ventral 
surface. There is never more than one row on the dorsal or 
ventral surface. The minor lie beyond the median coverts, and 
vary from 1 to 4-5 rows on both aspects of the wing. Beyond 
these are the marginal coverts; they occupy the pre-axial border of 
the wing, and help to clothe both the dorsal and ventral surfaces. 

A reference to Pl. 14. fig. 5 will make this much more clear 
than mere description. Questions concerning the overlap, 
variations in the number of rows of minor and marginal coverts, 
their length, the absence of more or fewer of these rows on the 
arm or manus in different groups, need not be discussed here. 
One point, however, is noteworthy. Sundevall (7), and later, 
and more correctly, Wray (8) pointed out that the major 
and median coverts of the ventral aspect of the wing turn their 
ventral surfaces downwards as do the remiges, and not upwards 
as do all the other coverts of the under surface. Wray’s inter- 
pretation of this was, that these feathers had been slowly carried 
round from the dorsal surface of the wing, whilst the tectrices 
of the under surface were separately derived from the ventral 
surface uf the body. 


Eutaxic and Diastataxic Wings. 

The Bird’s wing may assume one of two forms, known hitherto 
as (1) the Quintocubital, and (2) the Aquintocubital (p. 238). 
The two may readily be distinguished. In the former, each 
paic of secondary major coverts embraces a remex between 
them; in the latter, the remex from between the 5th pair of 
coverts is apparently missing—hence the name “ aquintocubital.” 
The perfectly regular arrangement of the coverts in such a wing, 
and the presence of a more or less distinct gap between the 4th 


238 MR. W. P. PYCRAFT ON SO-CALLED 


remex and that next succeeding it, seemed to justify the con- 
clusion that aremex had been lost—that the wing had undergone 
a reduction of the original number of its remiges by the loss of 
the 5th quill. The aim of the present paper is to show that no 
such loss has taken place. The 5th remex has lost its original 
relations, but not its existence. 


Before proceeding further, it will be well to say a few words 
concerning the terms which have been proposed as substitutes 
for the older and less exact ‘‘ Quinto-”’ and “ Aquinto-cubital.” 
At the time this paper was read the names Stichoptylic for the 
former, and Apoptylic for the latter were used. These were 
suggested to me by Prof. H. Ray Lankester, and were certaiuly 
preferable to those which we both desired to supplant. But it 
will be remembered that Mr. P. Chalmers Mitchell, in the course 
of his paper dealing with this same question, suggested the 
names Eutaxic for the quintocubital or stichoptilic form, and 
Diastataxic for the aquintocubital or apoptilic. His names not 
only have priority over mine, or, rather, Prof. Lankester’s (his 
paper having been read before mine), but they are, I think, 
actually preferable ; hence, throughout this paper, I shall adopt 
the terminology proposed by him. Moreover, by doing this I 
shall be rendering a service to my readers, by saving them the 
labour of keeping in mind the values of some half-dozen names 
for what may be called the positive and negative of one and the 


same thing. 


Lhe Embryo Wing. 


In all wings, the feather rudiments appear first along the 
post-axial border of the wing; those representing the remiges 
and their major coverts appearing simultaneously, and some- 
times together with very faint traces of one or more of the 
pre-axial rows representing the median and minor coverts. At 
this stage, it is often not possible to say whether a wing will 
ultimately prove eu- or diasta-taxic. The change takes place, 
however, generally at the close of this phase of development. 
As the rudiments of the median and minor coverts become more 
distinct, it will be noticed that the papille representing the 
remiges 1-4 no longer form an unbroken series with those 
running from this point inwards, but that they have moved 
backwards and downwards; sometimes this is very marked, 
sometimes only very slightly so. The hitherto unbroken series 


“ AQUINTOCUBITALISM ’’ IN THE BIRD’S WING. 239 


now becomes more or less distinctly divided into two portions, 
thus:—’ ***°*°.,., ,; at the same time, it is noticeable that 
this shifting of the outer remiges backwards may also be 
accompanied by an ontward movement towards the tip of the 
wing. Sometimes all four remiges participate in this outer 
movement, sometimes only the Ist or Ist and 2nd become 
notably disturbed. 

The movement of the remiges is in all cases accompanied by a 
corresponding movement on the part of the coverts associated 
therewith, from the post- to the pre-axial border. The result, 
when the wing is viewed as a whole, seems to show that a process 
of “faulting” has taken place, the major, median, and one or 
more rows of minor coverts from 1 to 5 in each row having 
slipped backwards so as to break the connection with their 
several rows proximad of this point ; each row—or, more correctly, 
the first 3 or 4 rows—now runs, not in a continuous line with 
that of its series, but between this and that immediately behind 
it (fig. 1,0). The disturbed rows, however, seem to readjust 


Fig. 1. 


ie Cc 
ae e 

f. ry lee. of 
! maj.c 


as tf doy 


PEIREGES 


rer ae 


Shifting of wing-coverts and remiges. a, before; 0, after. 


themselves very quickly so as once more to form continuous 
lines with the more proximal feathers; as, in the typical 
diastataxie wing, it will be found that, not counting the 
remiges, uniformity is regained at the 4th row, or rather a 
semblance of uniformity, inasmuch as this row is really made up 
of two rows. This is shown in the diagram (fig. 1,0). Here the 
3rd row of these downwardly shifted coverts appears as an inter- 
calary row, the 4th row becoming continuous with that of the 


240 MR. W. P. PYCRAFT ON SO-CALLED 


5th, thus affording at the same iime evidence in favour of the 
view adopted in this paper, that all these coverts have shifted 
backwards. Pl. 15. fig. 1 represents the wing of an embryo 
Pigeon showing this intercalary 3rd row very clearly. 

Only one row of ventral coverts appears to participate in the 
general disturbance which we have traced on the dorsal surface 
in connection with this backward and downward motion of coverts 
1-5 and remiges 1-4. This is well seen in fig. 3, Pl. 14, repre- 
senting a ventral view of the wing of an embryo Machetes pugnax. 

Before passing on to consider what is, apparently, the only 
possible objection to the explanation of the phenomena here set 
forth, I would draw attention to the accompanying diagrams. 

Fig. la (p. 239) represents the arrangement of the cubital 
coverts and remiges in the eutaxic wing as far inwards as the 
9th remex. Fig. 16 shows the effect of a backward and outward 
shifting of remiges 1-4 and coverts 1-5 from the major coverts 
forwards ; 7. e., a portion of each horizontal row of coverts from 
1-5 from the major coverts forwards to the tectrices minores, 
thus converting the eu- into the diasta-taxic wing. ‘The bending 
of the horizontal lines serves to indicate the amount of shifting 
which the remiges and coverts have undergone. The arrow “d” 
has been made to “ dip ’’ like the arrows a, 0, ¢, so as to indicate 
the amount of shifting of each row; it has also, by means of 
a dotted line, been made to pass straight outwards to indicate 
the restoration of parallel series. But it must be remembered 
that these restored rows are composed of feathers belonging to 
two different rows. ‘Thus coverts 1-5 of the 2nd row of minor 
coverts now become serial with the coverts of the Ist row from 
the 6th inwards, the coverts 1-5 of the Ist row having been 
cut off to form a series by themselves—the interealary row. 
This intercalary row actually obtains in a more or less well- 
developed form in all diastataxic wings, and this diagram enables 
us to see how it may have come into existence. In so far 
as the diagram is concerned, it is perfectly true that the 1-5 
coverts of the median row could equally well be regarded as 
an intercalary row. The size and position of these feathers in 
the adult probably account for their retention in the series to 
which they belong. The row (1-5) immediately in front are 
smaller and more easily isolated; hence these in the adult 
become the intercalary row. 

The diagram just described (fig. 1, a, 6), to show how the eu- 
may have been transformed into the diasta-taxic wing, can be 


“ AQUINTOCUBITALISM ”’ IN THE BIRD'S WING. 241 


readily constructed by the reader by laying over fig. 1 a a piece 
of transparent paper and marking over remiges 1—4 and coverts 
from 1-5 in each row. This being done, shift the paper back- 
wards aud slightly forwards so that the major coverts 1-5 come 
to lie parallel with the interspace between the remiges and major 
coverts proximal to 5, as has been done in fig. 16. This gives 
the two broken rows of coverts (major and median) and the 
intercalary row (3), which exactly agrees with actual wings 
(e. g., Pl. 15. fig. 1). 

Inasmuch as by this artificial mechanical shifting and re- 
arrangement of the feathers of the anterior end of the dorsal 
aspect of the forearm, ali the features of the diastataxic wing can 
be demonstrated, it may be reasonably contended that a strong 
degree of probability has been brought forward in support 
of the view that the phenomena of diastataxy are due to a 
backward and downward shifting of the remiges and their 
coverts. 


Summary of the foregoing Remarks. 


To summarize briefly, the contention of the present paper is :— 

(1) That there is no evidence in support of the hypothesis 
that diastataxy is due to the absence of a remex. 

(2) There is a very considerable amount of evidence to show 
that a process of shifting has taken place of the coverts and 
remiges at the distal end of the forearm. This has resulted in 
carrying remiges 1-4 and the first and each succeeding horizontal 
row of coverts from 1-5 backwards and slightly downwards and 
outwards. Thus the original relations between the 5th major 
covert and its remex have been disturbed, the covert having 
shifted away from its remex, which has now become associated 
with the 6th covert. Thus the 5th, together with its ventral 
covert, appears to have lost its remex. 

(8) The cause of this shifting is still a matter for investiga- 
tion; it is possibly due to a slight secondary lengthening of the 
forearm. 


Evidence in support of the above Conclusions. 


We will now proceed to review the evidence in support of the 
hypothesis just submitted. 

What follows has reference only to the developing remiges,. 
LINN. JOURN.—ZOOLOGY, VOL. XXVII. 18 


249 MR. W. P. PYCRAFT ON SO-CALLED 


and their coverts, of the forearm; those of the hand need not be 
taken into consideration in this connection. 

In the eutaxic wing of the Common Fowl! the earliest traces of 
feathers are those representing the remiges and their dorsal 
major coverts (Pl. 14. fig. 4). These form a double row 
along the post-axial border of the wing, the major coverts lying 
opposite the interspaces of the remiges, the ventral row of major 
coverts and the dorsal median and minor coverts appearing 
somewhat later, and the marginal last of all (Pl. 14. fig. 4). 
The arrangement of the median and minor follows that of the 
major coverts and remiges, so as to form a series of alternating 
rows, the feather rudiments of one row lying opposite the inter- 
spaces of the row in front of, and behind it. By the time the 
full complement of rows has been attained, however, this 
primitive arrangement in horizontal rows is somewhat masked, 
and the feather-papille appear rather to run in oblique rows 
sloping either from without inwards or vice versa, according to 
the view of the observer; but, as will be seen later on, when 
examining wings of other forms the obliquely-inward slope 
becomes finally adopted. 

The wing of the Lapwing (Vanellus cristatus) (Pl. 14. fig. 1) 
agrees with that of the first stage (described above) of the 
Common Fowl in having only two rows of feather rudiments 
along the forearm, and, like this, it is also eutaxic. That is to 
say, no shifting has as yet taken place. The fifth major covert is | 
not yet divorced from its remex. In a much later stage (PI. 14. 
fig. 2) this severance has taken place: the wing is now diasta- 
taxic. Unfortunately, I have no intermediate stages between 
this and fig. 1. It is of interest to note, that the downward 
shifting in this case must have been but slight, as the inter- 
ealary is the 2nd and not the 3rd row of coverts, as was the 
case in the typical wing described on p. 239. Moreover, the 
feather rudiments seem to have travelled forward, inasmuch as 
the 1st cubital remex now lies on a level with a line drawn 
through the proximal end of the manus in front of the carpus ; 
whilst in the younger stage the 1st remex lies proximad of this 
imaginary section. 

The wing of the embryo Guillemot (Lomvia troile) (Pl. 15. 
fic. 2) is one of the most valuable of the whole series now in 
my possession ; earlier stages than this are much to be .desired.. 
Here the remiges‘and major, median, and two (with faint traces of 


“ AQUINTOCUBITALISM ’ IN THE BIRD'S WING. 243 


more) rows of minor coverts are all represented, but, as yet, the 
wing is undoubtedly eutaxic, though soon after this stage it 
becomes diastataxic. A foreshadowing of this is plainly visible 
at the stage under discussion. A reference to Plate 15. fig. 2 
shows that the remiges 1-4 have already begun to move 
backwards through a small are of a circle, the movement having 
been greatest at the distal end of the row, whilst the proximal 
end, represented by the 4th remex, has as yet scarcely moved at 
all. It may be expressed thus ===, the biack line repre- 
senting the original and the dotted line the new position. The 
disturbance is much more obvious in the row of papillae imme- 
diately above the remiges—the rudimentary major coverts 1-5. 
These have become distinctly separated from the rest of the row 
proximad of this poiat. The disturbance of the rows preaxial to 
this of the major coverts is barely perceptible. There is yet no 
intercalary row. I have no stage between this and that of the 
nestling (woodcut, fig. 2). 


\ 

\\ 

+ 
Nie ae 
A=. 
Y, 


Right wing, dorsal aspect, of nestling Lomvia troile, to show the diastataxic 

arrangement of the coverts. 

The wings of the Pigeon (Columba domestica), Duck (Anas 
boscas), and Owl (Syrnium aluco) are selected as examples 
of typical embryonic diastataxic wings. In earlier stages of 

18* 


DAA, MR. W. P. PYCRAFT ON SO-CALLED 


these we must seek for confirmation of the course of development 
outlined in the case of the Plover and Guillemot. 

In the Pigeon’s wing (Pl. 15. fig. 1) the feather rudiments 
have arranged themselves in strongly marked transversely-oblique 
rows sloping inwards and presenting a strongly curved front 
towards the distal end of the wing. As yet, there are but faint 
traces of the marginal coverts. It is noteworthy that the trans- 
verse rows of coverts from 1-5 are more widely separated one 
from the other than is the case with the more proximal rows. 
The downward shifting of the anterior remiges and their coverts 
(1-5) is very marked. The intercalary row is the 3rd. This last 
is well seen in the wing of the nestling (Pl. 16. fig. 1). 

In the wing of the Duck (Anas boscas var. domestica), Pl. 15. 
fig. 8, the interealary is formed by the 3rd row of coverts, 
2. €. 1-5 of the minor coverts. There are faint traces of several 
more rows in addition to those in the figure. Compared with the 
adult wing, one very striking fact becomes apparent. It wilk 
be noticed that in the embryo wing (Pl. 15. fig. 3) the Ist 
median covert—that lying immediately above the first cubital 
remex and its major covert—lies over the base of metacarpal 
TI., whilst the 2nd median covert lies just below the angle 
between the carpus and the distal end of the ulna. In the 
adult this spot comes to be occupied by the Ist median covert,. 
that is to say, it apparently supplants the 2nd and takes 
its place. From this it would seem that we have indeed 
evidence of an increased lengthening of the forearm which can 
be measured by the distance from the 2nd to the Ist median 
covert. 


A possible Objection. 


Diastataxy, as we have endeavoured to show, is due, not as. 
was supposed, to the loss of the 5th cubital remex, but to the 
shifting of the remizes and coverts lying to the outer side of this. 
The ultimate fate, however, of the remex in question has so far 
only been hinted at. Hxactly what takes place during this 
shifting is difficult to make out, and will only be possible after a 
larger series of embryos have been examined. This much, how- 
ever, Seems certain,—that all the covert-feathers of the wing 
from the 6th inwards have moved outwards, one place, in the 
form of a series of obliquely transverse rows. Thus the trans— 
verse row which originally belonged to the 6th remex now 


‘* AQUINTOCUBITALISM ”’ IN THE BIRD’S WING. 245 


becomes associated with the 5th, that of the 7th remex with the 
6th, and so on. A reference to the diagrams will make this 
clear. 

At first sight, this outward movement seems to make rather a 
large demand upon the imagination, and to this extent to throw 
doubt on the interpretation of the facts recorded in this paper. 
The difficulty, however, is more imaginary than real. It simply 
means that the remiges in question become associated with 
transverse rows immediately in front instead of with those next 
behind. This must certainly happen in the case of the Guille- 
mot’s wing. Inthe Duck and Pigeon’s wings (PI. 15. figs. 1 & 3) 
this forward movement has already taken place. The ventral 
major coverts from the 6th inwards are subjected to the same 
forward movement as those of the dorsal surface. Thus the 5th 
remex lies between the 6th pair of major coverts, 6th remex 
between the 7th pair, and so on. The position of the ventral 
coverts 1-5 in these figures will illustrate the downward—ventral- 
ward—shifting of the feathers in this region. 


The Carpal Covert and Remex. 


A fixed point of no small value in the present connection is 
that afforded by the two feathers known as the carpal covert and 
the carpal remex (Pl. 14. fig. 2, ¢.¢., er); inasmuch as, since 
they occur in wings of both types, they serve as valuable land- 
marks, and show, moreover, that the disturbance is to be sought 
for proximad of this point, and thus help to confirm the conten- 
tions of this paper. The significance of these feathers has been 
discussed by Wray (8), the present writer (5), and by Degen (1), 
who bestowed on them the names Carpal covert and remexy, 
from their position on the carpus. Although feeling by no 
means certain on this point, I think the probability is that the 
““remex ” is really correctly so named, and that it represents a 
feather more or less completely dwarfed and in course of dis- 
appearance. Its office—as a remex—has not entirely ceased. It 
is probably being slowly crushed out of existence by reason of its 
position, which is in the angle of the wing caused by the folding 
of the hand on the forearm. The 1st cubital remex of the 
Galline is, like its carpal remex, and for the same reason, under- 
going a similar process of reduction. 


PAG MR. W. P. PYCRAFT ON SO-CALLED. 


Haplanations of Diastataaxy. 


The first recorded reference to diastataxy is that of Gerbe (8),. 
who describes it in the following words :—-“ Chez les Rapaces, les. 
Pigeons, les Echassiers, les Palmipédes, il y a atrophie compléte 
de Pune des remiges secondaires, et cette atrophie, qui parait 
étre originelle, porte invariablement sur la cinquieme. Ses 
satellites, c’est-a-dire, sa couverture supérieure et sa couverture 
inférieure, prennent un développement normal et occupent leur 
place respective, comme si elles accompagnaient la penne qui fait 
défaut. 

‘““Ni les vrais Passereux, ni les Zygodactyles (les Perroquets 
exceptés) ne présentent cette singuliére anomalie.” 

My friend and late colleague, Mr. E. 8. Goodrich, the 
Aldrichian Demonstrator of Comparative Anatomy at Oxford, 
took part with me for some time, in this investigation, and 
also formulated a theory of his own to account for the con- 
ditions which have been described and figured in this paper, 
and in justice to him, as well as because of its intrinsic value, I 
propose to endeavour to describe his theory here. 

Briefly, he holds that the phenomena of diastataxy are due to a 
bifurcation of a row of feather-papille, probably the second 
—major coverts—starting at what is now the 5th major 
covert. Thus a double row was formed representing the present 
major and median coverts 1-5. This theory does not demand 
either shifting of remiges or coverts. Supposing the shifting of 
the former be proved, the presence of this “intercalary row,” as: 
he termed what are now major coverts 1-5, is still more easily 
understood. They have appeared to fill up the space between the 
row immediately in front and the remiges behind. Bifurcation 
of this kind occurs in the seale-covered forearm of Reptiles for 
instance: or, again, in the form of additional rows of ossicles in 
the manus of Ichthyosaurs. 

Yet another attempt to solve this mystery is that of Degen 
(1). Though none will grudge this writer the credit of having 
evolved a very ingenious hypothesis, few probably will be found 
willing to adopt it. Degen carries us back to an imaginary 
quadri-dactyle manus in which each digit supported a set of 
remiges and major coverts. In course of time the 4th digit 
became suppressed and its remiges, 3 in number, migrated 


“ AQUINTOCUBITALISM ” IN THE BIRD'S WING. 247 


inwards on to the ulna—ousting the cubital remiges 1-3. Next, 
the remiges of Digit II. moved inward on the ulna. Originally 
there were five of these, but the 5th, lying in the carpal angle 
between the bases of Metacarpals III. and IV., became sup- 
pressed,—just as occasionally happens in the case of the “ carpal 
remex.” The coverts of this suppressed 5th remex were retained. 
Feathers 1-4 only remained to migrate on to the ulna. The 5th 
is now only indicated by its coverts,—hence the diastataxic wing. 
The carpal covert and remex of existing birds represents the short 
1st remex and covert of Digit III., which has travelled inwards 
along Me. III. to rest finally on the carpal joint at the base of 
Me. II. 


Haceptions. 


The wings of all birds are either eu- or diasta-taxic. More- 
over, there is no known exception to the rule that, though a 
genus may include both forms of wings, it will be found that 
the species constituting that genus will group themselves, in- 
variably, into two sections—those with eu- and those with 
diasta-taxie wings; for, as yet, individual variation in this 
particular is unknown. Therefore, the wing of any given species 
being found to be diastataxic, it may be certainly predicted that 
every individual of that species will also be diastataxic, and 
vice versa. 

Amongst the Carinate there are certain large groups every 
individual member of which, so far as is known, has diastataxic 
wings. These are :— 


Pygopodes = Divers, Grebes. 

Tubinares = Petrels, Albatrosses. 

Herodiones = Herons, Storks. 

Steganopodes = Cormorants, Gannets, Frigate-birds. 

Pheenicopteres = Flamingoces. 

Anseres = Swans, Ducks, Geese, Screamers. 

Accipitres = Hagles, Hawks, Vultures, and Secretary - 
bird. 

Rall = Rails. 

Limicole = Curlews, Plovers, Sandpipers, Auks, Gulls. 

Pterocletes = Sand-Grouse. 


Megapodes = Mound-builders. 


248 MR. W. P. PYCRAFT ON SO-CALLED 


To these may be added :— 


Psittaci = Parrots. 
Striges = Owls. 
Caprimulgi = Nightjars and Oil-bird. 


Similarly the following are eutaxic :— 


Tivami = Tinamus. 

Galli = Crax, Phasianus, Gallus. 
Turnices = Hemipodes. 

Opisthocomus = Hoatzin. 

Coccy ges = Cuckoos, Plantain-eaters. 
Coracize = Rollers, Bee-eaters, Motmots. 
Bucerotes = Hornbills and Hoopoes. 
Trogones = Trogons. 

Colu = Colies, or Mouse-birds. 

Pici = Woodpeckers. 

Passeres = Hurylemus, Pitta, Tyrannus, Menura, 


Atrichia, Corvus. 


The remainder of the Carinatze contain more or fewer ex- 
ceptions—both eu- and diasta-taxic forms :— 


Grues = Cranes, Trumpeter, Seriema. 
Columb = Pigeons. 


The Grues may be considered as a diastataxic group, the 
exceptions being :— 


Dicholophus = Seriema. 
Psophia = Trumpeter. 
Rhinochetus = Kagu. 
Eurypyga = Sun-bittern. 
Heliornis = Fin-foot. 


These are all very aberrant types, whose systematic position is 
still a matter for investigation. 

The Columbe, like the Grues, are a diastataxic group, ie only 
recorded exception being the genus Coluwmbula. 

The Coraciiform Alcedines (Kingfishers) and Macrochires 
(Swifts and Humming-birds) each contain genera the species 
comprising which include forms with both eu- and diasta-taxiec 
wings. Thus amongst the Alcedines, in the genus Ceryle, 


“ AQUINTOCUBITALISM” IN THE BIRD’S WING. 249 


C. torquata, C. rudis, C. alcyon, and C. maxima are diastataxic ; 
the remaining species of the genus are eutaxic. In the genus 
Halcyon, H. vagans, H. chloris, and H. sancta are diastataxic. 
D. gigas appears to be the only diastataxic member of the genus 
Dacelo. 

The Macrochires comprise the Swifts (Cypseli) and the 
Humming-birds (Trochili). 

The Trochili are all eutaxic. 

The Cypseli are mostly eutaxic, but contain at least two 
genera possessing both forms :— 


Dendrochelidon mystacea is diastataxic. 
Acanthyllis collaris 30 


No satisfactory explanation of these exceptions has yet been 
offered, though some sort of an attempt was made by the late 
Henry Seebohm (6). He suggested that some diastataxic species 
may have become eutaxic by elimination of the coverts belonging 
to the missing fifth remex, thus removing all traces of their 
former condition. From the facts already educed in the pre- 
ceding pages ot the present paper, this particular interpretation 
must now be regarded as probably discounted. Before anything 
like a final explanation can be hoped for, we must wait till more 
material is at hand. A large series of embryo and adult species 
of those genera containing both forms of wings will probably 
settle the question. For the present, perhaps, the few suggestions 
advanced on p. 252 may be acceptable. 


Facts correlated with Diastataay. 


According to Seebohm (6) :— 
1. No eutaxic bird has a webbed foot. 
2. Birds which have abnormal plantar tendons contain both 
eu- and diasta-taxic species *. 
3. There are very few diastataxic birds without an ambiens ; 
but there are no eutaxic families that contain birds both 
with and without it t. 


* Concerning 2 it may be remarked that this is equally true of birds having 
normal plantar tendons—Seebohm’s term for plantar tendons in which the 
flexor perforans supplies each of the front toes. 

+ This depends upon the individual taxonomer—as to whether he eliminate 
the discordant elements. 


250 MR. W. P. PYCRAFT ON SO-CALLED 


4, Pelargomorphe have normal plantars and are diastataxic. 

5. Aeithomorphe have normal plantars but are eutaxic. 

6. Coraciomorphe contain both diasta- and eutaxic forms. 
Of four diastataxic families two contain both eu- and 
diasta-taxic species. 

According to Degen :— 

“ Aquintocubitalism and Quintocubitalism seem to reflect 

on the presence or the absence of the 11th metacarpo- 


digital flight-feather.” (All 11-primaried birds are 
diastataxic according to this author.) 


According to Goodchild :— 
The diastataxic wing is characterized by a peculiar inter- 
ruption or faulting of the coverts of the dorsal 
surface. 


Degen’s contention is disproved by the fact that, as shown by 
Gadow (2), the followmg, though diastataxic, have only 10 
primaries :—Scopus, Eurypyga, Rallus, Ocydromus, Himantornis, 
Psittaci, some Cypselide, Oaprimulgus, and Megapodius. 

Again, many Cypselide, Hurylemus javanicus, Acanthyllis 
caudacuta, and Ceryla americana have 11 primaries, but are 
eutaxic. 


Goodehild’s observation refers only to the external phenomena 
of the relative length of the feathers composing the different 
rows in this region of the wing. This “faulting” is not always 
visible in diastataxic wings, as is well seen in many Parrots for 
instance. 


Some Degenerate Wings. 


In the present connection it will be sufficient to survey this 
subject briefly. In the most perfect form of wing it will be 
noticed (Pl. 14. fig. 5) that the manus is longer than the forearm, 
and that the angle which the primaries form with the skeleton 
‘changes more and more from within outwards; the innermost 
remex lying at a right angle to, and the outermost parallel with, 
the long axis of the wing. Thus it comes about that the wing-area 
of the hand is as great as, or greater than, that of the forearm. 
Correlated with the form of the wing is the nature of the flight. 
Thus, in the Swifts and Albatrosses the wing is ribbon-shaped— 
very narrow from the pre- to the post-axial border, and much 
produced outwards. In birds like the Heron, the wing is very 


‘¢ AQUINTOCUBITALISM ’’ IN THE BIRD’S WING. 25 


broad, and the flight, though strong and capable of being sus- 
tained for long periods, is not so rapid. 

It will be found that, the less the wing is used, the greater is 
the departure from this type. The manus shortens conspicuously, 
and the wing takes on a rounded form, making it difficult to 
distinguish primaries from secondaries in the outstretched wing, 
This is well seen in the wing of Opisthocomus (PI. 16. fig. 2). 
Again, compare the wings of the Kagu, Psophia, Ocydromus, or the 
Common Water-Rail, with that of the more perfect Cranes; or 
the wings of Coua, Phanicophaés, and Crotophaga, Turacus, and 
Musophaga, with that of the Common Cuckoo ; of Stringops with 
that of other Parrots, and so on. Im all, the wing-area of the 
hand is lessened, markedly so, and the relative length of the 
secondaries is increased, whilst the primaries and their coverts 
grow shorter from within outwards. On discussing this matter 
with my friend and colleague, Mr. Eugene Oates, he drew my 
attention to the fact that this shortening and widening of the 
wing obtains in two non-migratory Indian Ducks. 

The wings of the Rhea, Ostrich, Cassowary, and Apteryx 
afford evidence of still greater retrogression, passing from a 
relatively large wing, such as that of Rhea, in which may be 
distinguished primaries, secondaries, and coverts*, to the vestiges, 
more and more complete, in the Apteryx, Cassowary, 4pyornis, 
Moa, and Hesperornis. 

The wings, then, both of the Ratite and the Carinate, show 
that the reduction of the skeleton is soon followed by a reduction. 
in the size, and then in the number of the remiges, and that this 
latter takes place at the extreme distal end of the primary and 
proximal end of the secondary series, where they become shorter 
and shorter and finally disappear. 


* The pterylosis of the wing of (hea I hope to describe shortly. It differs 
‘markedly from that of all other wings in that the dorsal coverts and remiges)of 
the forearm are clustered together in strongly-marked obliquely-transverse rows: 
separated by deep furrows one from the other. Furthermore, there remains to be 
settled one or two points touching the nature and homology of the remiges, and 
the disposition of the feathers in the carpal region. The ventral surface of the 
wing is bare. 

In connection with the cubital remiges, it is interesting to note that these, in 
the Common Fowl, are, in the nestling, functionally preceded by their major 
coverts. ‘This appears to be the case also in the young Pigeon, as is well seen 
in Pl. 16. fig. 1. | 


252 MR. W. P. PYCRAFT ON SO-CALLED 


The probable Origin of the Diastataxie Wing. 

We may now turn our attention to a discussion on the 
probable origin of diastataxy. 

The primitive wing, I take it, was eutaxie, and resembled that 
of the Common Fowl in that it was clothed by numerous rows 
of covert-feathers; that of the Picariz and Passeres is a special- 
ization of a more primitive type, the number of rows of coverts 
having been reduced. Although, in these, the forearm may have 
increased in length, the remiges have decreased in number and 
become more widely spaced, and have developed broader vanes. 
Thus an equally efficient wing has been obtained with less 
expenditure of material. 

The diastataxic wing is a modification of the eutaxic, and is 
possibly due to an increase in the length of the wing accom- 
panied by a corresponding increase in the number of the remiges. 
It would seem more natural to assume, therefore, that all diasta- 
taxic wings have been derived from a common source; and thus 
this feature may be regarded as a sure sign of affinity, more or 
Jess remote, enabling us to classify all birds into groups eu- and 
diastataxic. 

The existence, however, of what we may term eutaxic genera 
amongst diastataxic families is certainly a serious difficulty in the 
way of this hypothesis. For instance, Columbula is the only 
known exception amongst the Pigeons, which are diastataxic, 
though other exceptional genera may turn up, and the numerous 
instances of diastataxy amongst the Kingfishers and Swifts. It 
might be pleaded that Columbula has re-acquired a eutaxial form, 
by reduction in the length of the wing, and a similar reduction 
in the number, accompanied by a readjustment of the feathers. 
Note the position, for instance, of the 5th and 8th remiges in 
the wines of Columbula and Columba. That this is problematical, 
however, is shown by the wing of Ocydromus, which, though very 
greatly reduced in size, still remains diastataxic, like the rest of 
the Rails. Again, it is probable that the Megapodes, which are 
diastataxic, are somewhat closely related to the Game-birds, 
which are eutaxic. Apart from internal anatomy, they present 
the following points in common :—The remiges, in the nestling, 
are well developed and functional before the pre-penne of the 
trunk are replaced by the definitive contour-feathers. The Ist 
cubital remex develops much later in life than the rest of the 


““ QUINTOCUBITALISM ”’ IN THE BIRD'S WING. 253- 


series, and is always much shorter than these. The Megapodes 
as we have just remarked, are diasta-, and the Game-birds 
eu-taxic. If the two are closely related, we might claim justifica- 
tion in holding that the arrested development of the 1st cubital 
remex was derived from a common source, and that diastataxy 
has been acquired by the Megapodes since then. If this inter- 
pretation be correct, it follows that we may hold it to be 
admitted that diastataxy may have arisen independently in 
different groups of birds,—a somewhat unlikely conclusion. The 
position into which we have drifted, then, may be stated as 
follows :— 

The Class Aves, very early in the process of its differentiation,. 
developed the phenomena of diastataxy, which has been retained 
by very different groups now regarded as only remotely allied. 
The presence of eutaxic forms in an undoubtedly diastataxic 
group, as in the case of Columbula, must be regarded as the 
result of a secondary re-arrangement of the wing-feathers, or as 
a reversion to the more primitive type of wing from which it was 
derived. Such admittedly aberrant eutaxic forms as Psophia, 
Cariama, Heliornis, must be regarded as more remotely allied 
to the diastataxic forms with which they are now associated 
than is generally believed. Diastataxy is probably an indication 
of consanguinity. 

There are certainly difficulties in the way of acceptance of this 
view, perhaps the most formidable being the case of the Swifts 
and Kingfishers, the majority of which are eutaxic; some 
genera, moreover, containing both eu- and diasta-taxic forms. 
We have to face two alternatives :— 


(1) That the group, whichever it may be, really belongs to 
the diastataxic stock, but that the majority of the species, 
like Columbula, have reverted to eutaxy ; or, 

(2) That diastataxy must be explained, in that and all other 
groups, as the result of the action of similar mechanical 
forces, upon a common type, and which may oceur inde- 
pendently in different groups, and even different species 
of the same genus. 


In spite of the objections which we may feel towards the first 
proposition, it seems more probable than the second, and more 
in harmony with the facts as a whole. 


MR. W. P. PYCRAFT ON SO-CALLED 


bo 
ii 
[es 


Diastataxy as a Factor in Olassification. 

Tf it be true that diastataxy is an indication of a more or less 
remote degree of consanguinity, as has just been hinted, or, in 
other words, if diastataxie forms are more nearly related one to 
the other than those which are not, we may find this character, 
used with discretion, no small help in systematic work. I say 
used with discretion advisedly ; for it is incontrovertible that the 
nature of the evidence from other sources makesit absolutely 
impossible to use this character as a primary factor, wherewith 
to divide the Class Aves into two great groups, eu- and diasta- 
taxic. But the presence of diastataxy in a little coterie of forms, 
admittedly related, but hitherto indiscriminately mixed with 
eutaxic, will be a sufficient reason to justify our separating them 
out to form a group by themselevs, on the assumption that the 
character was inherited from a common source, and that they are 
therefore more closely related one to the other than to the neigh- 
bouring eutaxic forms. The presence of discordant elements in 
the shape of eutaxic forms amongst our now diastataxic groups— 
such as the Kinefishers, Swifts, and Pigeons—must be attributed 
to reversion or secondary readjustment of the feathers resulting 
once more in eutaxy. This is not as convincing as it should be ; 
but it demands jess of us than the alternative hypothesis, that 
diastataxy has been independently acquired wherever it occurs. 

The result of the slight shifting here suggested is in no sense 
revolutionary in its tendencies. Amongst the Picarian forms it 
would bring together the Psittaci, the Striges, and the Capri- 
mulgine forms associated therewith, the Swifts and Humming- 
birds and the Kinefishers—all diastataxic, drawn from the ranks 
of eutaxic forms to constitute a little coterie by themselves. 
The Megapodes would be cut off from the remainder of the 
Galline forms, which are eutaxic, just as Heliornis, Psophia, 
Oariama, Rhinochetus, and Hurypyga remain as every modern 
systematist has left. them—as isolated and aberrant groups in 
the neighbourhood of the Grues. Cariama remaining as a sort 
of sign-post pointing the way, as Beddard has recently shown, 
from the Grues to the Accipitres. : é 

‘iThis scheme is doubtless open to criticism; but this may be 


said of every other. 


LINN. SOC. JOURN. ZOOL. VOL. XXVII. PL. 14. 


CSSBSs behets Sak GATE Paria Hae 


Morgan & Kidd, Collo. 


Pycraft del. 


DIASTATAXY (AQUINTOCUBITALISM). 


PYCRAFT. LINN. SOC. JCURN. ZOOL, VOL. XXVII. PL. 15. 


te ae rf a 


Pycraft del. rh = > i ‘ ' Morgan & Kidd, Collo. 


% DIASTATAXY (AQUINTOCUBITALISM). 


PYCRAFT LINN. SOG, JOURN, ZOOL VOL, XXVII, PL. 16. 


Cr 
Hae. 
ts / oy. C. 
Ue 
| A 
| 


Pycraft del. Morgan & Kidd, Collo. 


DIASTATAXY (AQUINTOCUBITALISM ) 


“ AQUINTOCUBITALISM ” IN THE BERD’S WING. 255 


Bibliography. 


(1) Degen, E. “ On some of the Main Features in the Evolution 
of the Bird’s Wing.” Bull. Brit. Ornith. Club, vol. ii., 
1894. 

(2) Gapow, H. Bronn’s Klassen und Ordnung. des Thier- 
Reichs. Vogel. Anatomische Theil, 1891. 

(3) Geree, Z. “Sur les Plumes du Vol et leur Mue.” Bull. 
Soe. Zool. France, tom. ii., 1877. 

(4) Goovcuiip, J. G. “The Cubital Coverts of the Huornithes 
in relation to Taxcnomy.” Proc. Roy. Phys. Soc. Edinb. 
vol. x., 1890-91, pp. 317-333. 

(5) Pycrart, W. P. “A Contribution to the Pterylography of 
Birds’ Wings.” Trans. Leicester Lit. and Phil. Soe., 
vol. 11. p. 123, 1890. 

(6) Szersoum, H. Classification of Birds. 1895. 

(7) SunpEvatt, C. J. On the Wings of Birds. (Translated 
from the Swedish.) Ibis, 1886, pp. 389-457. 

(8) Wray, R.S. “On some Points in the Morphology of the 
Wing of Birds.” Proc. Zool. Soc. 1887, p. 343. 


EPPLANATION OF THE PLATES. 


Reference letters. 


¢.¢. = carpal covert. 
c.7. = carpal remex. 
7.¢. = intercalary coverts, intercalary row. 
med.c. = median covert. 
my.¢. = major covert. 
v.mj.c. = ventral major covert. 
min.e, = minor covert. 
O11 Ce oh 


Puate 14. 


Fig. 1. Right wing, dorsal aspect, of an embryo Vanellus cristatus, showing the 
first appearance of the feather-papillze representing the remiges and 
major coverts of the forearm,—and indistinctly of the primaries. 

Fig. 2. The same in a more advanced stage of development. Several rows of 
coverts have now appeared, and “ faulting” has already taken place. 
The wing is now diastataxic ; in fig. 1 it is still eutaxie. 

Fig. 3. Portion of the right wing, dorso-ventral aspect, of an embryo Machetes 
pugnax, showing position of ventral major coverts. 

Fig. 4. Right wing, dorsal aspect, of an embryo Gallus bankiva, var. domestica. 
This is an eutaxic wing. The coverts have been marked as in the 
diastataxic forms for the purpose of comparison. 


256 MR. F. Je COLE ON THE DEVELOPMENT OF 


Fig. 5. Left wing, dorsal aspect, of an adult Asio accipitrinus, to show the 
typical, adult, diastataxie wing. Note the absence of a remex between 
the 5th pair of major coverts, and the marked gap between the 4th 
and 5th remiges. 

Puars 15. 


Fig. 1. Right wing, dorsal aspect, of an embryo Columba domestica, This is 
markedly diastataxic. The shifting of the coverts is very distinct. 
Compare Pl. 16. fig. 1. 

Fig. 2. Right wing, dorsal aspect, of an embryo Lomvia troitle, at present 
eutaxic; but a study of the coverts shows that a shifting has com- 
menced, the result of which ultimately reduces the wing to the typical 
diastataxic form. Compare this with the figure on p. 243, which 
shows the condition of the wing in the downy nestling. 

The figure immediately below is drawn from fig. 2 to show the 
effect of a slight increase in the shifting}of thezcoverts transforming 
the wing from the eutaxic to the diastataxic type, as seen in fig. 1. 

Fig. 3. Right wing, dorsal aspect, of an embryo Anas boscas, var. domestica, 

decidedly diastataxic. No earlier stages were procurable. 


Puate 16. 


Fig. 1. Right wing, dorsal aspect, of a nestling Columba domestica. Note the 
intercalary row of coverts, and compare with fig. 1, Pl. 14. ; also the 
large size of the major coverts of the forearm as compared with the 
cubital remiges (Ist ¢.r.), which have as yet only just begun to project - 
beyond the surface of the wing. 

Fig. 2. Right wing of adult Opisthocomus. 


On the Discovery and Development of Rhabdite-“ cells” in 
Cephalodiscus dodecalophus, McIntosh. By ({F. J. Cons, 
University College, Liverpool. (Communicated by Prof. 
G. B. Howes, Sec. Linn. Soc.) 


[Read 6th April, 1899. ] 
(PuateE 17.) 


A snort while back Professor Herdman was kind enough to 
place in my hands some small pieces of Cephalodiscus for treat- 
ment and sectioning by modern microscopical methods. As 
interest in this unique form has been again aroused by the 
recent work of Masterman *, it was proposed to revise the whole 
anatomy of the polypide besides investigating the few points 
which a consideration of the literature showed to be unsettled. 


* Q. J. M. S. vol. xl., 1897; Trans. R. 8. Edin. vol, xxxix. pt iii., 1898. 


RHABDITE-CELLS IN CEPHALODISCUS. 257 


So far, the results are the bodies described in the present 
communication, a possible true sense-orzan in the region of the 
gill-clefts, and a large and undoubted gland situated on the 
proboscis *. Descriptions of the two latter are left over for 
the present. Of the material at my disposal sections were cut 
in various planes and treated by various methods. The whole 
ceencecium with the contained polypides was sectioned, as well as 
also individual polypides orientated by the dissecting-microscope 
in paraffin. The latter process, owing to the size of the 
individuals, is by no means difficult, so that it is not necessary 
to resort to Patten’s method. So far as the purposes of the 
present paper are concerned, the only method of staining found 
to give really satisfactory results is Dr. Gustay Mann’s ex- 
cellent combination of methyl-blue-eosin +. The sections were 
stretched on a slide previously treated with Paul Mayer’s 
albumen-fixative and covered with a film of water, and then 
stained on the slide according to the directions given by Dr. Mann. 
Successful preparations that have not been over-stained (when 
properly decolorized for the rhabdites the general tissues are 
almost unstained) show a perfect differentiation of the rhabdite- 
“cells,” so that their structure is somewhat easily followed by 
examination with Zeiss’s 1°5 mm. apochromatic lens in conjunction 
with the compensating eyepiece No. 12. 


HiIstToRIcat. 


As is well known, after the return of the ‘Challenger’ 
Expedition, the bottle containing the specimens of Cephalodiscus 
obtained in the Straits of Magellan was sent, with the collection 
of Tunicata, to Prof. Herdman. It then bore a label in the 
writing of the late Prof. Moseley, stating that the animal was a 
“compound Ascidian.” Prof. Herdman examined it in the 
winter of 1879-80, and mounted some preparations in different 
ways (including the material referred to above), sufficient to 
determine that it did not fall strictly within the group Tunicata, 
and that its affinities were rather with what are now considered 
the other Protochordata. He returned the stock to Sir Wyville 


* This apparently is not the structure referred to by Harmer (Zool. Anz. 
1897), and I am not yet in a position to state its relation to the proboscis- 
gland of Balanoglossus. 

+ Journ. Anat. & Phys., vol. xxix. 

LINN. JOURN.—ZOOLOGY, VOL. XXVII. 19 


258 MR. F. Je COLE ON THE DEVELOPMENT OF 


Thomson, with a statement that he did “ not consider 1t a com- 
pound Ascidian, but rather an aberrant Polyzoan related to 
Rhabdopleura.” As a result the animal was at once forwarded 
to the late Mr. George Busk, and after that to Prof. McIntosh, 
as stated by the latter in the ‘Challenger’ report on Cephalo- 
discus (p. 8). The material I have used is a few fragments made 
use of by Prof. Herdman in his first determination of the probable 
affinities of the animal. 


As described by Professor M‘Intosh in his ‘ Challenger’ 
report, the branchial plumes ‘“‘are nearly of uniform size, and 
consist of a thickish central stem, occasionally slightly crenate, 
and furnished with a series of longitudinal fibres; while distally 
each is terminated by a peculiar bulbous enlargement, which at 
first sight resembles the tip of certain hydroid tentacles (e. g. 
Coryne or Syncoryne) bristling with dart-cells and pigment. 
The rugose appearance, however, is due to large gland-cells 
containing granules and globules, which are arranged in a some- 
what regular manner round a central cavity, and which present 
a deep yellowish tint in the preparations. This structure may 
perhaps be a further and special development of the somewhat 
large hypodermic granules of the tips of the pine.” Elsewhere 
MelIntosh states that the bulbous enlargements of the plumose 
arms may secrete the “ spinous processes or fimbrie” on the 
surface of the coenecium. Although McIntosh did not succeed 
in elucidating the nature of the “large gland-cells,” his description 
above, as far as it goes, and excepting perhaps the last statement, 
is quite correct. 

Masterman’s interpretation of the bulbous enlargements 
(which may be conveniently termed “ rhabdite-batteries”) is 
that they represent “a dozen large eyes of a very primitive 
compound type.’ As this conclusion is so directly opposed to 
the explanation given in this paper, it 1s perhaps as well that 
the evidence on which it is based were summarized. It is as 
follows :— 


1. “If the parts be subjected to partial maceration the clear 
globules [inside the gland-‘ cells’ ] can be obtained free, 
and they remind one irresistibly of a crystalline refractive 
lens.” 


RHABDITE-CELLS IN CEPHALODISCUS. 259 


2. “All [the gland-‘cells’] have fine pigment granules 
scattered throughout their interior, and a great number 
of them contain the crystalline lenses referred to.” 

3. The base of the ‘“‘ eye” is believed to have been “in some 
cases traced into the main nerve of the plume.” 

4. “The whole structure here described seems to indicate 
that these organs are rudimentary monostichous compound 
eyes, which bear a remarkable resemblance, both in 
appearance and structure, to the ‘branchial organs’ 
found in the sedentary Annelids, such as Potamilla and 
Sabella..... It seems most reasonable to regard them 
tentatively as primitive eyes, though the presence of 
compound eyes in the Chordata is rather remarkable.” 


First, as regards matters of fact, I find myself unable to 
confirm the statement as to pigment in paragraph 2, the existence 
of a nucleus as shown in the figure, and also the belief as to the 
nerve-supply in paragraph 3. Further, fig. 80 in Masterman’s 
paper is, I must confess, quite unlike anything I have seen. In 
all my sections the wall of the battery is considerably vacuolated, 
and I have never seen the cells closely opposed as shown in this 
figure (cf. my fig. 1). 

Second, as to matters of imterpretation, passing over the 
insufficient nature of the evidence on which Masterman bases so 
important a statement, the finer structure of the bodies in 
question as here described must, assuming the accuracy of the 
description, be held to negative the view which Masterman has 
stated. 

In his later paper (op. cié. p. 521) Masterman, in referring to 
the blastogenesis of the plumes and pinne, says: “The plumes 
arise throughout in pairs. They first make their appearance as 
a papilla, which elongates to a finger-shaped process, the distal 
extremity of which becomes slightly swollen, and then bulbous. 
The epithelium of this bulbous extremity then becomes modified 
to form the eyes. The cuticle of certain of the epithelial cells 
becomes thickened, and soon the thickening protrudes into the 
cavity of the cell as a lens-like body. Later it is detached from 
the cuticle and lies freely in the protoplasm. Here it becomes 
rounded off to form the lens.” In plate iv. figs. 75, 76, & 77, this 
process is illustrated, showing the nucleus of the “eye” at first 
peripheral in position, and afterwards thrust down to the basal 


260 MR. F. J. COLE ON THE DEVELOPMENT OF 


extremity of the cell by the ingrowing cuticle. As I have not 
investigated the blastogenesis of the rhabdite-“cells,”. I am 
not in a position to express an opinion as to the accuracy of the 
above statement. J can, however, assert that a true cuticle is 
not present on the free surface of the rhabdite-battery of the 
adult, although a peripheral deeply-staining membrane is often 
seen, but this is not a cuticle. That it would be noticed if 
present in my sections is shown by the fact that the axis of each 
branchial plume has a cuticle which is quite obvious, but with 
one exception, when it was traced on to the base of the battery, 
this is always seen to stop short of the knob of the battery. 
Indeed in some sections, which were kindly placed at my disposal 
by Prof. Howes, indications were not uncommon of a few rows 
of cells situated external to what I have supposed is the free 
surface of the battery, z. e. the surface bounded by the supposed 
cuticle. It is possible these may represent a true epidermis 
which has been lost by the maceration of the material, although it 
seems very improbable that it would have been lost in by far 
the greater majority of the batteries. The existence of such a 
layer is, however, rendered conceivable by the position frequently 
assumed jby the immature rhabdite-“ cells,” and also by the 
usually ragged and seemingly artificial free border of the enlarge- 
ments themselves. 


DESCRIPTION OF THE RHABDITE-“* CELLS.” 


The occurrence of the “cells” of the rhabdite-batteries on the 
branchial stems has been correctly described and figured by 
McIntosh, so that it is not necessary to recapitulate it here. A 
close examination of the rhabdite-“ cells” in a number of 
specimens discloses two important and significant facts: (1) No 
two “cells” are ever exactly similar—hence they are “cells” in 
which a great amount of metabolism is going on; (2) there are 
two extremes of position—(a) near the basal cells of the battery, 
and (0) absolutely outside the battery and lying on its surface. 
These two extremes of position are bridged over by the necessary 
intermediate stages. These two observations alone justify us in 
concluding—(1) That the “cells” are engaged, and actively 
engaged, in secreting something, and are hence of a sort gland- 
cells; (2) that that something is a body which, to serve its 
purpose, must be cast on to the exterior. Similarly, the observations 
at once preclude the possibility of the “cells ” beg sense-organs 


RHABDITE-CELLS IN CEPHALODISCUS. 261 


of any sort. They are not excretory organs, as their position and 
structural relations imply, and they do not store up food-material 
like the sacculi of the Crinoids, as the products of the “ cells” 
are (at least often) ejected on to the exterior. But since, on 
the other hand, the ultimate products of these bodies are a 
number of pointed or somewhat blunt rods, since the metabolism 
of the “cells” is always in the direction of the production of 
these rods, and since finally these rods can in some cases be seen 
in the various stages of being, by the rupture of the “ cell,” 
shot on to the exterior, the only structures with which the 
“cells” can be compared are clearly the rhabdite-cells of 
Turbellaria aud Trematoda, and the less specialized bodies found 
in the integument of Nemerteans. They will have the same 
function doubtless as the rhabdite-cells in the latter groups, 
whatever that is, and they are produced in Cephalodiscus by the 
following series of changes. 

It is first necessary to describe the structure of the wall of 
the battery itself (fig. 1). Such a description is necessarily 
based on the material as I found it, and is thus lable to a 
percentage of error due either to post-mortem changes or 
imperfect fixation or both. Figure 1 is a drawing of a portion 
of the wall of the battery which, after a prolonged search, seemed 
to have been most favourably preserved and cut accurately at 
right angles to the surface. The cavity of the battery (10) is 
lined by a series of occasionally nucleated fibres (8), which are 
doubtless the direct continuation of the longitudinal fibres of 
the stem described by McIntosh. Many of these fibres have 
snapped in fixation (9), giving the appearance under ordinary 
lenses of a row of large cilia projecting into the cavity. Situated 
on these fibres is a single row of large irregular cells (4), each 
containing at least one undoubted nucleus. In some preparations 
it can with certainty be made out, as shown in the figure, that 
the cells are continuous at the base, so that a perfectly con- 
tinuous layer of protoplasm surrounds the Jongitudinal fibres (4). 
From this layer of protoplasm there occasionally passes a long 
filament (7) which lodges one or more nuclei, and passes straight 
upwards to anastomose with the free surface of the battery (/). 
Similar filaments, which are however much more numerous, and 
also lodge nuclei, pass from the basal cells themselves to the 
surface (6). Whether either one or both series of filaments 
represent the narrow interstitial cells described by Burger as being 


262 MR. F. J. COLE ON THE DEVELOPMENT OF 


wedged in between the “ pseudo-rhabdites ” of Nemerteans is a 
possibility which only perfectly preserved material can determine. 
The three interesting features therefore about the basal cells 
are :—(1) Their protoplasm is continuous proximally; (2) they are 
separated by distinct intervals from each other, and present an 
almost amoeboid appearance; (3) they are connected by one or more 
nucleated filaments with the periphery of the battery. Lying in 
the spaces defined by the latter filaments are the rhabdite-“ cells” 
themselves (3)—to be described below. The varying position 
and structure of these bodies may be noted here. The periphery 
of the battery is often defined by a fine membrane (1), which 
seems to rupture to admit the passage of the rhabdites, whilst 
underlying this is a stratum containing two or three rows of 
undoubted nuclei (2). What this layer is the condition of the 
material did not permit me to determine, but many of the nuclei 
in the particular section shown in the drawing were certainly 
situated on the filaments described above. It is possible that 
this layer with its nuclei represents the epidermis, or, together 
with the basal cells, the dermis. 

Figure 2 shows what I conceive to be an early stage in the 
development of the rhabdites. The “cell” was projecting 
slightly from the surface of the battery (7), and contained two 
highly refractile rounded bodies—one being very much larger 
than the other. There can be little doubt, from their general 
appearance and waxy homogeneous structure, that these bodies 
are simply secretions of some sort, although of what nature 
I was unable to determine. This identification was at once 
independently suggested by the biologists to whom I showed 
the slides. Besides the larger secretions are two smaller aster 
secretions, which bear a superficial resemblance to centrosomes 
but which have not of course any relation to those problematical 
bodies. One of these asters is connected with the smaller 
secretion-sphere, which suggests that it may have originated by 
the fragmentation of that body. The asters are also shown in 
figs. 4, and 6a & 0b. Beyond that they appear to pursue the 
same course of development as the rhabdites themselves, that is 
to say they are formed by the breaking up of an originally 
spherical secretion-mass, of which the earliest stage I have seen 
is shown in fig. 4, I have failed to ascertain where they come 
from or what ultimate réle they fulfil. The presence, however, of 
other asters consisting simply of very fine rods with no central 


RHABDITE-CELLS IN CEPHALODISCUS. 263 


secretion-mass, as shown in figs. 2 and 6a, seems to indicate that 
they perform the same function as the larger rods, although 
such explanation does not meet all the facts. They are of fairly 
frequent, but not of universal occurrence, and they are the only 
bodies in the rhabdite-“ cells” which are of a fairly constant 
structure and position, and do not vary within wide limits. For 
the rest, the irregular outline or sac of the “cell” shown in 
fig. 2 contains a granular substance which seems to be either 
pure plasma or a more or less slight modification of that substance. 
It is to be noted that it is more densely granular at the two 
poles, the base, however, being more granular than the apex. 
Particular attention must be directed to the fact that in this 
“cell,” as in all the others examined, there is no structure 
whatever that can with any justification be called a nucleus. I 
have hence in describing the bodies as rhabdite-cells placed the 
word cell in inverted commas. 

Figure 3, drawn from a single field, illustrates three conditions 
well:—(1) the variation in position of the “cells” —d lying 
somewhat below the surface, a and 4 immediately beneath it, and 
c and f completely outside it; (2) the fragmentation of the 
secretion-mass as shown inf, d, g, and; and (3) the splitting- 
up of the secretion-mass to form rhabdite-rods, as shown in all 
except c. In the latter we have only the secretion-sphere and a 
small vacuole—the most undifferentiated rhabdite body that has 
been seen. In 0, however, the sphere has fragmented into two 
pieces—each piece lying apparently in a vacuole (an interesting 
feature also seen in the rhabdite-cells of certain Turbellaria), and 
just commencing to split up. Hence the serrated appearance of 
the periphery of the two pieces. In g, d, e, and f, successive 
stages in the splitting up of the secretion-mass are seen, whilst 
in a the splitting has proceeded so far that a number of rods 
have been formed connected by a central mass of hitherto un- 
differentiated secretion. The latter contains a small vacuole (see 
also fig. 4), whilst outside the clump of rods are two small 
secretion-masses, which may either be the two aster anlagen or 
fragments of the original secretion-sphere. 

In figure 4 we have a condition intermediate between f and a 
of the preceding figure. The secretion-sphere lies in a large 
vacuole, and the splitting up, though somewhat far advanced, has 
yet not proceeded as far asin 3a. The central vacuole has been 
already noticed. Above, at the apex of the “cell,” is a bent 


264 MR. F. J. COLE ON THE DEVELOPMENT OF 


plate of secretion which I have not seen in any other “cell,” 
unless it is comparable to the rod in the same position seen in 
fig.5. Below are two large asters of a comparatively simple type, 
and of the usual structure and position. As shown in figs. 3a and 
6 6, the split secretion-sphere does not always lie in a vacuole. 

The rhabdite-“ cell” shown in fig. 5 is lying practically outside 
the battery, the rhabdites have broken away from the central 
mass, leaving the latter lying in the middle of the “cell.” At 
the base is another mass of secretion, formed doubtless by an 
antecedent fragmentation of the original sphere. At the apex is 
a thin transverse rod, perhaps comparable to the curved plate in 
fig. 4, which I have not seen in any other “cell.” The “ plasma” 
does not completely fill the latter. The rhabdites are here 
practically ready to be discharged, and to the left are seen several 
free rods formed by the discharge of an adjacent “ cell.” 

Figure 6 shows two contiguous “cells” (lying immediately 
under the surface) from one field. Jn 6 we have a stage similar 
to that in 3a, except that in the former two small asters are 
present, each lying in a clear space, whilst the splitting of the 
secretion-sphere has not proceeded so far. In 6a, however, the 
splitting has resulted in the formation of two kinds of rhabdites 
—stout and fine. The latter are still adherent to the residue of 
the sphere and are the more numerous, whilst the former have 
broken off, may project through the wall of the “ cell,” and bear 
evident traces of their origin from such stages as those figured 
in 606 and 38a. The whole, together with a small aster of 4 rays, 
lies in a clear space in the “cell,” two small asters being em- 
bedded in the “ plasma.”’ 

The “cell” shown in fig. 7 is an almost isolated example of its 
kind, and no other exactly resembling it was seen *. Empty sacs, 
however, having faint longitudinal folds, which have been iden- 
tified by others besides myself as similar bodies that have been 
discharged, are not uncommon. In the “cell” above, which lay 
immediately below the surface, the rhabdites were arranged in a 
definite axial bundle, no small asters were present, there were no 
clear spaces in the “plasma,” and the whole of the secretion had 
been used up in the formation of the rhabdites—a condition not 
often seen. This stage may be described as the final one imme- 
diately prior to discharge. 

* Since writing I have observed others in Prof. Howes’ material at the 
R. College of Science, Lond. 


RHABDITE-CELLS IN OCEPHALODISCUS. 265 


Finally it is necessary to mention another stage which was 
only of occasional occurrence in the material at my disposal, but 
which seems to be common in the sections of Cephalodiscus in 
the laboratory of the Royal College of Science. Here the 
rhabdite-“ cells” are sharply divided into two parts, so as to 
present a strong superficial resemblance to certain of the pseudo- 
rhabdites described by Biirger in Nemerteans. The upper or 
apical portion contains a body which undoubtedly corresponds 
to the secretion-sphere and its products described above. The 
lower or basal portion is of a uniform, structureless and highly 
refractile appearance, as if this portion of the “‘ cell ” were filled 
with a homogeneous waxy secretion. J am unable to connect 
this stage with the others with certainty, but it seems to be an 
early stage in the formation of the rhabdites, and may possibly 
be the earliest yet seen in the material. Its occurrence was of 
too occasional a character in my material to enable me to study 
it as carefully as the other stages. 

The question that now arises is from what source are the rhab- 
dite-“ cells ” of Cephalodiscus derived. That they are portions of 
cells, and not complete cells in themselves, must be admitted, in 
view of the fact that they do not possess any structure that may 
be justifiably interpreted as a nucleus. On the other hand, 
their position and mode of occurrence is strongly suggestive of 
the view that they are disassociated portions of the basal cells 
(fig. 1, 4) of the battery. These basal cells therefore, on this 
view, will be constantly giving rise to the so-called rhabdite- 
“cells,” and the latter will as constantly be discharged on to the 
exterior. The term rhabdite-cell will hence belong properly to 
the bodies described in this paper + also the basal cells of the 
battery, since the two together are the morphological equivalent 
of the Turbellarian rhabdite-cell and its contents. 

In order to justify the term I have applied to the structures here 
described, the following comparison is appended between these 
bodies and the cells in Turbellaria to which the term rhabdite- 
cell was first applied by Graff in 1882. Lang* in the Polyclads 
distinguishes two kinds: (1) Rhabdite-cells ; and (2) “Schleim- 
stibchenzellen”” or Pseudo-rhabdites. Of the former he says 
(pp. 51-52) :—“ Der Kern der Stabchenzellen liegt stets am 
basalen Ende derselben ; das freie distale Ende der Zellen ist mit 
Flimmerhaaren besetzt, wovon man sich durch Isoliren der 

* Fauna u. Flora d. Golfes y. Neapel, Monog. xi. pp. 51-55. 

LINN. JOURN.—ZOOLOGY, VOL. XXVII. 20 


266 MR. F. J. COLE ON THE DEVELOPMENT OF 


Stabchenzellen des lebenden Thieres leicht tiberzeugen kann.” 
The “ Flimmerhaaren ” I have not seen in Cephalodiscus, although 
it is possible that living material may disclose them. Lang’s 
account of the development of the rhabdites is too long to be 
quoted 7m evtenso, but may be summarized as follows:—Situated in 
between the mature rhabdite-cellshere and there are much smaller 
cells each containing a nucleus. Lying close to the nucleus is 
a single small, round, homogeneous, highly refractile body. This 
body Lang regards as essentially a secretion. It grows and frag- 
ments to form a number of small balls, and each ball becoming 
lengthened and spindle-shaped, forms a rhabdite-rod—the whole 
of the rods becoming subsequently arranged to form a pyramidal 
bundle with the base opposed to the nucleus. The rhabdite- 
cells therefore contain a nucleus, plasma, and the rods. Lang 
and Graff are agreed that they are gland-cells, and that the 
rhabdites are their secretion. 

The Pseudo-rhabdites of Lang have an uneven periphery. 
They are figured in some species as one or more tall columns of 
end to end secretion-fragments (Blockchen) of an irregular shape, 
with the nucleus and plasma of the cell lying at its base and under 
the pseudo-rhabdites. In Stylochus, Lang says (pp. 53-54) :— 
“Die einzelnen Blockchen entsprechen ihrem optischen Ver- 
halten nach sehr den Rhabditen, sie sind klar, homogen, stark 
lichtbrechend und verhalten sich Farbmitteln gegenitiber ganz 
wie diese. Die Saulen, die sie bilden, erfullen beinahe die ganze 
Epithelzelle, in der sie hegen, und lassen hochstens am basalen 
Theil, wo der Kern hegt, ein Kliimpchen feinkornigen Plasmas 
frei.’ Lang considers that the psendo-rhabdites are fully com- 
parable to the true rhabdites, and describes several stages 
intermediate between the Stylochus-type and the mature rhabdite- 
cell with its clump of rods. 

Just as the pseudo-rhabdite must be regarded as a compara- 
tively simple modification, in which the secretion has not under- 
gone such differentiation as in the true rhabdite-cell, so do 
certain gland-celis in Nemertea represent a condition antecedent 
to the pseudo-rhabdite form. Hubrecht* describes and figures in 
Cerebratulus and Hupolia unicellular glands which he considers 
correspond precisely to the pseudo-rhabdites of Lang. They have 
in Oerebratulus highly refractile uniform contents, not, however, 


* ‘Challenger’ Reports, vol. xix. p. 61. 


RHABDITE-CELLS IN CEPHALODISCUS. 267 


divided into blocks. Biirger* describes other homologous bodies 

Nemertea as flask-shaped gland-cells. 

According to these descriptions there can be little doubt that 
the bodies described in this paper are similar, in all their essential 
points of structure and development, to the rhabdites, pseudo- 
rhabdites, &c. of Turbellaria and Nemertea. We have in fact 
a complete and gradually ascending series commencing with the 
comparatively simple cells in Nemertea, and terminating in the 
complex structures of Cephalodiscus. In Cephalodiscus they are 
more differentiated (1) as regards accessory secretions in the 
cells; (2) as regards the details of the development of the rhab- 
dites ; and (3), provided the view stated in this paper be correct, 
in the separation of the portion containing the secretion from 
the mother-cell, so that the two become quite distinct. 


SUMMARY. 


Cephalodiscus has a lophophore of 12 branchial plumes, each 
of which consists of a central stem or axis with its associated fila- 
ments. Hach axis becomes enlarged at its distal or free extremity 
so as to form a conspicuous hollow bulb, the cavity of which is 
continuous with the cavity of the stem. As the rhabdite-cells 
are entirely confined to these bulbs, the latter may be con- 
veniently termed rhabdite-batteries. Histologically, the wall of 
the battery is greatly vacuolated, and contains essentially two 
series of bodies: (1) a series of large nucleated basal cells ; 
(2) above these a series of non-nucleated bodies lodged in the 
vacuoles and termed rhabdite-“ cells.” 1 and 2 together, how- 
ever, are the equivalent of a rhabdite-cell of a Turbellarian, 
since 2 possesses no nucleus, and 1 contains no secretion. In 
fact 2 must be regarded as a disassociated portion of 1. The 
secretion of 2 is primitively a large homogeneous sphere. 
This may or may not become fragmented. Subsequently there 
often arise two small aster secretions of unknown origin, fate, 
and significance. The sphere of secretion afterwards splits 
peripherally so as to form, first a star-shaped structure, and then 
a number of stout free rods. The splitting usually leaves a 
residue, but the whole of the sphere may be used up in the 
formation of the rods. The rods have been observed arranged 
in a definite bundle parallel to the long axis of the “cell.” The 


* Bronn’s Thier-reichs, Bd, iv. 


268 DEVELOPMENT OF RHABDITE-CELLS IN CEPHALODISCUS. 


rods and sometimes the whole “ cell” may be found lying out- 
side the battery. No two cells are ever structurally idéntical, and 
their position varies from one in close proximity to the basal 
cells to one in which they lie completely outside the battery. 
Besides the secretion-spheres and their accessories, they contain 
a granular substance in varying quantity identified as plasma or 
a modification of that substance. In all essential respects they 
are comparable to the rhabdite-cells of the Turbellaria (and are 
hence of the same nature as the pseudo-rhabdites of Nemertea), 
since their prime object is to secrete rods and then to discharge 
them on to the exterior. 


EXPLANATION OF PLATE 17. 


All figures drawn with Zeiss’s 1:5 mm. apochromatic lens with the 
oc. compens. 12. 


Fig. 1. Portion of a median longitudinal section through the rhabdite-battery 
and stem of a branchial plume, showing the structure of a piece of the 
wall of the battery. Reduced. 

1. Peripheral membrane. 

2. Peripheral nuclei situated in the superficial layer of the wall of 
the battery. u 

3. Rhabdite-“ cells” (semi-diagrammatic ; note position). 

4. Basal cells. 

§. Stratum of protoplasm placing all the basal cells in communi- 
cation proximally. 

6. Nucleated filaments from basal cells to periphery. 

we ‘ ee from 4 to periphery. 

8. Layer of longitudinal fibres (here appearing transverse). 

9. Fibres of 8 snapped in fixation. 

10. Cavity of rhabdite-battery (continuous with that of stem). 

Figs. 2 to 7. Stages illustrating the development of the rhabdite-rods 
(arranged as far as possible in order). The ‘‘cells” are in all cases 
represented in the same position, 2. ¢., J is the peripheral membrane 
and above it is the exterior. The “cell” shown in fig. 5, therefore, 
lies practically outside the battery, and those of fig. 6 lie immediately 
under the surface. 


Linn Soc. Journ. Zoou. Vou. XAVIL, Pr. 17. 


Cole. 


MtFarlane & Erskine, Lith.Edin? _ 


Cole, del. 


RUSSO S iN (Cis PIVNLO US CUS). 


ON THE EDWARDSIA-STAGE OF LEBRUNIA. 269 


The Hdwardsia-stage of the Actinian Lebrunia, and the Forma- 
tion of the Gastro-celomie Cavity. By J. E. Durrpsn, 
Assoc. Roy. Coll. Sci. (London), Curator of the Museum of 
the Institute of Jamaica. (Communicated by Prof. Howes, 
Sec. Linn.Soc.) 

[Read 15th June, 1899.] 


(Puates 18 & 19.) 


CoNTENTS. 
Page 
10, PS SRSA ee opmponesndngundoa soueroononde As qoonbadacedaseobauadadcABueedes 269 
Meerexternal Characters) “cessaccssessee ace eee eee ae eee 271 
DevelopmentiofMentaclestioncesnansseestetee eet eeraesen ee 273 
TIT. Internal Anatomy and Histology ...............ceeccseeeeee saeeiae 276 
NEM CHOR CLINI LcPes ccc see iacteeunecee seer ee eee eas 278 


2. The Archenteron and Formation of the Gisophagus... 281 
3. Larval Coelomic Spaces and Formation of the Gastro- 


ColomichCavitype nesses. eescicanscerer ss reece eee 285 

AG AMIESentericst ne chaceacccseesoesaeh cece contac Lee 291 
HaViesenterialuilamentapsessccstcaesscceeececn eee eee eee eee 292 

IV. Relations of the Tentacles and Mesenteries ...........0.0..0-60 297 
Wo: (Glaravollinerigt avs ane aaron conanncoodatn oO ICeCEeeR Oo ROE eA cH Sone cnatacane 300 
Wilke ibliographyin.cs.s.cnacsntcncascscascescane reas seaccuscaudasscascnsene oll 


Durine the temporary establishment in November, 1898, of a 
marine biological laboratory at Bluefields *, Jamaica, in connection 
with the Institute of Jamaica, several specimens of a Lebrunia 
were found in the act of extruding larve. An examination of 
these, while in the living condition and when sectionized, discloses, 
amongst other characters, some very exceptional features in the 
development of the tentacles, and in the formation of the gastro- 
coelomic cavity of the adult. I am also able to add, by way of 
comparison, a few facts in connection with the freshly-extruded 
larvee of Aulactinia stelloides, McMurrich. 


I. Systematic. 


The specific determination of the first-mentioned Actinian 
calls for a few remarks. It is a small form which occurs in 


* Bluefields is the name given to a property, formerly a sugar estate, on the 
south-west coast of Jamaica, rendered classic to local naturalists as the residence 
of Philip Henry Gosse, F.R.S., during his collecting trip of eighteen months, 
from 1844-46. The principal features of its natural history, and those of 
Bluefields Bay, are most vividly portrayed in Gosse’s work: ‘ A Naturalist’s 
Sojourn in Jamaica’ (London, 1851), 


LINN. JOURN.—ZOOLOGY, VOL. XXVII. 21 


270 MR. J. E. DUERDEN ON 


considerable numbers all round the island, at a depth of two or 
more feet, within crevices of coral-rock. In ‘The Actiniaria 
around Jamaica” (1898), I identified it as the Hoplophoria 
coralligens of H. V. Wilson (1890), transferring it at the same 
time to the genus Lebrunia. 

The chief characteristic of this genus is the presence of four 
to eight dichotomously branched outgrowths (pseudo-tentacles) 
from the uppermost region of the column. They occur im- 
mediately external to the tentacles, and usually bear spheroidal 
batteries of nematocysts. 

' Dr. Wilson’s species was founded upon a single specimen, 
about 2 mm. in diameter, discovered in a hole in the coral-rock 
on the Bahama reefs. There appears to be the closest external 
resemblance between the Jamaican and the Bahaman examples; 
and anatomical and histological examination supports this, even 
to the peculiar forms of the nematocysts, and an exceptional 
arrangement in separate bands of the endodermal muscle of the 
columnar outgrowths. Comparisons made with the Bahaman 
Lebrunia described by Prof. McMurrich (1889), and with the 
Jamaican representatives of the genus, also show a close agree- 
ment; hence, in the paper above mentioned, I stated (p. 457) 
that “it is possible that it may be but a young form of 
L. neglecta.” 

When I first came upon the Jamaican polyps specimens were 
sent to Prof. Wilson for his opinion as to their similarity with 
his species, and he obligingly writes as follows:—“In your 
preserved specimens the pseudo-tentacles have a noticeably 
different appearance from those of my single specimen. The 
white nettle-batteries so conspicuous in my specimen are not 
obvious in yours. I note though that you find them conspicuously 
outlined in the living animal. The pseudo-tentacles in your 
specimens are much larger and more flaccid than in mine. All 
slight and variable points of difference. I make no doubt they 
are the same provided your sectional study shows the internal 
anatomy and histology to be alike in your specimens and in 
mine.” 

Prof. McMurrich (1898, p. 183) has expressed the opinion that 
Wilson’s species is identical with Viatrix globulifera, Duchassaing. 
But in the description and figures given by Duchassaing and 
Michelotti (1860) of this Guadaloupe species the columnar 


THE EDWARDSIA-STAGE OF LEBRUNIA. OHI 


eutgrowths are very numerous and vesicle-like, and the polyps 
float freely, characters never met with in the form under con- 
sideration. 

Within the past year, Prof. A. BH. Verrill (1898), without 
experience of either form, has cast doubts upon the identity of 
the Jamaican specimen with the true Hoplophoria coralligens, 
and also with the conclusion that it may be but a young form 
of Lebrunta. 

From the above facts, however, no other conclusion than 
that it is identical with Wilson’s species seems possible, and in 
the fuller description yet to be published I hope to show its 
agreement with the Jamaican Lebrunia. Meanwhile, it will be 
sufficient to refer to it as Lebrunia coralligens (H. V. Wilson). 

The adult polyps are hermaphrodite. Ova and spermaria 
occur in considerable numbers closely associated in the same 
mesentery. 


Tl. Exrernan CHARACTERS. 


In living polyps the larvee were to be seen through the some- 
what transparent tissues of the parent moving about within the 
celenteron, and especially within the tentacular cavities. As 
many as five were contained within a single tentacle. 

So far as could be observed, they possessed little or no motion 
of their own, but allowed themselves to be carried along by the 
circulation of the ccelenteric fluid. By a peculiar jerking action 
of the adult three or four larve at a time would be shot out 
through the mouth for some distance, and then sink to the 
bottom of the vessel. Here they would lie unattached for some 
time in any position, though usually on their side. Afterwards 
slow movements, both translatory and rotatory, would commence, 
but these at no stage became pronounced. Individuals might 
come to rest at apparently any point of their surface and again 
commence their swimming movements. 

When first extruded the larve measured about 1 mm. in length 
and were of a pyriform shape (PI. 18. fig. 1), divided by shallow 
grooves into eight, nearly equal, longitudinal regions. Sometimes 
they would take the form represented in fig. 2, or become short 
aud cake-like as in fig. 3. Viewed along the edge, the colourless 
ectoderm was easily distinguished from the endoderm, the latter 
dark brown from the presence of numerous zooxanthelle. In 

Zi 


Ne, MR. J. E. DUERDEN ON 


surface view eight longitudinal darker and lighter areas were 
clearly marked, the coloured areas corresponding with the 
mesenterial chambers, and the colourless with the line of origin 
of the mesenteries. The mouth was already indicated at the 
broader end as a slightly depressed, small, circular aperture 5. 
the aboral end was narrow and rounded (Pl. 18. fig. 4). 

The perioral ectoderm differed from the remainder of the 
outer layer in being dark brown in colour, owing to the inclusion 
of zooxanthelle. Some of these apparently worked their way 
out from the ectodermal cells and adhered superficially, or, in 
other cases, became free. Their presence at various heights: 
in this restricted region is also revealed in sections. 

Contrary to the usual condition in the Actiniaria, the ciliation 
was uniform over the whole surface. The larve progressed with 
the narrower, aboral end foremost, and also attached themselves 
finally by this extremity. Even during the first day their power 
of adherence to the bottom and sides of the vessel was considerable, 
sufficient to resist a strong current of water from a pipette. 

The description given applies to each’ of the larve extruded, 
all having apparently reached a similar stage of development. 
The subsequent growth, so far as followed, was likewise the same 
in each. No earlier stage could be seen through the transparent 
tissues of the adult, and when a polyp was cut in halves none 
floated out. It is evident, then, that in Lebrunia, as appears to 
be also the rule in many other Zoantharia, the ova are ripened 
and the embryos develop and are expelled in batches ; for, were 
it otherwise, different stages would be represented within the 
gastro-ccelomic cavity at one and the same time. 

Such, however, is not always the case. While at Port Antonio, 
on the north-east coast of Jamaica, working in the temporary 
Marine Laboratory established there in 1897, by the Johns 
Hopkins University, specimens of Awlactinia stelloides, McMur., 
were collected, and from these larve were also being setfree. Here 
the individuals, each with an aboral tuft of longer, less mobile 
cilia, were thrown out from the internal cavity of the adult by 
the same jerking motion as in Lebrunia; but many different 
stages were represented in one parturition, and a like variety of 
phases could be observed through the partly transparent tissues. 
Minute planule, showing what seemed to be the beginning of 
an invagination, would appear along with larve possessed of eight,. 


THE HDWARDSIA-STAGE OF LEBRUNIA. 273 


strongly marked ridges and furrows, and even, in a few cases, 
with as many as six or eight tentacles already developed. Only 
the younger stages here continued to swim about; the older 
straightway fixed themselves to the bottom of the vessel by 
means of a flattened basal disc. 


Development of Tentacles. 


Within the first twenty-four hours the larve of Lebrunia 
settled down, and indications of the future tentacles were pre- 
sented by a slight bulging from each of the eight intermesenterial 
areas. These were at first rounded as in Pl. 18. fig. 4a, but, on 
becoming larger, narrowed a little as in figs. 5 & 6. 

From the first a distinction into two alternating groups of 
four was obvious, one series being slightly larger than the other, 
though all constituted only one cycle. 

Many of the adult polyps are characterized by the occurrence 
of superficial, opaque white areas on the oral aspect of the 
tentacles, much more pronounced later on the columnar out- 
growths. The opacity on the tentacles is found to appear at the 
earliest stages, at first on the four larger prominences and later 
on the four smaller. 

For some time it was not possible to recognize any difference 
in size amongst the members of any one group of tentacles, the 
dise presenting a perfect tetrameral radial symmetry. Later, 
one tentacle became slightly larger than the rest. Following the 
conventional nomenclature proposed by Kolliker and now usually 
adopted, this will be the dorsal tentacle ; or, in the more morpho- 
logical terminology of Haddon (1889), the sulcular. The tentacle 
opposite will be the ventral, or sulcar, and the plane in which 
they are enclosed the dorso-ventral or sulculo-sulear. In the 
further increase in size the dorso-lateral pair of tentacles, that is 
the tentacle on each side of the dorsal tentacle, remained a little 
behind the ventro-lateral pair—these four forming the smaller 
series in the earlier stage. 

The larva thus changed from its primary condition of radial 
symmetry to a decided bilateral form, the sagittal plane passing 
through the large dorsal and the ventral tentacles. 

At the expiration of two or three days the larve had reached 
the stage represented by fig. 7 (Pl. 18). The aboral end is now 
flattened and disc-like, and even slightly larger in diameter than 


O74: MR. J. E. DUERDEN ON 


the column; the latter has narrowed and become cylindrical, but 
still shows only eight alternating dark and light areas. The 
eight tentacles are finger-shaped, rounded at their free extremity, 
and able to move about, but, as in the adult, are incapable of 
retraction. The oral disc is flattened, its ectoderm still retains 
the zooxanthelle, and the mouth is circular or oval-shaped. 

The dorsal tentacle is a little stouter and longer than the 
others, and the originally smaller ventro-lateral tentacles are now 
practically of the same size as the three remaining members of 
the first series, 7. e., the ventral tentacle and the two median 
laterals. The dorso-laterals, however, have lagged considerably 
behind in their development. At their base they are also pushed 
out slightly further from the disc than the other six, which now 
are beginning to form a cycle by. themselves. 

The larve are clearly sensitive: to the action of ight. When 
settling they would select the under surface or a crevice in any 
fragment of coral-rock placed in the vessel, and if the fragment 
were turned over would detach themselves and move to some 
less exposed spot. The adults usually live associated in shady 
erevices, the pseudo-tentacles and. tentacles protruding as a 
delicate fringe. During the ordeal of examination under the 
strong tropical light, the larve would slowly assume a more or 
less collapsed aspect, their healthy inflated appearance returning 
after being shaded for some time. This behaviour towards light 
I find also to be very general amongst adult Actiniaria and 
Madreporaria. 

Several examples were kept alive for six or seven days, but 
none passed beyond the stage represented in fig. 7. No trace 
was presented of the pseudo-tentacles. The development of 
the larve after extrusion was at first very rapid, the stage last 
mentioned being reached within forty-eight hours. The only 
external alteration during the next four or five days was a slight 
increase in dimension of all the parts, without any indication of 
additional tentacles or mesenteries. Unfortunately, circumstances 
did not permit of further observation. 

The chief external characters, other than that of size, in which 
the adults differ from the oldest larve are: (1) the presence 
of. several cycles of entacmous tentacles, arranged in the 
formula 6, 6, 12, 24, 48, the members of any cycle being equal ; 
and (2) the six (normally) large branching “ pseudo-tentacles,’ 


THE EDWARDSIA-STAGE OF LEBRUNIA. 275 


arising from the most distal region of the column and bearing 
spheroidal batteries of nematocysts. The mouth, instead of 
being circular, is very long and narrow. 

From what we already know of the development of other 
Zoantharia, the hexameral character of the adult tentacles is in 
most cases derived from the readjustment of primary tentacles 
of different sizes and origin. Thus, already in the larve the first 
cycle of six is indicated, constituted of the four larger of the 
eight primary tentacles and two of the smaller. The remaining 
two will probably form one of the pairs of the second cycle of 
six tentacles alternating with the first, in which case it is obvious 
that the second cycle will be composed of individuals of a very 
different order of development. 

As regards the tentacles, then, Lebrwnia coralligens passes 
through three very distinct stages:—(1) a tetrameral radial 
symmetry, with four large and four small tentacles; (2) a bi- 
lateral symmetry of the same parts, six tentacles being nearly 
equal and forming one cycle, and two remaining smaller and 
outside the others; (3) the approximate hexameral radia: 
symmetry of the adult, with numerous alternating entacmeous 
cycles. 

The simultaneous appearance and tetrameral radial arrange- 
ment of the tentacles here described in the first stage is in 
marked contrast with what is known of the tentacular develop- 
ment of the Zoantharia generally. 

Comparatively few observations on the order of appearance ot 
the tentacles have, however, been recorded. One of the fullest 
and best is the well-known account given by Prof. de Lacaze- 
Duthiers of the growth of Actinia equina, and supplemented by 
observations on other species. Not having the original mono- 
graph for reference, for what follows I am dependent upon the 
many figures and details reproduced by Prof. H. L. Mark in his 
“Selections from Embryological Monographs” (1884), and by 
Prof. A. C. Haddon in the concluding portion of his “ Revision 
of the British Actinie ” (1889). From these it appears that the 
dorsal tentacle is the first to arise, and for a long time this 
predominates over the others. The octoradiate stage quickly 
supervenes, the median lateral tentacles being the next largest 
to the dorsal, and the ventral protuberance one of the smallest. 
The great difference in size between the dorsal and ventrai 


276 MR. J. E. DUERDEN ON 


tentacles gives a strongly bilateral character to the larva, ex- 
ceeding that ever exhibited by Lebrunza. The latter seems more 
closely to resemble Bunodes verrucosa (B. gemmacea), in which 
Lacaze-Duthiers found the octoradiate stage to be of longer 
duration. 

The succeeding stages, showing the manner in which from 
tentacles of heterogeneous origin the hexameral cycles of the 
adult are obtained, do not now concern us, except to say that in 
all probability Lebrunia passes through similar phases. 

Aulactinia larve afford somewhat similar results. I obtained 
freshly-extruded examples with only six tentacles developed, one 
smaller and one larger tentacle on each side of the dorso-ventral 
tentacles. Others, again, show eight tentacles ; of the new pair 
one arises on each side of the dorsal tentacle. 

Among the Cerianthide MeMurrich (1891 a) and E. van 
Beneden (1891), in the same year, have confirmed the previous 
observations of Agassiz and Kowalewsky, that the first six 
tentacles arise in lateral pairs, and that then a single median 
tentacle—the sulear—appears. No radial phase is ever assumed 
in any of these earliest stages yet deseribed. Prof. G. von Koch 
(1897, p. 759), however, in the numerous larve of Caryophyllia 
cyathus which he had under observation, found the first twelve 
tentacles to appear apparently at one and the same time, six 
larger alternating with six smaller, just as in the adult. In one 
case they developed in pairs, the dorso-ventral pair appearing 
first. 

The relation of the different tentacles to the internal meser- 
terial spaces is described later (p. 297). 


IIL. Internat Anatomy anv HistoLoey. 


In all about thirty extruded larve were obtained, and reared 
to different stages within the course of a week. Among a 
number of adults collected in 1896, at the Port Royal Cays, a 
polyp with its interior crowded with preserved larve was also 
detected. Cutting this across upwards of thirty examples 
floated out freely, or were easily separated from among the 
tissues. These were devoid of tentacles, and internally all pre- 
sented precisely the same stage of development—a stage, it will 
be seen later, slightly in advance of that exhibited by the 
youngest of the expelled larve. In speaking, therefore, of early 


THE EDWARDSIA-STAGE OF LEBRUNTA. 277 


and late larve, I refer to phases reached in the development, 
independently of whether the examples were or were not ex- 
truded. Practically all the material was utilized for purposes 
of study. 

Succinctly the general facts of the anatomy and histology are 
as follows :— 

The ectoderm is a thick layer with the constituent elenents— 
ciliated supporting cells, nematocysts in all stages of develop- 
ment, gland-cells, and nerve-elements—well differentiated. The 
supporting lamella contains isolated cells, and a definite, though 
very weak, musculature is already formed on its inner surface, 
its fibres arranged in a circular manner. 

Eight mesenteries are present and extend the whole length of 
the larva; in the upper stomodeal region they are all perfect 
and arranged tetramerally *. 

The stomodeum at first is a very short, strongly ciliated, 
rounded tube, without any external aperture. Internally it 
communicates with four radiating canals, which are connected 
with two sinuses, and these terminate in a system of inter- 


* Such a stage in which only eight perfect mesenteries are present, and in 
which no further numerical increase takes place for some time, has been found 
to be very constant in the development of the Zoantharia. The mesenteries 
are those of the first four pairs, and consist of two lateral pairs and two 
median pairs—the directives. The retractor muscles on the two lateral pairs 
occur on the face of the mesenteries turned towards one pair of the directives, 
which, following the terminology of Haddon, is the sulcar or ventral. The 
other pair of directives is the sulcular or dorsal, and in these the retractor 
muscles are likewise on the face directed towards the sulear aspect of the 
polyp. 

The pair of lateral mesenteries next the sulcar end—sulco-lateral—are the 
first to appear in the embryo; then, in the majority of cases, the lateral pair 
next the sulcular end—sulculo-lateral ; the sulear directives constitute the third 
pair, and the suleular the fourth in the order of development. 

Lacaze-Duthiers first recognized the octoradiate condition as one of the 
resting stages in the development of the Actiniz. Haddon (1889) described. it 
as the third stage of mesenterial development, at the same time realizing its 
phylogenetic importance and similarity to the permanent condition in the 

- Edwardsie ; McMurrich (1889) obtained it ‘in Awlactinia stelloides, and later 
in others, and applied the term “‘ Edwardsia-stage,” an apt designation since 
adopted by all writers on the subject. The researches of these authors, and of 
H. V. Wilson, Boveri, van Beneden, G. von Koch, and others, demonstrate that 
the phase is repeated in the ontogeny of such diverse groups as the Ceriantheze, 
Zoanthes, Hexactiniz, and the Madreporaria. 


278 MR. J. E. DUERDEN ON 


mesenterial and mesenterial spaces. The spaces throughout 
have a distinct ciliated limiting layer, in continuity with the 
lining of the stomodeum. Otherwise the whole of the interior 
is filled with a vacuolated, undifferentiated tissue, provided with 
nuclei and zooxanthelle, but devoid of yolk. Such a larva may 
therefore be regarded as a nearly solid mass of cells. 

Between the earliest larve and the latest, changes of great 
significance have been effected, in addition to those involved in 
the production of the tentacles. 

The middle vacuolated tissue has broken down and a gastro- 
coelomic cavity of the usual Celenterate type is in process of 
formation ; into this the stomodeum, now provided with an oral 
aperture, depends and opens directly. ‘The primary spaces are 
in process of giving rise to the mesenterial chambers of the 
adult. 

One pair of the original eight mesenteries—the sulco-lateral— 
remains connected with the stomodeum longer than the rest, and 
each of the pair bears mesenterial filaments in direct continuity 
with the stomodzal lining. Only the median streak is as yet 
developed. 

- Of the many expelled larve secured, only two or three retain 
the solid phase in its perfect condition, with the stomodeum and 
its diverticula intact. The remainder, including the specimens 
preserved within the body of the adult, have passed into the 
second stage in which the middle tissue is becoming disorganized. 

The different structures will now be described in greater 

detail. 
1. EHetoderm. 


The ectoderm is an exceptionally broad layer, uniformly ciliated 
throughout, with its free margin very pronounced in sections 
stained in borax-carmine or hematoxylin(Pl.18.fig.8). This latter 
character is due to the occurrence of a very delicate rectangular 
enlargement at the base of each cilium; the closeness of these and 
the readiness with which they take up colouring-matter produce 
a strong delimitation from the rest of the layer. The Hertwigs 
(1879, Taf. iii. figs. 5-8) figure a similar appearance at the edge 
of the ciliated ectoderm of the external surface and stomodeum 
of Anthea cereus. 

Most of the nuclei of the various cellular constituents are 
arranged in a broad zone a little within the periphery, but a few 


THE EDWARDSIA-STAGE OF LEBRUNIA. 279 


appear in the deeper regions and as far as the mesoglea. A 
peripheral zone of the ectoderm is nearly colourless, being con- 
stituted mainly of the distal portions of the extremely long 
supporting cells, unicellular glands with clear contents, and a 
few gland-cells with fine granular contents. These last stand 
out very distinctly in sections stained with indigo-carmine. The 
nuclei of the principal zone are oval and closely arranged, stain- 
ing not nearly so intensely as those of the stomodeum and 
mesenterial filaments. The inner nuclei are rounded and much 
more scattered ; some approach the supporting lamella, and others 
appear partly or even entirely included within it. The meso- 
gloea of the adult polyps contains numerous isolated cells equally 
distributed throughout, and there is little doubt that these are 
derived both from the ectoderm and the endoderm. For in the 
larve nuclei in similar positions to those of the ectoderm already 
referred to also appear on the internal side, and the thin lamella 
of the mesenteries also displays included cells. Both in longi- 
tudinal and transverse sections a very delicate layer of nerve-fibrils 
can be distinguished next the mesogloea, much more obvious at the 
narrower end (PI. 19. fig. 21). Maceration-preparations of such 
delicate material in the preserved condition did not yield good 
results, but I was able in some instances to distinguish the usual 
nerve-fibrils of the sensory cells. 

No distinct evidence of muscular fibrils occurs on the ecto- 
dermal side, but in longitudinal sections such can be recognized 
on the inner border of the mesoglcea of all but the youngest 
larvee (fig. 21). 

The nematocysts are of two kinds. A long narrow form 
extends almost across the layer and is distributed throughout 
the ectoderm, even as far as the aboral region. They parti- 
cularly crowd the tentacular area. On maceration, a very 
small oval form is also obtained, often with the nucleus of 
the cnidoblast still connected. These are not early stages of 
the larger variety, as the developing forms of these latter are 
very conspicuous objects. 

The large nematocysts are extremely thin-walled, nearly trans- 
parent, and usually only a central, nearly straight, more highly 
refractive axis can be distinguished, extending from one end to 
near the other. In some conditions the presence of a very 
delicate, closely spiral thread can be observed in addition. Nearly 


280 MR. J. E. DUERDEN ON 


all my preparations reveal the curious fact that, in the process 
of sectionizing, many of the nematocysts become dislodged from 
the ectoderm, and dragged to what would be the margin of the 
paraffin block, without causing any apparent injury to the other 
elements. 

Here and there throughout the layer, more numerously in 
the tentacular region, occur nematocysts in various stages of 
development. They are at once recognized by their homogeneous 
and deeply-staining character. In this respect they contrast 
strongly with the mature forms, the walls of which are not 
affected by borax-carmine, though they become slightly blue in 
indigo-carmine. Most of the nematocysts originate in the deeper 
parts of the layer, and, at first, are a little concave or irregular 
in outline, and arranged obliquely or even tangentially to the 
_ surface. When mature they extend as far as the external sur- 
face and are disposed vertically. Both the wall and the axis 
take up the stain in the nearly ripe condition, and in trans- 
verse sections of earlier stages the contents are homogeneous in 
the middle but granular towards the margin. 

Around the aboral end the ectoderm is of a very different cha- 
racter from elsewhere (Pl. 19. fig. 21). The periphery is crowded 
with small nematocysts, provided with a very distinct spiral 
thread ; large, pyriform, deeply-staining bodies, perhaps glan- 
dular, occur more internally, and towards the supporting lamella 
the nervous layer is exceptionally well-developed. Such a special 
development of the protective, sensory, and probably glandular 
elements of the layer is, no doubt, in some way associated with 
the fact that in the free active stage this end of the larva is 
foremost, and that it is by this that it ultimately attaches 
itself. McMurrich (1891, p. 317, pl. xiii. fig. 18) describes and 
figures a somewhat similar exaggerated development of the 
nervous layer in the embryo of Rhodactis. 

As already mentioned amongst the external characters, and 
contrary to the usual condition in adult Zoantharia, the ectoderm 
contains zooxanthelle, restricted, however, to the central area 
of the oral dise. In the oldest larve they occur throughout the 
thickness of the discal ectoderm, but none remain in the adult 
polyps. 

Compared in other respects with the ectoderm of the adult, 
there is a great increase in the latter in the proportion of uni- 


THE EDWARDSIA-STAGE OF LEBRUNTIA. 281 


cellular mucous glands with nearly clear contents; cilia are not 
obvious in material preserved in the same manner and at the 
same time as the larvee; and practically no increase in thickness 
of the layer has taken place. A weak ectodermal musculature 
occurs on the tentacles, oral disc, and uppermost part of the 
adult column. 

The nematocysts of the columnar outgrowths (pseudotentacles) 
in the adult are of a different kind from those already described, 
being oval and much shorter, with a loose, irregular spiral 
thread. 


2. The Archenteron and Formation of the Esophaqus. 


Tt is remarkable that in all the larve taken from the preserved 
adult there is no external aperture (Pl. 19. fig. 21). This is also 
the case with the extruded larve at the time they are set free 
(fig. 15), but a mouth is established very early after as a result 
of an infolding of the discal ectoderm. From this it would seem 
that the structure in Lebrunia, hitherto spoken of as the 
stomodeum, must, prior to its communication with the exterior, 
be regarded as an endodermal-lined archenteron, for there are 
important morphological differences between its earliest and 
latest conditions. 

If the discovery of earlier stages should prove that at the 
blastula-stage of development a primary archenteric invagination 
is established, then its external opening—the blastopore—has 
become closed, and only later another aperture is produced, over 
evidently the same spot, and this persists as the mouth. The 
entire tube thus formed, partly from the archenteron and partly 
from the secondary invagination, persists as the cesophagus of 
the adult. 

Fortunately, a fairly complete series of preparations has been 
obtained showing the different stages in this latter transforma- 
tion. In the non-extruded larve (fig. 21) the blind archenteron is 
easily distinguished by means of its deeply-staining lining ; 
outwardly it extends as far as the external ectoderm, its narrow 
lumen recognizable all the way. The supporting lamella is seen 
to be invaginated for a considerable distance, but does not extend 
transversely. The ectodermal cells immediately over the arch- 
enteron are rather loosely arranged, and zooxanthelle occur 
amongst them. It appears as if all that is needed to bring about 


282 MR. J. E. DUERDEN ON 


the adult condition is for the archenteron to extend itself a 
little, and thus set up a communication with the exterior. 

Vertical sections of one of the freshly-expelled larve prove 
that this is apparently not the case, but that to form the adult 
oral aperture and cesophagus a distinct invagination of the super- 
ficial ectoderm takes place. This is very distinctly shown in the 
actual sections represented in Pl. 18. figs. 9 to 11, taken from a 
series of vertical sections through the archenteric region of an 
extruded larva, before disintegration of the central tissue has 
commenced. In the particular larva the transverse, separating 
tissue of the invagination has evidently just broken through, 
establishing communication, but the lateral walls still mdicate 
how the whole process has been carried out. 

Fig. 9 is from a tangential section through the wall of the 
archenteron and the invaginated region above. The ectoderm 
directed inwardly differs much in character from that beyond ; 
zooxanthelle are absent, and a greater number of deeply-staining 
nuclei cause it to stand out very distinctly. A straight, ciliated 
edge indicates the free surface of the floor of the ectoderm ; 
while a middle region below, nearly devoid of nuclei, represents 
where the lower surface of the invaginated end has come into 
contact with the expanded upper end of the archenteron. 

The free margin of the wall of the latter, bordering on the 
flattened archenteric or coelomic chamber, is also very strongly. 
ciliated. The floor of this chamber is at this stage entire and 
sharply defined; the limiting layer is ciliated and contains 
abundant nuclei, which easily distinguish it from the vacuolated 
tissue below and around. To the right the floor is continued 
further into the diverticular system (cf. left side of fig. 15, Pl. 19). 

Fig. 10 (P1.18) is from another tangential section, but nearer the 
lumen than the former. An external aperture is now presented, 
the outer ectoderm being seen as an inturned layer. The floor 
of the invagination is dome-shaped and extends some distance 
laterally. The area of its union with the roof of the archenteric 
cavity can even yet be distinguished owing to its diminished 
number of nuclei. The central coelomic space is still flattened, 
but the preparation does not permit of the lateral extensions 
being followed. 

Fig. 11 represents a section through the actual lumen. The 
central tissue, consisting of the floor of the ectodermal invagina- 


THE EDWARDSIA-STAGE OF LEBRUNIA. 283 


tion and the roof of the archenteron, is now absorbed or broken 
down. The floor of the invagination is still represented laterally 
by the very marked indentation on each side, the actual point of 
union of the two layers being in the neighbourhood of the lower 
edge. Neither in this section nor in any of the later stages can 
the actual union of the invaginated ectoderm and the archenteric 
endoderm be recognized by any histological difference. The 
lumen throughout is now much wider than hitherto, but soon 
narrows again. 

In all the other extruded polyps of which I possess vertical 
sections the stage reached is later than the above; the stomo- 
dzum has straight walls, and uo indication is afforded of any 
double origin of its lining. 

Strictly, then, only the upper, ectodermal-lined region of the 
gastric funnel in Lebrunia is homologous with the stomodsum 
or fore-gut of the Enterocela; the lower region, lined with 
archenteric endoderm, is equivalent to a portion of the mesen- 
teron or mid-gut of the higher Metazoa. While such conditions 
as those revealed in fig. 21 (Pl. 19), where the archenteric endo- 
derm is in absolute continuity with the filamental tissue, strongly 
suggest that the mesenterial filaments along the free edge of the 
mesenteries are but the banded continuations of the enteron. 

On account of the seeming double origin of the gullet in 
Lebrunia, the term csophagus is preferable to the term stomo- 
deum usually employed in Actinozoan literature for this structure. 
The former, as used in the Hnteroceela, implies merely the first 
portion of the adult gastric canal, without any reference to the 
embryonic origin of its lining, though usually it possesses both 
an ectodermal and an endodermal portion ; while the latter is 
the term employed to designate the extent of the invaginated 
ectodermal lining of the gastric canal. The justification for the 
employment of stomodzum in the adult Actinozoon rests upon 
the fact that, in general, the lining of the gullet is wholly 
derived from a primary ectodermal inturning, but it is obvious, 
from the details just given, that this may not always be the 
case. 

We may thus summarize the formation of the oral aperture 
and cesophagus:—An invagination of the external ectoderm 
takes place directly over the outer extremity of the archenteric 
tube, and at the same time the layer undergoes a considerable 


284: MR. J. E. DUERDEN ON 


histological alteration. Its floor presses upon the blind end of the 
archenteron, and the result is a mutual flattening, leading to a 
convexity in the former and a considerable lateral extension in 
each. The two ends ultimately break down in the middle, and a 
communication between the interior and exterior is thereby 
established. The lateral walls of the invaginated ectoderm and 
of the archenteric endoderm form the lining of the adult 
cesophagus. 

Once the esophagus is fully established, the lumen at first is 
extremely small, and circular in transverse section. Hven in the 
older larvee the outline may be circular in some places, but oval 
in others, the longer axis always corresponding with the sulco- 
sulcular axis of the mesenteries. No histological change sug- 
gestive of a gonidial groove ever takes place at either end, a 
condition not to be expected, as gonidial grooves are not present 
in the adult. 

The lining is more strongly ciliated than that of the columnar 
ectoderm ; indeed, it is the most strongly ciliated layer of the 
whole larva. As in the column a small, deeply-staining enlarge- 
ment occurs at the base of each cilium. 

The nuclear zone is very pronounced ; the nuclei are oval, 
closely arranged, and stain a brilliant red in carmine. Clear 
gland-cells appear to be absent, but granular gland-cells occur 
here and there, and long, narrow nematccysts similar to those of 
the column. The mesogloea is extremely thin. Only in some of 
the most favourable sections of the older larve can suggestions 
of a nervous layer be observed, though this is very pronounced 
in the adult. 

Aborally the cesophagus opens directly into the gastro-ccelomie 
cavity. In perfectly horizontal sections the sulcar and sulcular 
ends terminate at about the same level. At the inner extremity 
a slight reflection of the strongly-nucleated lining takes place all 
round, so that in transverse sections at this point the latter 
appears cut through twice (PI. 19. fig. 18). The reflected portion 
is more or less connected with the mesenterial filaments at their 
origin; it is further discussed under the section Mesenterial 
Filaments. 

In the adult polyp the esophagus is greatly elongated trans- 
versely, so that it stretches nearly across the ccelenteron, the 
directive mesenteries being much shorter than the four lateral 


THE EDWARDSIA-STAGE OF LEBRUNIA. 285 


pairs. Gonidial grooves are absent. An exceptionally well- 
developed nervous layer occurs, without any accompaniment of 
muscular fibres, and granular gland-cells are numerous. 


8. Larval Celomic Spaces and Formation of the Gastro- 
caclomic Cavity. 


Transverse sections through the upper archenteric region of 
the youngest larve, before any disorganization of the internal 
tissue has commenced, reveal very narrow, isolated, slit-like spaces 
within the intermesenterial, parenchymatous tissue, one near 
the middle of each face of a mesentery (Pl. 18. fig. 12, d,e,f). A 
little lower the two within each chamber become connected by a 
transverse channel, and still lower only a single median cavity 
occurs in each radial division (fig. 12, 9, 2, c), the lateral boun- 
daries always remaining close to the face of the two enclosing 
mesenteries. 

Approaching the inner termination of the archenteron, where 
three pairs of the mesenteries have become free, each cavity 
broadens somewhat, and communication is established between 
the five chambers on the dorsal or sulcular side of the sulco- 
jateral mesenteries, and also between the three on the ventral or 
sulcar side, two distinct sinuses being thus formed (fig. 13). The 
successive steps in the sinus formation, following the disappear- 
ance of the centripetal parts of the mesenteries, are shown in 
fig. 12, in connection with the intermesenterial spaces 6, a, h. 
In this particular instance the sulcar mesenteries are the first 
to lose their connection with the archenteron and permit of 
communication. 

Fig. 18 represents the arrangement of the various cavities at 
the inner end of the archenteron. It delineates the details 
observed in several sections of a polyp which happened to be 
eut slightly obliquely to the vertical axis. The archenteric tube 
terminates in four canals, one of which communicates with the 
sulcar sinus and three with the sulcular smus. Were the sulco- 
lateral pair of mesenteries developed only to the same extent as 
the others, it is obvious that a complete circumferential sinus 
would be established. 

For a short distance below, the two sinuses remain distinct, 
but later become sub-divided into eight V-shaped chambers, 
associated with the free edges of the mesenteries. 

LINN. JOURN.—ZOOLOGY, VOL. XXVII. 22 


286 MR. J. E. DUERDEN ON 


The various stages leading to this are represented in fig. 14, 
taken from a single oblique section in which the upper part is 
more aboral than the lower. Traces of the two sinuses are still 
seen in the latter region, and on the right side the last evidence 
of the sulcar canal; while, in the upper half of the figure, the 
V-shaped canals, embracing the free edge of the mesenteries, are 
distinct. This latter condition holds for all the mesenteries as 
far as their lower termination. 

A vertical section, such as would be obtained along the line 
z-y in fig. 12, is represented in fig. 15 (Pl. 19), taken from one of 
the youngest larve. The archenteron opens into a shallow central 
chamber, from which two canals radiate, one to the left and the 
other tothe right. Histologically the floor of the chamber differs 
considerably from the undifferentiated tissue below, more nearly 
resembling the endodermal lining of the archenteron. Vertical 
sections of one early larva show this chambered region very 
distinctly ; some (figs. 10, 11) with the right side terminating 
blindly, and others (fig. 9) with a canal-like continuation. The 
precise outline of the diverticulum to the left in fig. 15 is largely 
deduced from the various transverse sections, as only isolated 
parts are recognizable in the actual vertical preparations. 

The boundary of all the canals and spaces is regular and 
smooth, and cilia can be distinguished, not, however, so thickly 
disposed as in the archenteron. Hverywhere at its origin the 
limiting layer is in continuity with the archenteric lining, and 
throughout bears a close resemblance to it. The cells constituting 
it are more or less separable from the rounded, vacuolated cells 
below, being columnar and filled with finely granular, protoplasmic 
contents ; oval nuclei are much more numerous than elsewhere, 
and are arranged in a somewhat regular layer. 

Throughout the best-preserved larve the margins stand out 
prominently in sections stained in borax-carmine, owing to the 
greater abundance of nuclei which readily take up the stain. 

In sections the vacuolated tissue, both central and peripheral, 
most closely recalls the parenchymatous tissue of plants. The 
cells are somewhat polygonal in outline, their boundaries being 
extremely thin, and the contents have almost entirely disappeared. 
Here and there a nucleus occurs applied to the walls, and zoo- 
xanthelle are distributed throughout. No objects at all sugges- 
tive of yolk-granules are present in any of the stages. The tissue 


THE EDWARDSIA-STAGE OF LEBRUNIA. 287 


probably shares in the formation of the muscular system, for in 
connection with it delicate fibrils are recognizable on the column- 
wall (fig. 21) and mesenteries (fig. 16). 

The limiting cells next the supporting lamella of the column- 
wall are not arranged in any definite layer, and appear to differ 
in no respect from those more central. 

Nuclei sometimes occur here more abundantly than else- 
where, and often in a similar manner along the two faces of the 
mesenteries. They are probably connected with the formation of 
the mesoglea, and some seem to become included within it. 

To recapitulate: The archenteron terminates internally in a 
small central cavity, from which four canals radiate and open 
into two circumferential sinuses, extending slightly above and 
below. In the middle archenteric region of the larve the sinuses 
subdivide into eight distinct chambers, one for each radial 
division, and ultimately each of these terminates in two cornua, 
sixteen in all. Below the archenteric region the sinuses terminate 
im eight V-shaped canals, each of which is formed, as it were, by 
the union of two cornua from adjacent mesenterial spaces. 
Owing to this the lower terminal chambers are mesenterial in 
position, while the upper must necessarily be intermesenterial. 
A ciliated limiting layer closely resembling in structure, and in 
absolute continuity with, the archenteric endoderm is throughout 
easily distinguished from the general mass of undifferentiated 
tissue. This system of spaces I regard as a larval ccelome or 
body-cavity arising in connection with an archenteron, but not 
cut off from it. 

Instead of representing a definite and independent system of 
larval coelomic diverticula, it may be urged that the spaces are 
merely the initial clefts in the disorganization of the solid interior, 
to be completed later. Against this I would point out that the 
canals and spaces all exhibit the distinct limiting layer, very 
different from anything to be associated with disintegrating 
tissue; and that in the intermediate larve, where the process 
can be followed, the breaking down of the cells commences just 
below the internal opening of the archenteron, attacking first 
the central mass. It is not until very late that any alteration takes 
place in the more proximal and distal spaces. And further, there 
is evidence that the greater part of the limiting layer becomes the 
unilaminar epithelium of the adult cavity. Itis not disintegrated. 

22* 


288 MR. J. E. DUERDEN ON 


The internal appearances of the various regions of a later 
larva, one in which the disorganization of the central tissue is in 
progress, though the mouth is not formed, are represented in 
PI.19. figs. 16-21. The first five are taken from actual sections of . 
one of the non-extruded polyps, while fig. 21 is constructed so as 
to represent what would be obtained along the lines I-a, w-I in 
fig. 19. As a matter of fact, however, fig. 21 was obtained 
almost entirely from a single vertical section. The sections from 
which figs. 16-20 are taken would be along the lines 1-1, 2-2, 
3-3, 4-4, 5-5 respectively of fig. 21. 

Fig. 16 passes through the upper region of the archenteron, 
just below the sections showing the sixteen separate slit-like 
cornua. The spaces are very regular and crescent-shaped. 
Fig. 17 approaches the inner termination of the archenteron, 
only the sulco-lateral pair of mesenteries being still attached to 
the latter. The sulcular and sulcar sinuses are established) but 
neither here nor lower is there any trace of the archenteron 
dividing into canals. Four regions are indicated on its coolomie 
epithelium, in which the deeply-staining tissue in continuity with 
the archenteric lining is becoming concentrated. The two in 
the sulear sinus are distinct, but the sulcular pair are nearly 
united, though more distinct in sections above. 

The appearance of the terminal walls of the tube alone is 
represented in fig. 18. The deeply-staining endoderm is cut 
through twice as a result of its reflection (cf. fig. 21, 3—8). 
The reflected moiety is now separated into two parts by the con- 
nection of the sulco-lateral mesenteries, which are seen to extend 
as far as the actual termination. The same tissue, developed to 
a much less degree, can also be traced along the margin of the 
mesenterial epithelium. 

The next figure (fig. 19) is from a section below the archenteron, 
and shows the larger, filament-bearing, sulco-lateral mesenteries, 
and relics of the disorganized central tissue, in the gastro- 
celomic cavity now in progress of formation. Comparing this 
with figures 9-11 of the earlier stage, it will be seen that the 
portion to disintegrate is that central to the two sinuses, including 
both the limiting layer and the enclosed mass of parenchymatous 
tissue. 

The peripheral portion has not yet commenced the changes 
resulting in the unilaminar epithelium of the adult; it is still 


THE EDWARDSIA-STAGE OF LEBRUNIA. 289 


ae 


many cells in thickness, and its margin is strongly ciliated and 
nucleated. 

In sections for some distance below the middle of the larva the 
mesenteries are all of the same size, and there is a large central 
cavity, but towards the aboral region the cavity is again nearly 
obliterated. he details here represented indicate the stages 
followed during the progress of disintegration. The vertical 
section in fig. 21 will assist in making the relations described 
a little clearer. 

From these details it is evident that the only actual dis- 
integration as yet is that involving the central tissue internal to 
the two circumferential sinuses, and extending to the aboral 
region and some distance within the esophageal region. In the 
expelled larve the process, like that of the formation of the oral 
aperture, was set up immediately after liberation, and must 
have proceeded rapidly; for in only three specimens was I 
abie to obtain the stage in which the centre is still solid and the 
diverticula distinct throughout. On the other hand, it is well on 
the way in the non-extruded larve, which are still devoid of any 
external opening. 

Remains of the primary solid condition continue to be repre- 
sented for some time peripherally, by an enormously thickened 
ridge of parenchymatous tissue, connected with the columnar 
boundary of each radial chamber, and also by the ridge of similar 
tissue on each face of the first pair of mesenteries (fig. 19). 
And in none of the polyps has the disorganization proceeded so 
far that the central cavity extends to the aboral region ; but the 
process has progressed further at the oral extremity, whence the 
hollow tentacles arise. 

All the preparations reveal, from the beginning, only a thin 
epithelium on the three pairs of shorter mesenteries, except 
towards their connection with the column and with the archen- 
teron, where the layer joins that of the column-wall. 

As growth continues the columnar intermesenterial ridges 
must also become further diminished until the unilaminar epi- 
thelium characteristic of the adult polyp is obtained throughout, 
but the larve only present stages towards such a condition. 
There is every indication that it is the limiting layer of the larval 
spaces, and not the vacuolated tissue, which persists and con- 


290 MR. J. E. DUERDEN ON 


stitutes the lining of the adult cavity. In fig. 19, e, f, g, 2, the 
limiting-layer appears, as it were, creeping round the ridges stall 
remaining, and thus separating them for disintegration. 

The original colomic spaces in connection with each radial 
division, whether above or below the cesophageal region, enlarge 
as the vacuolated tissue disappears and as the polyps increase in 
size, and ultimately become the endocceles and exoceles of the 
adult. These latter are thus shown to have their origin 12 
primary spaces connected with an archenteron, exactly as occurs 
in the formation of the celome in the higher Metazoa, except 
that partial disorganization supervening, the spaces are never 
cut off from the central chamber. 

In all the larvee free. zooxanthelle and fragments of the dis- 
organized tissue occur in the gastro-ccelomic cavity. It seems 
likely that these are ultimately expelled through the mouth 
of the polyp. I did not observe such in Lebrunia; but in 
Aulactinia and other embryos of about the same stage I 
have watched extrusions of this character going on from the 
interior ; small irregular masses of mucus-like matter, mixed 
with what seemed to be yolk-particles and zooxanthelle, would 
at times be passed out through the oral aperture. 

The mesenterial filaments along the free edge of the mesen- 
teries probably represent a divided digestive tube. In which 
case the space central to the mesenteries, the result of the dis- 
integration, will be gastric, and the peripheral portion of the in- 
ternal cavity will be coelomic. Hence the term “ gastro-coelomic ” 
more nearly expresses the true morphological conception of the 
whole of the internal cavity of the Scyphozoa than either 
“‘colenteron” or “ gastro-vascular cavity.” 

It follows that the entire imperfectly-chambered internal 
cavity of the adult Lebrunia is ontogenetically both coelomic and 
gastric, and is a secondary formation having its origin in two 
very different phenomena :—Firstly, in a primary system of 
radiating, archenteric diverticula or ccelome spaces ; and 
secondly, in the disintegration of a primary, undifferentiated 
tissue. The former gives rise to the mesenterial chambers, 
distinct from one another in the esophageal region, but im- 
pertect below; the latter gives origin to the space (gastric) 
central to the free edge of the mesenteries, and results in 
the imperfect character of the mesenterial chambers below the 


THE EDWARDSIA-STAGE OF LEBRUNIA. 291 


cesophagus. The greater part of the limiting layer of the spaces 
persists as the unilaminar epithelium of the gastro-ccelomic 
cavity of the adult. 


4. Mesenteries. 


Hight mesenteries are present in each stage examined, all 
perfect in the upper cesophageal region ; but only two, the sulco- 
lateral pair, remain united as far as the lower edge of the cso- 
phagus. All may extend to the aboral termination of the 
internal cavity, but the sulcular directives sometimes cease a 
little in advance of the others. The four pairs subdivide the 
upper cesophageal region into eight chambers, four larger alter- 
nating with four slightly smaller, arranged so ag to present a 
perfectly tetrameral, radial symmetry (Pl. 19. fig. 16), in corre- 
spondence with that of the tentacles on their first appearance. As 
the three pairs become free the transverse sections assume a 
distinctly bilateral symmetry. In their free condition the sulco- 
laterals continue for a short distance still larger than the rest, 
but below all the eight are practically equal, and very short ix 
their centripetal extent. 

It is remarkable that no increase in the number of mesenteries 
takes place between the earliest and the latest larvee, represent- 
ing a period of about five days. This is in harmony with the 
numerical condition of the tentacles, but the development of 
the mesenteries precedes that of the tentacles in all knowr 
Cases. 

In some examples the two. sulcar directives, in the proximal 
region, become united along their free edges. 

The mesogloea of the mesenteries is usually broad at its origin 
in the column-wall, but is extremely thin beyond. In the 
earliest stages traces of muscular elements are indicated only in 
the basilar region, but later, transverse sections of very delicate 
longitudinal fibrils are seen along one face, just sufficiently well- 
developed to enable the paired arrangement of the mesenteries 
to be established (fig. 16). This follows the usual order, which 
is the same as that in Hdwardsia, namely, the retractor muscles 
on the directives are on the faces of the mesenteries turned away 
from one another, while in the two lateral pairs they are on the 
faces turned towards one of the pair of directives which, by this 
means, if by no other, can be distinguished as the sulcar 
directives. 


292 MR. J. E. DUERDEN ON 


5. Mesenterial Filaments. 


Mesenterial filaments are apparently undeveloped in the 
youngest larve, in which the whole of the vacuolated tissue is 
intact. In the earliest stages, however, in which disorganization 
is in progress, a filament occurs on the first pair of mesenteries, 
the sulco-lateral. These mesenteries, as already described, 
remain connected with the cesophagus as far as its lower termi- 
nation, are much larger than the three other pairs, and are very 
prominent objects in transverse sections below the cesophagus 
(EI 9: figs. 19; Zi): 

Of the filament only the middle Driisenstreif or glandular 
streak is yet formed. This structure stands out very conspi- 
cuously in sections stained in carmine, by reason of the deeply- 
staining character of its numerous close, oval nuclei, while the 
free surface is also strongly ciliated. Long, narrow nematocysts, 
similar 10 those in the superficial ectoderm and cesophageal 
lining, occur here and there in different stages of development, 
and occasionally granular gland-cells can be distinguished. 

The filamental tissue extends for some distance down the edge 
of the mesenteries, but ceases before the aboral termination 
is reached. Although probably a matter of no particular signifi- 
cance, the filament during these early stages is found to be 
rarely equally developed on each of the two mesenteries, and 
generally terminates on one at a level different from that of the 
other. 

In one larva sectionized transversely, the sulcular and sulculo- 
lateral meseunteries, in addition to the sulco-laterals, remain 
connected with the cesophagus as far as its aboral ending, and 
throughout their middle free course are proportionately larger 
than in other larve and also than the sulcar directives. The 
epithelium on each face is swollen a little, and towards the free 
edge stains more intensely than elsewhere, indicating that 
mesenterial filaments are in process of development. 

While in the oldest extruded larve, and in all the non-extruded 
specimens (fig. 21), the filamental tissue isin absolute continuity 
with the similarly deeply-staining lining of the cesophagus or 
archenteron, a break occurs in earlier stages of the first series, 
where disintegration has just been set up. 

A distinct bridge of unmodified tissue is seen to intervene 
between the cesophageal lining and the portion of the filament 


THE EDWARDSIA-STAGE OF LEBRUNIA. 298 


already developed below. ‘This is clearly shown in fig. 22, 
representing the centripetal, swollen portion of the sulco-lateral 
pair of mesenteries of an extruded larva of intermediate age. 
The section is taken transversely a little below the gullet. At 
this level the mesentery to the left presents the beginnings of a 
filament at its free edge, easily distinguished by the oval, 
deeply-staining, closely-arranged nuclei and fringe of cilia; the 
mesentery to the right possesses only unmodified epithelium. 
Only a few sections lower, however, the filament begins to appear 
on the right mesentery also, and still lower 1s nearly as strongly 
developed on both, as shown in fig. 19, taken from a non-extruded 
larva. Such a separation between the filaments and cesophageal 
lining occurs in several larve of which I possess transverse 
sections, and also in one longitudinal series. 

Any hiatus, however, is of a very temporary character. For 
in other early stages the filamental tissue already starts directly 
from the point at which the mesentery severs its connection with 
the cesophagus, at first very feebly developed, but becoming 
stronger below. Later, it is fully developed along the whole of 
its extent. 

In this connection the outward and upward continuation of 
the archenteric or cesophageal lining which takes place at the 
aboral termination of the cesophagus must be referred to (figs. 17, 
18, 21). Prof. H. V. Wilson (1888) found very pronounced 
reflections in the early stages of the development of the coral 
Manicina areolata. These push away the endoderm of the 
celenteric surface of the stomodsum, and Wilson considers 
them to be there concerned with the formation of the filaments, 
except those belonging to the first pair of mesenteries. These 
originated as direct downgrowths along the column-wall, to the 
mesogloea of which the stomodum at an early stage is apposed. 

Prof. McMurrich (1891, p. 320, pl. xiii. fig. 17) also describes 
and figures a reflection at one end of the stomodzum of the larva 
of Rhodactis Sancti-Thome. 

In several cases I obtain an appearance closely resembling 
that given by McMurrich ; and on following it down, section by 
section, the deeply-staining tissue is seen to enlarge until ulti- 
mately it extends all round the esophagus, with the exception of 
the two points from which the first pair of mesenteries are giver 
off (fig. 18). Atthese it passes along the edge of the mesenteries. 


294. MR. J. E. DUERDEN ON 


When the sections are perfectly horizontal the reflection at 
first is equally developed all round, except at the two points 
mentioned, and is continued above in four distinct bands (fig. 17). 
The extent of the reflection in the non-extruded larve (fig. 21) 
should be compared with the condition before and after disinte- 
gration in the others (figs. 11, 22). 

The sections of Lebrunia prove that such a condition as that 
figured by McMurrich—the stomodzum open at one end and 
the lips partly turned back, and with a free portion of the 
lining reflected outside the opposite end—is probably due to an 
obliquity in the sections to the vertical axis of the larva, but 
it would, of course, also be brought about if the stomodeum 
terminated at one end earlier than at the other. 

The reflection has the significance of constituting the point of 
continuity between the esophageal lining and the mesenterial 
filaments, and occurs also in the adult condition of most 
Zoantharia. 

At the present time one of the most important problems in 
Actinozoan morphology concerns the origin, from one or other of 
the two embryonic layers, of the mesenterial filaments. The 
facts observed in every case are as follows :—Histologically the 
lining of the cesophagus very closely, though perhaps never com- 
pletely, resembles the mesenterial filaments; and in the perfect 
mesenteries of all adult polyps the two are in absolute continuity. 
In so far as the filaments resemble the stomodeal lining do they 
differ from the ordinary epithelium of the gastro-ccelomic cavity ; 
everywhere they offer a very strong histological contrast to the 
latter. 

Embryological evidence is here of the greatest value. From 
this standpoint the question has been approached by Prof. H. B. 
Wilson (1884) for the Alcyonaria, by Prof. H. V. Wilson for 
the Madreporaria (1888), and by Prot. McMurrich (1891) and 
others for the Actiniaria. In his paper on “The Mesenterial 
Filaments of the Alcyonaria,” the first mentioned investigator 
arrives at the conclusion that the filaments of the two dorsal 
mesenteries in that group are ectodermal derivatives, that is, 
downgrowths of the lining of the stomodum, and are homologous 
with the Flimmerstreifen or ciliated streaks of the Actiniaria; 
while the six ventral filaments are endodermal in origin, and 
correspond with the Driisenstreifen of the Actiniarian filament. 


THE EDWARDSIA-STAGE OF LEBRUNIA. 295 


The former H. B. Wilson proves to be specially circulatory in 
function, while the latter are digestive. Studying the West- 
Indian coral Manicina, in which only the Driisenstreif is de- 
veluped, H. V. Wilson found that the filament of the first pair of 
mesenteries arises as a downgrowth of the stomodeal ectoderm, 
and the later ones from upward reflections of the same layer. 
He homologizes the simple Madreporarian filament with the whole 
of the trilobed Actinian filament. From his researches on Awlac- 
tinia and Rhodactis McMurrich is inclined to agree with EH. B. 
Wilson, and to regard the Driisenstreif as endodermal and the 
Flimmerstreifen as ectodermal in origin. The sections of Aulac- 
timia revealed a bridge of unmodified endodermal tissue between 
the early filament and the stomodeal ectoderm, but the results 
with Rhodactis were not so conclusive, although representing 
earlier stages. 

In this strictly limited aspect of the problem the evidence 
from Lebrunia at first sight appears incontrovertible. In having 
the four pairs of mesenteries already developéd, alone with a 
nearly solid interior, the larve present conditions very different 
from those studied by the previous investigators. As already 
described, no filament is discernible before the gastro-vascular 
cavity of the adult has begun to be established. And for the 
next early larve, the first portion of the free mesentery in the 
case of the sulco-lateral pair of mesenteries possesses only un- 
modified endoderm at its free edge, while a filament is well 
developed below. It is on evidence precisely of this character 
that E. B. Wilson and McMurrich affirm the endodermal origin, 
in the former case, of the six ventral filaments of the Aleyonaria, 
and, in the latter, of the median streak of the Actiniaria. 

Regarding as I do the lower region of the cesophagus in 
Lebrunia as a portion of the archenteron and therefore its lining 
as endoderm, the significance of its primary relationship with the 
mesenterial filaments becomes altered from that understood by 
the writers just mentioned. 

From such conditions as those revealed in fig. 21, before any 
ectodermal invagination appears, it is clear that the archenterie 
endoderm and the filamental tissue are morphologically one and 
the same. 

I venture to think that in Lebrunia the temporary discon- 
tinuity in the extruded larve between the cesophageal lining and 


296 MR. J. E. DUERDEN ON 


the developing filaments is a condition of no morphological im- 
portance, perhaps due only to the readjustment of the relations 
between the cesophagus and mesenteries consequent upon the 
disorganization of the central tissue and the formation of the oral 
aperture. Any subsequent discontinuity in such a larva as that 
from which fig. 21 is taken would clearly have no bearing on the 
morphological value of the filaments. 

From the details revealed by the larva of Lebrunia, I consider 
that we are justified in regarding the simple filament of the 
Madreporaria and Alcyonaria, and the glandular streak of the 
trilobed filament of the Actiniaria, as representing a continua- 
tion of the enteron; a relationship already suggested by other 
workers (p. 307). 

The fact that no such break in continuity as that referred 
to has ever been observed between the cesophageal epithelium 
and the lateral or ciliated streaks of the trilobed Actiniarian 
filament, has led E. B. Wilson and MeMurrich to regard these 
as ectodermal downgrowths from the stomodzal walls, and in 
this they are followed by most writers on the Actinozoa. My 
results, however, incline me to the opinion of H. V. Wilson, 
namely, that the simple Madreporarian filament is homologous 
with the complex Actinian filament. 

Ontogenetically the Flimmerstreifen appear later than the 
Driisenstreif, and, at any rate in Lebrunia, after the continuity 
of the latter with the cesophagus has been established. They 
are always highly specialized structures in that the constituents 
are wholly columnar ciliated cells, without an admixture of 
gland-cells and cnidoblasts, thus differing histologically from the 
cesophageal, lining or median streak of the filament. There is 
little doubt that they are, like the dorsal filaments of the Alcyo- 
naria, special circulatory organs. Usually they are most strongly 
developed in colonial Actiniaria, such as the Zoanthide. The 
so-called ‘“ Reflected Ectoderm ” of Haddon (1891, p. 619) met 
with in this family must, in all probability, be looked upon as a 
strongly developed portion of the ciliated streak. 

To anticipate results yet to be published, I find that along the 
two sides of the simple filament of the West-Indian species of 
Madrepora a very characteristic structure is developed, in no way 
histologically distinguishable from the Flimmerstreifen of the 
Actinie. 


THE EDWARDSIA-STAGE OF LEBRUNIA. 297 


Such an occurrence in a Perforate coral, in which the circu- 
latory system is very extensive and complex, seems to afford 
strong evidence for regarding the single filament as the homo- 
logue of the whole of the trilobed filament, and consequently 
for its origin as a whole from one and the same embryonic 
layer. 

In the adult Lebrunia the ciliated streaks are strongly de- 
veloped and continuous with the lining of the cesophagus at its 
inner termination. 


ITV. Reparions or THE TENTACLES AND MESENTERIES. 


The origin of the different tentacles in relation to the mesen- 
terial chambers discloses some unexpected conditions. And firstly 
a comparison may be instituted between the time of appearance 
of the tentacles as external organs of the polyp, and the stage 
reached in the development of the internal mesenteries and their 
corresponding chambers. 

Though the first eight mesenteries in all Zoantharia yet 
investigated develop successively in pairs, the early tentacles 
rarely do so. lLacaze-Duthiers, in his classic investigations 
already referred to, found the tentacles to appear in Actinia 
equina during the stage in which eight mesenteries were present, 
one from each mesenterial chamber, as is also the case in 
Lebrunia. Von Koch (1897) apparently found twelve tentacles 
to appear simultaneously in Caryophyllia cyathus, at a stage when 
twelve mesenteries were developed, eight only of which were 
perfect. In Haddon’s newly-hatched larve of Huphyllia no 
tentacles, however, were developed, although six pairs of mesen- 
teries were present, three pairs of which possessed filaments. 

On the other hand, both MceMurrich and van Beneden found 
that in the Cerianthid Arachnactis the appearance in pairs of the 
tentacles follows closely upon that of the formation of the mesen- 
terial chambers. But, in consequence of the stomodeum ex- 
tending right across the celenteron, from wall to wall, at the 
stage of development characterized by the presence of two, or 
perhaps three, pairs of lateral tentacles, there exists no trace of 
any median chambers, and it is only when these have been 
formed that a median tentacle appears. In this case it is the 
ventral tentacle; a median dorsal never occurs, the region being 


298 MR. J. E. DUERDEN ON 


one of apparently continuous growth in pairs of the tentacles 
and mesenteries. 

From the few examples given there is evidently little or no 
connection to be expected between the stage of internal develop- 
ment and the appearance of the tentacles. 

In viviparous polyps the formation of the tentacles rarely 
takes place until the larve are set free, whereas the internal 
development may have proceeded to almost any degree, though 
apparently not often beyond the Hdwardsia-stage. Yet in newly- 
hatched larvee of Awlactinia I found examples with six tentacles 
already protruding. 

The relation of the axis of symmetry of the tentacles in their 
bilateral stage with that of the mesenteries may now be con- 
sidered. 

In Actinia equina Lacaze-Duthiers found the first and largest 
tentacle to arise from the sulcular endoceele, a smaller and 
opposite from the sulear endoccele, and the three lateral pairs, 
varying somewhat in size, from the lateral mesenterial chambers. 
The sagittal axis of the tentacles, therefore, is the same as the axis 
of symmetry of the paired mesenteries, a relationship which would 
naturally be expected. The longer axis of the stomodeum is also 
im the same plane. ‘The figures of McMurrich and van Beneden 
show that in Arachnactis the plane of symmetry of the tentacles 
and of the mesenteric chambers likewise coincide, and von Koch 
represents the same relations for Caryophyllia eyathus. 

In one late larva of Lebrunia, in which a complete series of 
transverse sections enabled these relationships to be followed from 
one end to the other, a different relationship was encountered. 
This is represented in Pl. 19. figs. 23-25. 

The first section (fig. 23), passing through the basal tentacular 
region, exhibits the relative sizes of the tentacles such as has 
already been described among the external characters for the 
bilateral stage. Sections in advance of this leave no doubt as to 
the correctness of the dorgo-ventral relations here indicated. 
The longer axis of the oval stomodeum is in a plane at right 
angles to that of the median axis of the tentacles. Fig. 24, 
taken from a section through the middle stomodeal region, re- 
veals the same bilateral arrangement of the tentacles, but the 
dorso-lateral tentacles (f, 4) are now comparatively larger than 
before on account of their relatively lower origin in the polyp. 


THE EDWARDSIA-STAGE OF LEBRUNIA. 299 


Under a high magnification the very delicate retractor-muscle 
fibres on the mesenteries can be made out, enabling the two 
pairs of directives (III., III.; [V., LV.) to be determined. These 
are found to be arranged about an axis (a—e) at right angles to 
the median axis of the tentacles (g—c). I have represented the 
cut ends of the muscular fibrils in the figure, although they 
could not be discerned at such a low magnification as that 
given. 

The next figure is from a section a little below the stomodeal 
region, and, although no trace of any tentacles occurs, the 
section follows the others in regular sequence without any dis- 
turbance of the axes. The larva was sectionized in one con- 
tinuous ribbon and mounted accordingly. As in the previous 
figure, the sulco-sulcular plane of the larva (a-e) is at right 
angles to what would be the median plane of the tentacles 
(g-e); in other words, the two median lateral tentacles com- 
municate the one with the sulcar (@) and the other with the 
sulcular endoccele (e), while the two tentacles in the dorso-ventral 
plane communicate each with a median mesenterial space (¢, g). 

Another condition is disclosed by the slightly flattened larva 
from which fies. 26-28 are taken. The series indicates that on 
the left side the small tentacle next the large dorsal tentacle (/) 
communicates with the sulcular endocele (e), and the small 
tentacle (a) on the right side, next the ventral (6), communicates 
with the sulear endocele (a), so that the median: axis of the 
tentacles (/-0) is ina plane oblique to the suleo-sulecular axis 
of the mesenteries (a-e). Here, again, the longer axis of the 
esophagus is in the same plane as the sulco-sulcular plane of 
the mesenteries. 

A third larva was in the same condition as the first example 
described—the median axis of the tentacles at right angles to 
the axis of symmetry of the mesenteries. And, yet again, a 
fourth example, traced later, revealed the same oblique relation- 
ships as the second here recorded. 

In every case the longer axis of the cesophagus coincided with 
the axis of symmetry of the pairs of mesenteries, not with that 
of the tentacles. 

It is very evident, therefore, from these four cases, that the 
symmetry of the two sets of organs—tentacles and mesenteries 
—is independent in the bilateral stage of the larva. It may 


300 MR. J. E. DUERDEN ON 


perhaps be explained as a result of the late formation of the 
celenteric cavity, of the assumption of the bilateral stage of 
the tentacles before any decided internal bilateral symmetry has 
been established in the upper tentacular region, the nearly solid 
interior having no influence on the method of grouping of the 
tentacular outgrowths. 

Where, as in Arachnactis, the outgrowth of the tentacles 
closely follows the production of mesenterial spaces, it is obvious 
that the two must be directly related, and that the external sym- 
metry will be moulded upon the internal. Also in the species 
studied by Lacaze-Duthiers and others, where the ccelenteric 
chambers were formed in advance of the tentacles, the internal 
symmetry first assumed would be likely to impress itself upon 
the externa] organs arising later. 

The irregularity in Lebrunia is probably rectified in the sub- 
sequent rearrangement of the tentacles in hexamerous cycles, 
for, in the numerous adult specimens I have examined, the 
tentacles, cesophageal axis, and mesenteries exhibit the relation- 
ships usual in the Hexactinie. 


V. CoNCLUSIONS. 


From the foregoing account it is obvious that the larva of 
Lebrunia coralligens presents us with very unexpected conditions 
in Seyphozoan development; and this not alone m any one 
particular organ, but in almost every essential structure. The 
early tetrameral symmetry, followed by a bilateral phase, and 
that again by the hexamerous adult ; the system of ciliated 
coelomic spaces connected with a closed archeuteron, all embedded 
in a mass of undifferentiated tissue; the formation of the ceso- 
phagus by the breaking down of the floor of an ectodermal in- 
vagination in association with an archenteric tube; and the 
origin of the adult gastro-ccelomic cavity from a primary coelome 
and disintegration of the tissues, are all unique characteristics. 

The species seems to retain to a late period certain ancestral 
characters which in other forms are either passed over or dis- 
appear very early, so that other features dependent upon their 
presence are never exhibited. Thus, were the vacuolated tissue 
insufficiently developed, or to disappear early, we should have no 
clear evidence of a distinct larval ccelome. 

Some of the facts cbserved in the early larve appear to obtain 


THE EDWARDSIA-STAGE OF LEBRUNIA. 301 


their full significance only from a comparison with the tetrameral 
Scyphomeduse. Commenting upon the connection of this group 
with the Zoantharta Haddon, ten years ago (1889), wrote :— 
“The relationship of the Hydra-tuba and Scyphostoma stages 
of the Scyphomeduse (Acalephe) to the Zoantharia is now 
generally admitted, indeed a group (Tentolate) has been erected 
by Professor EK. Heckel to include them both. Later Professor 
A. Géotte has similarly proposed the term Scyphozoa for the 
same assemblage, but including the Ctenophora, as opposed to 
the remaining Celenterata or Hydrozoa. The Scyphostoma 
have an cesophagus lined by ectoderm (Stomodeum), four glan- 
dular mesenteries, the edges of which are true craspeda, and 
serve to digest food; in their upper portion nematocysts are 
present. The four tentacles are afterwards increased to eight, 
and finally to sixteen. It is especially noteworthy that at first 
there are only two tentacles: probably this is a reminiscence of 
a remote ancestor. The widespread occurrence of a symmetry 
of four amongst the larve of the Scyphozoa is very suggestive.” 

Discassing the Phylogeny of the Actinozoa McMurrich (1891 a, 
p. 149), two years later, also remarked :—‘‘ As regards the rela- 
tions of the Actinozoa to other Ceelenterates, there is little to 
be said ; the majority of authors who have committed themselves 
upon the subject, agree in tracing the Actinozoan stem back to 
a form similar to the Scyphistoma. The evidence we have seems 
to point in that direction ; but it must be acknowledged that it 
is exceedingly scanty, and there are many points of difference 
between any Scyphistoma of which we have a description and 
the simplest Actinozoa. It seems probable, however, that the 
Actinozoa are to be traced back to an ancestor possessing ouly 
four mesenteries. The occurrence of an octameral symmetry in 
the simplest Actinozoa seems to point in that direction, as well 
as the fact that, in the development of the Hexactinix, the 
stage with four mesenteries seems to mark an epoch, much less 
distinct, however, than that indicated at the close of the 
Edwardsia stage.” 

The tetrameral symmetry of the Lebrunia larva, perfect as 
regards the tentacles, and the mesenteries and coelomic diverti- 
cula of the upper archenteric region, seems to be explicable only 
on such a throwing back of the ancestry of the Zoantharia. The 
two alternating series of four tentacles exactly recall the eight- 

LINN. JOURN.—ZOOLOGY, VOL. XXYVIL. 23 


302 MR, J. E. DUERDEN ON 


armed stage passed through by most Scyphostoma, and the 
arrangement of the four canals, with the circumferential sinus 
terminating in eight divisions bifurcated at the end, resembles 
a gastro-vascular system such as that of the Ephyra. 

Though in the particular case of Lebrunia the eight tentacles 
develop simultaneously, it does not follow that they may not 
have had an ancestry in which they arose in pairs, as happens in 
the Seyphostoma. For there is evidence that such fundamental 
structures as the primary mesenteries develop successively in 
pairs, as is the case in all other Zoantharia. The two consti- 
tuting the fourth pair (sulcular directives) in most cases disappear 
at the aboral end before the others, and in every case the first 
pair (sulco-lateral) are larger than the others, and are the only 
ones which bear mesenterial filaments. 

The results appear to justify to a greater degree than has 
before been possible Heckel’s and Gotte’s union of the Scypho- 
meduse and Actinozoa under the term Teniolate or Scyphozoa ; 
a relationship first rendered probable by the discovery of the 
ectodermal character of the stomodzal liming of the former, and 
the presence of mesenteries and gastric filaments. The strong 
objection founded on the tetrameral character of the Scypho- 
medusx, as compared with the hexameral nature of the Acti- 
niaria, disappears on the demonstration that the larve of the 
latter may pass through a tetrameral stage, directly comparable 
with that permanent in the former. 

Further, where the transition from one stage to the other can 
be followed in the ontogeny of a single form, we have another 
to the many objections against the separation of the Tetracorallia 
or Rugosa from the Hexacorallia. ‘There can be no fundamental 
distinction on the ground of symmetry alone. 

The very marked arrest of the mesenterial and tentacular 
development in Lebrunia at a tetrameral or octameral stage, and 
the fact that such occurs to a greater or less degree in apparently 
all other Zoantharia (Hdwardsia-stage), suggest this as either a 
permanent stage or as a starting-point for various modifications, 
and emphasizes with how little reliability adult structure can be 
accepted as evidence of phylogenetic relationship. 

The Rugosa and Aleyonaria retain approximately a tetrameral 
or octameral type of symmetry, while the Hexacorallia and 
Actiniaria, by the further addition of two lateral pairs of 


THE EDWARDSIA-STAGE OF LEBRUNIA. 303° 


mesenteries, give rise to another—the hexameral. Even amongst 
the Hexactinie, however, octameral or tetrameral examples may 
occur; occasionally in such a form as Azptasia annulata, but 
apparently always in most species of the genus Corynactis 
(Duerden, 1898, p. 649). 

MeMurrich (1891, p. 311) is inclined to regard the differences 
of arrangement of the retractor muscles of the mesenteries, 
such as one sees in the directives of the Aleyonaria and in the 
Zoantharia, as of secondary importance, in comparison with the 
order of development and number of mesenteries. 

Undoubtedly the character of greatest concern is the occurrence 
im an Actinian of what appears to be a system of primitive 
ceelomic spaces connected with a closed archenteron, within an 
otherwise solid interior. And it remains to be seen what support 
there is for the views already advanced in regard to such. 

Until the earlier stages of the embryo have been obtained, it 
is impossible to determine the manner in which the conditions 
have arisen, and therefore to homologize them with certainty 
with what occurs in other groups. It will be necessary to ascer- 
tain whether an actual primary invagination takes place before 
or after the appearance of the supporting lamella, and if the 
spaces themselves are primitively formed as evaginations of this, 
the limiting layer being a direct continuation of its walls; or, 
whether the spaces originate independently as splittings within 
the solid tissue, their limiting layer then artsing as a modifica- 
tion of the marginal cells, and later entering into communication 
with the internal end of the archenteron. 

The evidence obtainable from the development of other Scy- 
phozoa assists but little in the elucidation of the actual facts 
presented by Lebrunia at the stages under consideration. From 
the fertilized ovum a blastosphere results, and, according to the 
accounts of some observers, the two-layered planula is formed by 
invagination, and, according to others, by delamination. Review- 
ing all the known cases, MeMurrich (1891), in the light of his 
own results with Metridium, comes to the conclusion that, with the 
probable exceptions of Pelagia and Nausithoé, in every case the 
endoderm in the Scyphozoa is produced by delamination ; that 
the results of Kowalewski with Actinza (sp.?) and Cerianthus, 
of Jourdan with <Actinia equina, as also of Heckel with the 
Aleyonarian Monoxenia.in that they ascribe the production of 

23* 


304 MR. J. E. DUERDEN ON 


the gastrula, and consequently of the endoderm, to invagination, 
are a misinterpretation of the appearances. 

In the course of development a primitive mouth or blastopore 
appears to be always formed, and in some instances becomes 
closed; but in any case the permanent mouth and stomodeum 
are considered to arise by a later infolding of the ectoderm. 

At the conclusion of the delamination in Metridium MceMurrich 
(1891, p. 308) observed a slight depression at the posterior pole 
of the larva, which soon after breaks through, a communication 
of the interior cavity with the exterior being thus established. 
His latest stages of this species did not permit of his 
ascertaining the manner of formation of the stomodeum. A 
primary mouth, with a later, invagination of the two-layered 
wall to form the stomodeum, is figured by Kowalewsky for 
Actinia (sp.?) and Certanthus, and by Jourdan for <Actinia 
equina and apparently for the coral Balanophyllia (1881, 
‘Embryological Monographs,’ pls. xi.—x1il.). 

In Aurelia, Gotte found that after the planula settled down 
an ectodermic invagination is formed, its lower end breaking 
through, thus establishing a communication of the interior with 
the exterior and constituting the cesophagus. First two and 
then four diverticula or gastric pouches of the general cavity 
are formed and arranged around the cesophagus, and a reflection 
of the lining of the latter also appears to take place. 

I believe it will ultimately be found that the early stages in 
the development of Lebrunia will be better understood by a 
comparison with what occurs in the Scyphomedusz than by what 
happens in the Actinozoa. 

In Lebrunia, we are confronted with the facts that a stomo- 
dal-like funnel already exists in association with a considerable 
mesenterial development, and that up to the time of liberation 
of the larve no oral aperture is present, but that this is formed 
shortly after extrusion. 

Can we then, in face of what is already known in Seyphozoan 
embryology, assume that the first mentioned structure is an 
endodermal-lined archenteron, the blastopore of which has be- 
come and is still closed, and that the later invagination is the 
one corresponding with that which gives rise to the mouth and 
ectodermal stomodzum in all other known Scyphozoan larve? 

Until the study of earlier stages can settle the question 


THE EDWARDSIA-STAGE OF LEBRUNIA. 305 


absolutely I can come to no other conclusion, and, of course, the 
correctness or otherwise of the explanation of many of the other 
peculiar features is determined thereby. The view has the 
merit of placing the conditions in strict harmony with what 
occurs in many Enteroceela. 

The limiting layer of the larval ccelomic spaces in continuity 
with the lining of the archenteron must be considered as the 
equivalent of the wall of the ceelomie pouches or ccelomie split- 
tings of the higher Metazoa. Here it is usually regarded as 
mesoderm or mesothelium, the latter term being employed in 
order to distinguish it from the mesoderm—“ mesenchyme ”— 
arising by immigration. 

The tendency, however, is to regard still as endoderm the 
walls of all the outgrowths from the archenteron and their de- 
rivatives. Thus in the latest important work on the morphology 
of the Echinoderms, Dr. H. L. Clark (1898) describes as meso- 
derm only the tissue of mesenchymal origin, and this plays a 
very insignificant part in the adult structures. Under endoderm 
he includes all the derivatives of the archenteron and its hydroen- 
teroceel, embracing ataongst other structures the lining of the 
digestive tract with most of the cesophagus; all the muscles of 
the body-wall and gut; the peritoneal lining of the body-cavity 
and epithelial covering for the various organs contained in it, as 
well as the genital organs and their ducts. 

Homologizing the limiting layer of the cclomic pouches 
in Lebrunia with the peritoneal epithelium of the body-cavity 
of the bigher Metazoa, we may, with Clark, still regard it as 
endoderm, or, following the more usually accepted terminology, 
speak of it as mesoderm ; and, as in all the Enteroceela, it 
gives origin to the principal muscular system and the gonads. 
The portion adjacent to the column-wall and that covering 
the two faces of the mesenteries would correspond with the 
somatic, and the remainder with the splanchnic layer of the 
higher Metazoa, and it is the former which remains as the 
epithelial lining of the major part of the adult mesenteric 
chambers. Below the cesophageal region it is the splanchnic 
layer which disappears during disintegration, but above, this 
constitutes the coelomic lining of the gullet. 

What, then, of the vacuolated mass of tissue which probably 
entirely disappears in the adult? It is the tissue within which 


306 MR. J. E. DUERDEN ON 


the ceelomic spaces are formed. To continue the comparison 
with the Tripoblastica, may we not regard it as the equivalent of 
the “mesenchyme” of the higher Metazoa? It is evident that 
in Lebrunia it is little more than a larval * nacking-tissue, ” 
disappearing as the adult Scyphozoan characteristics are taken 
on. As already mentioned, however, it probably shares in the 
formation of the muscular system. 

Even though beyond the esophagus no separate digestive tract 
with closed walls remains in the adult, there seems little doubt 
that the mesenterial filaments are to be looked upon as repre- 
senting an endodermal-lined digestive gut, continuity with the 
cesophagus being either original or established at a very early 
stage. As the primary distinctness of the ccelomic pouches has 
broken down at an exceptionally early stage, so the walls of the 
enteron come to be represented only by thickened ridges along 
the free edge of the septa of the celome. Or the same result 
may thus be conversely stated, and perhaps with greater morpho- 
logical truth, that as the lower portion of the enteron beyond the 
point at which diverticula are given off is not formed early 
enough, so the mesenchyme and ccelomic epithelia, which on 
their part are developed and would have surrounded it, beeome 
disintegrated, leading to the imperfect condition of the ccelomic 
chambers in the adult. It only remains to conceive of the adult 
mesenteries uniting along their lateral edges and we should 
have a closed gut as in the higher Metazoa, lined above by the 
invaginated ectoderm and below by the archenteric endoderm. 
The mesenterial chambers thus distinct from one another would 
constitute a true ceelome or body-cavity, exactly as in the ceso- 
phageal region and in the Enteroceela. As it is, the Seyphozoa 
are distinguished by having the lower portion of the enteric 
system in separate longitudinal bands, its cavity in communication 
laterally and below with the chambered body-cavity. 


Were the explanations here offered to be confirmed, the occur- 
rence in the Scyphozoa of an archenteron with distinct radiating 
coelomic diverticula would be recorded for the first time, but the 
broad relationships of the group with the higher Metazoa thereby 
implied have been already surmised by various workers. 

Prof. KE. B. Wilson (1884), in his paper on “ The Mesenterial 
Filaments of the Alcyonaria,” devotes a section to the relations 


THE EDWARDSTA-STAGE OF LEBRUNIA. 307 


of the Anthozoa to the Enterocela. Having established, both 
experimentally and histologically, that the digestive functions of 
the Alcyonaria are confined to the six ventral mesenterial fila- 
ments, he regards the latter as the representatives of the alimen- 
tary canal of higher animals, and suggests that “ they are not only 
physiologically but also morphologically the equivalents of the 
enteron of the Enterocela”; and continuing, atfirms that 
“morphologically we may regard the radial chambers as diverti- 
cula from the primitive enteron.” 

Van Beneden (1891), from his study of the development of 
the larval Cerianthid Arachnactis, likewise comes to the same 
conclusion in reference to the mesenteric chambers of the 
Anthozoa and the coelomic diverticula of the higher animals. 
He devotes special attention to a comparison of the origin 
and arrangement of the mesenteries and their chambers in the 
Cerianthide with paired coelomic diverticula of the segmented 
Metazoa. 

According to a preliminary notice appearing in ‘ Nature,’ 
March 2, received when this contribution was nearly completed, 
Mr. J. Stanley Gardiner, studying a supposed new species of the 
coral Cenopsammia from Lifu, has also come to practically the 
same conclusion as Prof. E. B. Wilson in regard to the enteron 
and mesenterial filaments. The notice contains the pertinent 
sentence: ‘It was further contended that the stomodseum 
together with the mesenterial filaments is homologous with the 
whole gut of the Triploblastica, and that the so-called endoderm 
is homologous with the mesoderm. The Actinozoon polyp then 
must be regarded as a Triploblastic form.” 

In the larva of Lebrunia we appear to have the actual embryo- 
logical proof of these surmises, founded mostly upon a con- 
sideration of the adult anatomy. And it is clearly such a problem 
as can only be established on embryological grounds. 

Whether the larval spaces are derived originally as paired or 
radiating evaginations of the terminal region of an archenteron, 
and their walls are then to be regarded as mesothelium or endo- 
derm, or whether they originate as splittings within the solid 
undifferentiated cell-mass, matters but little. Both processes 
occur in the higher animals: in some the celome originates from 
endodermic diverticula, e. g. Echinodermata, Amphioxus ; in 
others from mesoblastic splitting, e. g. nearly all Vertebrates. 


308 MR. J. E. DUERDEN ON 


The main point sought to be established is that the larval spaces 
of Lebrunia represent a paired coelome embryologically equivalent 
to that of the higher Metazoa, and, consequently, that the 
chambers of the adult are the same. For there is no doubt as to 
the manner in which these latter arise from the primary spaces. 

It remains to discuss what support there is otherwise for such 
a change in conception of the layers and internal cavity of the 
Seyphozoa. 

Prof. J. P. MceMurrich, in his most valuable series of ‘** Contri- 
butions on the Morphology of the Actinozoa,” devotes Part II. 
(1891) to the development of the Hexactiniew, and therein 
describes an almost complete series of stages from the egg to 
the adult, taking his examples from the genera Metridium, 
Rhodactis, and Aulactinia. In reviewing all the known cases of 
early development, he concludes, as already mentioned, that the 
so-called endoderm of the Actinozoa is derived from the hollow 
blastula by the process of delamination, not by invagination, the 
extent of its cellular development being mainly dependent upon 
the amount of yolk present. 

Following partly the results of Metschnikoff, MceMurrich 
regards the process of delamination as a modified form of the 
more primitive process of immigration. In groups higher than 
the Ccelenterata the products of immigration, whether from 
the ectoderm or endoderm or from both, are regarded as meso- 
dermic (ea. the “mesenchyme” of Echinoderms). There is little 
doubt that the internal parenchymatous tissue of the Lebrunia 
larva has arisen by delamination in the same way as in other 
Actinozoa: hence, if an independent endoderm, as from an 
archenteric invagination, can be established, there seems no 
reason why the first-mentioned tissue should not be regarded as 
the homologue of the mesoderm of higher Metazoa. May we 
not in Lebrunia have ‘“‘ mesenchyme” arising by delamination 
(in migration), and ‘ mesothelium” (endoderm) from an arch- 
enteric invagination, as in the Echinoderms ? 

To the solid larval stage in the Actinozoa, where segmentation 
is completed and before the formation of the stomodeum, 
MeMurrich (p. 310) applies the term “ Sterrula.” 

In many Alcyonaria, and apparently in some Madreporaria 
(Manicina and Balanophyllia), this is a solid mass of more or 
less definite cells ; whereas in Metridium, so far as McMurrich’s 


THE EDWARDSIA-STAGE OF LEBRUNIA. 309 


embryos allowed the study to be continned, the sterrula was 
provided with only a comparatively narrow layer of endoderm, 
the remainder of the cavity being filled with yolk-granules. In 
the solid sterrula of the Aleyonaria the middle cellular tissue 
early begins to disorganize as the growth of the larva proceeds, 
giving rise, of course, to the usual ceelenterate gastro-ccelomic 
cavity with a unilaminar epithelium; while in the other cases it 
seems that the ccelenteron is produced by the absorption of the 
yolk, or some of the latter may be extruded through the mouth 
of the embryo. 

The earliest larva of Lebrunia I possess has passed beyond the 
sterrula stage, the mesenteries and the archenteron being already 
formed. It is in the late stage to which the solid cellular tissue 
persists, that the species appears to differ from other examples 
yet recorded. And it would appear to be this entire retention 
of the central tissue, as compared with its early disappearance 
or absence in other Scyphozoa, which may enable the morpho- 
logical conceptions of the other systems to be obtained in such a 
way as is known for no other species. 

Is there any evidence that such a system of diverticula occurs 
at any stage in other Actinozoa? For it can scarcely be supposed 
that such an apparently fundamental phenomenon is restricted to 
an isolated type. 

Few Zoauntharia larve of exactly the same stage as the earliest 
Lebrunia have been minutely described. In some respects, as in 
the stage reached in the development of the mesenteries, the 
Lebrunia larva is far advanced ; but in others—the persistence 
ot the vacuolated tissue and non-formation of oral aperture—it 
is somewhat early in its development. McMurrich (1891) 
found in the youngest embryos of Rhodactis Sancti-Thome that 
the so-called endoderm-cells completely fill the central cavity, 
and show little or no arrangement into a definite layer. At the 
stage, however, where only two mesenteries are present, a well- 
marked central cavity already existed below the upper region of 
the body, though the endoderm above was yet solid, no inter- 
mesenterial cavities having appeared. In this species then the 
solid endoderm begins to disorganize at a much earlier stage 
than in Lebrunia, in fact before the mesenteries, with which the 
spaces are associated, are formed. MceMurrich’s earliest stages 
of Aulactinia possessed eight perfect mesenteries, the first only 


310 MR. J. E. DUERDEN ON 


being provided with mesenterial filaments. They therefore 
correspond with the oldest stage in Lebrunia. The endoderm, 
however, had already arranged itself into a somewhat definite 
layer, but lying scattered about in the body-cavity of the embryo 
were numerous, somewhat large, cellular elements and yolk-— 
eranules. 

The Zoanthid larva which van Beneden (1890) identified as 
closely related to Semper’s larva presents three pairs of complete 
mesenteries and four incomplete pairs, and a very thick endo- 
derm, with small ccelenteric spaces within the stomodzal region. 
Below, however, a considerable gastro-vascular cavity is formed. 
The same author (1891) found the larve of the Cerianthid 
Arachnactis, at the stage with only one pair of mesenteries and 
two pairs of tentacles, to possess a ccelenteron fully formed with 
an endodermal lining of only a single layer of cells. Prof. G. von 
Koch observed a central cavity in Gorgonia and in Caryophyllia 
cyathus (1897) before the formation of any mesenteries or the 
production of an oral aperture and stomodzeum. 

In the newly-hatched larva of Huphyllia rugosa, Haddon (1890) 
found three of the twelve pairs of mesenteries already bearing 
filaments, and alternating with the mesenteries were “large 
ridge-like vesicular outgrowths from the endoderm.” There can 
be little doubt that, as in Lebrunia, these latter are the detached 
survivors of a more or less solid vacuolated tissue in the earlier 
stages of the larve. Prof. EH. L. Mark (‘Selections from 
Embryological Monographs,” pl. xii. fig. 82) figures similar inter- 
mesenterial protrusions, thoroughly vacuolated, in Hdwardsia. 
The appearances should be compared with fig. 19 and with the left 
side of fig. 20 (P1.19) in the present paper. The phenomena are 
remarkably alike. On plate xii. fig. 15 of the same publication, 
Prof. Mark reproduces the transverse section of the larva of 
Balanophyllia regia given by EH. Jourdan, in which the embryo 
is still solid, though the central portion is indicated as yolk. 
Six mesenteries are here shown and the internal yolk seems 
clearly separated from the endoderm, which latter is still many 
cells in thickness. The conditions appear to be somewhat com- 
parable with those in fig. 14 (PJ. 18) of Lebrunia. I regret ex- 
ceedingly that, not having Jourdan’s original memoir for reference, 
I can do no more than merely draw attention to the possibility of 
larval coelomic spaces being present in such a case. The thick 


THE EDWARDSIA-STAGE OF LEBRUNIA. oll 


vacuolated endoderm persistent in Euphyllia and Edwardsia just 
mentioned recalls so strongly the conditions in the late larve of 
Lebrunia, that it seems not improbable similar diverticula may 
also have occurred in these species at an earlier stage in their 
development. 

It is evident from all this, that in most Scyphozoa the forma- 
tion of the gastro-ccelomic cavity of the adult takes place at a 
much earlier stage than in Lebrunia, so that probably no complete 
and distinct larval coelomic system is ever formed. This in no 
way disproves the view here presented. All that can be asserted 
from the résumé just given is that the disorganization to form 
the secondary body-cavity usually originates at such an early 
stage that no opportunity exists for the primary enteric and 
ceelomic system to establish itself. From such conditions as 
those revealed by Lebrunia in fig. 15 (Pl. 19), there seems no 
reason why at some time a species may not be found in which the 
archenteron is prolonged centrally beyond the origin of the 
diverticula, in which case we should have a temporary digestive 
cavity with closed walls. In this connection one is tempted to 
recall the ramified digestive tract described by Bourne (1887) as 
occurring in Huphyllia. 

The larva of Lebrunia is in many respects of an exceptional 
character. If the interpretation of its tetrameral primary 
tentacles be correct, it shows that phylogenetically it is, in 
regard to these organs, at a stage earlier than other Zoantharian 
larvee yet described, and perhaps the same may be said of much 
of its internal condition. No doubt the two sets of phenomena 
are In some way interdependent. 


VI. BretiogRarHy *. 


1860. Ducuassarne, P., et Mrcuenorti, J.—‘‘* Mémoire sur les 
Coralliaires des Antilles.’ Mem. della R. Accad. Sci. 
Torino, ser. II., xix. 

1879. Herrwie, O. & R.—‘ Die Actinien.’ Jena. 

1884. Marx, H. L.--‘“‘ Selections from Embryological Mono- 
graphs, III. Polyps.” Mem. Mus. Comp. Zool., vol. ix. 


* Under this heading I include only the few original works which, cut off 
from any fully-equipped scientific Library, I have been able actually to consult. 


1888. 


1889. 


1889. 


1889. 


1889. 


1890. 


1890. 


1890. 


1891. 


1891. 


1891. 


MR. J. E. DUERDEN ON 


Witson, HE. B.—‘ The Mesenterial Filaments of the 
Aleyonara.” Mitt. a. d. Zool. Stat. zu Neapel, Bd. v. 
Bourne, G. C.—‘‘ On the Anatomy of Mussa and Eu- 
phyllia, and the Morphology of the Madreporarian 

Skeleton.” Quart. Journ. Mier. Sci., xxvin. 

Witson, H. V.—“ On the Development of Manicina 
areolata.’ Journ. Morph., vol. i. 

McMourriceu, J. P.—“ On the occurrence of an Hdwardsia- 
stage in the free-swimming embryo of a Hexactinian.” 
Johns Hopkins Univ. Circulars, vin. 

Happon, A. C.—“‘ A Revision of the British Actinie, 
Part I.” Sci. Trans. Roy. Dublin Soe., 2 ser., vol. iv. 
McMourrico, J. P.—‘‘ The Actiniaria of the Bahama 

Islands, W.I.” Journ. Morph., vol. ii. 

von Kocn, G.—Uber Caryophyllia rugosa, Moseley.” 
Morph. Jahrbuch, xv. 

van Benepen, E.—“ Les Anthozoaires pélagiques recueillis 
par M. le Professeur Hensen dans son Expédition du 
Plankton.—I. Une Larve voisine de la Larve de Semper.” 
Pull. Acad. Roy. Belgique, 3me sér., xx. 

Happon, A.C.—‘ The Newly-hatched Larva of Euphyllia.” 
Sci. Proc. Roy. Dublin Soe., vol. viii. 

Witson, H. V.—“* Ona new Actinia, Hoplophoria coral- 
ligens.” Studies from the Biol. Lab., Johns Hopkins 
Univ., vol. iv. 

van BunepENn, E.—‘ Recherches sur le Développement des 
Arachnactis. Contribution a la Morphologie de Cérian- 
thides.” Archives de Biologie, Tom. x1. 

Hannon, A. C., & SHackieton, A.—‘‘ A Revision of the 
British Actinia.—Part II. The Zoanthee.” Sci. Trans. 
Roy. Dublin Soe., 2 ser., vol. iv. 

McMoretcn, J. P.—‘‘ Contributions on the Morphology 
of the Actinozoa.—II. On the Development of the 
Hexactinie.” Journ. Morph., vol. iv. 


1891 a. McMurricu, J. P.—‘ Contributions on the Morphology 


1893. 


of the Actinozoa.—III. The Phylogeny of the Actinezoa.” 
Journ. Morph., vol. v. 

McMoratcn, J. P—‘ Report en the Actiniz collected by 
the United States Fish Commission Steamer ‘ Albatross’ 
during the winter of 1887-1888.” Proc. U.S. Nat. Mus., 
vol. xvi. 


THE EDWARDSIA-STAGE OF LEBRUNIA. 313 


1897. von Kocu, G.—“ Entwicklung von Caryophyllia cyathus.” 
Mitt. a. d. Zool. Stat. zu Neapel, 12 Bd., 4 Heft. 

1898. Durrven, J. E.—‘“ The Actiniaria around Jamaica.” 
Journ. Inst. Jamaica, vol. 1. No. 5. 

1898. CruarK, H. L.— Synapta vivipara: A Contribution to the 
Morphology of Echinoderms.” Mem. Bost. Soc. Nut. 
Hist., vol. v. 

1898. Duerpen, J. E.—“ On the Relations of certain Sticho- 
dactyline to the Madreporaria.” Linn. Soc. Journ, 
Zool., vol. xxv. 

1898. Verrint, A. H.—‘‘ Descriptions of imperfectly-known and 
new Actinians, with critical notes on other species, II.” 
Amer. Journ. Sci., vol. vil. 


EXPLANATION OF THE PLATES, 


Reference figures and letters. 


The Roman numerals I. to IV. are throughout opposite the four pairs of 
mesenteries, and also represent respectively the order in which they usually 
appear in the development of the Zoantharia. 


I. I. Sulco-lateral pair. 
II. I. Sulculo-lateral pair. 
III. I1I. Sulcar pair of directive mesenteries. 
TY. LV. Sulcular pair of directive mesenteries. 


The letters a—A correspond with the mesenterial chambers: a, sulcar 
endoceele ; 0, left sulco-lateral chamber ; c, left median lateral chamber; d, left 
suleulo-lateral chamber = sulcular exocele; ¢, suleular endocele; f, right 
sulculo-lateral chamber = sulcular exoceele; g, right median lateral chamber ; 
h, right sulco lateral chamber. Both for the mesenteries and the chambers 


the terminology here adopted must necessarily become slightly altered as other 
miesenteries appear. 


arc.w., wall of archenteron. mes.fil., mesenterial filament. 
el.gl.c., clear gland-cells. mus.2., muscular layer. 

cw.s., ccelomie space. ner.l., nerve-layer. 
d.nem., developing nematocyst. ws., oesophagus. 

ect., ectoderm. or., oral aperture. 
ect.f., floor of ectodermal invagi- s.nem., small nematocyst. 
nation. sp./., supporting lamella. 

gr -gl.c., granular gland-cells. sup.c., supporting cells. 


lnem., large nematocyst. 
Llay., limiting layer. 


ve.t., vacuolated tissue. 
zoox., zooxanthellee. 


ol4 MR. J. E. DUERDEN ON 


Prare 18, 


Figs. 1-7 are not drawn to any scale; 1-6 are representations of the living 
larvee as seen under a low power of the microscope, and 7 as seen with 
a hand-lens. All the others are from sections, and were drawn with 


the assistance of a camera. 


Fig. 1. The usual form of the freshly-extruded, free-swimming larva. 

Fig. 2. A form occasionally assumed. 

Fig. 3. The contracted cake-like condition. 

Fig. 4. Oral aspect of a free-swimming larva a few hours after extrusion. 

Fig. 4a. Lateral aspect of a larva a few hours after extrusion. The tenta- 
cular protuberances are beginning to appear. 

Fig. 5. Oral aspect of free-swimming larva. 

Fig. 6, The same seen from the side. 

Fig. 7. A larva three or four days old. (See Plate 19.) 

Fig. 8. Vertical section through the middle region of the column-wall of a 
non-extruded larva in which the central tissue has already become 
disoryanized. xX 450. 


Figs. 9-11. Vertical sections through the archenteric region of a larva very 
shortly after liberation, showing the formation of the oral aperture and 
cesophagus. x 320. 

Fig. 9. Tangential section near the periphery of the archeuteric wall. The 
flattened inturned ectoderm is in close contact with the flattened 
archenteric wail below. The upper narrow slit indicates the upper 
outer surface of the floor of the former, and the broad slit below is 
the central celomic space, to the right continued peripherally and 
below. 

Fig. 10. Tangential section near the centre. The actual oral aperture appears ; 
the floor of the invaginated ectoderm is convex and produced 
laterally. 

Fig. 11. Radial section. “The floor of the invagination and the roof of the 
archenteron are here broken through and complete communication is 
established between the interior and exterior. The invaginated 
ectoderm forms the greater part of the cesophageal wall, but the 
innermost portion is derived from the archenteric endoderm. ‘The 
union between the two is indistinguishable, but from other sections 
it is clear that it occurs about halfway below the lateral inden- 
tations. 


Figs. 12-14. Transverse sections of a freshly-extruded larva before dis- 
organization is set up. x 250, 

Fig. 12. ‘Through the upper archenteric region. ‘The section is taken slightly 

obliquely to the vertical axis and serves to represent the various 


THE EDWARDSIA-STAGE OF LEBRUNIA. 35) 


appearances of the ccelomic spaces. d, e, f, g, present the successive 
appearances from above downwards, showing how the single 
mesenterial space ends in two distinct cornua. Between a, b the 
separating mesentery has broken down and the two chambers are in 


communication one with the other, and a little lower also with the 
space in h. 


Fig. 13. The figure is made up of the appearances presented by several oblique 
sections through the region of the termination of the archenteron. 
Four radiating canals communicate with the two ccelomic sinuses, 
the latter separated by the first pair of mesenteries—sulco-lateral, 
developed to a greater extent than the three other pairs. The 
limiting layer of the sinuses is both somatic and splanchnic, 


Fig. 14. A little below the archenteric region. The middle of the larva is 


filled with vacuolated tissue; towards the upper side the celomic 
canals are completely separated one from the other, while on the 
lower side the last traces of the connections are seen, and to the 
right the last trace of the sulcar canal. 


Puate 19, 


Fig. 15. Vertical radial section of a freshly-extruded larva such as would be 
obtained along the line x, y in fig. 12. x 120. 


Figs. 16-20. Successive transverse sections through one of the non-extruded 
larvee where disorganization of the central tissue is in pregress. The 
sections are such as would be obtained along the lines 1-1, 2-2, 3-3, 
4-4, 5-5, respectively in fig. 21. x 120. 


Fig. 16. Section through the upper archenteric region. The mesenterial 
chambers are founded on a radial plan, four large and four small. 


Fig. 17. Section towards the termination of the archenteron. In the ventral or 


sulear sinus the splanchnic portion of the central tissue shows two 
separate regions of more deeply-staining tissue continuous with the 
archenterie lining and with the limiting layer, while in the sulcular 


sinus the reflected ectoderm is an almost continuous limiting 
layer. 


Fig. 18. Section nearer the termination of the archenteron than in the last 
figure. Owing to its reflection the lining of the archenteron is cut 
through twice. The lumen of the archenteron is very small and 
circular. x 320. 

Fig. 19. Section below the archenteron. The splanchnic portion of the 

limiting layer and the tissue enclosed by it have become disorganized, 

and only loose fragments remain. The somatic layer and the 


vacuolated tissue between it and the supporting lamella are as yet 
unchanged. 


Fig. 


Figs. 
plete series through a larva several days old, showing the relation of 
the plane of symmetry of the tentacles to the sulco-sulcular plane of 
the larva. x 50. 


Fig. 


ON THE EDWARDSIA-STAGE OF LEBRUNTA. 


. Section towards the aboral region. Stages im the disorganization of 


the central tissue are represented. 


. Vertical section through a larva of tke same stage, such as would be 


obtained along the lines I-2, 2-I in fig. 19. The section passing 
through the pair of mesenterial filaments shows their continuity 
with the lining of the archenteron. 


. Transverse section through the free edge of the sulco-lateral pair of 


mesenteries of a larva one or two days old, taken a little below the 
cesophagus. ‘The mesentery to the left shows the beginning of the 
formation of the mesenterial filaments, while that to the right is as 
yet unaltered. A few sections lower the filament appears strongly 
developed on each.  X 320. 


93-25. Transverse sections selected at. different levels from a com- 


23. Section through the tentacular region and the cesophegus. The 


tentacles towards their origin are in the same bilateral relation 
as in fig. 7. The mesenterial chambers are further developed than 
in any previous section represented. The larger axis of the 
cesophagus is at right angles to the dorso-ventral axis of the 
tentacles. 


. 24. Section towards the lower end of the cesophagus representing still 


the same relations. The suleo-sulcular axis of the mesenteries (a-e) 
and the longer diameter of the csophagus are at right angles to the 
axis of symmetry of the tentacles (g-c). 


r. 25. Section below the cesophageal region and the tentacular prominences. 


The axes are in the same relation as in the two previous figures, 
The axis of symmetry of the mesenteries is at right angles to that 
corresponding with the dorso-ventral axis of the tentacles. 


Figs. 26-28. Transverse sections through a slightly compressed larva of about 


the same age as the former. ‘The series show that the sulco-sulcular 
plane of the larva (a-c) is oblique to the dorso-ventral plane of 
the tentacles (f, 0). x 50. 


if 


.--2ct. 


2 


1 


JE.Duerden del 
R.AHammond | 


A 


2 
— 
ae 
: 
Ps 
. 
te 
al 
(e) 
(@) 
N 
2 
2 
Bae) 
(o) 
_ ©) 
z 
Sl 


SS a(obagy nibs 


West, Newman amp. 


i 


X . 
ctr, 


Pl. 18, 


Laxw. Soc. Journ. Soon. Vor. XXVII 


: 
2 
a 
4 
é 


Ure th 

ee 

ihe “el 
iy 
hey 


J.F.Duerden del. . 
AR. Hammond lith. 


THER EDWARDSIA 


West, Newman imp. 


OF LEBRUNIA. 


EK 


r 


Linn. Soc. Journ. Zoon. Vou. XXVIII. Pl. 19 


COS, ~ 


ON THE MYOLOGY OF ANOMALURUS. 317 


The Position of Anomalurus as indicated by its Myology. 
By F. G. Parsons, F.R.C.S8., F.L.8., Lecturer on Comparative 
Anatomy at St. Thomas’s Hospital and Hunterian Professor 
at the Royal College of Surgeons. 

[Read 4th May, 1899.] 


For several years I have been anxious to dissect the muscular 
system of Anomalurus, and to compare it with that of other 
rodents ; | am therefore especially grateful to Professor Howes for 
kindly placing at my disposal a young specimen which originally 
came from the Congo. Personally I hold that muscles, if judi- 
ciously used, are capable of giving a great deal of information 
about the relationship of animals, because they do not readily 
adapt themselves to changed conditions of life. This opinion is 
the result of a systematic survey of the muscles of several orders 
of mammals, a survey which has occupied me for several years ; 
and although, in the opinion of many anatomists, muscles are 
very unstable structures, I would submit that both in Dobson’s 
hands, and to a lesser degree in my own, a certain amount of 
definite assistance has been afforded systematists by them in 
classifying animals whose position had previously been doubtful. 

Anomalurus gives a very good opportunity for testing the 
resources of myology, since its position is so uncertain. A study 
of Oldfield Thomas’s paper ‘“‘ On the Genera of Rodents’’* will show 
how many different positions have been assigned this animal, and 
one can therefore enter upon the task of finding out what its 
muscles have to tell with a perfectly free and unbiassed mind. 

It may be asked why I have elected to prefer the muscles to 
any of the other systems of the body as an index of relationship. 
It is chiefly a matter of convenience for comparison. The 
nervous, vascular, or alimentary systems may well have many 
secrets to tell, but in order to understand them it is necessary 
that the details of these parts in a large and representative series 
of other rodents should be recorded and collated. This work 
I have already done for the muscles t, and I am therefore able 
to describe these structures in Anomalurus more briefly than I 
should otherwise feel justified in doing, premising that those 
muscles which are not mentioned are fairly stable in all rodents 
and, in my opinion, of little or no value for classificatory 
purposes. 

* P. Z. 8. 1896, pp. 1012-1028. tT P. Z. S. 1894, p..251, & 1896, p. 159. 

LINN. JOURN.—ZOOLOGY, VOL. XXVII. PA 


318 PROF. F. G. PARSONS ON THE 


Muscles of the Head and Neck. 


Temporal.—This muscle is quite small, as is usual in rodents. 
The three parts described by Kunstler *—parietal, temporal, and 
zygomatic—are present, but not very clearly defined. The muscles 
of opposite sides do not nearly meet in the mid-line; but it must 
be remembered that the specimen is that of a very young animal, 
and there is reason to believe that, with the growth of the teeth, 
the masticatory muscles become more developed and the temporal 
occupies more of the side of the cranium. 

Masseter.—This is a muscle of great interest. In previous 
paperst I have pointed out that the hystricomorphine and 
sciuromorphine types of masseter are quite distinct, and it is well 
known that this difference causes a complete alteration in the 
shape of the skull. In most squirrel-like rodents no part of 
the masseter passes through the infraorbital foramen, but a 
special bundle, which has been spoken of as the sciuromorphine 
anterior deep part, occupies a nearly vertical groove in front 
of the zygomatic process of the maxilla. In the porcupine- 
like rodents a considerable part of the masseter, called the 
hystricomorphine anterior deep portion, passes through the 


MASSA cut./ OS eee, 
i 

ANTS DEEP PARY of 

MASSETER . 


The Masseter of Anomalurus. 


oreatly enlarged infraorbital foramen; while the mouse-like 
rodents show the transition between these two arrangements, 
and often, as in the case of the Hamster (Cricetus frumen- 


* Annales des Sciences Naturelles, sér. 7, t. iv. p. 150. 
+t P. Z. S. 1894, p. 251, and 1896, p. 159. 


————- $$$ a 


MYOLOGY OF ANOMALURUS. 319 


tarius)*, both the sciuromorphine and hystricomorphine anterior 
deep parts of the muscle are found in the same individual. 

Anomalurus differs from all the sciuromorphine rodents I have 
hitherto examined in having a small anterior deep portion of the 
masseter passing through the infraorbital foramen, as in many of 
the Myomorpha, and this arrangement I cannot help regarding 
as a myomorphine tendency on the part of the animal. 

Depressor mandibule (Digastric).—The great difference between 
the Sciuromorpha and Myomorpha on the one hand, and the 
Hystricomorpha on the other, is that in the former suborders this 
muscle is really digastric, and the tendons of opposite sides are 
connected across the middle line by a fibrous arcade with its 
convexity forward ; from this convexity the anterior bellies spring 
in such a manner that the mesial borders of the two are in 
contact in the middle line, and the anterior attachments of these 
anterior bellies are close to the symphysis mentit. In the 
Hystricomorpha, on the other hand, the muscle does not deserve 
the name of digastric—the division between the anterior and 
posterior bellies is only indicated by a few tendinous fibres on 
its surface, there is no tendinous arcade, and the anterior attach- 
ment is some distance from the symphysist. Anomalurus agrees 
entirely with the first-mentioned arrangement. 

Transversus mandibule.—This muscle connects the two halves 
of the mandible, just behind the symphysis, lying deep to the 
depressor mandibule and superficial to the mylohyoid; it is 
present in all the Myomorpha and Sciuromorpha except Castor. 
In Anomalurus it is well developed. 

Sterno-cleido Mastoid.—The sterno-mastoid and cleido-occipital 
elements of this compound muscle are present in Anomalurus, 
the former running from the presternum to the base of the 
paroccipital process, the latter from the inner half of the clavicle 
to the curved line of the occipital bone; as is always the case in 
rodents, the spinal accessory nerve passes deep to both parts. In 
many rodents the cleido-occipital portion is often overlapped by 
the clavicular fibres of the trapezius, but this arrangement is not 
confined to any one suborder, and it does not exist in Anomalurus ; 
but I do not at present regard the sterno-cleido mastoid as of 
much value from a classificatory point of view. 

* See figs. 2 & 3, P. Z. S. 1896, pp. 161, 162. 

T See fig. 1, P. Z. 8. 1894, p. 255. 

} See fig. 9, Journ. of Anat. & Physiol. vol. xxxii. p. 439. 
24* 


820 PROF. F. G. PARSONS ON THE 


Omo-hyoid.—This muscle I have hitherto found in every 
specimen of sciuromorphine and myomorphine rodent which I 
have dissected, but it is absent in many of the Hystricomorpha 
and in the Lagomorpha. In Azomalurus it is absent, and, if this 
should prove constant, it will be a mark of distinction between 
that animal and other sciuromorphine rodents. 

The Omo-trachelian (Levator clavicule) rises from the anterior 
arch and transverse process of the atlas and is inserted into the 
acromion and metacromion ; these are its usual attachments m 
Sciuromorpha and Myomorpha, though in the Hystricomorpha 
and Lagomorpha it often rises from the basioccipital. __ 

Rhomboidei.—The rhomboideus capitis in Anomalurus forms a 
continuous sheet with the cervical and thoracic parts of the 
muscle : this is generally the case in the Sciuromorpha, but in the 
Myomorpha the rhomboideus capitis is usually a distinct muscle. 

Splenii.—The splenius capitis is always well developed in 
rodents, and in Anomalurus it has the usual arrangement; the 
splenius colli, on the other hand, is a rare muscle, and Anoma- 
lurus differs from most other rodents in having it well marked. 

Trachelo-mastoid is present as in most rodents. 

Scaleni.—A scalene muscle passing ventral to the subclavian 
artery and brachial plexus is never found in the Sciuromorpha, 
very rarely in the Myomorpha, but often in the Hystricomorpha 
and Lagomorpha. Anomalurus has no scalenus ventralis or 
anticus, as this muscle is usually called ; it has, however, a scalenus 
longus passing to the anterior four ribs and, dorsad to that, a 
scalenus brevis inserted into the first rib. 


Muscles of the Anterior Extremity. 


The Pectoral Muscles.—I propose to adopt the same method 
of dividing this group that I have found to work fairly well in 
other rodents*. (a) The superficial manubrial fibres: these also 
rise slightly from the clavicle and pass superficially to all the 
other fibres to be inserted lowest of all, opposite the lower part 
of the insertion of the deltoid. (3) The greater portion of the 
muscle rises from the whole length of the manubrium and gladi- 
olus and passes, deep to the last, to be inserted into the pectoral 
ridge. (y) Theabdominal fibres, or Pectoralis quartus, rise from 
the linea alba as far back as the umbilicus and are inserted into 


* P, Z. 8. 1894, p. 259. 


MYOLOGY OF ANOMALURUS. e221 


the neck of the humerus. (é) The deep portion, or Pectoralis 
minor, rises from the 3rd, 4th, and 5th costal cartilages at their 
junction with the sternum, and is inserted into the head of the 
humerus and capsule of the shoulder. 

Subclavius.—This muscle is well developed, and is inserted into 
the posterior border of the outer third of the clavicle. Thisis the 
arrangement which is always present in sciuromorphine rodents. 
In the Hystricomorpha the subclavius is continued on to the 
spine of the scapula, covering the supraspinatus, and the whole 
muscle is called the sterno-scapularis. In the Myomorpha the 
arrangement is usually as in the Sciuromorpha. 

The Deltoid, Teretes, Supraspinatus, Infraspinatus, and 
Subscapularis have the typical sciuromorphine arrangement and 
resemble the same muscles in Scdwrus. 


Fig. 2. 


\ Z Veg 
Ls: li, W 
iN WN 
F))), SOSON 

h, Thane 


yt. SUBCUAN. 
I), s 


20 Mbt noo 


—= 


}() 
= SSS ~ 
: = SS 
< SSS SSN 
> SS . \\ 
<Q 
~ 3 


Ny, i 
“an 


= My 
Y} 4% re” 


i “tr;= - TENS -PAT 
ye 
Wore 
Wen, 
inne a 
ANY , 
\ Yj G 
iN =S 
i(X) 33 
WINN) 7 


Muscles of Patagium and Arm. 
Pect., Pectoralis; Bi., Biceps; Subclav., Subclavius; Tens. Pat., Coracce 
patagialis. 


322 PROF. F. G. PARSONS ON THE 


Flexor Longus Cubiti (Biceps).—This muscle has both heads 
well developed, it is inserted into the radius. 

Flexor Brevis Cubiti (Coraco-brachialis):—The frequency with 
which all three parts of this muscle are present seems to be one 
of the characteristics of sciuromorphine rodents, and Anomalurus 
is no exception to this rule. The three parts are closely blended 
near their origin, and the upper part or rotator humeriis sepa- 
rated from the middle, as is usual, by the musculo-cutaneous 
nerve. The middle and lower portions (medius and longus) are 
closely united in their whole extent and are inserted from the 
upper third of the humerus as far as the internal condyle. 

Coraco-patagialis—A. muscle with a small short fusiform belly 
and a long delicate tendon rises from the coracoid process, and 
is inserted into the patagium midway between the spur from the 
elbow and the trunk. Its action seems to be to act as a stay to 
the membrane, and keep its edge down during flight, thus: 
rendering the part of the patagium between the arm and the 
body more coneave and parachute-like. As I have never seen 
anything homologous with this muscle in other mammals, I have 
suggested the name “ coraco-patagialis”’ for it. 

Brachialis anticus.—This muscle resembles that of most sciuro- 
morphine rodents in having the inner and outer heads so closely 
connected as to be inseparable. 

The Extensor Cubiti (Triceps), Anconeus, and Epitrochleo- 
anconews call for no special remark. 

Latissimo-olecranalis (Dorso-epitrochlearis).—This rises from 
the tendon of the latissimus dorsi as well as from that of the 
teres major. It is inserted into the inner side of the olecranon 
process. 

Serratus ventralis (Serratus magnus and Levator anguli 
scapulx).—This continuous sheet rises from the transverse pro- 
cesses of the posterior five cervical vertebra and from the anterior 
ten ribs; it is inserted as usual into the vertebral border of the 
scapula. I only know two other rodents, Georychus and Bathy- 
ergus, in which the rib origin is so extensive. 

The Pronator Radii Teres is inserted below the middle of the 
radius ; it rises, as in Sccurus and Pteromys, from the lower part 
of the arch over the supracondylar foramen. 

The Flexor Carpi Radialis and Palmaris Longus have the usual 
human attachments and relations. 


MYOLOGY OF ANOMALURUS. 323 


The Flexor Sublimis Digitorum divides into three tendons for 
the index, medius, and annularis. There is no slip for the 
minimus; and I have called attention to this arrangement * as 
constantly occurring in the Myomorpha. 

In the Flexor Carpi Ulnaris both olecranal and condylar heads 
are present, they join high up, andthe usual insertion into the 
pisiform bone occurs. 

The Flexor Profundus Digitorum has the five parts of the 
typical muscle—1, radial; 2, ulnar; 3, radio-condylar; 4, ulno- 
condylar ; 5, centro-condylar. The latter joins the rest of the 
muscle about the wrist-joint. 

The Lumbricales are peculiar; there were six in each hand 
in my specimen. Que rose from each side of the two middle 
tendons, one from the ulnar side of the tendon to the index, and 
one from the radial side of the minimus tendon. They were too 
small for the nerve-supply to be made out with certainty. 

The Pronator Quadratus is only attached to the lower quarter 
of the forearm. The Sciuromorpha are remarkable for the feeble 
development of this muscle. 

The Supinator Longus is absent. Up to the present I have 
looked upon the absence of this muscle as a myomorphine 
characteristic. 

The Hatensor Carpi Radialis, Hxtensor Carpi Ulnaris, and 
Extensor Ossis Metacarpi Pollicis; have the usual mammalian 
attachments. 

The Extensor Longus Digitorum divides into four tendons, as in 
all Sciuromorpha; often, in the Myomorpha, the slip to the 
minimus is absent. 

The Hutensor Minimi Digiti divides for the annularis and 
minimus. 

The Extensor Indicis only goes to the index. 

The thumb is too rudimentary to require any short thumb- 
muscles. 

The first row of palm-muscles consists of an Adductor Indicis 
and Adductor Minimi Digiti, which have the usual rodent dispo- 
sition. Deep to these is a layer of double-headed Flexores breves, 
one for each of the four digits, and no muscles dorsad to these 
were found. 


* P. Z. 8. 1896, p. 188. 
t The pollex is rudimentary. 


324 PROF. F. G. PARSONS ON THE 


Muscles of the Posterior Extremity. 

Sartorius and Tensor fascia femoris.—These two muscles are 
feebly developed even for a rodent. 

Ecto-gluteus and Oaudo-femoralis (Agitator caudz).— When I 
described the muscles of the Sciuromorphine and Hystricomor- 
phine Rodents in 1894, I had not learned to draw any distinction 
between these two. The fact, however, was recorded that “in 
Sciurus, Spermophilus, and Pteromys the gluteus maximus (ecto- 
gluteus) is inserted by two slips, one into the third trochanter, 


Fig. 3. 


\ 

Satie 

WV Sz 
kL- 


-TENUISMS_ 


Muscles of Thigh (outer side). 
8.T., Semitendinosus; G.M., Ectogluteus; A.C., Caudo-femoralis (Agitator 
caudee); Bi., Flexor cruris lateralis (Biceps). 


the other into the lower part of the femur.”’ I have now no doubt 
that the upper of these is the true ecto-gluteus, the lower the 
caudo-femoralis or agitator caude. Anomalurus resembles these 
animals in the low insertion of the latter muscle ; indeed the ecto- 
gluteus and caudo-femoralis together are inserted into the whole 
length of the femur (see fig. 4.) 

The Meso-gluteus, Ento-gluteus and Gluteus ventralis (Scanso- 
rius) were present ; the latter I have not hiterto seen in sciuro- 


es 


MYOLOGY OF ANOMALURUS. 325 


morphine rodents, but it is seldom such a distinct muscle as 
to be unmistakable. 

Nothing wortby of special remark was seen in the examination 
of the short rotator muscles of the hip. 

The Biceps femoris (Flexor cruris lateralis) has only one head, 
and that from the tuberosity of the ischium (fig. 4). It is 
inserted into the fascia of the upper third of the leg. In most 
rodents a superficial head from the spines of the anterior caudal 
vertebre is also present, but in Sciwrus and Pteromys no head 
from this origin was found. 

The Zenuissimus (Bicipiti accessorius) rises from the sacral 
vetebre deep to the ecto-gluteus and runs down to be inserted 
with the posterior fibres of the biceps (fig. 3). 

The Semitendinosus, as is usual in rodents, rises by two heads, 


ii} 
Sp SA Bre! | 
ti; Bi.cut 


\ cc 


f—- ~~ —-C.FEM. PROF. 


Deep Muscles of Thigh. 
The references in this figure explain themselves. 
one from the anterior caudal vertebra, the other from the tuber 
ischii; these join in the upper third of the thigh, and are inserted 
into the junction of the upper and middle thirds of the internal 
surface of the tibia. 


326 PROF. F. G. PARSONS ON THE 


When the semitendinosus, caudo-femoralis, and ecto-gluteus are 
cut and reflected, the great sciatic nerve is exposed. Deep to 
this is a muscle which rises from the same origin as, but deep to, 
the semitendinosus, and after passing obliquely downward and 
forward, is inserted into the middle of the femur. Probably the 
name of caudo-femoralis profundus would best describe this 
(see fig. 4). 

The Semimembranosus rises only from the tuber ischii, as is 
usual with rodents, and is inserted into the tibia deep to the 
internal lateral ligament of the knee. 

The Presemimembranosus is quite distinct from the last and 
is closely connected with the adductor mass. This I have already 
pointed out (P. Z. 8. 1894, p. 286) is a sciuromorphine charac- 
teristic. 

The Quadratus Femoris is entirely fleshy, as in other Sciuro- 
morpha. 

The Pectineus is inserted just below the lesser trochanter of 
the femur. . 

The Adductor longus continues the plane of the last downward 
and is inserted just below it into about a third of the femur. 

The rest of the Adductor mass rises from the ramus and tuber 
ischii, and is inserted into the lower two-thirds of the shaft of 
the femur. 

The Gracilis (Adductor cruris) is a single broad muscle rising 
from the symphysis and ramus of the pubes and being inserted 
into the upper third of the shaft of the tibia. This single gracilis 
has been shown to be characteristic of the Sciuromorpha as 
opposed to the Myomorpha. 

The Tibialis anticus rises from the tibia only, and is inserted 
into the innermost of the five metatarsal bones. 

The Extensor Longus Digitorwm has the usual femoral origin 
and is inserted into the outer four toes. 

The Extensor Proprius Hallucis rises from the middle third of 
the fibula, and is inserted into the terminal phalanx of the hallux. 
The full complement of Peroneal muscles is present; viz., Pero- 
neus longus, brevis, quarti digiti and quints digiti. 

The Gastrocnemius and Soleus have the usual rodent appear- 
ance, they form a twisted Tendo Achillis *. 


* See Journal of Anat. & Phys. vol. xxviii. p. 414. 


MYOLOGY OF ANOMALURUS. 327 


The Plantaris expands in the sole into a muscular Flewor 
Brevis Digitorum. 

The Flexor Tibialis does not join the Flewor Fibularisin the 
sole. This is a sciuro- and myo-morphine characteristic. 

The Accessorius is present, and is inserted into the plantar 
surface of the flexor fibularis. In the Myomorpha the acces- 
sorius is absent, but it is usually present in the Sciuromorpha. 

The Lumbricales closely resemble those of the fore limb, but 
there are seven instead of six; one rises from each side of the 
tendons to the index, medius, and annularis toes, while the seventh 
comes from the fibular side of the tendon to the hallux. 

The Deep Muscles of the Sole are arranged in two layers: the 
superficial consists of an oblique adductor hallucis and minimi 
digiti, while the deeper is formed by the double flexores breves 
to each toe. The abductor hallucis and minimi digiti are the 
enlarged and somewhat displaced marginal members of this series. 


Having given a brief survey of the myology of Anomalurus, 
it remains to contrast these muscles with those of other rodents, 
and to see whether they throw any light on the position and 
relationship of this animal in the order. In the first place, it 
will be well to review the chief myological characteristics of the 
four suborders, and contrast them with those of Anomalurus. 

From experience gained in former dissections, 1 believe that 
the following are the chief myological characteristics of the 
Sciuromorpha :— 

1. The anterior deep part of the masseter lies in a groove in 

front of the zygomatic process of the maxilla. 

2. The digastric muscles have a central tendon from which a 
fibrous areade stretches across the middle line; to this the 
anterior bellies are attached and they are in contact in 
the mid-line of the chin. 

. The transverse mandibular muscle is usually present. 

. The omo-hyoid is present. 

5. The omo-trachelian muscle (levator clavicule) always rises 
from the atlas. 

6. The subclavius is never continued over the supraspinatus as 
a sterno-scapularis muscle. 

7. The coraco-brachialis brevis is always present, as are 
usually the medius and longus. 


He OO 


328 PROF. F. G. PARSONS ON THE 


8. The supinator longus is generally present. 
9. The pronator quadratus is never attached to more than the 
lower third of the forearm. 

10. The presemimembranosus is closely connected with the 
adductor mags, instead of being separate or fused with the 
semimembranosus. 

11. The gracilis is a single muscle. 

12. The flexor tibialis does not join the flexor fibularis in the 
sole. 

13. The accessorius pedis is present. 

14. The rhomboideus capitis forms a continuous sheet with 
the other rhomboids. 

15. The rectus ventralis (abdominis) does not decussate with 
its fellow at its origin from the symphysis pubis. 


The foregoing are not all found in the Sciuromorpha alone, 
though all are points of distinction between the Sciuromorpha and 
one or more of the other three suborders. It is only by taking 
a large number of characters that one can hope to neutralize the 
effects of individual variation or to reduce its disturbing influence 
to a minimum. 

It will be noticed that Anomalurus agrees with the Sciuro- 
morpha in the following points :—Nos. 2, 8, 5, 6, 7, 9, 10, 11, 12, 
13, 14, and 15, but differs from them in Nos. 1, 4, and 8. 

In reviewing the myological characteristics of the Myomorpha 
it will be evident that in a great many points they resemble the 
Sciuromorpha. 


The following are characteristics of the Myomorpha:— ~* 

1. A certain amount of the masseter always passes through the 
infraorbital foramen. | 

. The digastric muscles have the sciuromorphine arrange- 
ment. 

3. The transverse mandibular muscle is present. 

4, The omo-hyoid is present. 

5. The omo-trachelian always rises from the atlas. 

6 

7 


i) 


. The subclavius does not usually form a sterno-scapularis. 
- The coraco-brachialis brevis is seldom present, and the 
three parts of the muscle never occur together. 
8. There are always two heads to the biceps cubiti. 
9. The supinator longus is absent. 


Ww) 


MYOLOGY OF ANOMALURUS. 329 


10. The presemimembranosus is quite distinct from the 
adductor mass. . 

11. The gracilis is usually double. 

12. The flexor tibialis does not usually join the flexor fibularis 

in the sole. 

13. The accessorius is absent. 

14. The rhomboideus capitis is usually distinct from the rest 

of the rhomboid sheet. 

15. The rectus ventralis often decussates with its fellow at its 

origin. 

Anomalurus agrees with the Myomorpha in the following 
characters—Nos. 1, 2, 3, 5, 6, 8, 9, 12. But it must be borne in 
mind that Nos. 2, 3, 5,6, 8, and 12 are points which are common 
to both Sciuromorpha and Myomorpha. 

From a comparison of the foregoing, it will be seen that in the 
greater number of characteristic muscles Anomalurus agrees with 
both the Myomorpha and Sciuromorpha, but that, where these 
muscles vary in the two suborders, it agrees with the Sciuro- 
morpha in six and with the Myomorpha in two. 

The six points which mark its sciuromorphine affinities are :— 


1. The presence ofall three parts of the coraco-brachialis. 

2. The close connection of the presemimembranosus with the 
adductor mass. 

The single gracilis. 

The presence of the accessorius pedis. 

. The rhomboids forming one sheet. 

6. The non-decussation of? the rectus ventralis. 

Of these I regard Nos. 1, 2, 3, and 4 as of great importance. 


or sR 99 


The two points of myomorphine affinity are :— 

1. The passage of a small part of the masseter through the 

infraorbital foramen. 

2. The absence of the supinator longus. 

The second of these is a negative point, and probably too much 
stress should not be laid upon it; but the first is a point 
of great importance and to my mind shows undoubted myo- 
morphine tendencies. 


With regard to the Hystricomorpha there is no need to go into 
so much detail because this suborder has some very sharply 


330 PROF. F. G. PARSONS ON THE 


marked characteristics, aud without the presence of some of these 
no animal could be said to have hystricomorphine tendencies. 

The chief of these characteristics are :— 

1. A large part of the masseter passes through the infra- 
orbital canal. 

2. The depressor mandibule (digastric) has no well-marked 
central tendon, and the anterior bellies are not in contact 
in the mid-line. 

3. The subclavius is continued past the clavicle to the supra- 
spinosus fossa to form a sterno-scapularis muscle. 

4. The flexor tibialis joins the flexor fibularis in the sole. 

Anomalurus differs from the last three entirely, and from the 
first in degree, but this is a point in which the Myomorpha 
approach the Hystricomorpha. 

Tt will have been noticed that, in the absence of the omo-hyoid 
Anomalurus differed from both the Sciuromorpha and Myo- 
morpha, while in the Hystricomorpha it must be stated that the 
omo-hyoid may be present or absent. This may be looked upon 
as a feeble hystricomorphine tendency, but it is just as strong 
or as feeble a lazomorphine one. 


It seems unnecessary to take up space by a detailed com- 
parison of the muscles of the Lagomorpha with those of Anoma- 
lurus ; firstly, because no one has suggested any relationship 
between the two, and, secondly, because I have never had the 
opportunity of completing my knowledge of the myology of this 
suborder by the dissection of a Pica. A general idea of the 
relationship of Anomalurus to the four suborders of rodents as 
illustrated by their muscles, may be gained by referring to the 
accompanying table, and, in criticising it, it should be borne 
in mind that the muscles have not been selected with any 
reference to Anomalurus. They were chosen as the result of 
previous dissections of many rodents, because they were found 
to vary with the different suborders. It cannot be too strongly 
insisted on that, if muscles are used for classificatory purposes, 
they should not be chosen haphazard, nor should one or two be 
used as crucial tests; but, by careful comparison of a large 
number of animals, some fifteen or twenty muscles should be 
picked out which vary more or less constantly with the different 
groups. The whole of these muscles should then be examined 
in the animal whose affinities are to be tested, and the average 


301 


MYOLOGY OF ANOMALURUS. 


——_—$— S$ —————— ——————— ———————  — — — —————i sv SSSSSSSSSSSSSSSSSSSSSSSSSSSSSSSSSSSSSSSSFFe 


17. 


Anterior deep part of 
Masseter. 
Anterior bellies of 


Digastric. 
Transversus mandibule. 
Omo-hyoid ............... 
Omo-trachelian (Levator 
‘clavicule). 
Rhomboids ............... 


Splenius colli ............ 
Recti ventrales (abdom- 
inis) origin from 
pubes. 
Sterno-scapularis..... ... 


Coraco-brachialis brevis. 
Biceps cubiti ..... ...... 
Supinator longus......... 
Pronator quadratus ... 


Presemimembranosus... 
(EnACHbE  soooccooon00000000 
Flexor tibialis and 


Flexor fibularis. 
Accessorius pedis......... 


Sciuromorpha. 


In groove. 
In contact, 


Usually present. 
Present. 
Always from atlas. 


Continuous sheet. 
Absent. 
Separate. 


Only subclavius. 


Always present. 

Two heads. 

Generally present. 

Lower 4 of forearm or 
less. 

Fused with the Adduc- 
tor mass, 

Single. 

Do not join in the sole. 


Present. 


Anomalurus. 


Through infraorbital 
foramen (small). 
In contact. 


Present. 
Absent. 
From atlas. 


Continuous sheet. 
Present. 
Separate. 


Only subclavius. 


Present. 
Two heads. 
Absent. 
Lower +. 


Fused with the Adduc- 
tor mass. 

Single. 

Do not join in the sole. 


Present. 


Myomorpha. 


Through infraorbital 
foramen (small), 
In contact. 


Present. 
Present. 
Always from atlas. 


R. capitis separate. 
Absent. 
Frequently decussate. 


Usually only sub- 
clavius. 

Rarely present. 

Two heads. 

Absent. 

Variable. 


Separate. 


Double. 

Do not usually join in 
the sole. 

Absent. 


FHystricomorpha. 


Through infraorbital 
foramen (large). 
Separate. 


Absent. 

Present or absent. 

Sometimes from basi- 
occipital. 

Continuous sheet. 

Sometimes present. 

Sometimes decussate, 


Sterno-scapularis 
present. 

Rarely present, 

One or two. 

Generally absent. 

Often to 4 forearm or 
more, 

Separate. 


Often double. 
Join in the sole. 


Present, 


Lagomorpha. 


Absent. 
Separate. 


Absent. 
Absent... 
From basioccipital. 


Continuous sheet. 
Present. 
Separate. 


Sterno-scapularis 
present. 

Present. 

One head. 

Absent. 

Absent. 


Separate. 


Single. 
Do not join in the sole. 


Absent. 


832 PROF. F. G. PARSONS ON THE 


result will, in my experience, show a decided indication towards 
the group with which that animal is most nearly connected. 

Asa result of the application of this process to Anomalurus, 
I should say that its affinities are decidedly sciuromorphine in the 
main, though it shows certain definite myomorphine character- 
istics. It is extremely difficult to give an idea of the relationships 
of animals in a linear manner, and I submit a diagram giving my 
present ideas of the position of Anomalurus. 


Fig. 5. 


LAGOMORPHA 


HYSTRICOMORPHA 


SCIURO- 
MORPHA 


maturi dz 


MYOMORPHA 


Diagram of the affinities of Anomalurus. 


Winge* has urged that Anomalurus is closely connected with 
Pedetes. Fortunately I have lately had the opportunity of 
dissecting the latter animal, and have come to the conclusion 
that it was rightly placed by Oldfield Thomas + between the 
Hystricomorpha and Myomorpha. It may be advantageous to 
apply the same muscles used for determining the position of 
Anomalurus for a comparison of the two animals. 


* “ Jordfundne og nuleyende Gnavere fra Lagoa Santa,” E Mus. Lundii, 
iii, 1887. 
Tt P. Z. 8. 1898, p. 858. t P. ZS. 1896, p. 1012. 


MYOLOGY OF ANOMALURUS. 335: 


Comparison of Muscles of Anomalurus and Pedetes. 


Anomalurus. Pedetes. 
1. Anterior deep part of | Through infraorbital | Through infraorbital 
Masseter. foramen (small). foramen (large). 
2. Anterior bellies of | In contact. In contact. 
Digastric. : 
3. Transversusmandibulz.| Present. Absent. 
4, Omo-hyoid ..............- Absent. Absent. 
5, Omo-trachelian ......... From atlas. From atlas. 
Gaehomlboidsiersssssescee Continuous sheet. Continuous sheet. 
7. Splenius colli .-.....5...: Present. Present. 
8. Recti ventrales (abdom-| No decussation. No decussation.. 
inis) origin from pubes. 
9. Sterno-scapularis ...... Subclavius only. Subclavius only. 
10. Coraco-brachialis brevis.) Present. Absent. 
WIS Biceps cubity ss-..seeese: 2 heads. 2 heads. 
12. Supinator longus......... Absent. Absent. 
13. Pronator quadratus ...| Lower 3. Lower 4. 
14, Presemimembranosus...) Fused with adductors.| Fused with semimem- 
branosus, 
Us), Gmeroilish, “Geecgsaaneenocdocs Single. Single. 
16. Flexor tibialis and | Do not join in the sole.} Do not join in the sole. 
Flexor fibularis. 
17. Accessorius pedis ...... Present. Absent. 


In these 17 points there are 12 in which the arrangement is 
the same in both animals, but it must be remembered that many 
of these simply denote myomorphine tendencies on the part of 
both ; and in the five points in which they differ (Nos. 1, 3, 10, 
14, and 17) the divergence is always towards the Hystricomorpha 
on the part of Pedetes, and the Sciuromorpha on the part of 
Anomalurus. 

There are, however, three points (Nos. 4,7, and perhaps 15) in 
which both animals wander away in the same direction from the 
arrangement which one might have predicted for them: it may 
be worth while considering these points in detail. 

The first is the case of the omo-hyoid, which is absent in both 
animals, although it is constantly present in the Sciuromorpha 
and Myomorpha. In the Hystricomorpha it is present or 
absent, and its absence in Pedefes does not surprise us when the 
many hystricomorphine affinities of that animal are remembered. 
Its absence in Anomalurus is unexpected, and may be the 
result of an individual variation in the animal I dissected, or 

LINN. JOURN.—ZOOLOGY, VOL. XXVII. 25 


304 SIR JOHN LUBBOCK ON 


it may be that I have framed the generalization that the omo- 
hyoid is constantly present in sciuromorphine and myomorphine 
rodents on the observation of insufficient material. To the 
presence of the splenius colli, I am not disposed to attach too 
greatimportance. When the splenius capitis is largely developed 
there is not room for all the fibres to be attached to the skull, 
and some of the posterior ones become inserted into the transverse 
processes of the anterior cervical vertebre to form the splenius 
colli; still in the Sciuromorpha and Myomorpha this muscle is of 
rare occurrence. 

The single gracilis is capable of another explanation than that 
of pointing to a relationship between Pedetes and Anomalurus. 
The muscle is not constantly double even in the Myomorpha, 
while in the Hystricomorpha it is more often single than double, 
and in the Sciuromorpha always single. We should, I think, 
expect that animals on the sciuromorphine or hystricomorphine 
borderland of the Myomorpha would be more likely to have a 
single than a double gracilis. 

Similarities between Anomalurus and Pedetes in any one of 
the three muscles discussed would have been hardly worthy of 
notice, and it has been shown that no one of them is by itself 
of first-rate importance, but the three occurring together do: 
perhaps furnish a somewhat feeble plea fora connection between 
the two animals. Possibly Winge’s and Oldfield Thomas’s views 
might be brought more into harmony by the use of a diagram 
such as I have suggested (fig. 5, p. 332), in which Anomalurus 
and Pedetes are not so very far asunder. 


On some Australasian Collembola. By the Right Hon. 
Sir Joon Luszzock, Bart., M.P., F.R.S., LU.D., F.L.S. 


[Read Ist June, 1899.] 


Tur following paper contains descriptions of some new species 
of Collembola from New Zealand and Tasmania, kindly sent me 
by Mr. A. Dendy of Christchurch, N.Z. The collection also 
comprised some specimens, representing one or two other 
species, which however were not in a condition enabling me to 
describe them satisfactorily. They have been for some three 


SOME AUSTRALASIAN COLLEMBOLA. 339 


years in my possession, and I have tried, but in vain, to obtain 
more, in hopes of being able to give a more complete list of 
Australasian Collembola. Possibly the publication of this paper 
may induce entomologists to devote some attention to this in- 
teresting, though inconspicuous, group. 


ANOURA TASMANIA, sp.n. (Fig. 1.) 


Body long elliptical, dark purplish. Ocelli on a short pear- 
shaped bulb, about ten in number, on a reddish-black granular 
pigment, which covers most of the bulb, leaving only narrow 
interspaces. The antepenultimate segment of the antenna is 
quite short and almost covered by the overlapping edges of the 
2nd segment. The body-segments present a series of arching 
mamunill, and the skin is covered with scutellated, stiff, curved, 
whip-like hairs. Foot without tenent-hairs. Claw single, 
without teeth. 

Length ‘25 in. Breadth 1 in. 

Hab. Tasmania ; found under a fallen log (A. Dendy). 


Anoura tasmanie. X 10. 
The colouring-matter is arranged in meshes, leayine more or 
8 . . . =) 
less circular colourless spaces. The skin itself is covered with 
minute granules and rather long whip-like hairs. 


336 SIR JOHN LUBBOCK ON 


This species is closely allied t0 our Anowra muscorwm, but is 
much larger, and differs in the number and arrangement of the 
mamumille, of the eyes, and of the stiff sete. 


Awnoura Denpyi, sp.n. (Fig. 2.) 

Body long elliptical, darkish purple; sides and back bearing 
a number of tapering projections (fig. 4), coloured at the base like 
the rest of the body, white towards the ends. Foot (fig. 5) without 
tenent-hairs ; claw with a small tooth. 

Length ,4 in. Breadth 3; in. 

Hab. Tasmania (A. Dendy). 


Anoura Dendyi. x 8. 


The head has one spine in the centre of the upper part, the 
next segments have each a pair, one being at the margin, the 
other a little way from it. The second, third, and fourth 
abdominal segments have a pair of smaii processes, one on each 
side of the median line. 

The terminal segment of the antenna bears (1) scattered, 
curved sete ; (2) numerous, close, very short hairs ; (3) numerous 
close, broader, and somewhat conical bodies, probably sense- 
organs. 


a  _- 


SOME AUSTRALASIAN COLLEMBOLA. 300 


Figs. 8, 4, & 5. 


S 


Anoura Dendyi.—8. Section through the tip of the antenna, x 200. 4, One 
of the processes, X 250. 5. Foot of Ist pair, x 250. 


Fie. 7. 


Anoura spinosa, dorsal aspect. X12. Anoura’spinosa, ventral aspect. x12 
sp p ‘ pi 


338 MR. W. T. CALMAN ON THE CHARACTERS 


ANOURA SPINOSA, sp.n. (Figs. 6 & 7.) 

Body elliptical, bluish purple; body covered with spines, 
bluish purple at the base, and the rest yellowish brown. Skin 
granular. Antenne four-jointed ; basal segment large, but 
short ; second segment longer; each with 1 large and 2 or 3 
smaller spines; third segment small, without spines; fourth 
segment longer, also without spines. Feet without tenent-hairs ; 
¢law with a small basal tooth, but covered nearly to the tip with 
granules resembling those of the skin. 

Length 2; in. Breadth 75 in. 

Hab. Near Hobart, Tasmania ; found under a decaying log of 
wood (A. Dendy). 


On the Characters of the Crustacean Genus Bathynella, 
Vejdovsky. By W. T. Catman, B.Sc., University College, 
Dundee. (Communicated by Prof. D'Arcy W. THompson, 
C.B., F.L.S.) 


| Read Ist June, 1899. ] 
(PLATE 20.) 


Iv a memoir on the fauna of certain wells, published in 1882 *, 
Prof. Vejdovsky described under the name Bathynella natans 
a minute crustacean of very remarkable characters, of which 
two examples were obtained from a well in the city of Prague. 
Prof. Vejdovsky placed it at the end of his memoir as a species 
imcerte sedis, merely noting its superficial resemblance to a 
minute gammarid and giving no further discussion of its syste- 
matic position. No additional specimens of this interesting 
form have been obtained, and I am not aware that, with the 
exception of a passing remark by Moniez, any reference has been 
made to it by subsequent writers or any opinion expressed 
regarding its affinities f. 


* ‘“Thierische Organismen der Brunnenwisser von Prag, 1882. I am 
indebted to my friend Dr. Chas. Chilton for calling my attention to this 
memoir. 

t Since the above paper was read I have learned that Prof. Vejdovsky 
published in 1898 a short paper in the Bohemian language dealing with the 
systematic position of Bathynella (S.B. K. bohm. Ges. Wiss., math.-nat. Cl. 


— 


OF THE CRUSTACEAN GENUS BATHYNELLA. 389) 


By the great kindness of Prof. Vejdovsky I have been enabled 
to make a re-examination of the unique existing specimen of 
Bathynella, which he has sent to me for the purpose. The 
specimen, which is mounted as a microscopic preparation in canada 
balsam, is, unfortunately, not in a good state of preservation, 
being much shrivelled and lacking the greater part of antennules 
and antennze as well as part of the uropods. The following 
account is taken from Prof. Vejdovsky’s description, supplemented 
in one or two points from my own observations. 

Bathynella (Pl. 20. fig. 1) is deseribed by Prof. Vejdovsky as 
resembling in general appearance the minute copepods of the 
genus Canthocamptus. The body is, in the present state of the 
specimen, about ‘75 mm. long*, and consists, according to the 
original account, of (1) a cephalic region bearing two pairs of 
antenne, mandibles, and at least two more pairs of mouth- 
appendages ; (2) a thoracic region of seven free somites each 
bearing a pair of biramous appendages; and (8) an abdomen of 
six somites, of which the first and second bear small appendages 
while the last carries two pairs of caudal plates. It appeared to 
me, on examining the original specimen, that there were indi- 
cations of an additional somite in the abdominal region, and on 
mentioning my belief to Prof. Vejdovsky, he informed me that 
he had actually seen this somite in the living animal and sent 
me one of his sketches in which it was plainly figured. The 
presence of this somite, as Prof. Vejdovsky himself remarked in 
sending me the sketch, is very important for the settlement of 
the systematic position of the animal, for it enables us to readjust 
the delimitation of the regions of the trunk in such a way as 
to bring their segmentation into precise accord with the normal 
arrangement for the Malacostraca. 

Prof. Vejdovsky implicitly referred the segmentation of the 
body to the type characteristic of the Arthrostraca, in which 
the first thoracic somite, bearing the maxillipeds, is fused with the 
head. Were this the case we ought to find four pairs of buccal 


1898, xiv. 2 pp.). The author informs me that he has there expressed the 
opinion “‘das Bathynella eine besondere Stelle zwischen den Arthrostraken 
einnehmen diirfte, und vielleicht eine neue Ordnung ‘ Pleopodophora’ reprii— 
sentirt.” 

* Vejdovsky gives the total length as about 1 mm. 


340 MR. W. T. CALMAN ON THE CHARACTERS 


appendages, namely, mandibles, first and second maxille, and 
maxillipeds. As a matter of fact, however, the closest scrutiny 
of the undissected specimen shows only two pairs of appendages 
(maxille) behind the mandibles, and I believe accordingly that 
the maxillipeds are represented by the succeeding pair of con- 
spicuous appendages, those, namely, which are borne upon the 
first free somite. Assuming that these represent the first 
thoracic appendages, the eighth and last pair will then be 
represented by what Vejdovsky has considered to be the first 
abdominal pair; that is to say, precisely those on which the 
male genital ducts appear to open, as in the great majority of 
the Malacostraca. And the additional abdominal somite, of the 
presence of which we are now assured, still leaves us with the 
normal number of six abdominal somites. 

The two pairs of antenne are short, composed of few joints 
not differentiated into peduncle and flagellum. The first pair, 
or antennules, are uniramous and consist of eight joints. The 
second pair are seven-jointed, and bear a small unjointed exopod, 

-on the second joint. There is no trace of eyes. 

The mandibles have a serrated cutting-edge and a strong two- 
jointed (or perhaps three-jointed) palp tipped with two stout 
curved spines. The two pairs of mawille are large, but nothing 
can be seen of their structure. 

The first seven pairs of thoracic limbs (Pl. 20. fig. 3) are of 
nearly uniform structure. The main axis of the limb in each case 
is six-jointed. The small coxal joint bears on its outer face a 
small flattened vesicular appendage interpreted as a branchial 
plate or epipod. The epipod appears to be borne on a short 
peduncle, from which it is separated by a transverse articulation 
or suture. The second or basal joint is long, and carries near its 
distal end an exopod *, composed of two joints in the case, pro- 
bably, of the first five pairs and of one joint on the sixth and 
seventh. The main axis of the limb is continued by a slender 
endopod of four joints, scantily supplied, like the exopod, with 
sete. I cannot now find an epipod on the first pair of thoracic 
limbs (maxillipeds), but it is figured by Vejdovsky, and these 
appendages do not appear to differ in any other respect from the 
succeeding pairs. The eighth segment behind the head, described 


* Vejdovsky, by an oversight, refers to this as ‘‘ein innerer Ast.” It is in 
fact, as his figures show, on the outer side of the limb. 


OF THE CRUSTACEAN GENUS BATHYNELLA. 341 


by Vejdovsky as the first abdominal, but here regarded as the last 
thoracic somite, bears a pair of short appendages. These are 
described as consisting of two branches, an inner two-jointed 
stem, and an outer “ penis-like process,” the latter being appa- 
rently connected with the male genital duct. The appendages 
in question are now, unfortunately, so shrivelled that nothing 
can be made out as to their structure, but the position of the 
«¢ nenis-like process” on the outer side of the limb is so unusual 
that we are led to suspect some error in this part of the 
description. 

The appendages of the first abdominal somite (second ab- 
dominal of Vejdovsky) are uniramous and two-jointed. The 
four succeeding somites appear to lack all trace of appendages. 
The last segment of the body bears terminally two pairs of 
“ Schwanzplatten,” a dorsal and a ventral. The ventral pair are 
described as consisting of two joints, each with a row of stout 
curved sete along its distal edge, and of a small unjointed 
exopod inserted near the distal end of the first joint. These 
appendages, of which only the basal parts are now preserved, are 
clearly the uropods or appendages of the sixth abdominal somite ; 
but it is not quite so easy to interpret the dorsal pair of “ caudal 
plates.” These might be regarded as the two halves of a deeply 
divided telson, such as occurs in many Amphipods, but their 
cylindrical form and the fact that each is very distinctly divided 
into two segments seem rather to suggest a comparison with the 
“caudal furca” characteristic of Phyllopods, Copepods, and 
other Entomostraca, and which also occurs in Nebalia and in 
some larval Malacostraca. . 

The presumed gonads, which have been already referred to, are 
described as a pair of tubular organs lying in the abdomen, each 
expanding in the last thoracic somite into a vesicular dilatation 
which in turn appeared to communicate with the “ penis-like 
process.” There can be little doubt that these organs constitute 
the male generative system. No other internal organ could be 
distinguished save a pair of oval “ glandular”’ bodies of unknown 
significance, lying in the last somite of the abdomen. 


Reviewing the characters here summarized, it seems plain, in 
the first place, that Bathynella, in spite of its minute size, must 
be referred to the Malacostraca. The number of the somites, 

LINN. JOURN.—ZOOLOGY, VOL. XXVII. 26 


842 MR. W. T. CALMAN ON THE CHARACTERS 


the position of the genital apertures, the characters of the 
thoracic limbs, and the presence of appendages on the terminal 
somite of the abdomen, appear to afford conclusive evidence on 
this point. The possibility that it may be a larval form, as 
Moniez * has suggested, is excluded if the identification of the 
genital apparatus be correct; but it may be remarked that, as a 
larva, Bathynella would be no less unique than as an adult 
crustacean. 

Assuming for the present that we have to deal with an adult 
form, it is clear that it cannot be received into any of the 
divisions of the Malacostraca as commonly defined. The appa- 
rent similarity to the Arthrostraca in the segmentation of the 
body, which led its discoverer to compare it to an amphipod, 
disappears when it is recognized that the first thoracic somite is 
free from the head and that the appendages carried by it are not 
specialized as maxillipeds but resemble the succeeding thoracic 
limbs. 

The presence of natatory exopods and of external epipods on 
most of the thoracic limbs are further characters not shared by 
any of the Arthrostraca. The other divisions of the Malaco- 
straca agree in possessing a carapace with which one or more 
of the thoracic somites are usually united dorsally. While some 
of them show isolated characters in common with Bathynella, 
as, for example, the free first thoracic somite of Nebalia and 
some Stomatopods, the natatory thoracic exopods of Schizopoda, 
Cumacea, and some Decapods, the undifferentiated maxillipeds of 
Euphausiide, and the plate-like epipods of Stomatopods, these 
resemblances are accompanied by differences so profound that 
there can hardly be any question of immediate affinity. 

While our ignorance of many points in the structure of 
Bathynella, more especially as regards the mouth-parts, pre- 
cludes for the present any definite conclusion as to its precise 
systematic position, the characters already ascertained suffi- 
ciently show that we have to do with a very peculiar and isolated 
type without close affinities to any of the recognized divisions of 
the Malacostraca. It appears to me, however, that some light 
may be thrown on its relationships by a comparison with the 
anomalous “ schizopod ” Anaspides tasmanie >of Mr. G. M. 


* Rey. Biol. du Nord de la France, i. 1888-89, p. 253. 


OF THE CRUSTACEAN GENUS BATHYNELLA. 343 


Thomson *, the characters of which I have lately discussed fT at 
some length, calling attention to its affinity with the Eel ESO 
Gampsonychide and their allies. 

Anaspides (P1. 20. fig. 2) agrees with Bathynella in having no 
carapace, in possessing natatory exopods and plate-like epipods 
on the thoracic limbs, and in the fact that the maxillipeds are not 
greatly different from the succeeding appendages. Like Bathy- 
nella, Anaspides was originally described as having eight free seg- 
ments in the thorax ; but I bave pointed out that the so-called 
first thoracic somite is marked off from the cephalic region not 
by an articulation but by a groove on the integument, and that 
this groove probably represents-the line of junction of the man- 
dibular somite with that of the first maxille. -Anaspides there- 
fore differs from Bathynella and agrees with the Arthrostraca, 
in possessing only seven free somites in the thoracic region. In 
many other characters Anaspides shows important differences 
from Bathynella: the antennules are biramous, there are two 
epipodial lamellae on each of the thoracic legs (Pl. 20. fig. 4) 
(except the last pair), the second or basal joint of the thoracic 
legs is unusually small, while it is large in Bathynella, the 
abdomen carries a complete series of appendages, the Eisen 
is simple and undivided. 

Assuming, however, as I think we may fairly do, that such cha- 
racters as the small size of Bathynella, the absence of eyes, the 
simple form of all the appendages and the reduced number of 
joints in most of them, are due to degeneration correlated with 
its subterranean habitat, there would seem to be adequate 
grounds for suggesting that Bathynella is at all events less 
distantly related to Anaspides than to any other existing 
Crustacean. 

Many of the characters in which Anaspides agrees with the 
Paleozoic Gampsonychide are also shared by Bathynella; and in 
some points, such as the presence of a free first thoracic somite 
and of only a single series of epipods, the latter may even 
approach some of the fossil forms more closely than does 
Anaspides. 


* Trans. Linn. Soc. (2), Zool. vi. (3). 
t Trans. Roy. Soc. Edin, xxxviii. pt. iv. no. 23. 


344 


ON THE GENUS BATHYNELLA. 


EXPLANATION OF PLATE 20. 


Fig. 1. Bathynella natans, Vejd. (Altered from Vejdovsky.) x 100. 

This figure was constructed from Prof. Vejdoysky’s published}figure 
and from observations on the preserved specimen. It conforms accu 
rately (except that the abdominal segments appear a little shorter) with 
an original sketch (taken from the living animal) subsequently sent me 
by the author. 

2. Anaspides tasmanie, G. M. Thomson. X38. (Original.) 
3. Third thoracic limb of Bathynella, (Original.) 
4, Third thoracic limb of Anaspides. (Original.) — 


Reference Letters. 


. First and second antennz. 

. Basal joint of thoracic leg. 

. Coxal joint of thoracic leg, 

. ‘Cervical groove” of Anaspides, marking off so-called “ first 


thoracic somite.” 


. Hye. 
. Endopod of thoracic leg. 
. Epipod of thoracic leg. 


Exopod. 
Caudal furca of Bathynella. (Dorsal caudal-plate of Vejdovsky.) 


. Supposed testis. 

. Mandibular palp. 

. First and second maxille. 

. “ Penis-like process” of last thoracic limb. 
. Telson. 

. Uropod. 

. First thoracic somite, bearing maxillipeds. 
. Second thoracic somite. 

. Last thoracic somite. 


Abdominal somites. 


Calman. - Linn Soc. Journ. Zoor. Vou. XXVII Pu. a0. 


W.T.C. del. M.P Parker lth. 


1,38, BATHYNELEA NATANS, Vesdovsky. 
2,4, ANASPIDES TASMANIA, Thoms. 


LINN.SOC. JOURN.ZOOL .VOL.XXVII. PL.21. 


(fie A5 4 AG a7 
—_————_——__—_ nnn Ein [—S=== ——————SSSS==— = re 


THE URMI LAKE BASIN 


(AZERBAIJAN) 
By R.T.GUNTHER M.A. 


Compiled partly from Russian and English maps 
and partly from route surveys by 
General Houtum Schindler. 
Scale of Miles 
20 40 
Natural scale 1:1,000.000 or 15:78 miles = Linch, 


Heights iw feet. 


nw? 2 
x \* 2 
he ee 


Contributions to the Natural History of Lake Urmi, N.W. 
Persia, and its Neighbourhood. By Roserr T. Giyruer, 
M.A., F.R.G.S., Fellow of Magdalen College, Oxford. 
(Communicated by the President.) 

[Read 15th June, 1899. | 


(Map Pu. 21 & Piarus 22-30.) 


CoNnTENTS. 
Page 
General Remarks. By Rosrrt T. Gtwrunr, M.A., F.R.G.S. ............ 345 
List of Animals distinguished by Syrian Names ..................sceececeeees Byer 


Reports on the Specimens collected :— 
The Wild Sheep of the UrmilIslands. By Dr. A. Gintusr, F.R.S., 


IBresusMimman SOC ea tnan tas caceniae cca neeaecswSeeaon elas cao eee 374 

The Pliocene Mammalia of the Bone-beds of Maracha. By 
Rosert T. Guntuer, M.A., F.R.G.S.  ..........cccecccccececcee 376 
Reptilia and Amphibia. By G. A. Bounzencsr, F.R.S., F.Z.S.... 378 
Fishes. By Dr. A. Gunrurr, F.R.S., PLS. ...........0ccsecesenees 381 
Land and Freshwater Mollusca. By Epear A. Smiru, F.Z.8. ... 391 
Orustacea. By Rosert T. Guwruer, M.A., F.R.GS................ 394 
Chilopoda and Arachnida. By R. I. Pococks ...................0.00. 399 
Acari. By Ausert D. Micuant, F.L.S., F.R.M.AS.  ............... 407 
Lepidoptera Rhopalocera. By A. G. ButnEr, Ph.D................ 408 
Lepidoptera Phalenz. By Sir Guoren F. Hampson, Bart. ...... 411 
Neuroptera and Diptera. By Rosert T. Gonruer, M.A. ...... 414 
Orthoptera. By Matcozm Burr, F.Z.8., F\ELS. .........000...00.00- 416 
Note on a Jurassic Ammonite. By G. C. Crick, F.G.S8. ......... 418 
Fossil Hchinoidea. By J. W. Gregory, D.Sc., B.G.S. ............ 419 
Fossil Corals. By J. W. Gregory, D.Sc., FIGS. .......0..0....00- 424 

Marine Tertiary (Miocene) Mollusca. By R. Burin Newton, 
BEGG eel attra Ae on RR Bi Ma ait ati aon ee 430 


Note on a Palxozoic Limestone. By R. Butten Newton, F.G.S. 452 


GENERAL REMARKS. 


Jr is now more than three years since Mr. P. L. Sclater showed 
me a letter from my friend the Rev. F. I. Irving, dated Urmi, 
Oct. 6, 1895, in which the organism dwelling in the Salt Lake of 
Urmi, which Lord Curzon of Kedlestone had previously referred 
to as a jellyfish, is compared to “a tiny shrimp.” Further 
searches in the literature showed that Abich had already recorded 

© presence of a Crustacean, but that the majority of travellers 
wud writers described Lake Urmi as a “ Dead Sea,” totally devoid 
of hic on account of its excessive salinity. 

I received private information that edible fish were to be caught 
in the rivers ; but I was unable to obtain any further information 

LINN. JOURN.—ZOOLOGY, VOL. XXVII. 27 


346 MR. R. 1. GUNTHER ON THE 


concerning the aquatic fauna of the Urmi basin, although its fossil 
fauna had been frequently described. 

The fascination of an apparently entirely unknown zoological 
region near the waterparting of the hydrographie systems be- 
longing to the Atlantic, Arctic, and Indian Oceans, induced we 
to devote a summer vacation to visiting the plateau of North- 
western Persia. 

The chief problems which attracted my attention were naturally 
those which centred around the salt lake and its fauna, and its 
tributaries and their faunas; but incidentally I was able to 
collect a few terrestrial animals, which are described in the Reports. 
The results obtained during a few weeks of summer show, I think, 
how much still remains to be done by a naturalist-explorer who 
may be able to work the Urmi district during those months of the 
year which are most favourable for the purpose of collecting. 

The present communication is chiefly concerned with observa- 
tions and collections made between the middle of July and the 
middle of September, 1898. Wherever I have profited by the 
work of others I have endeavoured to make due acknowledgment 
but, in addition, I should like to record my especial obligations 
to a more extended circle of helpers. Jam indebted to the Royal 
Society for a grant in aid of my expedition; to the Foreign Office 
for most useful letters of introduction ; to Prince Imam Guli 
Mirza, at the intercession of the Amir Nizam, for the loan of the 
‘Nahangk,’ a vessel of some 20-tons burden, upon the Lake of 
Urmi; to Mr. E. Chapman ; and to the past and present members 
of the Archbishop of Canterbury’s Mission to the Assyrian 
Christians at Urmi. Lastly, my sincere thanks are due to my 
friends and coadjutors, who have unselfishly helped on my own 
studies at the expense of their own, and who have contributed the 
most valuable part of the present paper. My readers are indebted 
to the Council of the Royal Geographical Society for permission 
to republish my map of the neighbourhood of Urmi, originally 
contributed to the Journal of that Society. 

Before proceeding to the more purely biological characteristics 
of the region, it may be well to recall to mind its main physio- 
eraphical features—its “ external conditions of existence.” 

The geographical and, more particularly, the hydrographical 
relations of Azerbaijan, the N.W. province of Persia, are of 
interest because the highlands of this province are part of the 
system in which the most important waterpartings of the Old 


NATURAL HISTORY OF LAKE URMI. 347 


World meet. The divides which separate three great systems of 
rivers—namely, the river-systems of the Indian Ocean from those 
of the Atlantic, the river-systems of the Atlantic from those of 
the Arctic, and the river-systems of the Arctic from those of the 
Indian Ocean—meet in the highlands of Armenia and Kurdistan, 
between the massif of Asia Minor and the Persian plateau. 
Notwithstanding the altitude of the region, the meteorological 
conditions tend towards those of a desert, and produce features 
ebaracteristic of deserts. On the whole the annual evaporation 
is so much in excess of the annual precipitation that the lakes 
do not overflow, and their water is more or less salt. Similarly, 
in’ Turkish territory we have Lake Van, in Russian territory 
the Caspian Sea, and in Persia Lake Urmi. 

Climate-—The climate of the Azerbaijan plateau may be fairly 
classed among the continental, or “ excessive” climates as they 
have been termed by Buffon; but at the same time it is the 
healthiest in Persia, because the summer heat is more tempered 
than in the other provinces and there is generally a sufficiency of 
water for man and beast. 

The seasons are extreme, but very variable both as regards 
duration and time of year. This variability is largely due to the 
fact that these uplands of N.W. Persia, of 4300 feet altitude and 
upwards, form part of a tongue of high land which runs from 
N.W. to 8.E., and divides the warm depression of Mesopotamia, 
in which the palm-tree flourishes, from the deserts of Persia and 
from the hot South-Caspian provinces of Ghilan and Mazenderan. 
Thus we find, in close proximity, the snow-mountains of Kurdistan 
and tropical lands warm enough for the growth of the date-palm. 
It is a pity that methodical meteorological observations in so 
interesting a region should be practically non-existent. In 
default of the more accurate indications of the meteorological 
conditions, | am compelled to supplement my remarks by such 
observations on animal and vegetable life as seem to afford 
information on the climate. 

The winter snows may commence towards the end of November 
(23rd in 1890), or they may not fall until February 1st (1893), 
There is often fine weather between the first snows and those of 
January and February. The last snows may fall as late as Lady- 
day. In cold January weather the minimum thermometer some- 
times falls below 0° Fahr. (12° F. is a common temperature), and 
the snow does not melt in the middle of the day. As a general 

Pate? : 


348 MR. R. T. GUNTHER ON THE 


rule, snow may be expected to lie on the ground in the plain for 
three or four months, and on the hills (6000 feet) in streaks until 
well into July. In the winter of 1895 there is said to have been 
no sunshine for five weeks. 

When spring comes the snow melts, often with surprising 
rapidity, and a few days’ warm sunshine produces a marvellous 
change in the aspect of the country. Irises of a delicate purple 
colour, and white and pink crocuses are in flower before the end 
of March ; the plum blossoms by the Ist of April, and sometimes 
even in March, with snow on the ground. The quince blossomed 
on April18thin 1891. At the same time the storks and hoopoes 
return from their winter pilgrimages; the earliest record of 
the stork at Urmi seems to be March 14th, in 1898. 

Summer commences about St. George’s day, on May 6th, and 
usually lasts unbroken until late in October; popular tradition 
prolongs it until the second St. George’s day in autumn, on the 
first Monday in November (O.8.). Rarely, as in 1895, a few 
rain-storms interrupt the continuity of the hot weather. 

During the warmest months it is hot even at 6 o’clock in the 
morning. The glaring, eye-dazing sky is unrelieved by any 
clouds. The shade temperature rises to 90° F. in the open, and 
much higher among the sun-baked mud-walls of the villages. The 
nights are generally cool, and become uncomfortably cold during 
the latter months of summer, owing to unchecked radiation. 
The greater number of the fruit-trees are in blossom in the first 
week of June. The first fruits to ripen are the white and morella 
cherries and plums, which are followed by peaches and apricots, 
and in August by an abundance of excellent nectarines, melons, 
grapes, and also of cucumbers, vegetable-marrows, black and red. 
tomatoes, &c. 

No climate would seem better adapted for the growth of the: 
vine. The grapes of the plain of Urmi are magnificent, both in 
flavour, size, lusciousness, and variety. On the other hand, the 
severe winters do not permit of the cultivation of the orange and 
the lemon. 

Rainfall.—The rainfall seems to have been recorded during 
the year 1853-54*, but I have been unable to discover any 
subsequent series of observations. During that year 547 milli- 
metres of rain seem to have fallen, and the monthly rainfall was. 


* Supan, in Petermann’s ‘ Mittheilungen,’ Erganzungsheft No. 124, 1898. 


NATURAL HISTORY OF LAKE URMI. 3849 


as follows :—January 43, February 72, March 103, April 183, 
May 62, June 11, July 0, August 12, September 16, October 38, 
November 24, December 33: total 547 millimetres. 

Irrigation and Agriculture.—On the Urmi plateau, wherever 
there is water there is agriculture. Without artificial irrigation 
' the greater part is a sterile waste of gravel, capable of supporting 
little but thistles, Centaurea, Carduus, Achillea, Pyrethrum, and 
other plants characteristic of the Persian steppes. In spring, 
the snow has hardly had time to melt away before this steppe- 
vegetation covers the plains and hillsides with a gyeen mantle; 
but soon the water in the soil commences to give out, and then 
the brown-burnt stalks and leaves demonstrate the cessation of 
vegetable growth. 

The chief objects of the agricultural engineer are first to 
convey water, with as little loss from evaporation as possible, from 
the valleys near their emergence from the hills to the most fertile 
soll of the plains, and, secondly, to distribute the water over as 
great an area as possible. The Persian poets praise King 
Menucheher as the inventor of the system of subterranean 
canals which have become so universal in Persia, and which 
have converted many a wilderness into a fertile plain. Sub- 
terranean aqueducts or kanats, sometimes many miles in 
length, which are employed to convey water beneath arid tracts 
of land, have a double advantage: they not only reduce loss from 
evaporation to a minimum, but, inasmuch as they are nearer 
the underground water-level, little of their water is lost by 
drainage into the ground. It is always curious to note how 
similar inventions have been independently made in different parts 
of the world. The ancient Peruvians constructed subterranean 
watercourses on a noble scale in connection with their gigantic 
works of irrigation (one traversing the district of Condesuyu 
measured 400 tv 500 miles), and, like the Persians, had laws 
prescribing the quantity of water which individual landowners 
might be permitted to draw (Prescott). The waters of the 
Persian kanats contain the ordinary chub, Capoéta, and Leuciseus 
of the rivers ; and the dark-vaulted spaces are often the abode of 
great numbers of bats. 

The chief crops cultivated are cereals, vines, castor-oil, tobacco, 
melons, rice, and cotton. ‘The fields are so level, and so well 
trenched, that they can be watered by a simple diversion of an 
irrigation channel. 


300 MR. R. T. GUNTHER ON THE 


Vegetation.—Perhaps the most noticeable feature of the 
vegetation of the Urmi plateau is the almost total absence of 
indigenous, uncultivated trees. And yet in some parts the 
landscape can only be described as thickly wooded. The villages 
are often completely hidden by trees grown for fruit or fuel. 
There is no lack of all manner of fruit trees: apples, pears, 
quinces, apricots, peaches, nectarines flourish in the gardens round 
the villages; willows and poplars line all watercourses ; planes, 
walnuts, and jujubes form avenues along roads; but all indi- 
genous trees have long since been cut down. It is not the case 
that the conditions would be unsuited to the growth of arboreal 
vegetation, because they are much the same as those prevailing 
in the oak-forest districts in Turkish Kurdistan. 

By roadsides and in villages the plane (Platanus orientalis, L.) 
is often planted, for the sake of the shade afforded by its spread- 
ing branches. Here and there a giant has been spared, and has 
become regarded as a holy tree. Chardin refers to one in a garden 
near Shiraz upon the branches of which amulets, rags, and other 
votive offerings were hung. The Rev. 8. J. Daltry observed a 
similarly decorated tree near the sulphur-springs on the hills to 
the north of the plain of Gavilan. There is an enormous plane 
in the village of Dekhargan growing over a water-tank, which is 
reached by some steps descending between the buttresses of the 
tree. Close by are three ancient Mussulman tombstones rudely 
carved like the quadrupeds of our Noah’s-arks. Beneath the 
same tree stands a stone bench used as a castor-oil press. It 
is a pity that more of these magnificent trees are not planted by 
the present generation, who, although ready enough to gain 
advantage by the good deeds of their forefathers, are too idle 
to follow their good example. 

I did not see any specimens of the Oriental plane or of the 
walnut (Juglans regia) which could unhesitatingly be described 
as wild or indigenous in the Urmi district. I am therefore 
inclined to consider that Dr. Radde’s* dictum that all planes and 
walnuts in the Caucasus have been planted by man, with the 
possible exception of the walnuts of Gilan, is equally true of the 
Urmi district. I saw a few walnut-trees on the islands of 
Koyun and Arzu, in the Lake, which might be regarded as 
indigenous. The largest was a stunted tree of about 15 feet 


* G. Radde, ‘ Reisen an der Persisch-Russischen Grenze,’ 1886. 


NATURAL HISTORY OF LAKE URMI. 301 


in height, with a stem 2-8 feet in diameter near its base. It 
is, however, equally possible that the trees were planted when 
the islands were inhabited and joined to the mainland. Formerly 
the larger islands seem to have been fairly thickly wooded, 
but the fuel-collector bas not only cut down the trees with an 
unsparing axe, but now the sailors of the ships earn a living by 
eradicating all trace of their existence by laboriously digging- 
out their very roots. Walnut-wood is in great request for the 
best joinery and cabinet-making. 

The banks of the watercourses are recognizable from afar on 
account of the poplars and willows planted along them. The 
willows are generally pollarded at regular intervals of time. 
The long straight poplar poles are used as rafters for the roofs 
of houses and balconies. Split laths are laid transversely from 
rafter to rafter, matting is laid on them, then hay, and finally a 
thick layer of mud and chopped straw. 

Geology of Urmi Basin.—The Azerbaijan lake-basin occupies an 
area of some 20,000 square miles; its greatest depression is more 
than 4000 feet above sea-level. The mountains on its periphery 
separate its water-system from the circumjacent basins of the 
Tigris, the Aras, and the Kizil Uzun, the latter two of which 
flow into the Caspian Sea. Several peaks rise above 10,000 feet, 
and the voleanic cone of the Savalan reaches 15,000 feet. Out- 
crops of voleanic rock occur in many localities, and probably 
had their origin during the vast upheaval of land which occurred 
after the Miocene age. 

The geological record of Lake Urmi is still but very partially 
read. The oldest rocks are Paleozoic and probably of Car- 
boniferous Limestone age, because the genus Hndothyra is to 
be detected among their Foraminifera (p. 452). The Jurassic 
age is represented by a solitary Ammonite in my collections 
(p. 418); but Rodler, von Borne, and others have noted several 
localities near the west coast of the Lake of Urmi where Jurassic 
rocks are to be found. For the paleontology of the later 
deposits, I would especially refer the reader to Mr. Bullen 
Newton’s comprehensive retrospect on p. 430. 

The Carboniferous rocks of the Lake itself are overlaid by a 
ereat thickness of Miocene chalk, an important section of which 
ig referable to the Helvetian period, and which contains the 
remains of organisms identical with those of the Lower Miocene 
(pp. 480-452). The North Persian Tertiary rocks are divisible 


302 MR. R. T. GUNTHER ON THE 


into three well-characterized equivalent zones, which have been 
defined by Pohlig as :— 

Zone 1. The marginal zone of sandstones, conglomerates, and 
gravels, often containing organic remains from earlier 
strata. The Seir hill probably belongs to this zone. 

Zone 2. Light-coloured marls, generally bright red, with beds 
of rock-salt and gypsum. Both zones 1 and 2 take a large 
part in the formation of the hills of the Persian plateau. 

Zone 3. Limestones and caleareous marls; best seen in the 
islands in the Lake of Urmi, but also in the mountains to 
the south-west, and which are very rich in molluses, reef- 
building corals, and silicious sponges. 

At Guverchin Kala, near the northern end of the lake, the 
entire series of Miocene strata rest almost horizontally and 
directly on red granites traversed by felspar-dykes, which are a 
very conspicuous feature in the landscape. 

Urmi must therefore have been covered by a sea of normal 
salinity in Miocene times, which was apparently an extension of 
the Miocene Mediterranean Sea. ‘Then commenced those vast 
changes of level which resulted in the upheaval of a Miocene 
sea-bottom some 5000 feet to its present position as part of the 
Persian plateau ; this vast earth-movement is still clearly recorded 
by the numerous volcanoes and outbursts of volcanic rock which, 
monument-like, mark the spots where the earth-crust gave way 
before overwhelming strains and irresistible pressures. The 
next chapter in the story of Urmi tells us how at a later date the 
dry land about Maragha supported a mammalian fauna, which 
must have been very similar to the almost contemporaneous fauna 
of Samos and of Pikermi in Greece. The bones of herds of 
elephants, antelopes, and horses, as well as those of many other 
mamials, are to be found preserved in the pumiceous tufa- 
deposits of Kirjawa near Maragha (p. 376); it is still uncertain 
whether or not the animals perished in consequence of a volcanic 
eruption cr from some other cause. 

During Pliocene and post-Pliocene times the land undoubtedly 
underwent considerable changes both of level and of contour. 
There is no evidence that the present lake is the remains of a 
Miocene sea, and therefore it is not to be regarded as a “ Relikten- 
See,” a view which is also shared by Rodler. The excellent Ameri- 
can monograph on Lakes Bonneville and Lahontan is an example 
of how the physical geographer has been able to reconstruct 


NATURAL HISTORY OF LAKE URMI. 353 


former lake-basins by an examination of terraces and of sinter 
and travertine formations; but in the case of Lake Urmi neither 
were the investigations of Loftus or Rodler rewarded by the 
discovery of lake-terraces, nor do the travertine deposits of 
Dashkiesen appear to be of lacustrine origin. The travertine 
deposits round Lake Urmi seem to be the result of mineral 
springs. Pohlig mentions Pleistocene flood-zones of Neritina 
and Dreissena on the islands and on the Shahi peninsula; but 
although I saw a single specimen of a Dreissena-travertine mass 
in Urmi, I was not able to discover where it had been found, and 
consequently I was not able to confirm Pohlig’s theory. 

Lake of Urmi.—The lower parts of the depression of the 
Azerbaijan plateau are covered by the great salt-lake of Urmi. 
The length of this sheet of water, as measured from north to 
south, does not fall short of 80 miles, and its breadth is about 
25 miles at the wider parts, but is far less where the Shahi 
peninsula, jutting out, lessens the distance to ten miles. In 
former days the mountain of Shahi seems to have been entirely 
surrounded by water, but now it is counected with the eastern 
shore by dry land except when the spring floods convert the 
isthmus into a marsh. The area covered by the lake is about 
1750 square miles. 

The shores of the lake, in some few places where the hills come 
down to the water’s edge, are rocky, and falling away abruptly 
exhibit low cliffs of erosion; but for the most part the land 
slopes so gradually that the bather may have to wade for a mile 
or two before he reaches water which is out of his depth. The 
more notable elevations on the coast-line besides Mt. Shahi are 
at Guverchin Kala, a promontory of granite with pink felspars 
and of Miocene limestone with shells and echinoderms, near 
Gavilan ; at St. George’s Hill, Superghan; and at the Bezau 
Daghi, which are at any rate partly of voleanic origin, since 
their lower flanks are composed of pumiceous hornblende-biotite- 
andesite. 

The islands will be described in greater detail below. There 
are afew small islets off Guverchin Kala, at the northern end 
of the lake, which I was unable to visit; but the more important 
group of islands is situated in the southern half of the lake, 
though its exact geographical position has yet to be defined. 

The depth of the lake is inconsiderable. At no place does 
this huge expanse of water appear to be more than 40 feet in 


354 MR. R. T. GUNTHER ON THE 


depth, and it is doubtful whether the average sounding 
would be as much as half this; indeed, it seems to me to be 
probable that 15 feet would more nearly represent the mean 
depth. As in other salt lakes the height reached by the water is: 
lable to considerable seasonal variation ; the difference of level 
seems to bear such a large proportion to the average depth of the 
lake that the composition and specific gravity of the water must 
undergo very considerabie alteration during the change from the 
dry-season level to that of the wet-season. It might be quite 
possible to compute the total volume of the lake from the data 
supplied by gravimetric measurements considered in relation 
to the easily measurable increase of volume after the rainy 
season. 

The specific gravity of a sample of water obtained on Sep- 
tember 16, 1898, was 1:1188 at 15°C. Its salinity was equal to 
about 3 of that of the Dead Sea (sp. gr. 1:2225 at 120 metres, 
Lartet). 

A chemical analysis of the filtered water, undertaken for me 
by my friend Mr. J. J. Manley, gave the following proximate 
results, calculated in parts by weight in 100 grammes of the 
solid salts :— 


Sodiumarchloride imeem cence: 86:203 
Wikcanesnmin Calorie 354000540000 6:3816 
Magnesium sulphate .........-.. 3915 
Calciumiysulphatekeere mcr neisr ee eis 
Potassimmsullpiateln eae 1-741 

99°826 


"017 per cent. of free carbonic acid gas was also present, as well: 
as an unweighable trace of barium. ‘The weight of the total 
solids came to 14°893 parts in 100 of the water *. 

The average temperature of the water during the month of 
August was about 80° I. at the surface and some two degrees: 
lower at the bottom (depth 25 feet). The very shallow water 
very near the shores was of course heated up to a higher tem- 
perature (82° F.), and was far more concentrated by evaporation 
than the open lake-water: in many places along the margin the 


* For further details concerning the chemical and physical characters of 
the water, see a paper by R. Giinther and J. J. Manley, ‘On the Waters of the 
Salt Lake of Urmi,” in Proc. Royal Society, 1899, vol. lxv. pp. 312-318. 


NATURAL HISTORY OF LAKE URMI. B00 


concentration of the saline solution had become extreme, and the 
salts were being deposited upon the surface of the loathsome 
black mud, reeking of sulphuretted hydrogen and probably 
yielding marsh-gas as well, which seems to be such an inseparable 
feature of salt lakes. For some distance from the water’s edge 
the foreshore was covered with a dazzling crust of white salt. 

The salt water has a very unpleasant physiological action upon 
all the mucous membranes, and produces nausea if swallowed, 
but is otherwise innocuous to the bather. The Syrians are in 
the habit of bathing in the lake upon St. Thomas’s day (July 8rd 
O.S.),im order to commemorate the tradition that the Saint 
erossed the lake on his way to India. On emerging from the 
water the skin becomes rapidly covered with a thin crust of salt, 
unless the water be rapidly removed with atowel. But although 
harmless and even invigorating to man, the salinity is fatal 
to any freshwater fish of the rivers which may happen to swim 
out too far; wherefore at the present day the lake forms a very 
efficient barrier to the distribution of fish from one river to the 
next. Quantities of dead fish may sometimes be seen near the 
mouths of some of the rivers. I tried the experiment of putting 
freshly-caught chub into the salt water, and found that they 
died in three and a half minutes. When the salt water was 
gradually substituted for the fresh, the fish died when the 
mixture contained about a third of the salt water, which was at 
the end of forty minutes. On the 2nd of August a specimen 
of the green tree-frog was found vainly endeavouring to escape 
from the salt water into which he had accidentally jumped; but 
his rapidly weakening efforts showed that he too would soon 
have become a victim of the salt water, and have involuntarily 
verified Semper’s observations (‘ Animal Life,’ p. 150), had he 
not succumbed to the collector’s alcohol. 

Plankton.—lt might very naturally have been supposed that 
so hostile an environment as a strong saline solution, surrounded 
by a zone of a still stronger one almost saturated with sulphuretted 
hydrogen, would have been incompatible with organic life. The 
Lake of Urmi, however, is in no sense of the word a Dead Sea: 
it is simply teeming with living organisms, both animal and 
vegetable. Whether near the shore or miles from it, the clear 
water may be seen to sparkle in the sunlight owing to the 
enormous numbers of organisms which constitute its plankton. 
Throughout all the vast volume of water the distribution of life 


356 MR. R. T. GUNTHER ON THE 


seemed to be fairly uniform, for even water drawn from a depth 
of 28 feet contained its due proportion. 

In the month of August the vegetable portion of the plankton 
consists of small green masses, either of a globular or of a 
membranous, flat and irregularly expanded form, of soft or 
gelatinous substance, and varying from ;/, to ? of an inch in 
diameter. I at first regarded them as simple colonies of alge ; 
but Mr. G. Murray, who has been kind enough to examine my 
all too scanty material, assures me that their structure is that of a 
bacterial zoogleea of micrococei invested by a number of small 
diatoms. 

Their presence in such enormous quantities in the lake makes 
me suspect that there may possibly be some more intimate vital 
relations between the two organisms than would appear at first 
sight. Their abundance in parts of the lake where there is 
unlikely to be a proportionately large food-supply for plants of 
holozoic nutrition, seems to indicate symbiotic relations between 
the chlorophyli-containing diatoms and the bacterial colonies, of 
a nature very similar to those which enable the constituents of 
the lichen to maintain life in situations where life would be im- 
possible without such a symbiosis. At the same time, it is 
possible that the colonies may feed upon matter brought down 
by the rivers, and that they may owe their universal distribution 
to the surface-drift of the waters caused by the wind. 

The vegetavle portion of the plankton affords nutriment for 
the fauna. As I have already stated in my letter to ‘ Nature’ of 
Sept. 8, 1898, the so-called “jellyfish” alluded to by Lord 
Curzon of Kedlestone and Mr. P. L. Sclater is a species of 
brine-worm allied to Artemia salina, Leach. In the shallows 
near the muddy shores are to be found the aquatic larve of a 
Dipterous insect not unlike the larve of Hphydra riparia. As 
in the rat-tailed larvee of Hristalis and of Ptychoptera paludosa, 
the respiratory tubes are prolonged posteriorly and admit of con- 
siderable extension, so that the larva is able to draw air into its 
tracheal system while crawling in search of food beneath the 
surface. The extremity of the respiratory tube is forked, and 
each branch is tipped with small hairs which naturally increase 
the clinging power of the apparatus to the surface-film of the 
water. The larvee were about 10 mm. in length. 

The little brine-worms (Artemia urmiana) were as a rule uni- 
formly distributed throughout the lake, but clouds and streaks 


NATURAL HISTORY OF LAKE URMI. 357 


of Artemia are occasionally to be seen at certain places near the 
shore. In such streaks in which the Artemias were more than 
ordinarily numerous, there were always a large proportion of dead 
individuals or of individuals with impaired swimming powers ; 
the greater density of their aggregation seemed to be the 
result of local currents. In the clear green waters brightly 
illumined by the sun, the delicate cuticles of the Artemias sparkle 
like star-spangles. In order to obtain data for comparing the 
density of population of Lake Urmi with that of other lakes, a 
tow-net of silk bolting-cloth (meshes -25 mm. square) was slowly 
drawn horizontaliy through about 35 yards of water in the middle 
of the island archipelago, and again in a vertical direction through 
25 feet (the utmost obtainable at the station) ; the organisms 
captured were put into alcohol, their numbers were counted and 
their apparent volume measured. The two horizontal fishings 
gave results not differing from the mean by more than 8 per cent. 
The cubic metre of water was found to contain 1577 individual 
Artemias—or roughly, an Artemia to every pint of water. The 
preserved material, after being allowed to settle for 24 hours, 
occupied an apparent volume of 42°5 cubic centimetres. The 
vertical fishings gave rather lower results. Six hauls of the 
plankton-net showed that the average vertical distribution was 
about 1203 Artemias to the cubic metre of water. The inference 
is that, although the population is less dense near the bottom 
than near the surface, yet on the whole the organisms may be 
said to be fairly uniformly distributed throughout the lake. My 
observations are therefore in accordance with those of Reighard 
made during a biological examination of Lake St. Clair *. 

The fishings recorded above were made at 8 o’clock in the 
morning, but others made at midnight (with strong moonlight) 
and at midday showed that there does not seem to be any 
extensive diurnal migration of the kind which Forel has 
shown to be characteristic of the pelagic fauna of deep fresh- 
water lakes, and which is also usual with many marine pelagic 
animals. Indeed, during the day, owing to the shallowness of 
Lake Urmi, the bottom is not much less brightly illuminated 
than the surface, and a fauna of negatively heliotropic nature 
would search in vain for twilight in its inconsiderable depths. 
I am of opinion, therefore, that 1200 Artemias per cubic metre 


* Bull. Michigan Fish Commission, 1894, No. 4. 


308 MR. R. T. GUNTHER ON THE 


is a fair estimate of the density of the population. If it be 
assumed that the average depth of the lake is 6 metres, the 
total population of the 1750 square miles of lake must be at least 
39 x 10” adult individuals. 

The eggs of the Artemia may often be seen floating on the 
surface in long, interlacing streaks of a brown colour. The 
Artemias and their eggs are indubitably the food of the great 
concourse of water-fowl which have been remarked by all travellers 
as dwelling on the shores of the lake. In August I saw no 
flamingoes, although quantities of their pink feathers on the 
beaches testified to their presence earlier in the year. Gulls 
and ducks were very abundant. 

Influence of a Saline Environment.—lt appears that although 
the water of Lake Urmi is not incompatible with the well- 
being of all organic life, yet its salinity is too great to admit of 
' the existence of more than a few species. They are on the whole 
similar to the species described from other salt lakes in desert 
regions, but the fauna is rather more limited. Lake Urmi, in 
respect of its fauna, is very similar to the Adschidarja, near the 
Caspian Sea ; if does not contai so many species as the less saline 
pools of the neighbourhood of Odessa (Schmankewitsch), nor do 
any Coleopterous insects appear to have become acclimatized to 
its waters as has happened in certain American and European 
salt waters. It is possible that a worm may yet be found living in 
its waters, as Pachydrilus does in the strong brine-springs of 
Kissingen (Semper) and Kreuznach (Claparéde). The affinity of 
the Urmi fauna is undoubtedly with that of the fresh-water 
rather than with that of the sea; the fauna is therefore not to 
be regarded as halolimnic in Mr. J. BH. 8S. Moore’s sense of the 
word: it is decidedly halophilous. 

The influence of a strong saline solution upon the struc- 
ture and habits of an organism has engaged the attention 
of several naturalists. The early researches of Plateau and 
Beudant proved that many freshwater animals will live in sea- 
water and vice versa, so long as the change from one to the other 
ve not effected too suddenly; and also that different animals 
have different powers of resistance to such a change. In 1889 
Boas * described the remarkable changes of structure which 
occur in Palemonetes varians in accordance with its growth 
in fresh or salt water. The last few years have produced 

* Zool. Jahrb. iv. 


NATURAL HISTORY OF LAKE URMI. 359 


an abundant crop of literature dealing with the effect of 
salinity upon the early stages in the development of organisms 
and especially of Echinoderms (Morgan, Loeb, Rawitz, Norman, 
Vernon, and other students of ‘‘ Entwickelungsmechanik ”). 
None, however, are of more importance than the classical 
treatises of Schmankewitsch upon Branchipus, Artemia salina, 
and A. Miilhausenit. By some accident a tank near Odessa 
became filled with salt water of a density of 1:0567 (8° Beaumé) 
in 1871. The water in the tank became concentrated by very 
gradual evaporation until in 1874 it reached a density of 1:2015 
(25° B.). The Artemia salina which populated the tank in 
1871 underwent a gradual change as the salt water became 
more and more concentrated. The set of the furecal lobes of 
the tail became fewer in number and dwindled in size, the eill- 
lamelle enlarged and altered in shape, the abdomen tended 
to alter its seymentation, and finally the entire species became 
metamorphosed into A. Mulhausenii, a species only known from 
the most saline waters. 

The Artemia urmiana lives in a far more saline water than 
the A. salina of Schmankewitsch did, but notin such a saline 
solution as A. Mulhausentt. Consequently we should expect 
A.urmianato be in ermediate in structure between these forms in 
those characters w ich are determined by the strength of the saline 
environment ; and this is precisely what I have found to be the 
case with respect to the tail-lobes and their sete. Many of the 
Artemia urmiana «xhibit clearly the intermediate condition figured 
by Schmankewitsch on plate vi. fig.5*, The specifie gravity of 
the water which produced this condition in Schmankewitsch’s 
experiment was about 1:1373, while the specific gravity of the 
water of Lake Urmiis 1°1188, or 0235 lower. It must be remem- 
bered that the salinity in the ponds of Schmankewitsch was 
increasing at avery rapid rate as compared with the rate of 
change of the salinity of Lake Urmi. It is therefore likely that 
if the Artemias of Odessa had had more time, as measured by 
Artemia-generations, in which to adapt their structure toa salinity 
indicated by a specific gravity of 1:1138, the resemblance between 
the Russian and Persian Artemias would have been even yet 
more striking. Even as it is, the Artemia seems to act as a 
hydrometer with an error of less than ‘03 in the determination 
of specific gravities. 


* Zeitschr. f. wiss. Zool. 1875. 


2860 MR. R. T. GUNTHER ON THE 


Islands.—In the southern half of the lake is a small group of 
rocky islands which from a distance present a rounded appearance, 
like the knolls on our chalk downs, but from a nearer point of 
view their precipitous cliffs and rugged hill-sides testify to the 
erosive powers of the heavy salt waves in stormy weather. Their 
geological structure has been investigated by Abich, who pro- 
nounced them to be built of Miocene chalk resting upon Paleo- 
zoic calcareous strata which are to be seen near the north end of 
Koyun Daghi. The Miocene chalk is divisible into three main 
divisions. The uppermost is porous and contains Ostrea Virleti, 
Deshayes; the middle division is of a more compact nature and is 
rich in corals. In the lower deposit bivalve and coral fragments 
are found, together with Turritella Archimedis, Brongniart, 
T. turris, VOrb., and 7. gradata, Menke. 

On many of the islands are beaches of shells, coral fragments, 
and echinoderms, organisms which could have lived only in 
a sea of marine salinity and in connection with the ocean. 
These marine reliquiz, now for a second time rolled by salt waves, 
tell the tale of a Miocene sea of normal salinity which has been 
supposed by Pohlig to have been a northerly continuation 
of the Persian Gulf from the Indian Ocean, but which, as 
already stated, was more probably an extension of the Mediter- 
ranean. This Miocene sea, like the Red Sea of to-day, was a 
coral sea. Upon its floor were laid down the chalk and lime- 
stone formations of the Urmi Archipelago, as well as those of 
the calcareous mountains to the south of the lake. 

The largest of the islands are Koyun and Arzu, of which the 
former measures between 3 and 4 miles in length and rises to a 
height of about 1000 feet above the level of the lake. 

The islands appear to have been formerly inhabited, and there 
may still be seen foundations of houses near a spring at the 
south-eastern end of Koyun Daghi. At the present time 
they are uninhabited, but are often visited by sailors, who 
turn out goats and sheep to pasture on them during certain 
months in the year, and who not only cut down the trees, but even 
dig up for fuel the roots of trees felled by their predecessors. 
There is a tradition that some eighty years ago the islands were 
connected with the mainland by a causeway which can, it is said, 
still be detected by sounding. This tradition is certainly con- 
firmed by the zoology of the islands. There are at least five species 


NATURAL HISTORY OF LAKE URMI. 361 


of land-shells and two species of lizards (Humeces Schneidert 
and Ophiops elegans), which would be very unlikely to have 
crossed the broad expanse of water separating the islands from 
the mainland. ‘here were also mollusks, woodlice, scorpions, 
Galeodes, and several wingless insects. 

The most interesting feature of the fauna of these islands is 
that upon them once lived a wild sheep, a distinct variety, appa- 
rently allied to the Cyprian Ovts ophion. Unfortunately, I 
brought home only one skull, although there was an entire 
skeleton upon Koyun Daghi. It is possible that this sheep, 
too, may belong to a “ Relikten-fauna,’ if I may be permitted 
to apply Credner’s term to a terrestrial area. 

Some eagles (probably Aguila chrysaétus, L.) breed among 
the highest crags of Koyun Daghi, and the lower crannies are 
the homes of countless rock-pigeons. The shores are often lined 
with ducks and gulls, which feed on the crustaceans (Artemia) 
and bacterio-diatom colonies of the Salt Lake. On the pebble- 
beaches I picked up many pink feathers of the flamingo, but I did 
not see any of the birds during my stay; they are abundant in 
winter, and are snared on the mainland in spring. Some of the 
beaches are covered with reed-stems, which have probably been 
drifted across from the mouths of the great southern rivers by 
southerly winds. Partridges and quails were occasionally heard, 
and magpies were to be seen at the spring disputing with the 
pigeons for the right of drinking first. 

Upon the small gragss-grown Shazalan Island, to the north of 
Arzu, I saw two young specimens of Humeces Schneideri as well 
as many Ophiops elegans. Myriads of rock-pigeons which nest 
in the crevices of the rocks are regarded by the sailors as an 
unfailing source of pigeon dinners. The rocks consist of a dark 
slate-coloured limestone, with many crevices and cavities con- 
taining secondary depositions of calcium carbonate. The lime- 
stone is very similar to that of the N.W. end of Koyun Daghi, 
and is of Carboniferous age. 

The vegetation growing on the islands at the time of my visit 
was mostly burnt up by the sun except near the springs. On 
Koyun and Arzu there are a few stunted walnut-trees, the largest 
of which measured between two and three feet through at a yard 
from the ground. ‘The calcareous soil favours the growth of the 
karuan plant (Artemisia): the effect of its aromatic odour as an. 

LINN. JOURN.—ZOOLOGY, VOL. XXVII. 28 


362 MR. R. T. GUNTHER ON THE 


insectifuge is as well known as it was in the days of our fore- 
fathers, when the housewife was taught that— 


‘‘ Where chamber is swept, and wormwood is strewn, 
No flea, for his life, dare bide or be known!” 


Plains.—The shores of the lake are for the most part sur- 
rounded by gently sloping gravel or alluvial plains, which extend 
as far as the feet of the hills. They are traversed by the 
numerous rivers which flow into the lake, and which distribute 
their waters among innumerable villages and over vast tracks of 
cultivated land. When seen from a distance the irrigated fields 
look like extensive forests, owing to the poplars and willows 
which line the watercourses, and which doubtless have given rise 
to the misleading statement that Lake Urmi is “surrounded by 
wooded shores and hills ’’ (Curzon, ‘ Persia,’ p. 532). 

The country which is too far from water to be irrigated is of 
the nature of a thistle-covered steppe, whereas the lower marshy 
portions exhibit rank growth of rushes and blue irises. 

My first impressions of the Zoology of the Urmi plains were 
of disappointment, and for two reasons—firstly, because the 
general facies of the fauna is pre-eminently Huropean, and, 
secondly, because I found the country much more densely popu- 
lated than I had expected. There are villages wherever sufficient 
water is obtainable to make agriculture possible; and as these 
villages have existed for very long periods of time, within their 
spheres of influence, Nature has been tamed and the fauna pro- 
foundly changed. 

Among domesticated animals the most striking are the camels, 
buffaloes, and fat-tailed sheep (Ovis steatopygus). Buffaloes only 
do really well where they can spend several hours a day in the 
water. When water deep enough for complete immersion cannot 
be found, a small boy is often told off tor an hour or so to bale 
up water in a pot and to pour it over their backs. The winter 
on the Urmi plateau is so severe that the buffaloes have to be 
kept in stables until the return of warm weather. 

The gazelle (Gazella subgutturosa) doubtless once lived on 
the plains of Urmi, but has now become exterminated by the 
spread of cultivation. The badger (eles canescens) is rare. The 
rats and mice are unknown to science, although their holes are 
common in the fields and along watercourses. A species of 
gerbille oecurs, but is not very common. 


NATURAL HISTORY OF LAKE URMI. 363 


Quails and partridges are common, and afford a favourite sport 
to native falconers, who use sparrow-hawks. The bee-eaters 
(Merops apiaster) were extremely plentiful near the northern end 
of the lake ia July, but were not to be seen further south in 
August and early September. 

Storks are abundant; their twig nests are to be found in trees 
or on the roofs of the houses in most of the villages. The Syrians 
have learnt the Persian story that the stork visits Mecca during 
his winter migration, and doubtless that is why “ Hajji Laqlaq”’ 
is regarded as bringing good luck and is attracted by nesting- 
baskets put up in trees. Storks return about March 14th. 

The hoopoe is common. The Syrians consider them unclean 
birds (Lev. xi. 19, R. V.; Deut. xiv.18), and call them “ Pupu” 
and also “ Birds of Solomon,” from the old legend that they got 
their crests from him. “ A flock of them once sheltered him from 
a burning sun. In gratitude the king asked them what he should 
do for them, and they asked for crowns of gold. But finding 
that they were being killed by greedy men for the sake of the 
crowns, they begged Solomon to change them for the crest which 
they retain to this day” (Maclean). 

In addition to the reptiles mentioned below, Clemmys caspia, 
de Filippi, is common. Many may be seen walking along the 
banks of the watercourses, ready to dive into the stream on 
being disturbed. After lying still for three to five minutes in a 
hole or beneath some water-weed, they cautiously come up to 
see whether the danger is past. All the specimens examined 
had the yellow sternal shields with black patches characteristic 
ot C. caspia as opposed to the European C. leprosa. 

Testudo ibera is common at Seir, as indeed it is on the mountains 
to the east of the lake. On Koyun Daghi I found the fragment 
of a carapace which probably belonged to this species. 

Fish.—Inasmuch as the rivers which flow into the lake are 
separated from each other by a medium, the salt water, in which 
no fish can live for a period long enough to enable it to swim 
from one river to the next, the characteristics of the fish fauna 
of the individual rivers are deserving of the most careful investi- 
gation, because they may supply us with a clue to the physical 
conditions which obtained during a particular epoch in the iake’s 
history. If it can be shown that the faunas of all the rivers are 
identical, or that no local variations or species are distinguishable 
in the different rivers, then it must be inferred that the rivers 

28* 


364 MR. BR. UT. GUNTHER ON THE 


have not been isolated from each other long enough for such 
faunistic differences to have appeared. Butif, on the other hand,. 
it can be proved that there are faunistic differences between the 
rivers, then it must be inferred either that the rivers have been 
populated from different stocks, or that they have been populated 
from the same stock at so remote a period that their isolated 
faunas have had time to become modified in different directions. 

My collections and observations are by no means as complete 
as could be desired for the solution of the problem, but still they 
would seem to afford a strong indication of the existence of local 
faunistic differences in the rivers. I think they show that there 
are not only differences of species and varieties, but that the 
relative abundance of the common species varies greatly in the 
different rivers. 

As I have already pointed out, the rivers which flow into Lake 
Urmi may be divided into three groups*. The Zola, Nazlu, 
Shaher, Barenduz, and Gader rise in the mountains to the west 
of the lake; the Tatawa and the Djaghatu enter the lake from 
the south; and the Murdi, Safi, and Adji receive the eastern, 
drainage. 

Probably all the rivers contain the chub, Capoéta gracilis, 
and an Alburnus. The Gader, Tatawa, and Djaghatu are 
noted for the Silurus glanis which is absent in all the western 
series of rivers except the Gader Chai, which opens near 
the Tatawa, and which probably often mingles its flood-waters 
with those of the latter stream on the flat land between their 
mouths. Leuciscus ulanus seems to be peculiar to the waters of 
the Plain of Salmas, and Leuciscus gaderanus to the southern 
rivers. I succeeded in catching Barbus caucasicus and Nema- 
chilus persa only in the more northern streams, although the 
former is common all over Persia. 

Most of the fish procured were taken by means of poison, a 
method of fishing which, so far as its efficiency and unsportsman- 
like character are concerned, is second only to the method of 
fishing with dynamite. A mixture of flour and the pounded 
berries of Cocculus indicus is mixed with sufficient butter to 
make a stiffish paste. A backwater or reach where the stream 
does not flow too rapidly is then ground-baited with small pellets 
of the paste. If the fish are feeding, the poison will begin to 
work in about ten to fifteen minutes; and the poisoned fish will 


% 


‘Proc. Roy. Geographical Society,’ vol. xiv. pp. 504-523. 


NATURAL HISTORY OF LAKE URMI. 365 


begin to swim in small circles at the top of the water, and 
may be taken in a landing-net. Many which have eaten much 
of the poison will swim into the bank and lie helpless in the 
shallow water. If the river is at all full of fish, a great number 
may be caught with a surprisingly small quantity of poison. On 
September 9 I mixed about six ounces of the poisoned paste 
and ground-baited the Tatawa Chai near Sujbulak. We took 
38 Capoéta and chub of five ounces and upwards, and both sides 
of the river were silvered by lines of poisoned smaller fish for a 
distance of about a hundred yards. 

The fish which have not eaten much of the poison gradually 
recover, and regain the power of coordinating their movements. 
The dead fish which are too small to be worth picking up by the 
villagers, who always turned out in crowds at the mere mention 
of fishing, are soon devoured by the crabs. If the poisoned fish 
be carefully cleaned, they have no bad effect when cooked and 
eaten. 

The large rivers which flow into the lake from the southern end 
—the Djaghatu Chai, Tatawa Chai, and Gader Chai—are stated by 
the natives to contain “ whales” whose ancestor swallowed Jonah. 
The “whales” turned out to be Silurus glanis. They are caught 
either by being driven into nets or by the gaff. A Jewish fisher- 
man accompanied me to a place near Ocksa, where the Gader Chai 
has cut a deep channel! under a bank overgrown by willow-trees. 
After divesting himself of all needless clothing, he dived into the 
deep water with his gaff, and hooked about among the willow-roots 
at a depth of about 4-5 feet. After eight minutes he succeeded 
in gafling a large female Silurus 3 feet 9 inches in length, and 
before a quarter of an hour had elapsed he had three fine fish 
out on the bank. He told me that the largest “‘ whale” he had 
ever seen was about five feet long. They spawn in deep pools 
below the willows, where the water runs slowly, about one and 
a half months after the melting of the snow. The Siiurus is 
eaten by Armenians, but is unclean to Jews and Mahomedans on 
account of the absence of scales on its body. 

I was informed that the Governor, when he wishes to do 
honour to a guest, organizes a“ whale” hunt, and all the villagers 
assist at the ‘‘tamasha.” The buffaloes of several villages are 
turned into the river and are made to walk upstream for a couple 
of miles, with the result of driving the fish before them. Ata 
suitable place, a line of men with nets stand prepared to catch 


3866 MR. R. T. GUNTHER ON THE 


all fish driven in their direction; and in this way several hundred 
‘whales’? may be caught amid the din of the exhortations of 
the enthusiastic onlookers, who often cannot refrain from rushing 
into the water and from joining their struggles with those of the 
splashing fish. 

Invertebrata.—-The most conspicuous member of the freshwater 
invertebrate fauna is the freshwater crab, Telphusa fluviatilis, 
which may be found under stones in burrows, part in and part 
out of the water. Always on the look-out for carrion, these 
scavengers were frequently seen trying to drag poisoned fish 
from the margin of the stream to some more retired nook in the 
deeper water. 

In clear spring-beds Gammarus pulex was often met with, 
but Asellus seemed to be entirely absent ; and by its absence the 
pond-life afforded a very striking contrast to that of Europe 
_within the range of Asellus. Leeches and planarians were 
common. 


My small collections of terrestrial invertebrata have been 
examined by Messrs. Edgar Smith, Butler, Pocock, Burr, and 
Sir G. Hampson, and are described below. 

The traveller in Persia is always interrogated about “ insect” 
pests, and consequently a few words upon them may not be out 
of place. Mosquitoes are painfully abundant in the Araxes 
plains both in Russian and in Persian territory. I was not 
troubled by them at all in August and September, either at Urmi 
or in any of the villages at some distance from the lake, although 
I always slept on balconies or housetops without any protection 
to the face. On the other hand, in the country near the lake, 
as at Superghan and Ardishai, they are said to be intolerable 
during September. 

The much-dreaded Argas persicus is not rare, but its venomous 
bite does not appear to be followed by the fatal or fabulous 
consequences often attributed to the bug of Mianeh, where, 
according to Maurice Kotzebue, a victim ‘‘ éprouva bientot dans 
tout son corps une chaleur violente, tomba dans une espéce de 
délire, et expira enfin au milieu d’épouvantables convulsions.” 
The native cure is the application of the still smoking skin of 
a newly-flayed ox to the seat of the evil. 

Scorpions are common, but are rather local. Many Persians 
are familiar with the old story of the scorpion which, when 


NATURAL HISTORY OF LAKE URMI. 367 


surrounded by a circle of live coals and unable to escape, stung 
itself in the head. Dr. Wills (‘The Land of the Lion and the 
Sun,’ 1891, p. 249) even asserts that he has witnessed the suicide 
of fire-girt scorpions on more than one occasion. Unfortunately 
Dr. Wills does not designate the particular kind of scorpion with 
which his experiments were performed. A solitary experiment 
of my own, made with Buthus caucasicus subsp. persicus at 
Maragha, proved as unsuccessful as those which were under- 
taken by Prof. Ray Lankester. 

The sting of scorpions is greatly feared, and peculiar pre- 
cautions are taken against them in many places. Many stories 
are told of deaths attributed to scorpions, but it is always difficult 
to ascertain whether the venom was the sole cause or merely ar 
accompanying complication. However, it is certain that scorpion 
stings are better avoided. In some parts dried thistles, spread 
loosely under the beds, are employed in order to keep scorpions 
away: indeed, many of the mountain folk of Kochanes hold that 
a goat-hair blanket will suffice for this purpose; since scorpions 
have such a tender skin on their stomach that if they can be 
made to walk over the hair of a goat, they will receive a “ mortal 
wound.” 

Mimicry.— On August 19 I noticed some interesting cases 
of what appeared to be protective coloration. Near the spot 
where the Shaher Chai leaves the hills and enters upon its 
course across the plain of Urmi, numerous poplar trees have been 
planted. The bark of the Populus alba is on the whole of a grey 
colour, the ground being dark and sprinkled with light-tinted, 
almost white stippling. Upon the bark of the trees were several 
species of insects which so exactly mimicked the bark upon which 
they were sitting that it was extremely difficult to distinguish 
them from their background (PI. 27. fig. 1). The insects found 
upon the poplar bark were Yponomeuta padellus, L., Pentatoma 
baccarum(?), and Bathyoscopus poceilus, H.-Sch. There were 
also numerous spirally coiled shell-like larva-cases or cocoons of 
Apterona crenuiella. No other insects except ants were seen. 
The Pentatoma, with its grey speckled thorax and wing, and 
with its antenne barred alternately white and black, was more 
invisible against the grey bark than its emerald-green relative 
was upon the fresh green leaves of herbaceous plants near the 
stream. 


368 MR. R. DT. GUNTHER ON THE 


Seir.—The chief locality in the hills at which collections were 
made was near the small village of Seir, 5-6 miles W.S.W.of Urmi. 
It is situated at a height of some 1460 feet (Loftus) above the level 
of the lake, on the eastern slopes of the Seir Dagh. Here a gravel, 
conglomerate and sandstone hill rises to a height of 7260 feet 
above sea-level; its strata dip to the east and overlie the lime- 
stone of the plain of Mergawar. From Seir there is a fine view 
over the wide cultivated plain of Urmi to the deep blue lake, in- 
terrupted only by the triple-crested Bezau Daghi and the conical 
hill of Superghan ; and when the mountains on the further side 
of the lake are covered with snow, the prospect must be indeed 
superb. Here, as it were on the dividing-line between the rich 
cultivation of the plain of Urmi and the barren wildness of the 
Kurdish hills, the Christian Missionaries seek the healthful 
fresh mountain air when the noisome summer heats of the city 
become unendurable, and exchange an artificial environment for 
a natural one. 

The hillsides have been dissected by the rain-water torrents, 
and many a section has been laid open for geologists. The 
uncompacted sands and gravels which intervene between the 
more coherent conglomerates and sandstones are rapidly removed 
by the spring torrents, and the compacter rocks fall away and 
litter the stream-beds with great boulders which may measure a 
score of feet square and half as many through. 

The pebbles in the conglomerates, consisting of both igneous 
and calcareous rocks, have not as a rule been very thoroughly 
rounded, and therefore appear to be of a fluviatile rather than of 
a sea-worn origin. I was quite unable to discover any organic 
remains coeval with the conglomerates ; but many of the pebbles 
are of older, fossiliferous formations. Especially abundant were 
limestone fragments containing three or four species of corals, 
while others with shells were occasionally met with. The 
igneous rocks consisted of granites and felsites. 

Seir Vegetation.—In August the vegetation covering the hillsides 
above the village of Seir was burnt to a dull yellow hue by the 
summer drought. It was always a subject of wonderment to me 
how flocks and herds pasturing on such barren-looking slopes 
could manage to pick up a living. Earlier in the year the hillsides 
are green and gay with white and pink crocuses and irises; but 
in August the most conspicuous plants were large blue thistles, 
dwarf yellow hollyhocks, yellow-green euphorbias, not unlike our 


NATURAL HISTORY OF LAKE URMI. 369 


own species but larger, and yellow everlastings, and here and 
there a few Daphne bushes *. 

Tam indebted to Dr. O. Stapf, the authority on Persian Botany, 
for the identification of the following species. The list has no 
pretence to completeness ; it is merely intended to give an idea 
of the general aspect of the flora in the latter half of summer :— 


Farsetia suffruticosa, DC. 

Scabiosa olivieri, Coult. 

Cephalaria sp. 

Xeranthemum squarrosum, Boiss. 
Helichrysum armenium, DC. 
auchert, Boiss. (?). 

Centaurea virgata, Lam., var. squarrosa. 
Lactuca orientalis, Boiss. 
Campanula sp. 

Podanthum sp. 

Acantholimon sp. 

Odontites aucheri, Boiss. 

Ziziphora clinopodioides, M. B., var. 
rigida, Boiss. 

Thymus kotschyanus, Boiss. & Woh. 
Teucriwm polium, L. 


In moister situations on the sides of a small stream from a 

spring were :— 
Glycyrrhiza glabra, L. 
Eryngium billardieri, Laroch, var. meiocephalum, Boiss. 
Echinops sp. 
Pulicaria dysenterica, Gaert., var. microcephala, Boiss. 
Centaurea solstitialis, L. 

virgata, Lam., var. squarrosa. 

Mentha tomentosa, Urv. 

Marrubium crassidens, Boiss. 

Buphortia cheiradenia, Boiss. & Hoh. 

Daphne acuminata, Boiss. & Hoh. 


Seir Zoology.—Foxes are said to be common in the valleys 
near Seir. The only individual which I saw was of a much paler 
colour than our English species. 


* Of the bulbs from Seir which haye reached this country alive, Puschkinia 
scilloides 1s the only one up to the present time which has flowered. 


370 MR. R. T. GUNTHER ON THE 


Wolves occasionally visit the villages by night, when they are 
received like Kurds, with gunshots fired in no particular 
direction from the roofs of the houses. 

Two bats (probably Rhinolophus hipposideros, Bechst.) were 
seen flying about the houses in the evening. 

The birds most in evidence were magpies, hoopoes, small 
insectivorous hawks which nest in great numbers in the higher 
trees in the city, tits, and numberless sparrows. In the hills a 
few coveys of the red-legged partridge are to be seen, but they 
are difficult to approach. Quails frequent the cultivated fields 
and are taken by hawking, a very popular and fashionable amuse- 
ment in Persia. Quails are also caught by taking advantage of 
their stupid curiosity. The fowler and his assistants walk out 
into the field where the quails are supposed to be and hold up a 
large white sheet, often decorated with painted snakes, birds 
of prey, or other unpleasant but conspicuous creatures. The 
inquisitive quails cautiously walk towards the lure, and. finally 
approach near enough for the patient fowler to put a net over 
them. 

Testudo ibera is common in the dry torrent-beds on the hill- 
sides. I examined about a couple of dozen and found only three 
which were free from parasitic Acari, which Mr. A. Michael has 
kindly identified for me as Rhipicephalus simus, or sanguineus. 
The parasites are always attached to parts where they are in no 
danger of being rubbed off. One tortoise had four of these 
bloodsuckers, one in the “armpit” of the right hind foot and 
three on its tail. 

Neither lizards nor snakes were as common as I should have 
expected. Phrynocephalus seemed to be unknown to the natives, 
who, however, described a lizard which seemed to be Stellio 
caucasicus. Hremia was fairly abundant. 

A list of the insects collected at Seir is given on p. 408. The 
general character of the Lepidoptera struck me as being remark- 
ably European at first sight; but many of the species exhibit 
a rather more sandy or desert type of coloration. In the 
early morning the blues were extremely common on the Mentha 
tomentosa growing near the little stream, 4 mile to the S.W. of 
the village. Several fell a prey to the voracity of a large black- 
and-yellow spider (Lycosa). The only beetles which I obtained 
were Julodis levicostatus, L. & G., and Lixus bardane, F., 
covered with yellow pollen. The higher slopes of the hills were 
poor in butterflies during the middie of the day, but rich in 


NATURAL HISTORY OF LAKE URMI. Syidl 


Orthoptera (Decticus and Gdipodide), which, almost invisible 
when at rest, used to get up in blue clouds before the feet of 
our horses. Towards evening many wall butterflies come out 
of their hiding-places under stones and fly about during the 
short twilight. 

Mole-crickets (Gryllotalpa gryllotalpa, L.) are common in the 
moister localities. Ants and ant-lions are to be found up toa 
height of 6000 feet. 


List of Animals distinguished by special names by the Syrians 
living in the Urmi Basin. 


In the compilation of the following list I owe much to my 
interpreter friends, Shamasha Josip of Superghan and Pepino 
Sadok or “ Popina.” The list is confined to those animals with 
which my informants professed to be personally acquainted, and 
will enable the reader to form some idea of the state of their 
zoological knowledge, even though the English equivalents of 
many of the names are very doubtful. My heartiest thanks are 
due to Professor Margoliouth and to his talented wife for their 
help in the revision of the list and in the identification of the 
languages to which the words beiong, whether Old Syriac (O.S.), 
Neo-Syriac (N.S.), Armenian (Arm.), Arabic (Arab.), Persian 
(P.), or Turki (T.). (Stoddard) indicates that the word is to be 
found in his Modern Syriac Vocabulary, of which the manuscript 
is now in the possession of Professor Margoliouth. 


MamMatta. 

Bait pateesssaseesees- praka 1élé, O.S. ; Zit.=“ flier by night.” 
Caiieiescinecseeenee: catou, Arm. 
Leopard ............ nemra, O.S. 
ID@ES — scaconosc0c00se kalba, O.S8. 
Witenes kalba dmia, O.S. “ Water dog.” 
\WVOLE Aa ceaBeancndaee deva, O.S. 
BOR esc sexecones tala, O.S. 

Bn cash eselareeines vachaque, P. 
Bears ei Assicetncsaee débba, O.S. 
Badger ............ kouja, N.S. (Stoddard). 
TEREST, sacconococoe hotare. Rare in plain of Urmi, common at Mosul. 
Hedgehog ......... kadouda, koutoulta, koubda. (?=the name of a bird.) 
Squirrel ............ sennara. 
WIG) Mendneconnes ukb’ra, O.S. (?=jerboa.) 
Water-rat (?)...... kora mechk. 
Gerbillus or Dipus kaecha. 

Sa age meies garoun. 

Flare pin sccuwensics kirviche, N.S. (Stoddard). 

(siibiesatoatonacatcon qara bache. “Black head,” T. (Perhaps the caracal, or 


desert-lynx *.) 


* According to Prof. Blochmann our word caracal is derived from the Turki 
gara-gqolag, signifying “ black-ear.” 


372 MR. R. T. GUNTHER ON THE 


Bull eee eseaea tora, O.8. 
TBE Gasdecooosee kalla, N.S. (Stoddard). 
pSHVS(2) 0). sa denaceouscbad vana,O.S. Generally Ovis steatopygus. 
ao War ade seer vana dtoura, O.S8. 
am bweeceetonsercicc: péra, O.S., N.S. (Stoddard). 
Groat retaass Petaza Ose 
» Mountain ... azadtoura, O.S. 
DD) COr sr ecer eccscincsc khzura. 
Chine Secccdencdounes gamla, O.S. 
LOC Pea eser see ctr surni, O.S. 
HEVORSO Ve iosciis sci sussya, O.8. 
Donkey, ass ...... khmara, O.8, 
Mule (Horse 9 
x Ass Go.) ...00 cavédna, O.S. 
AVES. 
Haley .dvssseceene nishra, O.S. = Persian bargut. 
Hawk, ? sp. ...... bashuka, Arab. 
allconwageseeeere ... tocha. =Persian bahri. (See below, bahra.) 
Re aces oscnactoce nessa, O.8. 
Falco venaticus .... djoudjna, P. 
IDERAOVESE — Goooacveces baziga, P. 
Sparrow-hawk ... baza, Arab. =Persian basha. 
Goshawk ......... tarlane, N.S. (Stoddard). Used for hawking ; =Persian 
taigun, and tarlin male and female. 
Owvligee tees peste oae buma, Ar. or O.8. 
Goldfinch ......... sagga, Arab. 
Nightingale ...... andali, Arab. 
(MAvASIO, Tocaccconeec khazel. 2 
TaI@O]NOS. soooesodcoos vadvade. 
JOpRAK @) cococooonasc djidjerta. 
Blackbirdaeeese ee shoukrta. Cf. shakhroura, O.8. 
Fire-crested Wren. gorguma, P.; azel doukbé, P. 
AA VET Lee tentccnie ourva, O.S8. 
Lal emaueaemenaner djara. 
Moet are ey a djekha. Cf. djakha, below. 
Swallow, Swift ... snonita, O.S. 
OMORD® | oodcqnoon00. koukou. 
IPASEONM Geaaconcoce. kavédare. 
Turtle-dove ...... shoufnina, O.S. 
Pheasant (?) ...... djourda, P. 
Raniridoeleera cs: qiqvana, N.S. (Stoddard); gaqbana in O.S. 
ey nine eats S10 zarkha. 
Quail eee sescsase ss. goupshina, O.S. 
isu a agdodonaneenee kheta, déchta. ‘‘ Hen of the plain.” 
S18) Gosonoquenpanes laqlaq, Arab. 
Blamingo) <.....-.. smouqta, baqlane, O.S. 
Jealligein” coscoscdeooe qotan qaqa, O.S. 
AU eseeeneeenise se qakhouka, NS. (Stoddard). 
OM, loaanostesede sevanoga, Arm. =blackbird. 
IDI scasoaccnGa000 tona. 
Ba) Ae ame ae ourdak, T. 
PA | Goce ocaaecoee jandjz. 
Goose Gacessce nese. gaza, T. 
Wild Goose ...... laqlaq quissé. 
SKWEWE | coocasoannde ‘qiqanousse. 


The following Bird names are of uncertain signification :—juka; kourta ; 
dédé, or korkore; sissiarek; djarguna; métlédérou; shagra; djita; maria 
toré; poupou; bahra; djoulourda; djapdgepa, N.S. (Stoddard) (“ flapping 
wings”); chavere chagane, O.8.; shavere balga, O.8.; souravéle; gatou 


NATURAL HISTORY OF LAKE URMI. 373 


margué (‘cat of the marsh”); sharvélta, gaza lake; garkha diama; khazel, 
separmari; jeho; gogtatan; djaroura; alidji; djelkima; kadare; angourte ; 
djakha, P. (sparrow or owl); hogare; vaga, P.; metloue. 


REPTILIA. 


Lizards belonging) masousta, N.S. (mésazé in Stoddard), 
to various genera: | jdelourin. 
Humeces,Phryno- { mgadia goudayé. 


cephalus, &c....... kémkéma. 
Nnakorswecncecane khouvé, O.S. 
pps poBoeaKOSGAdGBO khouvé dkanouchta, O.S. “Serpent du ballet,” grey, 
poisonous. 
Petes iSeca ccs a\ete api. (?=hyzena.) Said to bleat like a goat, 
PM ie eccceoecos cits jira mar. Long, thin, crawls very fast. 
BME een Sakisiene khélda. Very poisonous. 
6p. «_ eapesaoaunadabe sola. Pointed head, creeps in a straight line. 
MOrtoisekesseesses qaraia, N.S. (Stoddard). 
AMPHIBIA. 
TRIROS?  GanpbedaBoseode piqqa, pagé, N.S. (Stoddard). 
PiscEs. 


Fish with barbels. mnuna oursa. ‘“ Male fish,” with moustaches. 
Fish without 

barbels ......... nuna niqua, O.S. ‘Female fish,” without moustaches. 
Silurus glanis...... mage, N.S. (Stoddard). ‘‘ Whales,” occur at Solduz. 


Insecra. 
lNSEChS i enecieecocss bajouji (or rakhsha), N.S. (Stoddard). From rkhish, to 
creep. ; 
Caterpillar ......... terterra, N.S. (Stoddard). 
Locust, Cricket, or 
Grasshopper ... kamsa, O.S. 
Mole-cricket ...... qerwa khmareé (lit.=‘“ Ass’s scorpion ”), 
Harwig ....... waeees karsu. 
i Cateeepereececcensi: perta’né, O.8. 
Myriopods ......... madala, O.S. 
Hi livptanenovascnee sere didwa, N.S, (Stoddard). 
ARACHNIDA. 
SYONCII casoacaoa000000 zakra gardé, O.S., shikra dmaia, 
SOHO — seooae000 agerwa, O.S. 
Galeodes............ harapasse. =‘ Reptiles” in O.S. 
CRUSTACEA. 
Crabs, Crayfish ... kedjala. 
Woodlouse......... ‘arsha. 
VERMES. 
Intestinal worms . kurkana, N.S. (Stoddard). 
WWESEN, cocooseasnonsa: zalu. 
Harthworm or sektéara, O.S., or sikta dar’é. Jit., a pointed stake or 
Long worm. ploughshare of the ground in Old Syriac. 
Hairy worm ....... spadita dkhouvé, O.S. Spadita is derived from O.8. word 


for “pillow.” 


374: DR. A. GUNTHER ON THE 


THe Witp SHEEP or THE UrRmti ISnANDs. 
By Dr. A. GinrHer, F.R.S., P.L.S. 


(PLATE 22.) 


A cranium in tolerably good condition with the skin and hairs 
adhering to the face and forehead, with perfect horns, but with- 
out lower jaw, was picked up on Koyun Daghi, the largest 
island of the Urmi Archipelago. The head is that of an adult 
yam. There is no other wild sheep (at least not in the very rick 
collection of the Natural History Museum) to which this head 
gomes nearer, as regards size of cranium and form of the horns, 
than the Cyprian Mouflon (Ovzs ophion), Yet there is a striking 
difference in the sweep and direction of the horns; but without 
further information it would seem to me premature to introduce 
this sheep as “a distinct species. It certainly must appear very 
singular that a sheep from a lacustrine island of Western Persia 
should be more nearly allied to the distant and local form from 
Cyprus, than to the typical Ovzs orientalis which is reported 
from the Elburz and Armenian mountains * and other parts 
of Asia Minor. 


Ovis OPHION, var. URMIANA. 


I have no materials to demonstrate any craniological cha- 
racters by which this form may differ from either O. orientalis 
or O. ophion, but the size of the skull (and by inference of the 
whole animal) may be conceived from the following measure- 


ments :— 
millim, 

Distance between end of intermaxillary and upper 

rim of occipital foramen ....-....-++-+-+.--- 230 
Distance between end of intermaxillary and palatal 

THO EG lee eee eas Tae cease ce ee Rarah NTL AU acai San eet 120 
Length of molar series ......-- ++ ++. se esse ees 65 and 69 
Distance between the two seriesin front .......... 15 

a a5 Ay behinds eyeqstsiscioe 50 
‘Distance between lower rims of orbits ............ 125 

a Ms styloid) processes) =<). = 26.526 a0. 63 
Greatest width of occipital condyles .....-........ 87 


* W.'T. Blanford, ‘ Hastern Persia, ii. p. 88; Danford & Alston, Proce. 
Zool. Soc. 1880, p. 55. 


Gunther. 


J.Green del et hth. 


OV USMo® 


Tate 


Pp 


JE 


mM. 


Linn, Soc. Journ Zoor, .VYou.X> 


LD. 


Mintern Bros .i 


IRMIANA. 


Len Ce 


Sri mtn per aint 


6 Linn.Soc. Jounn. Zoon Vou. XXVIII. Pr. 2! 
Gunther. 


Mintern Bros 4 


WILD SHEEP OF THE URMI ISLANDS. 375 


The horns are bent outwards in a regular curve, describing a 
semicircle, without any trace of that spiral twist at the extremity 
which seems to be constant in the adult Cyprian Mouflon; in 
fact the whole of their posterior surface, which is broad and flat 
(or partly concave), lies in the same plane; and the horns are so 
little turned backwards, that this plane would make an angle of 
about 70° with a plane vertically bisecting the cranium. The 
horns measure round the outer curve 500, round the inner 305 
millim., their circumference round the base being 190 willim. 
and the distance from tip to tip 360 millim. They are re- 
markably flattened and compressed in a vertical direction, with 
an obtuse upper and a sharp lower ridge; a fronto-orbital 
ridge, which I observe to be still very distinct in O. gmelint 
and O. cycloceros, has nearly disappeared on the left horn, 
and is very obtuse near the base of the right horn. A trans- 
verse section through the left horn about two inches from 
the base would represent one half of an irregular oval (fig. a 
on Plate 22), and one about the middle of the horn would be 
still more compressed (fig. 6). The end of the horn is almost 
knife-shaped. The transverse wrinkles are blunt, coarse, and 
rather distant. Besides these transverse wrinkles, there are at 
irregular intervals five deep grooves penetrating through the sub- 
stance of the horn, and dividing each horn into six sections of 
unequal length. These grooves are, as regards position, perfectly 
symmetrical on both sides, and evidently indicate periods of 
growth (probably annual) ; and if this be the case, the skull would 
be that of an animal six or seven years of age. The terminal 
(oldest) section measures (along the middle) 70 mm., the next 70 
mm., the third 40 mm., the fourth 115 mm., the fifth 75 mm., the 
sixth (during which the animal was killed) 33 mm. Abundance 
or scarcity of food, or other physiological causes, may account 
for the want of regularity in the growth of the horn. 

No trace of the fronto-orbital ridges is visible on the bony 
core, a transverse section of which represents one half of a 
regular oval. 


The colour of the hair attached to the head is now a uniform 
light isabelline, but no importance can be attached to this, as the 
colour may have been bleached by exposure; the horns are also 


similarly bleached, traces of the normal dark colour being still 
visible in some parts. 


376 MR. R. T. GUNTHER ON THE 


If the specific limits of the Asiatic Mouflon (Ovis orientalis 
s. gmelint) be so far extended as to include the Cyprian 
Mouflon*, our sheep should be named Ovis orientalis, var. 
urmiana; but in the:present very fragmentary state of our know- 
ledge of the wild sheep of Western Asia, I think it best to 
associate it with the Cyprian Mouflon, to which it seems to be 
more closely allied than to any of the other forms from Asia 


Minor. 


Tue Priocene MamMatia or THE Bone-BEDS OF MARAGHA. 
By Rozerr T. Ginter, M.A. 


On arriving at Maragha my first enquiries were about the 
fossil bones found in the neighbourhood. I stayed at the house of 
Quasha Mushi, a Christian preacher, who informed me that he had 
already excavated and despatched 14 loads of Mammalian bones 
and two boxes of other stones (? Jurassic fossils) to Dr. Pohlig at 
Vienna. He also told me that his paleontological researches 
had been stopped by a suspicious government who believed that 
all digging was for hidden treasure, and therefore was an illicit 
interference with the rights of the mineral monopolists. 

On Sept. 5th I started soon after sunrise on a three-hours’ 
ride to Kirjawa, the village nearest the bone-beds. The road 
from Maragha crossed a succession of gravelly hills largely com- 
posed of pebbles of andesites of varying basicity, from hornblende- 
andesite to basalt, which shortly before Kirjawa are succeeded by 
hills of pumiceous tufa. Mr. Prior, who has been kind enough 
to examine my specimens, assures me that this volcanic deposit 
is remarkably similar to the tufa of the bone-beds at Samos of 
similar age. 

During the day the villagers and myself were able to pick up 
the bones in the following list. In their identification I have 
been greatly helped by the experience of Dr. C. I. Forsyth Major, 
who has recently studied the contemporaneous fauna of Samos. 
T am glad to take this opportunity of thanking him for the kind 
way in which he interrupted his other researches to help on mine. 
The species in ¢talics have been described from Maragha by other 


paleontologists T. 


* A view taken by Lydekker in ‘ Wild Oxen, Sheep, and Goats.’ 
t+ Cf. Forsyth Major, ‘Le Gisement ossifére de Mitylini.” Lausanne, 1894. 


PLIOCENE MAMMALIA OF MARAGHA. 377 


MAMMALIA. 
CARNIVORA. 


Hyena evimia, Roth. et Wagn. 

Ictitherium hipparionum, Gaudr. 

Meles maraghanus, Kittl. 

ARTIODACTYLA. 
Fam. ANTILOPID#. 

Gazella deperdita, Gaudr. 

Gazeila brevicornis. Horn-core. 

Prostrepsiceros (?) sp. 

Tragoceros amaltheeus, Gaudr. Upper left m', m’. 

Paleoreas lindermayeri, Gaudr. 

Antilcpe sp. Right upper molar. Metacarpal about 1 ft. 2 in. 

in length. 

Paleoryx pallasi, Gaudr. 

Protoryx longiceps, Maj. 

Protoryx gaudryt, May. 

Helicophora rotundicornis, Weith. 

Fam. Ovip2. 
Criotherium argalioides, Maj. Left upper m’. 
Fam. GirAFFID. 
Samotherium boissieri, Maj. 2 calcanea. 
Fam. Suipz. 
Sus erymanthius, Roth. et Wagn Right upper molar. 
PERISSODACTYLA. 
Fam. Equip2. 

Hipparion mediterraneum, Hensel. 14 upper molars and pre- 
molars ; 5 lower molars and premolars; 1 incisor ; 
right and left astragali; fragment of splint bone. 

RHINOCEROTIDA. 
Rhinoceros sp., probably blanfordi, Lyd. Fragments of teeth 


and of lower jaw; patella. 
PROBOSCIDEA. 


Fam. ELEPHANTID2. 
Mastodon pentelici, Gaudr. et Lart. Molar. 
Mastodon sp. Tusk. 
EDENTATA. 
Orycteropus gaudryt, Maj. 


It will thus be seen that the presence of Tragoceros amaltheus, 
Gaudr., at Maragha is confirmed, and that the known fauna has 
been enriched by the discovery of a sheep which Dr. Major con- 
siders identical with his Criotherium argalioides from Samos, and 
of which there is an exceedingly fine skull in the British Museum. 
The presence of Gazella brevicornis is also indicated. 

LINN. JOURN.—ZOOLOGY, VOL. XXVII. 29 


878 MR. G. A. BOULENGER ON THE 


The bones are well preserved in a deposit of light brown 
pumiceous tufa, stratified in horizontal beds of unequal hardness, 
but they are difficult to get out entire without fracture. The 
village people regard the Mastodon bones as the remains of the 
big men who lived before the flood; but the chief man of the 
village classified the bones under the heads elephant, deer, swine, 
unicorn, and “dévy,”’ or men with horns like genn. 


REPTILIA anp AMPHIBIA. 
By G. A. Bouneneer, F.R.S., F.Z.S. 


LACERTILIA. 

}. PHRYNOCEPHALUS HELIoscoPus, Pall. 

Khoi: 1 specimen. North of Lake Urmi: 2 specimens. 

These specimens agree with the Persian variety described 
by De Filippi and by Blanford under the name of P. persicus, 
having a pair of pink spots, edged with blue, on the nape. One 
of the specimens from L. Urmi differs, however, from Blanford’s 
description of P. persicus im having the larger scales along the 
spine bluntly but very distinctly keeled. The large series 
preserved in the British Museum removes all doubts as to the 
specific identity of P. persicus with P. helioscopus. The bright 
nuchal markings are often present in Central-Asian examples. 

An excellent description of the coloration from living speci- 
mens has been given by L. v. Méhely, Zool. Anz. 1894, p. 82; 
but I can see no sufficient ground for the establishment of his 
yar. Horvath. 

9. LACERTA VIRIDIS, Laur. 

Bash Nurashin, Sept. 23. 

A young specimen of the var. strigata, Eichw.: uniform dark 
olive-brown above, with five yellowish-white longitudinal lines. 
Ventrals in 6 longitudinal series; 38 scales across the body; 
17 pores under each thigh. 

3. OpHIoPs ELEGANS, Weénétr. 

Numerous specimens of the typical form, all with two super- 
posed postnasals. Considering the wide range of variation of 
the numbers of scales and femoral pores, it may be of interest 
to record the figures occurring on the specimens collected by 
Mr. Gunther. Under A is given the number of scales round the 


REPTILIA AND AMPHIBIA OF LAKE URW. 3/9 


middle of the body, ventrals included ; under B the number of 
femoral pores on both sides.. 


AX, 13, A. B. 
Aeznlislands et eee 39 9-9 Sein n ouaernrce sanctions 33 = 10-11 
» » Ona ae 31 8-9 5 Gh iahaceocseneneeens 32 10-10 
” soe yah eae 32 9-8 SOME ORR Seeman stance es 32 12-12 
» 3 bets poaeect 32 8-9 AY deers tba Macey 30 9-9 
Shazalan Island, ¢ .... 33 8-8 FEM et doeoeesucdeaqeose 3l 10-11 
Swiyowilels, SS scososseacoe 34 = =11-12 ip SVfBS ccocooabosoe ee SLO =n 
St. George’s Hill, 3. UN Barstontian sans seats 34 10-10 
Superghan ...... 3 31 10-10 KGirjaiwale OMimeep ee eeeee 30 9-9 
» 55 OFS eles) sh FRR eaten ee 34 10-10 
» g 2 8s OA ih syed aaa 32 10-10 
Between Superghan. 
enol Wirear, 6P  Gosons OS 8-8 


4, MABuraA SEPTEMTMNIATA, Reuss. 

A single half-grown specimen from Koyun Daghi. 

34 scales round the body, dorsals indistinctly tricarinate. As 
in one of the Persian specimens in the British Museum (Teheran), 
the four black dorsal stripes of the typical form are present on 
the nape, whilst on the back they are broken up into spots. 


5. HuMEcES ScHNeIDeRI, Daud. 

Two specimens from Koyun Daghi agree entirely with the 
typical, widely distributed form, as figured by Anderson in his 
‘Reptiles of Egypt,’ pl. xxv. Greyish olive above, with small 
orange spots and. an orange lateral streak. 28 scales round the 
middle of the body. The distance between the end of the snout 
and the fore limb is contained once and four-fifths in the distance 
between axilla and groin. 

A remarkably large specimen from Vizastra (17 centimetres 
from snout to vent) is brownish above, without orange spots, 
but with a darker dorsal stripe occupying the two median rows 
of scales, and a still darker stripe, two scales wide, un each side 
above a yellow lateral streak which extends from the mouth, 
through the ear, to the hind limb. The body is more elongate, 
the distance between end of snout and fore limb being contained 
twice in the distance between axilla and groin; and there are but 
24 scales round the body. This lizard approaches the form 
figured by Geoffroy in the ‘ Description de/’Egypte,’ Rept. pl. iii. 
fig. 3, of which we have a specimen from Jerusalem in the British 
Museum, and agrees with specimens from Cyprus (Scincus 
cyprius, Cuy.), Transcaspia, and Persia. 

20% 


380 REPTILIA AND AMPHIBIA OF LAKE URMI. 


A young specimen from Shazalan Island (73 millim. from 
snout to vent) agrees in the proportions and the number of scales 
(28 round the body) with the Koyun Daghi examples, but the 
back is marked with four blackish stripes, in addition to which 
every other scale in every longitudinal series bears a central 
yellowish spot. This system of coloration is closely approached by 
that of a young specimen from Jerusalem in the British Museum, 
which differs only in having a blackish line bordering the belly 
below the yellow lateral stripe. 

A study of the juvenile livery of this species would be a great 
help towards the establishment of geographical varieties ; but it 
is a most curious, and to me unaccountable fact, that young 
specimens of the common Palearctic Humeces are extremely rare 
in collections. I have only seen three of HL. Schneideri and not 
one of #. algeriensis. 


OPHIDIA. 
6. Eryx gacutus, LD. 


A single specimen from Seir, near Urmi. It agrees in every 
respect with the typical or Western form, the habitat of which 
is known to extend from Greece and the Ionian Islands to Lower 
Egypt and Persia; the Eastern form, var. mzdiaris, with smaller 
head-shields, inhabiting Transcaspta, Turkestan, and Afghanistan. 
The Urmi specimen has 5 scales between the eyes, 3 between 
posterior nasal and eye, 9 round the eye, and 9 upper labials 
on each side. 48 scales across the thickest part of the body. 
Ventrals 186; subcaudals 18. eid 


7. TROPIDONOTUS TESSELLATUS, Laur. | 

Eastern shore of L. Urmi, Seir, Maragha. Four specimens. 

This is acommon and widely-distributed snake ; but one of the 
specimens is of interest in having the upper labials almost 
excluded from the eye by the suboculars, merely the point of the 
fourth labial penetrating between the latter. Specimens of this 
species may probably turn up some day with the labials entirely 
excluded from the eye, asin Tropidonotus ferox, anoscopus, and 
cyclopium. 


BATRACHIA. 
8. Rawa EscuLenta, LZ. 


Several specimens, half-grown, young, and larve, belonging to 
the var. ridibunda, Pall. Ocksa, Superghan, Maragha, Kirjawa. 


FISHES OF LAKE URMI. 881 


9. Rana Camerant, Blgr. 
Banda, Urmi River. ‘Two young specimens. 
10. Buro viripis, Laur. | 


“young specimen from Sujbulak and several tadpoles frem 
the town-ditch at Urnii. 


11. Hyza arporza, Z. 

Several specimens, half-grown, young, and larve, belonging to 
the var. Savignyi, Aud. Bash Nurashin, Urmi town, Banda, 
Superghan; also a young specimen from Lake Urmi, “in salt 
water.” 

The tadpoles have the black lines on the muscular part of the 
tail to which attention has been drawn by Camerano. 


12. PELOBATES FuUSCUS, Laur. 

Three large tadpoles from Superghan, one with well-developed 
limbs. This species was known to occur at Lenkoran on the 
Caspian, but had not yet been recorded from Persia. 


FISHES. 
By Dr. A. Gtnrner, F.R.S., P.L.S. 
(Piares 23 & 24.) 


Tue general character of the Freshwater-fish Fauna of Western 
Asia, as a part of the Huropxo-Asiatic region, is well known 
through the researches of Russell, Heckel, de Filippi, Keyserling, 
Kessler, Radde, Lortet, Sauvage, and myself *; and in this respect 
the small collection from the Urmi basin does not contribute 
any additional facts of great novelty. On the other hand, our 
acquaintance with the local faunas of the various districts of 
Western Asia is more or less fragmentary, and a comparative 
study of the several species of which they are composed is a great 
desideratum. Of the Fish-fauna of the Urmi district we had no 
positive knowledge whatever; and the materials available at 
present for a useful direct comparison of the species inhabiting 
the rivers which take their rise in the Kurdistan and Armenian 
mountains, viz., the Tigris, Euphrates, Kur, Araxes, and the Lake 
Van and Urmi rivers, are extremely scanty and insufficient for the 
purpose. 


* Consult more especially H. H. Sauvage, “ Notice sur la Faune ichthyologique 
de l'Ouest de l’Asie,’”’ Nouy. Arch. Mus. vii. 1884. 


382 DR. A. GUNTHER ON THE 


In the consideration of the Urmi fauna we have to bear in 
mind that the rivers of this basin do not communicate with the 
sea, and, consequently, that migratory fishes are absent: like the 
Jordan, they are not inhabited by shad, eel, sturgeon, or lamprey. 
Neither is there direct communication between the rivers them- 
selves in spite of the close proximity of their mouths, the water 
of the lake being of such concentrated salinity as to prevent the 
passage of fish from one river into another. Such an isolation of 
the inhabitants of the several rivers must have been a favourable 
factor for the development of differential specific characters. The 
great severity of the climate in winter will account for the absence 
of warmth-loving types such as Oyprinodon. 

If we are permitted to draw conclusions from so small a col- 
lection, we may say that the fauna bears distinctly the character 
of that of Central Europe, so far as the genera are concerned 
(with the addition of Oapoéta); and although the majority of the 
Species seem to be sufficiently differentiated local representatives, 
two of them, namely, the Silurus and Chub, are identical with the 
European forms. Probably further investigations may reveal 
the presence of certain other fishes, such ag stickleback, trout, 
Rhodeus, which Mr. Gunther has not been able to find; but the 
fish-fauna as a whole is certain to prove to be poor in species, 
much poorer than that of the Kurand Araxes, to which, however, 
it is most closely allied, as is evidenced by the identity (real or 
supposed) of five out of the ten species enumerated below. 

The specimens were collected,—- 

1. In the Tatawa Chai near Sujbulak, entering the south end 
of the Lake. 

2. In the Gader Chai near Ocksa, likewise at the southern end 
of the Lake. 

3. In the Urmi River (Shaher Chai). 

4. In the Nazlu Chai, entering the north-western part of the 
Lake, at Superghan near the mouth, and at Tergawar in its upper 
courses. 


5. In the Zola Chai, near Ula. 


SILURUS GLANTS, D. 
It might have been supposed that the Silurus of Western 
Persia would prove to be identical with the form which Sauvage * 


* Bull. Soc. Philom. 1882, p. 163; Nouv. Arch. Mus. 1884, vii. p- 19, pl. 
1K, I 


FISHES OF LAKE URMTI. 383 


has described from the River Kur under the name of Silurus 
chantret. But this supposition is not borne out by a specimen 
obtained in the Gader Chai at Ocksa. It is the skin of an adult 
individual much cut about and without pectoral fins. However, 
the number of anal rays can be ascertained to be 84, thus nearly 
approaching in this respect our European 8S. glanis, whilst 65 is 
the number given for 8. chantrei. Algo in other respects, espe- 
cially with regard to the composition of the dorsal fin and the 
length of the barbels, it proves to be a typical S. glanis. I have 
some doubts as to whether S. tréostegus of Heckel, from the 
Tigris, can be maintained as a distinct species. The dorsal fin 
of Sz/wrus is a rudimentary organ, and therefore may be expected 
to vary in the number and development of its rays. Even in 
Kuropean specimens the fourth dorsal ray is not constantly 
branched, but may be simple and reduced in size, and I have 
found it so also in a specimen from the Tigris. However, the 
barbels of Mesopotamian specimens seem to be shorter than in 
European. 


CAPOETA GRACILIS. 

Scaphiodon gracilis, Keyserling, Zeitschr. ges. Naturw. xvi. 1861, p. 9, 
tab. 4. 

Scaphiodon sieboldi, Steindachner, Verh. zool.-bot. Ges. Wien, 1864, 
p. 224. 

? Capoéta sevangi, De Filippi, Viaggio in Persia, p, 812 (1865). 

Capoéta gracilis, Ginther, Fish. vii. p. 80 (1868). 

D. 3/8. A. 3/5. IL. lat. 58-55. LL. transv. 84/6-7. 

The height of the body is one fourth or two ninths, the length 
of the head two ninths of the total length (without caudal) ; the 
diameter of the eye is one fifth of the length of the head and 
‘contained 12 in the length of the snout, and 24 in the width of 
the interorbital space. Barbels rather shorter than the eye. 
Origin of the dorsal fin midway between the end of the snout 
and the root of the caudal; distance between the first dorsal ray 
and occiput rather less than that between the last dorsal ray and 
root of the caudal. Least depth of the caudal peduncle about 
one half of the length of the head. Third dorsal ray feeble, soft 
jn its terminal portion, and distinctly serrated, the serrature 
being hidden below the skin. Pectoral rather shorter than the 
head, not extending to the ventral. Caudal deeply emarginate. 
Scales in the anal region not distinctly enlarged. Coloration 
uniform. 


384 DR. A. GUNTHER ON THE 


This fish was originally described from the neighbourhood of 
Ispahan, and, if my identifications are correct, at later periods 
from Amasia (Asia Minor) by Steindacbner, and from Lake 
Gokcha by De Filippi. Thus it seems to have a considerable range 
in Western Asia, and the present collection contains specimens 
from Sujbulak, from the Urmi River, and the Nazlu Chai. 
The largest specimen is 213 millim. long. The species is subject 
to a considerable amount of variation, not merely dependent on 
local influences, but apparently quite individual, as it obtains in 
specimens captured at the same spot and at the same time. In 
specimens from Sujbulak the cleft of the mouth is quite straight, 
extending the whole width from side to side, the dorsal spine is 
very feeble, and the crown of the head flattened. In a specimen 
from the Urmi River the crown of the head is more convex, the 
dorsal spine sensibly stronger, the mouth gently crescent-shaped, 
and there are only six longitudinal series of scales between lateral 
line and ventral fin. The form of the mouth varies in three 
specimens from Superghan (Nazlu Chai), from straight to a gentle 
crescent, and to a distinct crescent; the strength of the dorsal 
spine is intermediate between the Sujbulak and Urmi specimens, 
and the longitudinal rows of scales between lateral line and 
ventral fin varies from six to seven; they have altogether the 
appearance of being less well-fed than the specimens from Suj- 
bulak. Two specimens from Ula represent a dwarf form with 
very feeble and indistinctly serrated dorsal spine; the larger, a 
male, is only 125 millim. long, yet fully mature, with developed 
testicles and with a seasonal growth of minute tubercles along 
the whole side of the body. 

These fishes, for which I have adopted Keyserling’s name, 
belong to a group of the genus which is distinguished by the 
feeble development of the dorsal spine and by scales of moderate 
size. The fishes of this group show a considerable amount 
of modification of the characters of various organs, such as I 
have pointed out in the specimens from the Urmi district. Con- 
sequently they have been described under numerous specific 
names; but without long series of specimens with exact localities, 
it is impossible to form an opinion as to which of these names 
deserve recognition. 

Thus, Capoéta steindachneri, Kessler (Izvest. obshchest. Lynbit. 
estestv. x., Mose. 1872, p. 47, pl. 6. figs. 8-5, or Fedschenko, Fauna 
of Turkestan, Pisces, St. Petersb, 1874, p. 7, pl. 1. figs. 3 & 4), 


FISHES OF LAKE URMI. 3385 


seems to be distinguished (if we judge from the figure) from our 
C. gracilis by a greater number of scale-rows between lateral 
line and ventral fin. It is said to have one or two pairs of 
barbels. The specimens were from the River Sarefschan. 

Filippi’s description of his Capoéta sevangi from Lake Gokcha 
contains nothing by which we could distinguish this fish from 
C. gracilis. But the fish described and figured under the name 
of C. sevangi by Kessler * is certainly distinct from our C. gracilis. 
As, according to Kessler’s investigations tT, Lake Gokcha is in- 
habited by several species of Capoéta, we are left in uncertainty 
as to whether the fishes named sevangz by Filippiand Kessler are 
specifically the same. 

Scaphiodon sieboldii, Steindachner, is described from a speci- 
men 133 millim. long. The eye is noted as of somewhat larger 
size than in our specimens, but the comparative size of this 
organ is subject to the usual changes with the growth of the 
individual. 


BaRBus CAUCASICUS. 

? Barbus caucasicus, Kessler, Aralo-Caspio-Pontine Ichthyol., in Grimm, 
Aralo-Casp. Exped. p. 102. 

D.10. A.8 UL. lat. 90. L. transy. 18-14/18-20. 

The height of the body is contained 53 or 53 times in the total 
length (without caudal), the length of the head 42 or 43 times. 
The diameter of the eye is one half of the length of the snout 
and two elevenths of that of the head; interorbital space not 
quite twice as wide as the eye. Anterior barbels a little shorter 
than posterior, which are not twice as long as the eye. Caudal 
peduncle longer than deep. Dorsal spine rather feeble, but 
finely serrated behind, somewhat nearer to the root of the caudal 
than to the end of the snout, opposite to the root of the ventral. 
Anal fin not reaching to the caudal. Caudal slightly emarginate, 
but in young examples (of 110 millim.) the excision is deeper. 
Pectoral shorter than the head, rounded. Eleven longitudinal 


* “ Wish of the Aralo-Caspio-Pontine region,” in Suppl. Trans. St. Petersb. 
Nat. Hist. Soc. 1877, p. 81, fig. 18. 

t L.c.; Sauvage, Nouv. Arch. Mus. vi. 1884, pp. 5, 22. In the figure of 
C. gotschaica (pl. 3. fig. 3) the scales are represented much larger than in the 
description, 

{ 90, counting the transverse series above the lateral line; the perforated 
scales of the lateral line are somewhat larger and irregular, and about 80 in 
number. 


356 DR. A. GUNTHER ON THE 


series of scales between the lateral line and ventral fin; pharyn- 
geal teeth 5 | 3 |1. Sides of the body, dorsal and caudal fins 
speckled with greyish. 

Three small specimens were obtained in the Zola Chai at Ula, 
and a larger one in Tergawar, in the upper waters of the Nazlu 
Chai. The latter is 153 millim. long, and presumably young. It 
shows some larger scales on each side of the vent, and therefore 
the question arises whether our fishes are the young stage of a 
species of Schizothorax. I have no materials which would assist 
me in ascertaining whether the peculiar anal sheath of Schizo- 
thorax is developed with age. The two genera, Barbus and 
Schizothorax, are so closely allied that in all probability species 
exist in which the initial stages of the development of an anal 
sheath may be traced either as a distinctive specific character or 
ag an individual, more or less abnormal condition. 

The determination of these Persian specimens as Barbus cau- 
casicus is not by any means satisfactory. I should not have 
recognized them from Kessler’s description, who gives as scale- 
11-12 

8-9 
larger; but the Natural History Museum has received from 
Russian sources, two larger specimens named B. caucasicus. 
These specimens approach the Persian closely enough to be 
referred to the same species. That collection possesses also 
specimens of Barbus ciscaucasicus and Barbus goktschaicus, two 
other species described by Kessler in the work quoted. These 
also are most closely allied to B. caucasicus, but B. goktschaicus 
has the dorsal spine extremely feeble. I have finally to add that 
neither of the two specimens of B. caucasicus in the Natural 
History Museum has enlarged anal scales, and that in one of 
them the anal fin reaches to the caudal, when laid backwards. 

Our fishes differ from Barbus miliaris (de Filippi, Viaggio, 
p- 358), from Teheran, in having only 13 or 14 series of scales 
between the dorsal fin and lateral line, the Teheran species 
having 18. 


formula 60, > 65, indicating a fish with scales considerably 


GoBio PERSA, sp. n. (Pl. 23. fig. B.) 

D.10. A.8. L. lat. 43. L. transv. 6/9. 

The height of the body is contained 53 times in the total length 
(without caudal), the length of the head 4 times; caudal peduncle 
slender, but slightly compressed, its greatest depth being two 
fifths of its length. Snout not greatly elongate, the diameter of 


‘FISHES OF LAKE URMTI. 387 


the eye being two thirds of its length and one fourth of the 
length of the head ; cnterorbital space broad and fiat, its width 
being a little more than the diameter of the eye. Upper jaw 
overlapping, but not much projecting beyond the lower ; maxillary 
barbel reaching to the hind margin of the eye. Dorsal fin with 
seven, anal with six branched rays; pectoral reaching the ventral, 
ventral the anal. Caudal moderately excised, with pointed lobes. 
Five series of scales between lateral line and ventral fin. Back 
and sides irregularly speckled with black; a series of from seven 
to nine larger ovate spots along the lateral line. Dorsal and 
caudal rays speckled with black. 

Seven specimens, 70 millim. long, from Ocksa in the Gader 
Chai. 

This form of Gudgeon comes nearest to G. wranoscopus, having 
a slender peduncle of the tail, but slightly compressed; and com- 
pared with specimens of the same size, it shows a distinctly 
shorter snout and a greater width of the interorbital space. I 
have not the means of comparing it with Gtobio kessleri from the 
Dniester, which, however, is described as possessing eight branched 
dorsal rays. 

Levciscus cEPHALUS, LZ. 

The Chub is one of the most common fishes in the rivers 
falling into Lake Urmi. Specimens of small and moderate size 
were obtained at Sujbulak, in the Gader Chai and Urmi Rivers, 
and in the Nazlu Chai. I consider them specifically ideutical 
with the European Chub; their head is equally broad, but 
rather more elongate or depressed than is usually observed in 
British specimens. But Continental specimens frequently show 
the same degree of elongation and the same form as the Persian 
fishes. 

The short description given by de Filippi of his Squwalius 
turcicus (Viagg. Pers. p. 859), from Erzeroum, applies very well 
to these Persian specimens. 


LEUCISCUS ULANUS, Sp. n. (PI. 24. fig. A.) 
D.10-11. A.18. L. lat. 44. L. transv. 8/5. 

The height of the body is contained from 33 to 43 times in the 
total length (without caudal), the length of the head 4 or 43 times. 
The diameter of the eye is one fourth of the length of the head 
and a little less than the width of the interorbital space, which ig 
transversely convex. Snout obtuse, as long as the eye; upper 


388 DR. Ae GUNTHER ON THE 


jaw slightly overlapping the lower; cleft of the mouth oblique, 
the maxillary not quite reaching the vertical from the front 
margin of the orbit. Nape of the neck but little raised above 
the level of the head. Extremities of the fins obtusely rounded 
off. The origin of the dorsal fin is nearer to the root of the 
caudal than to the end of the snout, and behind the vertical from 
the root of the ventral; it is higher than long. Anal fin rather 
lower than long. Caudal excision moderate. Pectoral shorter 
than the head, not reaching the ventral. Caudal peduncle nearly 
or not quite twice as long as deep. ‘There are three series of 
scales between the lateral line and ventral fin. Back bluish, sides 
silvery, both colours separated by a narrow straight black band 
running from the upper half of the eye to the end of the lateral 
line. Pharyngeal teeth 5.2-2.5. 

Two specimens, 105 and 83 millim. long, from Ula on the Zola 
Chai. 


LEUCISCUS GADERANUS, sp. n. (Pl. 24. fig. B.) 
D.11. A.12. UL. lat. 40. L. transv. 64/4. 

The height of the body equals the length of the head and is 
one fourth of the total (without caudal). Diameter of the eye 
one fourth of the length of the head and a little less than the 
width of the imterorbital space, which is transversely convex. 
Snout obtuse, as long as the eye; upper jaw slightly overlapping 
the lower; cleft of the mouth oblique, the maxillary not reaching 
the vertical from the front margin of the orbit. Nape of the 
neck but little raised above the level of the head. The origin of 
the dorsal fin is nearer to the root of the caudal than to the 
end of the snout, and behind the vertical from the root of the 
ventral; it is higher than long. Anal fin a little higher than 
long. Caudal excision moderate, lobes pointed. Pectoral shorter 
than the head, not reaching the ventral. Caudal peduncle twice 
as long asdeep. There are two and a half series of scales between 
the lateral line and ventral fin. Back bluish, sides silvery, dotted 
with numerous minute pigment-spots; a narrow straight blackish 
band runs from the upper end of the gill-opening to the end of 
the lateral line. Pharyngeal teeth 5.2-2.5. 

Three specimens, the largest 90 millim. long, from the Gader 
Chai; three young specimens from near the mouth of the Nazlu 
Chai at Superghan. 

The propriety of distinguishing this form from L. ulanus under 


FISHES OF LAKE URMI. 389 


a separate name may be questioned. However, the larger size 
of the scales is so striking a feature, that, without having inter- 
mediate forms, I think it better to keep the two forms distinct. 
Telestes leucoides, Filippi, Viagg. Pers. p. 359, from the Batoum 
River, seems to be also a closely allied species, but is described 
as agreeing in form with Z. aula, which is a fish with a much 
higher body. 


ABRAMIS URMIANUS, sp. n. (PI. 23. fig. A.) 

D.11. A.14-16*. IL. lat. 58-62. LL. trangv. 12/8. 

The height of the body is two sevenths, the length of the head 
one fourth of the total (without caudal). Snout neither pointed 
nor obtuse, equal to the diameter of the eye, which is one fourth 
of the length of the head ; interorbital space transversely convex, 
scarcely wider than the orbit. Cleft of the mouth slightly oblique, 
with the jaws equal in front, the maxillary not extending to the 
vertical from the front margin of the eye. Abdomen rounded in 
front, and compressed behind, the ventrals. Caudal peduncle 
longer than deep. Pectorals not reaching the ventral, shorter 
than the head. Origin of the dorsal midway between the end of 
the snout and the root of the caudal. Caudal excision moderate. 
Five series of scales between the lateral line and ventral fin. 
Pharyngeal teeth 5/2, hooked. Silvery, greenish-olive on the 
back; sides with numerous minute brownish pigment-spots ; 
they are more crowded above the lateral line, producing 
an imconspicuous darker band along the whole length of 
the side. 

Five specimens from the Gader Chai and two small ones from 
the Urmi River; the largest is only 144 millim. long, but 
specimens of this size are mature, showing not only fully de- 
veloped sexual organs, but also some of those deep-black spots 
which appear in so many Cyprinoids during the breeding-season. 
The gill-rakers are very short, triangular in shape, and widely 
set, as is characteristic of the genus Abramis, in opposition to 
Alburnus. 

This species resembles Alburnus punctulatus, Kessler (Aralo- 
Caucas.-Pont. Ichthyol. p. 159), but has a shorter and smaller 
anal fin, whilst in the former species this fin is composed of 17 


* 14 in two, 15 in five, 16 in one specimen ; the first three rays being simple 
in all. 


390 FISHES OF LAKE URMTI. 


to 20 rays. The gill-rakers of A. punctulatus are very short ; 
and therefore this species should be referred to Abramis. A 
specimen from Tiflis is in the Natural History Museum. 


ALBURNUS FILIPPII. 

Alburnus filippit, Kessler, in Grimm’s Aralo-Caspian Exped., Pisces: 
1877, p. 153. 

D.11. A.18-14*. IL. lat. (54) 56-60. LL. transv. (9) 10/6. 

The height of the body is contained 42 times in the total length 
(without caudal), the length of the head 43 times. Snout rather 
pointed, equal to the diameter of the eye, which is scarcely one 
fourth of the length of the head. Interorbital space transversely 
slightly convex, scarcely wider than the orbit. Cleft of the 
mouth oblique, with the lower jaw slightly the longer, the max- 
illary not extending to the vertical from the front margin of the 
eye. Abdomen rounded in front, slightly compressed behind, 
the ventrals. Caudal peduncle at least twice as long as deep. 
Pectorals not reaching the ventral, shorter than the head. Origin 
of the dorsal nearer to the root of the caudal than to the end of 
the snout. Caudal excision moderate. Four series of scales 
between the lateral line and ventral fin. Pharyngeal teeth f 4/2, 
looked. Silvery, olive on the back; a narrow, straight, well- 
defined blackish band from the upper end of the gill-opening to 
the middle of the caudal fin. 

Three specimens from Sujbulak, the largest 113 millim. long ; 
ohe specimen from Superghan near the mouth of the NazJu 
Chai. 

The gill-rakers are lanceolate, closely set, but the longest 
scarcely half as long as the eye. 

The specimen from the Nazlu Chai differs slightly from the 
others, inasmuch as the scales are apparently a little larger; I 
count only 54 in the lateral line, and 9 in the transverse series 
between dorsal fin and lateral line. 

Kessler states that his specimens were obtained from the Kur 
at Tiflis and at Borjom; in the fin-formula the number of dorsal 
rays is stated to be 1/6-7, which would be so abnormal in this 
genus that I suspect it to be due to some inadvertence. 


* 13 in one, 14 in three specimens, the three anterior rays being simpl>. 
+ Examined in one specimen only. 


ies States Gia Dc ASRSNC RMON (0h a) “SONVINUN SINVUEV'V “aH 92 Tsp e250 ¢ 


"@@ Td TAKX- TO/_ 1007 Naf (* 2 00G “NNIT “eyquns) 


Gunther . Ln. Soc. Journ. Zoot Vou. XXVII.Pi.24. 


J.Green del et lth. ier eee ate 
AAG UCISCOS WLANUS - BEE UCISiCUS GCADERAN USE 


MOLLUSCA OF LAKE URMI. 891 


NEMACHILUS PERSA. 

Cobitis persa, Heckel, in Russegger’s Reis. i. 38. p. 266; de Filippi, 
Viaggio, p. 360. 

Nemachilus persa, Giinth. Fish.’ vii. p. 347. 

ID ys ae ieaeles MOE We Me 

Scales minute, but conspicuous. Caudal fin distinctly emar- 
ginate. Origin of the dorsal fin nearer to the root of the caudal 
than to the end of the snout. The height of the body is less 
than the length of the head, which is contained 41 times in the 
total (without caudal). Head rather narrow ; snout somewhat 
pointed, nearly as long as the postorbital portion of the head ; 
eye small, Pectoral as long as the head, its length being two 
thirds of the distance between its base and that of the ventral. 
Caudal peduncle nearly twice as long as deep. Whitish, densely 
reticulated and speckled with greyish. Dorsal and caudal fins 
speckled with greyish. 

Four specimens, 65 millim. long, from the Zola Chai; others 
from the Hlinja Chai, a tributary of the Araxes. 

The specimens on which this species was founded were obtained 
at Persepolis; de Filippi found it generally distributed in 
abundance in all Persian rivulets. 


EXPLANATION OF PLATES 23 & 24. 


Pl, 23. Fig. A. Abramis urmianus, sp. n. 
Fig. B. Gobio persa, sp. w. 

Pl. 24. Fig. A. Leuciscus ulanus, sp. n. 
Fig. B. Leuciscus gaderanus, sp. n. 


LAND AND FRESHWATER MOLLUSCA. 
By Epvear Surra, F.Z.S. 


1. Herrs (HELICOGENA) FIGULINA (Parreyss). 
Hab. Seir. 


2. HELICELLA ACUTISTRIA (Bottger). 
flab. Koyun Daghi. 


3. HELICELLA PARABLETA (Bottger). 
Hab. Urmi district (?). 


392 MR. E. SMITH ON THE 


4, HELICELLA, 0. sp. 

Hab. Seir. 

A single specimen only, with the peristome immature. Prof. 
Dr. O. Bottger very kindly examined this and several other species 
enumerated in this list, and considers it a new species “aus der 
Gruppe apicina, Lmk.” He also observes, “ Aehnliche Arten sind 
meines Wissens aus Asien noch nicht beschrieben.” It has very 
much the form of the preceding species, but with the spire less 
elevated. It is white, prettily marked with about eight inter- 
rupted spotted brown spiral lines, whereof three are above the 
obtusely-keeled periphery, the rest and a few intervening series 
of minute dots being on the lower surface. Some of the brown 
spots are punctate, especially the row just above the peripheral 
keel. The two nuclear whorls are light corneous and smooth, 
the remaining three volutions being marked with strong, very 
oblique, incised lines of growth. 


5. HELICELLA PIsIFORMIS (Pfeiffer). 
Hab. Seir. 


6. Butiminus (Zesrinus) HomENacKeRrt (Arynickz). 
Hab. Ula, Plain of Salmas. 


7. BULTMINUS (CHONDRULUS) TRIDENS (JZuller). 

Hab. Ula?, Plain of Salmas. 

These specimens belong to the var. major, Kryn. (=bayernt, 
Parr.), and var. diffusus, Mouss. Of the latter variety Prof. 
Bottger possesses specimens without any trace of oral teeth from 
several localities. 


8. Buriminus (CHonpDRULUS) TETRODON, Wortillet. 

Hab. Seir. 

A rare species, occurring also in Armenia and Transcaucasia 
near Tiflis. 

9. Butiminus (CHONDRULUS) DIDYMODUS, Bottger. 

Hab. Koyun Daghi. 

The series of specimens exhibit considerable variation in size, 
the convexity or flatness of the whorls, and the general form, 
but the armature of the aperture is very constant. 


10. Butrminus (AMPHISCOPUS) CONTINENS, Hosen.* 
Hab. Koyun Daghi. il 
Three rather short examples, 6 millim. long. 


* Nachrichtsblatt deutsch. Malak. Gesell. 1892, p. 125. 


MOLLUSCA OF LAKE URME. 393 


11. Pupa eranum, Draparnaud. 
flab. Koyun Daghi. 


12. Pura stenata, Mousson. 
Hab. Koyun Daghi. 


13. Limy ma staenaris, Linn. 
fab. Urmi district (?). 


14, Taimnma panusrreis, Muller. 
Hab. Maragha. 


15. LivNwA TRUNCATULA, Muller. 
Hab. Seir. 


16. PLANORBIS MARGINATUS, Draparnaud. 
Hab. Maragha. 


17. SpH#@rium LacustrRe, Muller. 
Hab. Town ditch, Urmi. 


18. Unto Sinversi, Drouet, var. KoBELrt. 

Hab. River Gader near Ocksa. 

Prof. Bottger considers this form distinct from U. batavus, 
with which I had united it. 


The collection, presented by Mr. Gunther, although containing 
few specimens of each species, and mostly in poor condition, 
forms a useful addition to the Museum collection, which contains 
but very few specimens from the north-western part of Persia. 
Their value is also considerably enhanced by their having been 
partly named, or their names confirmed, by Prof. Bottger. Most 
of the species quoted are extremely common forms, having a wide 
geographical range in the South and South-east of Europe and 
Turkey in Asia. Their distribution is weli known and fully 
recorded in various works. Westerlund’s ‘ Fauna der paliarct- 
ischen Region,’ and Rossmassler’s ‘ Iconographie der Land- und 
Sitisswasser-Mollusken, ete.’ should be consulted for such 
information, references, and figures. 


LINN. JOURN.—ZOOLOGY, VOL. XXVII. 30 


394 MR. R. T. GUNTHER ON THE 


CRUSTACEA. 
By Roserr T. Gtnrunr, M.A., F.R.G.S. 


(PuatTe 25.) 


MALACOSTRACA. 
DECAPODA. 


Macrvura. 
IT was unable to discover that the Crayfish is known from any 
of the streams that flow into the Jake. It is abundant in the 
Kur. The Syrian zoologist does not differentiate between crabs 
and crayfish: to him both are ‘ Kédjala’ and unclean. 


BRACHYURA. 

TELPHUSA FLUVIATILIS, LD. 

This freshwater crab is extremely common in all the rivers. 
They are to be found beneath stones and in little burrows in 
wet banks. A favourite attitude is sitting half in and half out 
of the water, so that they are able to pass either air or water at 
pleasure through the branchial chamber. They seemed to be able 
to employ the openings into the branchial chamber either as 
exhalant or as inhalant apertures. Indeed, instead of continually 
propelling water through the branchial chamber in the same 
direction, they were continually changing the direction, and using 
the apertures at the anterior margin of the carapace and at the 


axils of the ambulatory appendages alternately as inhalant and 
exhalant apertures. 


When sitting regardant, their eyestalks and feelers are set at 
an angle of 45° with the antero-posterior axis of the body. In 
colour they are of a greenish-sepia hue, often tending more to 
ereen in older specimens, owing to the growth of a green a!ga on 
the carapace. The chele and ambulatory limbs are generally 
tipped with orange-red, and their sides are often streaked witha 
purplish tinge. 

The Persian specimens do not appear to differ essentially from 
those of Italy, Greece, and Palestine which are preserved in the 
National Collection in the British Museum. 


Isopopa. 
HEMILEPISTUS. 


Common under stones in damp places at Seir. 


CRUSTACEA OF LAKE URMI. 395 

Asutuus, which is so common in all European waters, seemed 

to be entirely absent in the Urmi basin. It is said to occur in 

the western tributaries of the Caspian, but not in Russian 
Turkestan or N.H. Persia. 


AMPHIPODA. 


GAMMARUS PULEX, De Geer. 

In the springs near Seir, and also on the other side of the lake 
in the clear water of a stream near Kirjawa. My observations 
confirm those of Dr. Walter in regard to the fact that G. pulew 
seems to prefer the cool, upper, narrow courses of the streams to 
the warmer, more sluggish, lower reaches in the plains. 


ENTOMOSTRACA. 
CoPEPODA. 


A Cyclopid was very common in a pond near Kirjawa, 
Maragha, but I was not able to determine the species. 


PHYLLOPODA. 
DAPHNIA Sp. 
Common in the stagnant water of the ditch surrounding the 
walls of the town of Urmi; also at Kirjawa. 


ARTEMIA URMIANA, sp. n. (PI. 25.) 

The general shape of the body is similar to that of A. salina, 
but stouter than in the American species. The males are some- 
what shorter than the females, in the proportion of 10 mm. to 
13 mm. The head and “ thorax,” bearing the eleven swimming- 
appendages, are very slender, and measured together are nearly 
equal to the abdomen in length. The segmentation of the abdo- 
men is so difficult to recognize that I am inclined to regard the seg- 
mentation as having been partially lost. The first two or three 
abdominal segments are the only ones clearly distinguishable. 
The labrum is well developed and is bent over the mouth so as 
to hide the mandibles. Its distal end is square-ended, with two 
papilliform processes at the corners; the processes bear small tufts 
of hairs at their apices, and are probably sensory in function. 

The ventral appendage of the genital segment differs greatly 
in the two sexes. In the female it has the function of an ovisac 
and usually contains ripe eggs. Its shape has been described as 
resembling that of broad flask. When quite ripe the eggs 

3U* 


396 MR. R. T. GUNTHER ON THE 


escape through a transverse slit just below the apex of the 
ovisac. The ovaries extend into the posterior thoracic and 
anterior abdominal segments. The eggs vary from about °25 to 
‘°3 mm. in diameter. 

In the male the vasa deferentia open at the slightly expanded 
extremities of a V-shaped organ, which probably contains paired 
sperm-sacs and ductus ejaculatoria. The distance between the 
external sexual apertures exactly corresponds to that between 
the two processes on the claspers, which will be described below, 
and which I am inclined to consider as being employed for the 
purpose of transferring the spermatozoa to the female. Near 
the base of each of the two processes, and on its inner aspect, 1s 
a small spinular process with a sharp tooth. 

The abdomen terminates in a slight dilatation, within which 
numerous rectal muscles arranged radially and obliquely traverse 
the space between the body-wall and the rectum. The anus is 
terminal and is flanked by two small lateral furcal lobes, upon 
which are implanted two simple non-plumose sete in the males, 
but none in the females. In respect of the scant development of 
these sete, A. urmiana approximates to the condition of A. mul- 
hausenti, as Schmankewitsch described it, from brine of sp. gr. 
1-2015, rather than the form from brine of 1:1878 sp. gr., which 
was provided with one or two, though seldom three, sete. At 
the same time it must be remembered that A. wrmiana has lived 
for a longer time in its water of sp. gr. 11188 than Schmanke- 
witsch’s type from water of sp. gr. 1:1373, and has therefore had 
a longer time to develop those anatomical characters which seem 
to be the direct result of life in brine. 

Appendages.—Vhe antenne are sometimes twice the length of 
the eyes and eyestalks. The joints seem to vary in number and 
relative proportion ; as many as five may be distinguished in some 
individuals. The terminal joint bears three sensory hairs. 

The claspers are enormously developed in the adult males, and 
may measure as much as 4 mm. across. The proximal joints are 
much thickened, and carry upon their inner faces two rounded 
processes which are used for clutching the females, and which are 
probably of service in transferring the spermatozoa (spermato- 
phores ?) to the females from the two lateral expansions of the 
male genital organs. The second joint is large, flattened and 
triangular in shape; it terminates in a pointed process which is 
often incurved. 


CRUSTACEA OF LAKE URMI. 397 


The female homologues of the claspers are insignificant in 
comparison, their width not exceeding 1°5 mm. across. 

Mandibles are powerful, with a finely serrated margin and a 
palp which does not appear to be always present. 

Maxille are in two pairs. The first is provided with a tuft of 
anteriorly directed sete. 

The eleven natatory appendages are all constructed en the 
same plan. ‘Their axes all bear two respiratory bracts upon one 
margin and five endites upon the other. Tne 6th endite is 
terminal. The 1st or proximal endite bears extremely numerous 
close-set sete upon its two lobes in all the swimming-appendages 
except the last, in which the sete of the distal lobe are longer 
and not so closely set as upon the longer proximal lobe. 

Endites 2, 3, and 4 are small and only bear 3, 2, and | set 
apiece, respectively. The 5th endite has a rounded margio 
provided with numerous sete armed with reflected barbs. Such 
barbed sete do not appear to be present on any specimens of 
A. salina which I have had the opportunity oi examining, aud 
they are certainly absent in some specimens from Guernsey 
which were given to me by the Rev. Canon Norman. The setz 
upon the distal portions of the margin are much longer than 
those on the proximal portion, and more nearly resemble the long 
swimming-sete borne by the terminal endite. 

The extraordinary numbers in which the Artemia is found ia 
Lake Urmi have already been referred to (p. 357). The females, 
as in Lake Utah, were present in greater numbers than the males, 
in the proportion of 5:3. Many of the males were holding on 
behind the ovisacs of the females by means of their claspers, and 
with such strength that immersion in alcohol did not cause them 
to separate, even after death. Consequently, although partheno- 
genesis may have been a mode of reproduction, it was by no 
means the only one in August. 

The Artemias swim by synchronous movements of the eleven 
natatory appendages, which are moved at the rate of 160 strokes 
per minute. 

In colour the males incline to a pale greenish, and the females 
to a more reddish hue. The alimentary canal is usually dark 
brownish green, owing to the food contained in it. 

It is a debatable point whether new specific names should 
be applied to newly discovered members of a group of animals 
of which the other species have been mainly diagnosed by 


3898 CRUSTACEA OF LAKE URMI. 


characters of doubtful taxonomic value. Artemia salina, A. 
miilhausenii, A. fertilis, A. gracilis, A. monica, and A. utahensis 
have all been distinguished by characters which vary to a greater 
or less extent with the salinity of the water in which they live. 
IT am inclined to agree with Packard that only two well-defined 
species of Artemia have been described, viz., the Old-W orld form 
A, salina, with which A. mulhausenii (or Artemia sp. gr. 1°2015) 
has been proved by Schmankewitsch to be identical, and the New- 
World A. gracilis, including Verrill’s other American species as 
synonyms. At the same time it is possible that, when a complete 
revision of the group is made and more minute details are taken 
into account, other and better specific differences may be found 
to exist. In the meantime, in order to attract notice to its 
peculiar features, | venture to propose Artemia urmiana as a new 
Old-World species, with the following diagnosis :— 


A. URMIANA, Sp. 0. 

Resembling A. salina, but with an meompletely-segmented 
abdomen ; furcal lobes bearing a single seta apiece in the male 
and none in the female ; claspers of male of larger size than in 
the male A. salina; labrum with two sensory setose papille, 
The margin of the fifth endites of the thoracic feet bordered 
with short sete bearing barbs of peculiar nature. 

Hab. Lake Urmi, in water of specific gravity 1:1188. 


EXPLANATION OF PLATE 25. 


Fig. 1. Artemia urmiana, sp.n. Ventral view of male. x 15, 

2. a. Ventral view of head of female. x 15. 6. Ventral view of ovisac of 
female. 

3. Lateral view, showing the position of the male when in the act ot 
clasping the female. Twice natural size. 

4. a. Pestabdomen of g Artemia urmiana, b. Postabdomen of 2 Artemia 
armiana. ¢. Postabdomen of 9 Artemia salina (Coll. Norman). - 

5. a. The ventral margin of the seventh thoracic natatury appendage. x 40. 
The flabellum and bract are indicated by dotted lines. 0. Barbed 
setze from margin of the fifth endite. x 400. c. Sensory seta. 


Lins Soc. Journ. Zoou. Vor, XXVII Pr 25. 


Ginther. 


Geo. West & Sons imp. 


M.P.Parker del. et lith. 


ZAIBE IO HD MUA UNE IMULA INU - 


aia 
Oe at 
rae 


‘ 


CHILOPODA OF LAKE URMI. 399 


CHILOPODA anp ARACHNIDA. 
By hk. I. Pocock, of the British Museum (Natural History). 


(Puatx 26.) 


Class CHILOPODA. 
Genus ScutieEra, Latr. 


SCUTIGERA COLEOPTRATA (Linn.). 

Syst. Nat. ed. x. i. p. 637. 

[For synonymy, see Latzel, Die Myriop. Oester.-Ungar. Mon. i. p. 25 
(1880). ] 

ocr Sell. 

Ranges from Madeira and Spain throughout South Europe. 


Genus Liruostius, Leach. 


LirHOBIUS PERSICUS, sp. 0. 

Colour a uniform reddish brown as in L. forficatus. Head 
smooth, furnished on each side with about 15 ocelli arranged in 
four rows. Antenne long, composed of about 42 segments, the 
exact number being doubtful on account of the indistinetness of 
the divisional line between some of the segments. Tergal plates 
smooth, the posterior sparsely punctured and hairy ; the posterior 
lateral angles of the 11th not produced beyond the level of the 
posterior border; those of the 18th produced and spiniform, as 
for example in L. forficatus; the posterior angle of the remaining 
terga not produced. Cowal teeth of external maxillipedes 2-2. 
Coxal pores of posterior four pairs of legs 3, 5, 4, 5. 

Anal leg with single claw; armed below with 0, 1, 3, 3, 1 
spines, the coxa with a single external lateral spine. Tibia of 
anal ieg and of preanal leg longitudinally sulcate above in the 
male; these appendages otherwise not modified. 

Total length 19 mm. 

Loc. Seir. A single male example. 

Very closely allied to the common European species Z. muta- 
bilis, L. Koch, but apparently distinct on account of the presence 
of the lateral spine on the anal coxa and of the deutitorm pro- 
longation of the 18th tergite. 


400 MR. R. I. POCOCK ON THE 


Genus ScoLOPENDRA, Linn. 


ScoOLOPENDRA CANIDENS, WVewp. 

Scolopendra canidens, Newp. Ann. Mag. Nat. Hist. xii. p. 98 (1844) ; 
id. Tr. Iiinn. Soc. xix. p.399 (1845) ; id. Cat. Myr. Brit. Mus. pt. 1. p. 48 
(1856). 

Scolopendra affinis, Newp. Ann. Mag. Nat. Hist. xii. p. 98 (1844) ; 
id. Tr. Linn. Soc. xix. p. 386 (1845) ; id. Cat. Myr. Brit. Mus. pt. 1. p. 33 
(1856). 

Scolopendra oraniensis, Lucas, Rev. Zool. 1846, p. 287 ; id. Expl. Sci. de 
VAigérie, Anim. Art. p. 344. 

Scolopendra dalmatica, C. Koch (1847) ; and recent authors. 

Loc. Seir. A single specimen. 

Although abundant in North Africa and South Europe, I am 
not aware that this species has ere this been recorded farther to 
the east than Egypt. 


Class ARACHNIDA. 


Order ARANER. 
Genus Aratorg, Sav. 


ARGIOPE BRUENNICHI (Scop.). 

Aranea Bruennichi, Scopoli, Obs. Zool. in Ann. V. Hist. Nat. p. 125 
1772). | 
[For synonymy, see Thorell, Remarks on Synonyms, &e. p. 518.] 

Loc. Seir. 

Widely distributed throughout Central and South Europe, and 
extending as far north as Paris, Hungary, &c. 


Genus Terraanatua, Latr. 
TETRAGNATHA EXTENSA (Linn.). 
Aranea extensa, Linn. Syst. Nat. ed. x. i. p. 621 (1758). 
[For synonymy, see Thorell, doc. cit. p. 459. ] 
Loe. Seir. 
A single male example, identical with British examples of the 
species of Tetragnatha referred to T. extensa (Linn.). 


Genus Laturopectus, Walck. 


LATHRODECTUS TREDECIM-GUTTATUS (/toss7). 

Aranea 13-guttata, Rossi, Fauna Etr. ii. p. 186 (1790). 

[For synonymy, see Thorell, loc. cit. p. 508. } 

Loc. Seir. 

A single specimen of the black variety of the species (var. 
erebus, Sav.) was found. 


ARACHNIDA OF LAKE URMI. 401 


Genus Lycosa, Latr. 

Lycosa GUENTHERI, sp.n. (PI. 26. figs. 1, La, 16.) 

2. Colour. Carapace with two broad brown bands extending 
from the eyes to the posterior margin on each side of the pedicel, 
separated by a flavous median band about equal to them in width ; 
a flavous band of about the same width running along the lateral 
border; abdomen yellowish white, with two fuscous bands ex- 
tending along each side of the upper surface from the anterior to 
the posterior extremity, and a median dark band between them, 
distinct on the anterior half of the upper surface but breaking up 
and becoming obsolete posteriorly; sides and lower surface of 
abdomen yellowish white, with narrow indistinct abbreviated lines 
behind the epigastric fold; legs yellowish, infuscate distally, 
sometimes mottled with fuscous spots above ; underside of femora 
yellow, of tibiz yellow at base, becoming fuscous distally ; tibia 
of 4th leg with two black bands, one apical and one basal, the 
basal sometimes obsolete ; tibia of 8rd lex sometimes with apical 
band; cox and sternum flavous; mandible black in the apical 
half, yellow above; palpi flavous, with tarsus black. 

Carapace about as long as patella and tibia of 1st leg and as pro- 
tarsus of 4th, barely as long as patella and tibia of the 4th; width 
of carapace about equal to tibia of 4th and exceeding that of 
Ist ; posterior median eyes not much more than half a diameter 
apart; eyes of anterior line narrower than those of the median 
line by about half the radius of one of the latter on each side, 
only slightly procurved, the upper edge of the laterals above the 
centres of the medians, the medians only slightly the larger; the 
laterals almost their own diameter below the posterior medians. 

Mandibles clothed above and externally with yellow hairs, black 
internally and at the apex ; posterior border of sulcus armed with 
3 subequal teeth. 

Legs 4, 1, 2, 8 in length, longish and slender; tibie of 3rd and 
4th with two spines above; patelle of 1st unarmed, of 2nd armed 
with a minute anterior spine, of 3rd and 4th spined in front and 
behind ; tarsal scopula of 3rd and 4th divided by a narrow but 
sharply defined band of sete. 

Vulva as in fig. 16, Pl. 26. 

3. Carapace barely as long as patella and tibia of 2nd leg, dis- 
tinctly shorter than those of 1st and of 4th and than protarsus 
of latter; patelle of all the legs armed with an anterior and a 
posterior spine. Palpus as in fig. 1a, Pl. 26. 

Measurements in millimetres:—@. Total length 18; length 


402 MR. R. I. POCOCK ON THE 


of carapace 7°8, width 5°5; length of 1st lee 21, of 2nd 19, 
of 3rd 18, of 4th 26 (all measured from base of femur); patella 
and tibia of lst 7°5, of 4th 8°2; protarsus of 4th 8. 

3. Total length 15; length of carapace 7°6, of Ist leg 24, of 
2nd leg 22, of 3rd leg 21, of 4th leg 29; patella and tibia of 1st 
leg 8°5, of 4th 8°85 protarsus of 4th 9-2. 

Loe. Seir. 

In size and colouring this species approaches L. ferox of Lucas, 
but may be at once recognized by its higher head, shghtly pro- 
curved anterior line of eyes, of which the medians and laterals are 
subequal. A considerable number of Transcaspian species of this 
genus have been established, and the species that has been here 
described as new may perhaps belong to one of them; but until 
reliable figures of the generative organs of both males and females 
have been published, or until the specific characters have been 
set forth in tabular form, the satisfactory identification of the 
species will remain an almost hopeless task. 


Order SoLIFUGS. 


Genus GaLEopEs, Oliv. 

GALEODES TRUCULENTUS, sp. n. (Pl. 26. figs. 2, 2 a.) 

@. Colour. Upper surface of head strongly infuscate, with 
median lanceolate pale stripe extending back from the black ocular 
tubercle; mandibles yellow, with a pair of faint fuscous stripes 
above ; femur of palp yellow, lightly infuscate above at the tip ; 
upperside of tibia infuscate, except for the two extremities which 
are pale; protarsus infuscate almost to its extremity; tarsus 
lightly infuseate above; distal half of femur and proximal two- 
thirds of protarsus of 4th leg, and in a lesser degree of the 2nd 
and 3rd legs, infuscate. 

Width of cephalic plate almost equal to length of tibia of 
palp and exceeding the protarsus and half the tarsus of that 
appendage, equal to the protarsus of the 4th leg and exceeding 
the protarsus and half the tarsus of that appendage. 

Mandibles with inferior fang armed with 5 teeth ; 3 small teeth 
between the two terminal large teeth. 

Legs and palpi short; palpus about three and a half times as long 
as the width of the head; 4th leg about four and a half times the 
length ; spine-armature of legs as in normal G. arabs (see Ann. 
Mag. Nat. Hist. (6) xvi. p. 77). | 

3. Coloured as in 9; inferior mandibular fang armed with 


ARACIINIDA OF LAKE URMI. 403 


5 teeth, asin that sex. lagellwm resembling that of G. citrinus 
and differing from that of the Egyptian form G. arabs (or Lucasii) 
in having the basal partion stouter and the distal sensory portion 
more expanded. Ocular tubercle of normal size, and about one- 
fifth the width of the head-plate. 

Measurements in millimetres:— 2 . Total length (not including 
mandible) 86; width of head 12°5; length of mandible 17, of 
palpus 48, its tibia 14, protarsus 11, lst leg 33, 2nd leg 30, 3rd 
leg 37, 4th leg 56, its tibia 13, protarsus 9. 

3» Total length 32; width of head 8, of ocular tubercle 1'5 ; 
length of palp 49, of 4th leg 60. 

Loe. & (type), Island of Koyun Daghion Lake Urmi; ¢, Su- 
perghan. 


The females of the three species of Galeodes from South Persia 
nay be recognized as toilows :— 


a. Width of cephalic plate almost equal to length of 
tibia of 4th leg, exceeding protarsus of 4th by 
more than half the tarsus and only a little 
shorter than protarsus and tarsus of palp ; 
palpus about three and a half, 4th leg about 
four and a half times the width of the cephatic 
plate; cephalic plate and legs more strongly 
TIOURDIS( NRO Veh Walp le cite 1s Osta Hain ain Cet iy ee truculentus, sp. 0. 
b. Width of cephalic plate much less than tibia of 
4th and not exceeding protarsus of palp and 
4th leg; legs and palpi much longer; cephalic 
plate and legs scarcely infuscate. 
a’, Legs and palpi shorter; palpus a little more 
than four and a half, 4th leg about six times 
as long as width of cephalic plate.......... darius, Poc.* 
6’. Legs and palpi longer; palpus a little more 
than five times, 4th leg a little less than seven 
times, as long as the width of cephalic plate.. evtrinus, Poe.t 


The males of the three South Persian species may be recog- 
nized as follows :— 


a. Ocular tubercle very large, its width about one- 
third that of the head-plate; palpus longer, 


* Ann. Mag. Nat. Hist. (6) xvi. p.81 (1895). A single female of this species 
from Fao on the Persian Gulf was sent to the British Museum by Mr. W. D. 
Cuming. 

t Loc. cit. The British Museum has received many examples of this species, 
collected at Jask, on the Gulf of Oman, from Messrs. Butcher, B. T. Ffinch, 
aud FE, W. Townsend. 


404 MR. R. I. POCOCK ON THE 


seven times as long as width of head; tibia, 
protarsus, and tarsus completely black ; distal 
tarsal segment of 2nd and 38rd legs with 
(CO CUI OMNES 4 oaagoroucscoccoceccuacc cyrus, Poc.* 
6. Ocular tubercle much smaller, its width about 
one-fifth that of the head-plate; palpus shorter, 
only about six times as long as the width of 
the head; the tibia pale at the apices; tarsus 
scarcely infuscate ; distal tarsal seement of 2nd 
and 3rd legs with one anterior spine. 
a’. Cephalic plate, legs, and femur of palpus 
scarcely noticeably infuscate ; lower fang of 
mandible with only 1 small supernumerary 
tooth between the two larger teeth ........ citrinus, Poe. 
6’. Cephalic plate, posterior legs, and femur of 
palpus distinctly infuscate; lower fang of 
mandible with 3 supernumerary teeth...... truculentus, sp. 1. 


Order SCORPIONES. 
Genus Butuus, Leach. 


Burnus caucasicus (fscher). 
Scorpio caucasicus, Fischer, Zoogn. p. 401, pl. iv. fig. 1 (1813) (= 
eupeus and thersites, C. Koch), é 


Subsp. PERSICUS, Nov. 

Colour. Tergites yellow with five black stripes, three marking 
the keels and one on each side between the lateral keels and the 
border: carapace correspondingly marked in its posterior half; 
in its anterior half the tubercle, the frontal keels, and the anterior 
border are black, and there is a black patch on each side between 
the ocular tubercle and the lateral margin; median and lateral 
inferior caudal keels black ; palpi yellow, with traces of black 
lines on the humerus, brachium, and hand; femur and patella 
(tibia) of legs also partially infuscate. 

Structurally this Scorpion much resembles B. afghanus, Poe. 
(Tr. Linn. Soc. (2) iii. p. 116, 1889), from Meshed in Afghanistan ; 
but the tail is considerably more powerful in B. persicus, the 
segments being relatively both higher and broader. For example, 
the height of the 3rd segment in the ¢ is about equal to the 
length of the inferior keel, and that of the 9 a little less, 
whereas in afghanus (3 2) the height is noticeably less. Again, 


* Ann, Mag, Nat. Hist. (6) xvi. p. 79 (1895). Based upon a single male 
example from Fao (W. D. Cuming). 


ARACHNIDA OF LAKE URMTI. 4D5 


although the tail is more powerful, the crests are very per- 
ceptibly less strongly granular. This is particularly noticeable 
on the inferior median crest of the 2nd and 3rd segments, which 
in B. afghanus are strongly elevated posteriorly, but are scarcely 
noticeably so in B. persicus. Similarly, the inferior lateral keels 
of the 5th segment are much less strongly denticulate, and the 
posterior lateral prominence is trilobate, not bilobate. 

Peetinal teeth 25-26, g; 20-21, 2. 

In the palpi the hands are large, smooth and rounded, not 
crested in either sex, larger in ¢ than 2. Fingers more strongly 
lobate in 3; in both sexes the movable digit is much longer than 
the hand-back, and slightly exceeds the length of the brachium; 
12 rows of teeth, as in afyhanus. 

Measurements in millimetres :—¢ (type). Total length 49; 
length of carapace 5°5, of tail 31; height of 3rd segment 39; 
length of inferior keel 3°8, width 4; width of hand 3°4; length 
of hand-back 4, of movable digit 6. 

2. Total length 61, carapace 6:2, tail 34. 

Loe. Seir; landing on east side of lake. 

According to Dr. A. Birula, who has examined many Trans- 
caspian Scorpions allied to afghanus and persicus, B. afghanus, 
Poe., is synonymous with B. thersites, C. Koch *, the latter 
being but a subspecies of B. eupeus of C. Koch. But since 
the locality of the original thersites is unknown, and the descrip- 
tion of the specimen or specimens to which Dr. Birula gave that 
name does not apply to the typical examples of B. afyhanus, at 
all events in the form of the 3rd segment of the tail, I prefer to 
cite the species or subspecies by the name under which I originally 
described it. 


Norr.—Subjoined is the description of a new species of the genus 
Buthus from Persia :— 

BUTHUS VESICULATUS, sp. n. (PI. 26. fig. 4.) 

Colour of trunk, chele, and tail entirely pale yellow. In structural 
characters, ¢. e., form of palpi, of cephalothoracic and abdominal keels, 
closely allied to B. parthorum, Poc., from near Meshed (Tr. Linn. 
Soe. (2) ii. p. 115, 1889), but at once recognizable by the form of the 
vesicle and the shortness of the aculeus of the tail. In B. parthorum the 
vesicle is small and piriform (Pl. 26. fig. 8), its width exceeding its 


* Die Arachn. vi. p. 51, fig. 466 (1859). 
t Die Arachn, y. p. 127, fig. 419 (1838), 


4.06 ARACHNIDA OF LAKE URMI. 


height, its height being much less than the width of the upper surface 
of the 5th caudal segment and only a little more than half the length 
of a straight line drawn from the outer side of the base of the aculeus 
to its point; the aculeus is very long and lightly curved, its length 
represented by a straight line, drawn as described above, considerably 
exceeding the width of the 5th caudal segment, exceeding also the length 
of the vesicle, equal to half the length of the movable digit and to the 
distance between the anterior edge of the median eye and the posterior 
border of the carapace. In B. vestculatus, on the contrary, the vesicle 
(Pl. 26. fig. 4) is large and globular, the height exceeding the width ; 
the aculeus is short and more strongly curved, its length represented by 
a straight line drawn as above being much less than, not much more than 
half, the leneth of the vesicle, less than the width of the 5th caudal 
segment, about one-third the length of the movable digit, and much less 
than half the length of the carapace. 

Pectinal teeth 20 in 9, 25 in S. 

Q. Total length 55 mm.; length of carapace 65, of tail 33; height of 
vesicle 3, width 2:8; length of aculeus 25. [In type(@ ) of B. parthorum 
the measurements, in mm., are:—Total length 74, carapace 8, tail 44 ; 
height of vesicle 2°5, width 3; length of aculeus 4:6. | 

Loc. Astracan, in Persia. 

There are three examples of this species in the British Museum: two, 
@ ad. and ¢ immat., from the above locality, and 1 9 ad. in bottle 
without locality but containing other typically Eastern Mediterranean 
specimens of Arachnida and Chilopoda. 

Prof. K. Kraepelin (Das Tierr., Scorpiones, p. 24, 1899) cites B. par- 
thorum as doubtfully synonymous with B. eaucasicus, Nordman (Demidoff, 
Voy. Russie, iii. p. 731, Avachn. pl. i. fig. 1, 1840). Judging by Kraepelin’s 
description of the latter species, the two are certainly allied; but since 
nothing is said about the form of the caudal vesicle and aculeus, there is 
at present no certainty on this point. In any case the name caucasicus 
cannot stand for the species, since it was previously used by Fischer, as 
Birula has shown, for another species of Buthus. Hence, if cancasicus, 
Nord.=parthorum, Pocock, the latter will be the name for the species. 


EXPLANATION OF PLATE 26. 


Fig. 1. Lycosa Guentheri, sp.n. Dorsal view, 3. 


I Gig ms wt Palp of 3. 

1d. a 4 Vulva of 9. 

2. Galeodes truculentus, sp.n. @, nat. size. 
2a. 45 ee Flagellum of ¢. 


3. Buthus parthorum, Poe. Vesicle and aculeus of tail. 


4. * vesiculatus, sp. n. ah Bo 


Linn. Soc. Journ. Zoon. Vou. XXVII. PI. 26. 


Pocock. 


West, Newman imp. 


F.0 Pickard-Cambridge del. et lth. 


URMI. 


KE 


ARACHNIDA FROM LA 


ACARI OF LAKE URMI. 407 


ACARI. 


By Auzert D. Micuaet, F.L.S., F.R.M.S. 
(Puats 27.) 


Amone the specimens collectéd by Mr. R. T. Ginther the 
following three species of Acari have been identified :— 
ASTOMA GRYLLARIA, le Baron. 


This species, common in the Urmi district, was found attached 
near the bases of the wings of Caloptenus italicus, L., in July. 
The geuus As/oma cannot be considered a good one, being founded 
entirely upon a larval type. The adult form would certainly be 
one of the Trombidide. Trombidium sericeum locustarum was 
described as the adult form by Riley, but as the Astoma gryllaria 
larva is totally different from the larva of 7’. sericewm, this iden- 


tification seems doubtful. Riley considered the larva very 


destructive to locusts and consequently beneficial to man, and 
brought it into notice in a series of articles (“‘ The Locust in 
1876,” New York Tribune, Aug. 16, 1876; ‘‘ Rocky-Mountain 
Locust,” Appleton’s Amer. Cyclop. 1875, pp. 371-874; and 
“Mite Transformations,’ Trans. Acad. Sci. St. Louis, vol. iii. 
Proc. p. 267). 

RuHIPICEPHALUS stuvs, C. LZ. Hoch; or saNnGuiINngEus, Latr. 

These are practically world-wide species» dogs and other 
animals carry them everywhere, and they will attach themselves 
to almost any creature which has blood to suck. 

The Persian specimens were all found upon the hind legs and 
tails of the Testudo ibera, in situations in which they are free 
from the risk of being rubbed off. All the older tortoises at Seir 
carried three or four of these bloodsuckers. 

ARGAS PERSICUS, Fischer. 

Hab. Seir Hill. 


This species I regard as synonymous with Argas reflexus of 
Fabricius. I have obtained specimens which are indistinguishable 
from A. reflexus from all quarters of the globe ; they are probably 
transported by pigeons and other birds. In a hot country the 
bite of this tick 1s believed occasionally to produce fever, 


madness, and even death (cf. p. 866). Mr. Giiuther found one 
specimen upon a tortoise. 


A bright orange-coloured species of Acarid was in two 
instances found attached to the nape of the neck of a species of 
Machilis (Pl. 27. fig. 4). 


408 MR. A. G. BUTLER ON THE 


INSECTA (Uerripoprera RHOPALOCERA). 
By A. G. Burrer, Ph.D., F.L.S. 


[For the specimens marked ‘“ Daltry collection” I am much 
indebted to the energy of the Rev. 8. J. Daltry.—R. T. G.] 


NyMPHADLID24. 


SaTYRINE. 
1. HrppaRCHIA BRISEIS, var. TURANICA. 
Satyrus turanica, Stgr.; (cf. Rihl, Paleearkt. Gross-schm. 1. p. 532 (1892). 
Seir, 8 miles west of Urmi, Aug. 13-19 (1898). 
2. HIPPARCHIA PELOPEA. 
Satyrus pelopea, Klug, Symb. Phys. 3, pl. xxix. figs. 5-8 (1882). 
Seir, 8 miles west of Urmi, Aug. 13-16 (1898). 


3, HIPPARCHIA CIRCE. 
Papilio circe, Fabricius, Syst. Ent. p. 495 (1775). 
3, Urmi (Daltry coll.). 


4, KPINEPHELE LYCAON, var. LUPINUS. 

Satyrus lupinus, Costa, Faun. Nap. (1835?); Staudinger, Hore Soe. 
Ent. Ross. 1870, p. 79. 

2, Seir, 8 miles west of Urmi, Aug. 13-16 (1898). 


5. EPINEPHELE,HISPULLA. 
Papilio hispulla, Esper, Kur. Schmett. 1. 2, pl. exix. figs. 1, 2 (1809 ?). 
@ 2, Urmi (Daltry coll.). 


6. EREBRIA AFER. 
Fapilio afer, Esper, Eur. Schmett. pl. lxxxiii. figs. 4, 5 (1783), 
Uimi (Daltry coll.). 


7. C&NONYMPHA PAMPHILUS. 
Papilio pamphilus, Linn. Syst. Nat. x. p. 472 (1758), 
Seir, Aug. 19, and Urmi (Daltry coll.). 


8. PYraMEIS CARDUL. 
Fapilio cardui, Linn, Faun. Suec. p. 276 (1761). 
6, Urmi (Daltry coll.). 


9. ARGYNNIS MATA. 
Papilio mata, Cramer, Pap. Exot, i. pl. xxv. B, C (1775). 
3, Urmi (Daltry coll.). 


10. MELIT®A AURINIA. 
Papilio aurinia, Rott. Naturf, vi, p. 5 (1775). 
@, Urmi (Daltry coll.). 


BUTTERFLIES OF LAKE URME. 409 


LYcHZNIDA. 
11. Curmpo DAMON. 
Papilio damon, Schiffermiiller, Wien. Verz. p. 182 (1776). 
3, Seir, Aug. 16, 1898. 


12. Curipo ADMETUS, var. RIPARTII. 

© Papilio riparti, Freyer, Beitr. Schmett. iii, pl. 133. fig. 3 (1830). 

3 3, 2 2, Seir, Aug. 16, 1898. 

The males have the veins of the primaries broadly bordered 
with woolly androconia from the base to beyond the middle. In 
the specimens from Seir these have been to some extent abraded, 


but are still easily discernible. 


13. CUPIDO ICARUS. 
Papilio icarus, Rott. Naturf. vi. p. 21 (1775). 
66, 2 9, Sar, Aug. 16, 1898. 


14. CUPIDO AGESTIS. 
Papilio agestis, Schiffermiiller, Wien. Verz. p. 184 (1776). 
3 3, QQ, Seir, Aug. 16, 1898. 


15. CuprIpo BELLARGUS, var. OCEANUS. 

© Papilio oceanus, Bergstrasser, Nomencl. iii. pl. 53, figs. 3, 4 (1779), 

36, Urmi (Daltry coll.). 

The female has the submarginal red spots well defined above 
in this variety, but the male only differs from our C. adonis 


below. 


16. CurpIpO ENDYMION. 

Papilio endymion, Schiftermiiller, Wien. Verz. p. 182 (1776). 
Papilio daphnis, id. ibid. 

3 3, 2 9, Seir, Aug. 16 & 19, 1898. 


17. CUPIDO DAMA, var. P 

Lycena dama, Staudinger; cf. Riihl, Palzearkt. Gross-schm. p. 287 
(1892). 

3 Sd, 2 @, Seir, Aug. 16 & 19, 1898. 

This species is new to the Museum series. Mr. Elwes kindly 
informed me that it was “probably OC. dama”’; the description 
in Rwhl’s book does not, however, represent a form like that 
before me; indeed, the specimens from Seir seem rather to 
resemble the description of C. aedon of Christoph, but both fore 
and hind wings have the veins entirely blackish, not merely with 
blackish tips. 

LINN. JOURN.— ZOOLOGY, VOL. XXYII. ol 


410 BUTTERFLIES OF LAKE URMI. 


18. CHRYSOPHANUS THETIS. 
Lycena thetis, Klug, Symb. Phys. pl. 40. figs. 17, 18 (1834). 
3 9, Seir, Aug. 16. 


19. CHRYSOPHANUS THERSAMON, var. OMPHALE. 

Tycena omphale, Klug, Symb. Phys. pl. 40. figs. 12-14 (1834). 

Q, Seir, Aug. 16. 

Tt seems hardly likely that this tailed form can be a mere 
variety of C. thersamon, but Dr. Staudinger regards it in that 
light. 


PAPILIONIDS. 


PIERIN#. 
20. Cotas EDUSA. 


Papilio edusa, Fabricius, Mant. Ins. ii. p. 28 (1787). 

3 3d, Urmi (Daltry coll.); 3 2, Seir, Aug. 13-19, 1898. 

Although the names hyale, electra, and croceus all take priority 
over edusa, the later name is so widely recognized that the use of 
any of them without the general consent of Lepidopterists is 
likely to lead to misapprehension*. If the name hyale be 
rejected on account of its long application to a different species 
in the genus, I do not see how electra applied to a common 
African sport of the species can be ignored with any fairness or 
regard to the law of priority. That C. electra is not a species 
distinct from the European insect is certain, inasmuch as typical 
forms of both insects with numerous intergrades reach us from 
the same localities. 


21. SYNcHLO# DAPLIDICE. 
Papilio daplidice, Linneeus, Syst. Nat. 1, ii. p. 760 (1767). 
3, Seir, Avg. 13-16, 1898. 


22. GANORIS RAPA, var. ERGANE. 
Papiho ergane, Hiibner, Eur. Schmett. i. figs. 904-7 (1827). 
Seir, Aug. 13-16, 1898. 


PAPILIONINE[. 
23. THars CERISYI. 
Phais cerisyt, Godart, Mém. Soc. Linn. Paris, ii. pl. 2 (1822). 
Urmi (Dalry coil.). 


* Another objection to the use of the name edusa for a Colias is that, in his 
‘Genera Insectorum,’ Fabricius used it for Synchloe daplidice. 


MOTHS OF LAKE URMI. 411 


HESPERIIDG. 

24. CHARCHARODUS ALTHER. 

Papilio althee, Hiibuer, Eur. Schmett. i. figs. 452, 453 (1798-1803). 

Seir, Aug. 13-16, 1898. 

25. ADOP@A LINEOLA. 

Papilio lineola, Ochsenheimer, Schmett. Eur. i. p. 230 (1808). 

3, Urmi (Daltry coll.). 

26. AUGIADES SYLVANUS P 

Papilio sylvanus, Esper, Eur. Schmett. i. 1, pl. 36. fig. 1 (1778 ?). 

2, Urmi (Daltry coll.). 

The single example is larger than any specimen of this species 
which I have seen. 


INSECTA (Leprrporrera PHALHN®). 
By Sir G. F. Hampson, Bart. 


Tur following species have been identified in the collections 
made by Mr. R. T. Gunther. 


SYNTOMID. 

Syntomis persica, Koll. Denkschr. Akad. Wiss. Wien, Math.- 
nat. Classe, 1. p. 53 (1850) ; Hmpsn. Cat. Lep. Phal. B. M. i. 
[Oe IOI, jolle tye te, Oe 

Urmi (Daltry),2$. Only known previously by the type o 
in bad condition in the Vienna Museum; the patagia are 


yellowish white. 
ARCTIADS®. 


Drrorera PULCHELLA, Linn. Syst. Nat. 1, il. p. 884. 
Urmi (Daléry), 19. 

Nocturp. 
AGROTIS YPSILON, Rott. Naturf. xi. p. 141. 
Urmi (Daltry), 12. 
Agroris Curistopnt, Sigr. Berl. e. Zeit. 1870, p. 110. 
Seir near Urmi (Gunther), 19; Urmi (Daliry),1¢. 
HapDENA BImacuLosa, Linn. Syst. Nat. xii. p. 856. 
Urmi (Daitry), 32. 


ACRONYCTA CENTRALIS, Stgr. 


Urmi (Daltry), 23,1°. 
31* 


412 SIR G. F. HAMPSON ON THE 


Unocuimna uirta, Hiibn. Hur. Schmett., Noct. f. 591. 
Urmi (Daltry), 1d. 


EpiseMA GLAuCINA, Hsp. Hur. Schmett. 81. 4, 5. 

Urmi (Daltry),1<é. 

XANTHIA OCELLARIS, Bork. Nat. Hur. Schmett. iv. 647. 
Urmi (Daltry),1¢. 

BryopHina PERLA, Schiff. Wien. Verz. p. 70. 

Urmi (Gunther), 13. 

TARACHE LucTUOSA, Schiff. Wien. Verz. p. 90. 

Urmi (Daltry),1¢. 


TARACHE SULPHURALIS, Linn. Syst. Nat. 1. 2. p. 881. 
Urmi (Daltry), 23. 


XANTHOPTERA TRIANGULARIS, Warr. P. Z. S. 1888, p. 309. 
Urmi (Ginther), 13. 

CarocaLa NEONYMPHA, Esp. Hur. Schmett. 198. 1, 2. 
Urmi (Daltry), 13. 

Carocana ELocata, Hsp. Hur. Schmett. 99. 1, 2. 

Jrmi (Daltry), 23; Seir (Gunther), 23. 


U 
Evcripra mt, Olerck, Icones, pl. 9. £. 5. 
Urmi (Daltry), 1¢. 


LYMANTRIADA. 
Lymantria pispar, Linn. Syst. Nat. x. 501. 
Urmi (Daltry), 16. 


SATURNIAD®. 


SATURNIA PYRI, Schiff. Wien. Verz. p. 49. 
Urmi (Daltry), 13. 


SPHINGID®. 


SMERINTHUS POPULI, Linn. Syst. Nat. x. p. 489. 
Urmi (Daliry),1¢. 


MAcroGLossa STELLATARUM, Linn. Syst. Nat. x. p. 493. 
Urmi (Daltry),1¢. 


NovroponrTip#. 


Dicranvra vinuta, Linn. Syst. Nat. x. p. 499. 
Urmi (Daltry),1¢. 


MOTHS OF LAKE URMI. 413 


GEOMETRID#. 


Macarta Murtnaria, Schiff. Wien. Verz. p. 105. 
Urmi (Giinther), 12. 


Awnarmis praciata, Linn. Syst. Nat. x. p. 526. 
Urmi (Daltry), 13. 


ANAITIS USGENTARIA, Ségr. 
Urmi (Daltry), 19. 


CATACLYSME BILINEATA, Linn. Syst. Nat. x. p. 525. 
Urmi (Daltry), 13,19. 


EupitHecra supumBrata, Schiff. Wien. Verz. p. 110. 
Urmi (Gunther), 23,19. 


RHODOSTROPHIA CALABRARIA, Zell. Stett. e. Zeit. 1852, p. 180. 
Urmi (Daliry), 23. 


ReopostRoPHIA INconsPicua, Butl. P. Z. 8. 1886, p. 391. 
Urmi (Daltry), 13. 


CRASPEDIA MARGINEPUNCTATA, Goeze, Beytr. ii. 3, p. 385. 
Urmi (Daltry), 1¢3. 


LASIOCAMPID®. 


METANASTRIA TRIFOLLL, Schiff. Wien. Verz. p. 53. 
Ab. terreni, H.-S. Eur. Schmett. ff. 120, 121. 
Transcaucasia, Kivorak (Géinther), 1d. 


CLISIOCAMPA CASTRENSIS, Linn. Syst. Nat. x. p. 500. 
Urmi (Dailtry), 22 of a very pale form. 


PsyYcHID”. 


APTERONA CRENULELLA, Bruand. Mon. Psych. p. 76, f. 49. 
Urmi (Giinther), 3 larva-cases on a piece of bark. 


ZAYGENIDEA. 


YyvexNA Cuviert, Boisd. Mon. Zyq. iii. 6, p. 53. 
Urmi (Daltry),1¢. 


PYRALIDE. 
HETEROGRAPHIS PYRETHRELLA, Herr.-Schiff. Neue Schmett. 80, 
5 
Seir, near Urmi (Gunther), 13. 


414 MR. BR. T. GUNTHER ON THE 


Actossa PIneuinatts, Linn. Syst. Nat. x. p. 533. 
Urmi (Giinther), 1¢é. 

Pyratis FaRINALIS, Linn. Syst. Nat. x. p. 226. 
Urmi (Daltry), 1d. 

Hyrsopyera costatis, Labr. Syst. Ent. p. 132. 
Seir, near Urmi (Gunther), 2d. 


ScopaRta CEMBR&, Haworth, Lep. Br. p. 498. 

Seir, near Urmi (Gunther), 23. 

NomopHILA NOCTUELLA, Schiff. Wien. Verz. p. 186. 
Seir, near Urmi (Gtinther),1¢. 

PyravstTa CESPITALIS, Schiff. Wien. Verz. p. 123. 
Seir, near Urmi (Gunther), 2d. 


Pyravsta AvRATA, Scop. Hnt. Carn. no. 565. 
Seir, near Urmi (Gunther), 13. 


ORNEODID2. 
ORNEODES sp. 
Seir, near Urmi (Gunther), 2 3. 


TINEIDA. 


Three species undetermined. 


NEUROPTERA (Hemerobiide) anp DIPTERA. 
By Rozert T. Ginturr, M.A., F.R.G.S. 


(Puate 27.) 


On Plate 27. fig. 2 is figured the larva of one of the Hemerobiude 
or Lacewing flies, which both in structure and habit is related to 
the larva of Chrysopa perla, and is not unlike the larva of an 
Hemerobius? figured by Sowerby in his ‘ British Miscellany,’ 
pl. 66. The entire animal was hardly larger than a hemp-seed ; 
the body is long and distinctly segmented; the head carries a 
pair of powerful mandibles, which are about one-third of the 
length of the body. The labial palps are at least 3-jointed, and 
the antenne are longer than the mandibles and are unjointed. 
Immediately behind the conspicuous eyes are two knobs, each bear- 
ing a tuft of some two dozen white hairs; to these succeed four 


Gunther . Lima. Soc. Journ. Zoot, Vor. XXVIl PL. 27, 


J.Green delwet lth Mintern Bros.Chromo 
INSECTA AND ACARID FROM LAKE URMI. 


DIPTERA OF LAKE URMI. 415 


other pairs of similarly tufted basal knobs borne by the succeed- 
ing tergal portions of the segments. The posterior tufts carry 
a large globular mass of white cottony threads,’ like American 
blight, in which are entangled the inedible portions of the car- 
cases of its prey. Upon a specimen captured on rocks near Seir 
the dry cuticle of a small spider was distinguishable. 


DIPTERA. 


EPHYDRA URMIANA, sp.n. (Plate 27. fig. 3.) 

The larve which are found in considerable numbers near the 
margin of the lake are so similar to the larve of Hphydra sali- 
naria which were observed by Klug in “‘Salzsiederei”’ in Silesia, 
ard described by Loew, and to the halophilous larve of Ephydra 
described by Packard, that I have no hesitation in referring them 
to the same genus. At the same time, the larva of the Ephydra 
of Lake Urmi does not agree with either Z. salinaria or with 
E. halophila, and it is to accentuate this fact that [have ventured 
to base a new species upon a larval form, 

The larve of H. urmiana caught on July 21st near Superghan 
were whitish in colour, and about 10 millim. long. The body is 
composed of eleven segments, nearly cylindrical, and pointed 
anteriorly but terminating bluntly behind, unlike the larva of 
E. salinaria (Westwood, Introduction, fig. 132,no.11). The last 
segment is prolonged dorsally into a long bifurcated respiratory 
process, three-sevenths the length of the body, and containing 
two tracheal vessels which open by two spiracles situated at the 
tips of the two branches. Mouth-parts present. All but the 
first three smaller segments are provided with two groups of 
chitinous bristles, mounted on inconspicuous tubercles. Abdo- 
minal appendages, like those of the larva of E. halophila, 
absent. 

The rarity of halophilous insects must enhance the interest 
attaching to the particulars of their life-histories,and consequently 
it is much to be desired that some naturalist should endeavour 
to rear the adult imago of this remarkable fly. The only other 
species of Diptera which have been recorded, so far as I am 
aware, from very salt water are :— 

Ephydra salinaria (Halmopota salinaria) in Silesia, Diirren- 
berg, &c. (Westwood, loc. cit.; Bouché, Naturg.; 
Loew, Zeit. Naturwiss, Halle, 1867). 


416 MR. MALCOLM BURR ON THE 


Ephydra halophila in Mlincis. (Packard, Proc. Essex Inst. 
vi. 1869; Verrill, Proc. Boston Soc. 1866.) 
Chironomus oceanicus in Salem Harbour. (Packard, loc. cit.) 
Chironomus sp. in Utah. (Stansbury, Report on Valley of 
Great Salt Lake of Utah.) 
Halophilus in Illinois and Salem Harbour. (Packard, loc. 
cit.). 
EXPLANATION OF PLATE 27. 
Fig. 1. Group of insects upon the bark of Populus alba (p. 367). 
a. Larva-cases of Apterona crenulella. 6. Bathyscopus pocillus. 
c. Pentatoma baccarum (?)*. d. Yponomeuta padellus, lL. 
e. Moulted skin of larval insect. 
9. Larva of Hemerobius sp. 
3. Larva of Ephydra urmiana. 
4. Acarid upon Machilis persa (?). 
* Or some allied species. The identification is that of Mr. C.O. Waterhouse. 


ORTHOPTERA. 
By Matcotm Burr, F.Z.8., F.E.S. 


Tur small collection of Orthoptera made by Mr. R. T. Ginther 
in North-west Persia contains twelve species, of which one is a 
new variety of a well-known species. The most interesting 
capture, perhaps, is the Decticus assimilis Fieb. 

It is unfortunate that they have been preserved in spirit, 
which has bleached the colours of all, especially of the Cidi- 
podide. 

FoRFICULARIA. 


ForFICULA AURICULARIA, L. 

Seir, N.W. Persia. 2 5,19. 

The two males represent the variety figured by Fischer (Orth. 
Eur. tab. vi. fig. 117). The Marquis Doria captured the same form 
in Northern Persia some years ago, and quite recently a similar 
specimen, undoubtedly British, was exhibited at the South 
London Natural History Society. It differs from the type in the 
narrowness of the dilated part of the male forceps above the tooth. 

In the typical form the part of the forceps from base to this 
tooth is contiguous, the sides of the branches being parallel. In 
this variety the inner margins of the basal part diverge, making 
roughly a triangular area between the pygidium and the teeth of 
the forceps. 


ORTHOPTERA OF LAKE URMI. 4AT 


MANTODEA. 


BonivaRIA BRACHYPTERA, Pall. 
Seir, 2 ¢,6 9; Koyuo Daghi, 1 ¢,1 Q. 
This species is common in Southern Russia, the Caucasus 
and Asia Minor. 
ACRIDIODEA. 


TRUXALIDA. 


TRUXALIS UNGUICULATA, Ramb. 
Seir. 19. A widely distributed species. 


(EprIeoDIpz. 

(Kpreopa Scnocut, Br. 

Seir, Aug. 16,1898. 1 9. 

This specimen, of which the wings have been bleached by the 
spirit, is 35 millim. in length; de Saussure gives 32 millim. as 
the dimensions of the female. Recorded from Aleppo and the 
Caucasus. 


SPHINGONOTUS SATRAPES, Sauss.? 

Seir, Aug. 16, 1898. 16. 

This specimen does not agree perfectly with de Saussure’s 
description, the posterior femora showing no black; but this 
character may have well been destroyed by the spirit, which has 
considerably bleached it. The colour of the wings has entirely 
gone, leaving a whitish base, with the black fascia. ' hesitate 
to regard it as distinct. 


(Epatius niGRorascratus, De Geer. 
seme, lunes, IG, Tei, IL @- 


Pyra@opera cristata, Ff. de W. 
Seir, Aug. 16, 1898. 16. 
A native of temperate Asia and Eastern Europe. 


ACRIDIIDE. 

CaLoPrENnus iTraticvs, L. 

Found at Superghan, July 22nd, infested with Astoma gryl- 
laria (see p. 407). 

PYRGOMORPHIDS. 

PYRG@OMORPHA GRYLLOIDES, Lafr., var. nov. GUENTHERI. 

Deira 40g 

A P. grylloide typico differt, elytris alisque, in feminis saltem 
abbreviatis, segmentum abdominale quintum attingentibus. 


418 MR. G. C. CRICK ON A FOSSIL AMMONITE. 


The shortness of the elytra and wings gives this form an ap- 
pearance different from the type, but the comparative develop- 
ment of organs of flight in Orthoptera is too slender a character 
on which to base a species. 


LOCUSTODEA. 


DEcTICIDz. 
PACHYTRACHELUS Sp. 


Seir, Aug. 16,1898. 19. 
This is very probably a new species, but it is impossible to 
describe it without the male. It is close to P. striolatus, Fieb. 


DEcTICUS ASSIMILIS, Fieb. 

Sema 

Both these specimens are considerably larger than an example 
in my collection from Tiflis, approaching more nearly to D. albi- 
frons, Fabr., in size and appearance. It has hitherto been 
recorded by Brunner and Fieber from “ Tiflis and Syria.” 


GRYLLODEA. 
GRYLLOTALPID #. 


GRYLLOTALPA GRYLLOTALPA, L. 

Seir, Urmi. Several specimens, in all stages of development. 

Mole-crickets were always abundant, but at the same time I 
never met the swarms which have been described from the neigh- 
bourhood of Ispahan and Shiraz, where one intelligent observer 
estimated their numbers at one to every square fathom over an 
area of many square miles.—R. T. G.) 


NOTE ON A JURASSIC AMMONITE. 


By G. C. Crick, F.G.S., of the British Museum 
(Natural History). 


THE only Jurassic fossil in Mr. R. T. Giinther’s collection is an 
Ammonite preserved on the surface of a small block of limestone 
of reddish-brown colour. It consists of a portion of the outer 
whorl that has been much flattened during fossilization, and of 
the impression of the greater part of the rest of the shell. 
Jurassic Ammonites have been recorded from N.W. Persia by 


FOSSIL ECHINOIDEA OF LAKE URMI. 419 


Weithofer* and by Borne tf; and the present specimen is doubt- 
less referable to Perisphinctes curvicosta, Oppel, sp.t, a species 
which Borne has recorded and figured (op. cit. p. 14, pl. i. fig. 1; 
pl. iv. fig. 14) from rocks of Callovian age in the neighbourhood 
of Maragha, whence the present specimen is believed to have 
come. 


FOSSIL ECHINOIDEA. 
By J. W. Gregory, D.Sc., F.G.S. 
(PLats 28.) 


Mr. Ginruer’s collection of LEchinoidea consists of five 
specimens all of which belong to the genus Clypeaster. 
According to Mr. Giinther, they are “said to have come from 
Guverchin Kala, at the extreme northern end of Lake Urmi.” 
This locality is no doubt the same as the Guverchine Kalak 
where the Hon. W. K. Loftus made a collection of fossils now 
in the British Museum. 

Miocene Hchinoidea from Lake Urmi have been described by 
Abich §, who has recorded thence eight species of Clypeaster. 


1. CLYPEASTER aff. IMPERIALIS, Michelin ||, 1861. (Plate 28. 
fig. 1.) 

The most massive Clypeaster in the collection is unfortunately 
so much broken on the margin and base that its positive deter- 
mination is impossible. But it clearly belongs to the group of 
Clypeasters of which C. olisiponensis, Mich.4, may be taken as 
the type. From that species the Urmi echinid differs by having 


* K. Weithofer, “Ueber Jura und Kreide im nordwestlichen Persien”: 
Sitzungsb. d. k.-k. Akad. Wissensch. Wien, Bd. xcviii. Abth. 1, Dec. 1889. 

t G. v. dem Borne, ‘Der Jura am Ostufer des Urmiasees.’ Inaugural 
Dissertation. Halle, 1891. 

¢ A. Oppel, ‘Die Juraformation,’ p. 555 (1857). See also J. v. Siemiradzki, 
“ Monographische Beschreibung der Ammonitengattung Perisphinctes” : 
Palxontographica, Bd. xly. p. 96, 1898. 

§ H. Abich, “‘ Ueber das Steinsalz und seine geologische Stellung im Russ- 
ischen Armenien,” Mém. Ac. Imp. Sci. St. Pétersb. ser. 6, vol. ix. pt. 1, Mém, 
Sci. Math.-Phys. vol. vii. 1859, pp. 111-114; and ‘ Geologische Forschungen in 
den Kaukasischen Landern, Abth. ii, Geologie der Armenischen Hochlandes,’ 
i, 1882, pp. 223-225, 270-283. 

|| H. Michelin, ‘‘ Monographie des Clypéastres fossiles,” Mém. Soc. Géol. 
France, ser. 2, vol. vii. p. 118, pl. xviii. figs. 2, a-d. 

{ Michelin, op. cit. p. 118, pl. xx. figs. 1, a. 


420 DR. J. W. GREGORY ON THE 


the height one half the length and not one-third ; moreover, the 
granules on the ridges between the poriferous furrows are less 
numerous. In these respects the fossil agrees more closely with 
O. imperialis, a Miocene species from Crete. Mlichelin’s figures 
do not show the side-view of the test, but in his description he 
gives dimensions which show the proportions. 

The following table illustrates the relations of the Cretan and 
Armenian specimens of C. imperialis and C. olisiponensis :— 


Oh CASE AU. C. olisiponensis. 
Michelin’s Specimen T 
type. from Urmi. yPe: 
leicht Wee tot eressoesocseorneas 75 mm. 47 mm. 40 mm. 
[GSIVERHOT Sodcasdquosscosooaqaecoso00000 ND. 5, 2 UPA as 
AS Vato tel he Gdonortaduneecostancaconecgadc 145 _,, Sines 105 4, 
Ratio of height to length ...... 1:2:3 1:1:95 1:3:0 
ms Pee wicit lameness 1:19 1:1:85 1:26 
Number of granules on ridge 
between interporiferous 5 3-4 10 
TUUPONYS ooassoneq6c0e0000K00 


These dimensions show that the specimen is far closer to 
CO. imperialis than to C. olisiponensis. As the elevation of the 
test is not illustrated in Michelin’s figures, that aspect is shown 
on Plate 28. fig. 1. The petals in this specimen are unusually 
flat, aud the test is very thick. 


2. CLYPEASTER GUENTHERI, n. sp. (Plate 28. fig. 2.) 

Diagnosis.—Test pentagonal, with well-rounded angles. The 
base is flat, with sharp, slightly sinuousambitus. Upper surface 
flattened. Anterior slope at an angle of 45°-50°, fairly regular ; 
posterior slope steep to a posterior foot-like projection. Length 
of test approximately equal to the width, and 33 times as great 
as the height. Apical dise excentric posteriorly. 

Petals very tumid, broad and completely closed externally ; 
the outer end is broad and well rounded. The petals are long, 
and reach about three-fourths of the distance from apical area 
to the ambitus. The anterior ambulacrum is longer than those 
of the parietal series. 

Periproct large, circular, and close to the margin. 

Granules of interporiferous ridges about four on each ridge. 


FOSSIL ECHINOIDEA OF LAKE URMI. 421 


Dimensions. 
Specimen Specimen 

P : figured by C. turritus, Phil. 

from Urmi. ad 

Abich. 
mm. mm. mm. | mm. | mm. 

TEIGHEING oosoovbco0aecde 23 45 37 3 67 
IOBNayEt As odesecdsodeG0 82 120 55 76) 119 
NATO Ge ee es eee: 81 112 52 64 112 


Affinities.—A large specimen of this species was figured by 
Abich as Clypeaster turritus, Philippi, a species which was 
admirably figured and described by its founder*. The specimen 
referred to O. turritus by Abich seems to me a very distinct form. 
In C. turritus the height is almost exactly half the length, 
whereas in Mr. Gunther’s specimen it is less than a third the 
length ; in Abich’s specimen it is almost a third the length. A 
still more important difference is in the length of the petals. 
In C. turritus the ratio of the length of the petal to the 
non-petaloid portion of the ambulacrum between the petal 
and the ambitus is as 2:1. In C. Guentheri the ratio is 
as 3:1. 

C. turritus is one of the conical, pyramidal species of Clype- 
aster. C. Guenthert, on the contrary, is one of the flat-topped, 
depressed species. Jts nearest ally is C. gibbosus (Risso) +, which 
differs in having a higher and longer test and fewer granules on 
the interporiferous ridges. 

Fischer £ has already suggested that Abich’s C. turritus should 
be included as a synonym of C. gibbosus. But Fischer figured 
as C. gibbosus an echinid which is distinct from Abich’s C. gib- 
bosus ; Hischer’s C. altus§ is, however, probably the same as 
C. Guentheri and C. turritus of Abich. 


* R. A. Philippi, “Ueber Clypeaster altus, C. turritus, und C. seille,” 
Palxontogr. vol. i. 18—, p. 323, pl. xxxviii. figs. 1-5. 

t Scutella gibbosa, Risso, Hist. Nat. Europe mérid. 1826, v. p. 284; Michelin, 
op. cit. p. 120, pl. xxii. figs. a-g, pl. xxiii. figs. 1, a-c. 

{ P. Fischer in Tchihatcheff, ‘Asie Mineure,” Paleontologie, 1866-69, p. 306, 

§ bid. p. 308, pl. vii. fig. 1. 


422 DR. J. W. GREGORY ON THE 


Pomel * has founded a species, C. suboblongus, on some spe- 
cimens from Corsica and Algeria, which are closely allied to 
O. Guentheri, but they appear to me to be closer to C. gzbbosus. 
O. suboblongus, at any rate, differs from C. Guenther by having a 
longer and more gradual posterior slope, without the separation 
into the steep upper part and thin basal foot. In C. Guentheri, 
moreover, the petals are proportionately longer, and the inter- 
poriferous granules less numerous, and there are fewer plates in 
the petaloid portions of the ambulacra (about 40 instead of 
about 60). 

Abich included C-. aléws in his synonyms of C. turritus; but in 
his explanation of plates he quoted the specimen as C. altus, 
TLam., var. turritus, Philippi. That he was right in identifying 
CO. turritus with C. altus I have urged previously t. Later on, 
Abich figured another specimen of C. altus; the specimen can- 
not be determined satisfactorily from the figure, but the echinid 
is certainly not a Clypeaster, and is probably a Conoclypeus. 


8. CiypeasTeR Martini, Desmoulins ¢. 

Abich, in 1882, figured an echinid from the Lower Miocene of 
Mamachatun which he referred to this species; the specimen is 
imperfect, and the ambulacra are so badly shown that the iden- 
tification might have been questioned. But Mr. Giinther’s 
collection includes three specimens which must, I think, be 
referred to C. Martinz, and thus support Abich’s identification. 
Mr. Loftus’s collection in the British Museum includes another 
specimen (H. 2446) which may be included in the same species, 
though the petals are more tumid than in the other specimens. 

The five specimens in question are by no means identical in 
form. One of Mr. Giinther’s specimens is too fragmentary to 
be of service; but the dimensions of the other four specimens 
and of Michelin’s figured specimen of C. Martini are given on 
the opposite page. The dimensions for C. melitensis, Mich., and 
C. Michelotti, Ag. (fide Michelin) are also added. 


* A. Pomel, ‘Paléontologie de l’Algérie: Zooph., Fase. 2. Echinodermes,’ 
Livr. 2, 1887, pp. 192-3, pl. B xxiii. figs. 1-6. 

+ J. W. Gregory, “The Maltese Fossil Echinoidea,” Trans. R. Soc. Edinb. 
vol. xxxvi. pt. 3, 1891, p. 594. 

+ C. Desmoulins, “Etudes sur les Echinides,” Mém. 3, Actes Soe. Linn. 
Bordeaux, vol, ix. 1837, p. 64; Michelin, op. cit. p. 134, pl. xxxv. fig. 1. 


23 


FOSSIL ECHINOIDEA OF LAKE URMI. 


Figured by Mr, Ginther’s Loftus Michelin’s Tes : ; 
Abich, specimens, Collection. type. C. melitensis. |C. Michelotts. 
ISIOHEING cocooooe écondecccunaadendote ‘eeeee|| 14) mm, 21 mm. 26 mm. 32 mm. 19 mm. 45 mm. 20-35 mm. 
IVGITY{010 Seceopesnace ee NeRe mane Secects 100 _,, i MOO), 101 _,, D0 TS op Om, 90-145 ,, 
Width at anterior angle....... Soaanaae Ss 5, SY) 5 S34, TBD 5, 84 C,, 
150 ,, 80-130 ,, 
= JOONWENOP 5 ccovspoononed Sol here — Gao W825; 
Ratio of length of petal to eg sa ; i a eis ¢ er 
Oli HN® HE) coosncosnensonos estat Dee Doe) 3:43 3:48 aoe 3:4] 
Number of granules on interpori- . 
ferous ridge Adasen ae Sete | He © 8 A os 10 9 


424: DR, J. W. GREGORY ON THE 


The main difference between the three specimens of C. Martini 
from Guverchin Kala is in the tumidity of the ambulacra. This 
character is least developed in the more broken of Mr. Ginther’s 
specimens, and most pronounced in the specimen in the Loftus 
Collection, which presents an approach to C. erassicostatus, 
A gassiz.* 

Among the specimens of Clypeaster from the Indian Cainozoic 
this species is closest to C. falorzensis, Dunc. & Sladen‘, which has 
the same long open petals and depressed test ; but in the Indian 
specimen the margin is tumid and the height of the test less. 


FOSSIL CORALS. 
By J. W. Greeory, D.Sc., F.G.S. 


(PuaTE 28.) 


TueE fossil corals in Mr. Giinther’s collection from Lake Urmi 
number 19 specimens, of which 6 came from the conglomerates 
at Seir, 6 or 8 miles west from the lake, and 8 came from 
the island of Koyun Daghi; the exact locality of the remainder 
is not stated. One of the specimens from Seir and one from 
Koyun Daghi are indeterminable. The rest may be divided 
among eight species, which have been previously recorded or 
described from Lake Urmi by Abich{. The corals are Miocene, 
and mainly Helvetian in affinities. The Ostrea-Virleti beds 
which yield LThamnarea polymorpha are shown dy the Mollusca, 
according to Mr. R. B. Newton, to be Tortonian in age. 


1. Orstcetta Derrancet (Hdwards §& Haime), 1849. 

One of the best-preserved corals in Mr. Giinther’s collection 
is a fragment of a flat tabular corallum, about 15 mm. thick, in 
which the septa number three orders with rarely a rudimentary 
septum of the fourth order; the septa are thick near the margin, 
but have no paliform lobes; the columella is parietal, but strong 

* Michelin, op. cit. p. 115, pl. xvii. figs. 1, a-f. 

+ Duncan & Sladen, ‘“ Fossil Echinoidea of Kachh and Kattywar,” Pal. 
Indica, ser. XIV. vol. i. pt. 4, 1883, p. 50, pl. xii. fig. 15. 

+ H. Abich, 1859, ‘‘ Ueber das Steinsalz und seine geologische Stellung im 
Russisenen Armenien,” Mém. Ac. Imp. Sci. St. Pétersb. ser. 6, vol. ix. pt. 1, 
Mém. Sci. Math.-Phys. vol. vii. pp. 89-102; and 1882, ‘Geologische Forsch- 
ungen in den Kaukasischen Landern, Abth. ii. Geologie des Armenischen 
Hochlandes, i. Westhalfte, pp. 270, 273, 274, 281-283. 


FOSSIL CORALS OF LAKE URMI. 425 


and rod-like; the exotheca develops as horizontal layers, and 
is not vesicular; the coste are long and prominent on the sur- 
face; the calices are 5-6 mm. in diameter, and are separated by 
exothecal bands froma 2-6 mm. in width. 

The coral is accordingly an Orbicella, and agrees very closely 
in structure with the corals from Urmi figured by Abich* in 
1859 as Astrea Guettardi. That the coral is identical with that 
so determined by Abich I have no doubt; but his naming of the 
species is open to question. Orbicella Guettardi was founded by 
Defrance + on a figure by Guettard{; the species has been 
admirably refigured by von Reuss §, and described by Edwards 
and Haime ||. Von Reuss’s figures confirm those of Guettard 
in showing that the exotheea is vesicular, that the septa are four 
cycles in number, that at least the primary septa have paliform 
thickenings, and that the diameter of the calices is over 10 mm. 
In all these characters the Urmi specimens are different; the 
septa belong to 3 cycles, there are no paliform lobes, the exo- 
theca is lamellar, and the calices are from 5-6 mm. in diameter. 
The Persian specimens must therefore be transferred to O. De- 
francet (Ed. & H.) 4, a Lower Miocene species from Dax and 
Northern Italy, which has been recorded from Asia Minor by 
Fischer **, whose figures agree with those of Abich and with 
Mr. Giiuther’s specimen. Michelintr has figured the species 
from Northern Italy under the name of 4A. argus, Lam., and 
von Reuss tf has given excellent figures of it. The specimens 
figured by Abich §§ as Astrea Defrancet are different, and one 
of them is here described as a new species. 


* H. Abich, op. cit. 1859, p. 89, pl. ii. fig. 4, pl. v. fig. 5. 

tT Defrance, 1826, Dict. Sci. Nat. vol. xlii. p. 379. 

t Guettard, Mém. sur différ. Parties Sci., vol. iii. 1770, pl. xlviii. figs. 2-4. 

§ Von Reuss, “ Pal. alt. Tertiarsch. Alpen,’ Denkschr. Akad. Wiss. Wien, 
vol. xxviii. 1869, p. 245, pl. xxiii. figs. 1, 2. 

|| Milne-Edwards & Haime, Hist. Nat. Cor. vol. ii. 1857, p. 462. 

9 Milne-Edwards & Haime, Mém. Astr. pt. 3, Ann. Sci. Nat., Zool. ser. 5, 
vol. xii. 1849, p. 106. 

xx P. Fischer in Tchihatcheff’s ‘L’Asie Mineure,’ Paléont., 1866-1869, p. 314, 
pl. xvi. figs. 4-6. 

tt H. Michelin, Icon. Zooph. 1842, p. 59, pl. xii. fig. 6. 

{tt Von Reuss, “ Foss. Kor. dster.-ung. Mioc.,” Denk. Ak.Wiss. Wien, vol. xxxi. 
1871, p. 239, pl. ix. fig. 3, pl. x. fig. 1. 

§§ Abich, 1859, op. cit. p. 93, pl. ix. fig. 6 ; 1882, op. cit. pp. 272, 273, pl. vii. 
fies. 15-20. 

LINN. JOURN.—ZOOLOGY, VOL. XXVII. 32 


426 DR. J. W. GREGORY ON THE 


2. OrBICELLA GUENTHERI, 0. sp. 

Diagnosis. Corallum’ massive, and growing apparently in 
tabular expansions. : 

Corallites of medium size; closely packed, and therefore 
appearing angular in shape, and usually hexagonal. Hxotheca 
narrow but dense, forming a stout wall. 

Septa three complete orders, and an imperfect fourth order 
represented. Seen from the exterior, the six primary septa are 
especiully conspicuous. 

Columella large, appearing substyliform. 


Dimensions. 
From Specimen 
Abich’s figure. from Seir. 
Inm mm 
Diameter of corallites ..........0.0scee00s. 8-9 5-7 
Diameter of calice ..............02-eceeees 5-6 3-4 
VIvieliday Oe WEIL cosnqssdooscoonacaccosaqade00 13-2 eile 


Affinities. This species is founded on three corals—one figured 
by Abich* as Astrea Defrancet, and the other two collected by 
Mr. Giinther in the conglomerate of Seir, 6 to 8 miles west 
from Lake Urmi. The corals agree very closely, the only differ- 
ence between them is the somewhat smaller diameter of the 
corallites in the specimens from Seir. 

The coral differs from O. Defrancet in three important cha- 
racters:—The corallites are smaller in diameter; the wall is 
narrower, denser, and not lamellar; the corallites are polygonal, 
and they are not divided by valleys or depressions between the 
raised calycinal margins. Hence the coral appears to me quite 
distinct from O. Defrancei. 1t may be distinguished from 
O. vesiculosus (Ed. & H.) + owing to the absence of the vesicular 
endotheca and exotheca. 

In 1882 Abich figured two additional Armenian astreans as 
Heliastrea Defiancei. Of these the first variety { is probably 


* Abich, 1859, op. cit. p. 93, pl. ix. fig.6; ? also 1882, op. cit., Heliastrea 
Defrance?, var. 1, p. 278, pl. vii. fig. 20. 

+ M.-Edwards & Haime, Mém. Astr. pt. 3, Ann. Sci. Nat., Zool. ser. 3, vol. xii. 
1849, p. 107. 

t Abich, 1882, op. cit. p. 278, pl. vii. fig. 20. 


FOSSIL CORALS OF LAKE URMI, 427 


a specimen of O. Guenthert ; the transverse section looks different 
from that of Abich’s figure (pl. ix. fig. 6 a) of 1859, but that figure 
was probably based on an altered specimen in which the true in- 
ternal structure was obscured. The second variety is no doubt 
a worn fragment of O. Haimez (d’Archiac) *. 


3. OrpicetLa Haimet (d’ Archiac) t, 1866. 

This species was founded for a specimen said to come from the 
Lower Tertiary of Thrace. D’Archiac’s figure agrees so closely 
with two specimens from the Seir conglomerate, that I feel doubt 
as to the correctness of the Thracian horizon. The two specimens 
agree in all essential characters; but the smaller specimen has 
been rolled and worn so that the corallites are left separated by 
a raised wall, instead of by intercalicular depressions. 

The dimensions of the type and of the specimens from the 
Seir conglomerate are given herewith :— 


Type. Specimens from Seir. 
mm. mm. mm. 
Length of corallum ............... 130 by 75 80 by 50 70 by 50 
Maiclaressh) |, \ sleeves ete — 70 30 
= o 
Diameter of corallites ......... { 2 uN 5). Be \ 8-15 15 


The coral figured by Abich in 1882 ¢ as Heliastrea Defrancei 
var. 2, is probably a specimen of this species. 


4. SOLENASTREA TURONENSIS (Michelin), 1847 §. 

This widely distributed Helvetian coral is represented by two 
specimens—a large mass 130 mm. in diameter from the conglo- 
merate at Seir, and a rolled specimen from Koyun Daghi. In 
reference to the latter, it should be noted that the lithological 
condition of the fossil is very different from that of the other 
specimens from that island. As usual in fossil specimens of Solen- 


* Abich, bid. p. 274, pl. vii. fig. 15. 

} Heliastrea Haimei, d Archiac in Tchihatcheff’s ‘ L’Asie Mineure,’ Paléont., 
1866-69, p. 191, pl. xv. figs. 5, 6. 

t Abich, 1882, op. cit. p. 274, pl. vii. fig. 15. 

§ Astrea turonensis, Michelin, 1847, Icon. Zooph, p. 312, pl. Ixxv. figs. 1, 2; 
Solenastrea turonensis, Hdwards & Haime, 1857, Hist. Nat. Cor. vol. ii, p. 498. 


o2* 


428 DR. J. W. GREGORY ON THE 


astrea, the septa are seldom preserved; but they are shown in 
a few corallites of the Koyun Daghi example. The species 
occurs in Egypt *, but has not, so far as I am aware, been pre- 
viously recorded from Asia Minor or Persia. Abich + quotes a 
Solenastrea astroites from Lake Urmi, and that record may have 
been based on 8. turonensis. Abich did nct give figures, and 
accepted the species for the Sarcinula astroites (Goldf.)t, which, 
according to Milne-Edwards & Haime§, is a synonym of Orbi- 
cella Ellist (Defr.). That determination is quite consistent with 
Goldfuss’s instructive figure. Abich quoted as a second refer- 
ence the Solenastrea columnaris (Rss.)||, which appears quite 
distinct both from “ Sarcinula astroites” and from Solenastrea 
turonensis. 


5. PRIONASTRHA IRREGULARIS (Defrance), 1826 4. 

This species has been recorded from Armenia by Abich, who 
has figured a specimen collected between Malu and Khoi, which 
shows the deep steep-walled calicular fossa of this species. 
Mr. Giinther’s collection includes two specimens from Lake 
Urmi, exact locality not stated. 


6G. Puyznocmnta Arcutract, Edwards jf Haime, 1848 **. 

This species was recorded by Abich from Koyun Daghi, but 
his figures alone are not convincing of the specific identification. 
Mr. Ginther has collected two other specimens from the same 
locality, which enable the characters to be more fully determined. 
The Armenian specimens differ from Milne-Edwards and Haime’s 
diagnosis in the more solid nature of the exotheca and in the 
apparent absence of the numerous granulations on the coste, 
which are said to be visible on worn specimens. The specimens 
from Koyun Daghi are extremely worn, and it may be that the 


* J. W. Gregory, “ Egypt. Foss. Madrep.,” Geol. Mag. dec. 4, vol. v. p. 247. 

t Abich, 1882, op. cit. p. 282. 

{ Goldfuss, Petref. Germ. vol. i. p. 78, pl. xxiv. fig. 12. 

§ Milne-Edwards & Haime, Hist. Nat. Cor. vol. ii. 1857, p. 467. 

|| Von Reuss, “ Pal. alt. Tertiarsch. Alpen,” pt. i. Denkschr. Ak. Wiss. Wien, 
vol. xxviii. 1868, p. 170, pl. xi. figs. 7-9. 

“| Astrea irreqularis, Defrance, 1826, Dict. Sci. Nat. vol. xlii. p. 381; 
Cellastrea irregularis, de Blainville, 1850, bid. vol. lx. p. 342, Figured by 
Michelin, 1842, Icon. Zooph. p. 61, pl. xii. fig. 9, but the figure is indefinite. 

** Milne-Edwards & Haime, Mon. Astr. pt. i., Ann. Sci, Nat., Zool. ser. 3, 
vol. x. 1848, p. 803, 


Gregory: Linsn. Soc. Journ. Zoon’ Vou. XXVIII. Pl. 28. 


E_Drake ad nat.lith. West, Newman imp 


FOSSIL HCHINOIDEA AND CORALS, FROM LAKE URM1. 


FOSSIL CORALS OF LAKE URMI. 4929 


eranulations have been thus obliterated. In the compactness of 
the exotheca the corals approach Abich’s Astrea grandistella*, 
which, owing to the absence of columella, is probably also a 
Phyllocenia; but as the naturé of the exotheca and union of the 
corallites is not shown by the figures, its generic position cannot 
be absolutely determined. 

The two specimens from Koyun Daghi are flat-topped, tabular 
coralla, and in the larger specimen the corallites show a tendency 
to become elliptical or subtriangular. 

There is a third specimen in Mr. Ginther’s collection from 
Koyun Daghi referable to Phyllocenia; it has well raised cali- 
cular margins, as in P. zrradians, Hd. & H. +, but the fragment 
is too small and worn for satisfactory determination. 


7. THAMNAREHA POLYMORPHA (Adich) ft, 1859. (Plate 28. 
fig. 3.) 

This interesting coral was well figured by Abich, whose speci- 
mens came from Lake Urmi and Malischent or Maku. Their 
specific identity with specimens collected on Koyun Daghi by 
Mr. Gunther is unquestionable. The horizon of the coral 
on this island is settled by the fact that one of them is at- 
tached to an Ostrea which Mr. R. B. Newton has identified as 
O. Virleti. 

The generic position of the coral is a matter of some interest, 
as Thamnarea has not previously been recorded from Miocene 
or even Cainozoic deposits. That the coral is not a Porites is 
shown by three characters: the absence of pali, the strongly 
developed, imperforate septa, and the presence of the conspicuous 
synapticular platforms, which are well shown in Abich’s figures. 
On casual inspection the coral appears spongiform rather than 
corallian in structure. 

Thamnarea was founded by Thurmann & Htallon§ in 1864, 
and has recently been discussed by Dr. Ogilvie ||, who includes 
it in the Madreporide; but, with Htallon, von AZittel, and 
Dunean, I include it among the Microsolenide. 


* Abich, 1859, op. cit. p. 92, pl. ii. fig. 3. 

t+ This species has been recorded from the Oligocene of EHrivan by Abich, 
1882, op. cit. p. 257, pl. vii. fig. 16. 

{ Porites polymorpha, Abich, 1859, op. cit. p. 100, pl. ix. figs. 1, a-e. 

§ “ Leth, Brunt.,” Neue Denk. schweiz. Ges. Naturw. vol. xx. p. 411. 

|| M. M. Ogilvie, “ Kor. Stramb. Sch.,” Paleontogr. Suppl. ii. pt. 7, p. 153. 


A430 MR. R. B. NEWTON ON THE 


8. P PoRITES LEIOPHYLLA, von Reuss *, 1847. 

The collection includes a large nodular corallum 100 mm. long 
by 80 mm. wide and 40 mm. thick, which is a cast of a Porites. 
It agrees in characters with Abich’s ¢ description of the coral 
from the islands of Lake Urmi referred to P. leiophylla. It 
differs from von Reuss’ type of that species by the greater 
size of the corallites. This feature may only be the result of the 
greater size of the corallum. The coral is only a cast, so that 
its determination is difficult, and it may be provisionally left in 
the species to which Abich assigned it. The corallites in the 
present specimen vary from 7-10 mm. in diameter. 


EXPLANATION OF PLATE 28. 


Hig. 1. Clypeaster imperialis, Mich., side view, nat. size. 


2. Clypeaster Guentheri, n. sp., side view, nat. size. 


3. Transverse and radial sections of Zhamnarea polymorpha, xX 10 diam. 
Yortonian, Koyun Daghi, Lake Urmi. 
a. Trarisverse section across a series of corallites, x 10 diam. 
b. Section across part of a transverse section of a branch, showing 
the corallites cut longitudinally, with the synapticular plat- 
forms. 


MARINE TERTIARY [MIOCENE] MOLLUSCA. 
By R. Butien Newton, F.G.S8., of the British Museum 
(Natural History). 


(Puatus 29 & 30.) 


INTRODUCTION. 


Turs communication deals with a number of Marine Tertiary 
Mollusca obtained from Lake Urmi and its vicinity by Mr. R. T. 
Giinther, who has generously presented the specimens to the 
Geological Department of the British Museum. 

The specimens are mostly casts and frequently much water- 
worn, yet of considerable interest since they serve to increase the 
number of species already recorded from this area. 


* Von Reuss, “ Foss. Polyp. Wien. Tertiarb.,” Haid. Naturw. Abh. vol. ii. 
pt. L, p. 28, pl. v. fig. 4. 
+ Abich, 1859, op. edt. p. 101. 


MIOCENE MOLLUSCA OF LAKE URMI. 431 


One Lamellibranch, which appears to differ from every known 
form, is here described (p. 447) as a new species under the name 
of Meretrix persiensis. 

Speaking generally, the collection consists of Miocene species, 
although a few of its members assume a facies indicative of the 
Aquitanian division of the Tertiary rocks, which by some authors 
(e.g. Giimbel and Fuchs) is regarded as Lower Miocene, and by 
Lapparent and others as Upper Oligocene. 

From palzontological evidence it would appear that the 
Tertiary rocks of Lake Urmi form part of an extensive series of 
deposits which is traceable through Asia Minor, Southern Europe 
(Greece, Vienna Basin, Italy, and Spain), Egypt (Siwa Oasis, 
Pyramids, Geneffe), Northern Africa, and Madeira. 

The following remarks on the localities represented in the 
collection may be of interest. 

The majority of the specimens were obtained from the beach 
and cliffs of the Island of Koyun Daghi, which, according to 
Abich, has an elevation of 5258 feet above sea-level, and in 
section is seen to be composed entirely of Miocene rocks resting 
on a Paleozoic base—(a) the highest bed contains Alectryonia 
Virleti ; (6) beds of marble appearance with Corals; (c) concre- 
tionary limestones with Turritella Archimedis, T. turris, T. gra- 
data, &c.; (d) Paleozoic rocks. 

The following species from this locality are represented in 
Mr. Giinther’s Collection :— 

Alectryonia Virleti, Ostrea pseudodigitalina, Ostrea lamellosa, 
Pecten suburmiensis, Pecten allied to Burdigalensis, Pecten 
(Flabellipecten) sp. indet., Chlamys (diquipecten) Mal- 
vine affin., Chlamys sp. indet., Cardita sp. indet., Strombus 
like Bonelli, Conus sp. indet., Turritella Archimedis, 
Turritella gradata var., Turritella rotifera affin., Latirus 
crispus. 

In considering the zone or zones of the Miocene system to 
which these forms should be referred, it may be observed that the 
presence of Alectryonia Virleti, Latirus crispus, Ostrea lamellosa, 
&c., would suggest a later age than that represented by Turri- 
tella Archimedis, T. gradata var., Pecten allied to Burdigalensis, 
&c. It is therefore quite possible that these older species are 
of Helvetian age, whereas the others are probably Tortonian, 
or even later, as they occur also in Pliocene deposits. The 
specimens from Koyun Daghi are in a cream-coloured marly 


432 MR. R. B. NEWTON ON THE 


caleareous rock; those found on the beach being very much 
rounded and thickly impregnated with salt. 

Another of Mr. Giinther’s localities is that of Guverchin Kala 
(see sections made by Mr. Loftus in 1855), from which the 
following species have been identified in his collection :— 

Pecten convexocostatus, Pecten suburmiensis, Venus Aglaure, 

Meretrix persiensis, Meretria like incrassata, Pyrula cin- 
gulata, Cassis sp. indet. 

This assemblage of forms is suggestive of two horizons—one, 
represented by Pecten convexocostatus and P. suburmiensis, which 
is probably Helvetian ; the other, on account of such species as 
Venus Aglaure, Meretrix Persiensis, M. like incrassata, Pyrula 
cingulata, Cassis sp. indet., is most likely of Burdigalian age or 
even older, as the shells referred to exhibit the Aquitanian facies 
before mentioned, besides being lithologically distinct. The 
Pectiniform species are in a light-coloured limestone; whereas 
the supposed older specimens are preserved as casts of reddish- 
brown colour, and came probably from near the base of the 
‘* Loftus Section” (P about no. 7 bed), having been collected and 
given to Mr. Giinther by the Rev. C. Labaree, of the American 
Mission Station. 

Some further specimens have been obtained from other 
localities close by, which are referred to in the body of this paper 
and do not call for any special reference now, except that, 
judging from their characters, they indicate a Lower Miocene age, 
and belong either to the Helvetian or Burdigalian stages of that 
period. 

A summary of previous work on the invertebrate paleontology 
of Lake Urmi and neighbourhood may now be given. 

In 1855 W. K. Loftus * published his geological researches on 
the Turko-Persian frontier, in which is described the ‘“‘ White 
Limestone”? promontory (regarded by Loftus as of Upper 
Nummulitic age) bearing the ancient fortress of Guverchin Kala 
(=Castle Rock), situated at the northern extremity of Lake 
Urmi. A detailed section of this point, where the rocks rise 
perpendicularly to a height of more than 400 feet, is given thus 
(taken from p. 305 of the paper by Loftus) :— 

“The following is a careful descending section of the Castle 


* Lorrus, W. K.—‘‘On the Geology of Portions of the Turko-Persian 
Frontier and of the Districts adjoining,” Quart. Journ. Geol. Soc. 1855, vol. xi. 
pp. 247-344, & Geological Map. 


MIOCENE MOLLUSCA OF LAKE URMI. 433 


Rock. The beds are all conformable to each other, and dip at an 
augle of 7° towards the H.S.E., which dip is of course due to the 
igueous rocks of the Wurgowiz spur on the north. 


‘““1. Compact, hard, crystalline, white limestone, be- 
coming concretionary in passing downwards, 
afterwards marly. 

2. Iight-blue matl, with hard flesh-coloured flints and 
nodules, and irregular fragments of limestone. 
It contains abundant Corals in sitw and in layers, 
below which are numerous specimens of Clype- 
aster, Echinolampas, Pecten, Serpula, and casts 
of various Univalve and Bivalve shells. 

3. Compact mass of highly crystalline coralline 
nodules in hard marls. 

The thickness of the above three bedsisabout.. 280 feet 

4, Fine reddish gravel, or coarse sand-conglomerate, 
much hardened, and filled with fragments of 


OSS SP atectea hed 5 oa Ge cnet tg cecratcay st «chia «oleae oe SW 
5. Friable yellow sandstone, very finely grained, 
WHE) ECMO AZ HONS Soleoodsonsesdnocenc 4 IGF 


This bed passes into 
6. Hard and compact, grey, marly limestone, filled 
with Corals and castsof shells .............. Gee 
7. Hard reddish marl, abounding in shells ........ Zieie 
8. Brownish-yellow, friable sandstones, with several 
thin layers of gravel and conglomerate, of 
variously-sized rounded pebbles ............ OO 


Total thickness of Section .... 410 ft.” 


Dr. H. Abich * issued his first paper on the paleontology of 
this region in 1858, accompanying it with an excellent geologi- 
cally coloured map taking in the whole of the islands on Lake 
Urmi, and recognizing them as belonging to the ‘“ Miocéne 
inférieur.” His. determinatious of the fossils inclined him to 
consider that they should be referred to the ‘‘ Molasse Moyen ”’ 


* Axsicu, H.—“'Tremblement de Terre observé 4 Tébriz en septembre 1856, 
Notices physiques et géographiques de M. Khanykof sur l’Azerbeidjan,” Bull. 
Classe Physico-Math. Ac. Imp. Sci. Saint-Petérsbourg, 1858, vol. xvi. pp. 840- 
341, pl. 111. (=Geological Map: coloured). 


434 MR. R. B. NEWTON ON THE 


or “ Falunien ” (Orbigny) age of the Mediterranean Basin, which 
he stated was identical with the “ Calcaire Moéilon” of Marcel 
de Serres and very similar in characters to the Leithakalk of 
Austrian geologists. 

This author also called attention to the great extent of the 
Miocene deposits, which he stated could be traced from Marocco 
through Greece to Lake Urmi. 

The following fossils, collected by M. Khanykof from the 
islands on Lake Urmi, were listed without descriptions or 
figures as follows :— 


Actinozoa (Corals): 
Porites dendroidea, n. sp. 
Astrea Guettardi, Detr. 
—— Lllisiana, Defr. 
—— Defrancei, M.-Edw. 
Phyllocenia Archiaci?, M.-Edw. 


EcHINODERMATA : 
Clypeaster altus, Lam. 
crasstcostatus affin., Ag. 


Hchinolampas complanatus, n. sp. 


Brrozoa: 
Cellepora gracilis, Miinst. 
Ceriopora palmata, Orb. 
anomala, Nn. sp. 
Polytrema spongiosa, Orb, 
Diastopora gemmifera, n. sp. 
Membranipora fenestrata, Hichw. 


| LAMELLIBRANCHTATA : 
Ostrea Virleti, Desh. 
excavata, Desh. 
lamellosa, Lam. 
Pecten benedictus, Lam. 
flabelliformis, Broechi. 
simplex, Mich. 
Spondylus bifrons, Goldf. (Minst.). 


GASTEROPODA: 
Flaliotis Philberti, Serres. 


Much more systematic work on the fossils of this Iceality was 


MIOCENE MOLLUSCA OF LAKE URMI. ABS. 


carried out by Abich* in the same year, when he figured and 
described the following species as having been obtained from the 
“ Bryozoen und Foraminiferen-Kalkstein” division of his “ Supra- 
Nummulitenkalk” series, at Urmi and neighbourhood. 


TFoRAMINIFERA : 
Polystomella quaterpunctata, Nn. sp. 
Actinozoa (Corals) : 
Astrea Guettardi, Defr. 
— Lllisiana, Defr. 
—— Defrancii, M.-Edw. 
—— grandistella, n. sp. 
Phyllocenia @’ Archiaci, M.-Edw. 
Porites polymorpha, n. sp. 
letophylla, Reuss. 
ECHINODERMATA : 
Clypeaster altus, var. turritus, Phil. 
diversicostatus, N. Sp. 
Echinolampas complanatus, nu. sp. 
Bryozoa: 
Ceriopora palmata, Orb. 
Polytrema spongiosa, Phil. 
Cellepora gracilis, Minst. 
Diastopora gemmifera, n. sp. 
Membranipora fenestrata, Kichw. 
Ceriopora anomala, n. sp. 
LAMELLIBRANCHIATA : 
Pecten planicostatus, n. sp., P. convexocostatus, n. sp., 
P. benedictus, Lam., P. maximus, Linn., 
P. burdigalensis, Lam., 
P. flabelliformis, Brocchi, P. simplex, Michelotti, 
P. varius, Linn. 
Spondylus bifrons, Goldfuss. 
Ostrea Virleti, Desh., O. excavata, Desh., 
O. lamellosa, Lamarck. 
GASTEROPODA: 
Haliotis Philberti, Serres. 


* Axsicu, H.— Ueber das Steinsalz und seine geologische Stellung im 
Russischen Armenien,” Mém. Ac. Imp. Sci. St. Petérsbourg, 1858, ser. 6, 
vol. vii. pp. 61-150, pls. 1-10. Although read December 14, 1856, and issued 
asan author's copy in 1857, this paper was not published until 1858. 


436 MR. R. B. NEWTON ON THE 


During 1873 Dr. W. T. Blanford * published his views 
regarding the Salt swamps and lakes of Persia, including the 
lake of Urmi; and in 1876 the same author 7+, in his ‘ Hastern 
Persia,’ gives a special chapter on the geology of Persia 
generally with an exhaustive réswmé of all previous work on 
the subject. 

A distinct advance was made by Dr. Abich } in his great work 
of 1882, which includes a geological protile section of the 
islands (p. 275) on Lake Urmi, representing them to be composed 
of Miocene rocks resting on a Paleozoic base, having the Ostrea 
Virleti-beds at the top of the series. The elevation of the 
principal island, that of Koyun Daghi, is stated to be 5258 feet 
above sea-level. 

In this work the following species of Miocene age are described 
and (mostly) figured as having been collected on the ‘ Urimia 
Plateau ” :— 


Actinozoa (Corals): 

Phyllangia grandis, Reuss. 
alveolaris, Catullo. 
Cyathomorpha gregaria, Catullo. 
conglobata, Reuss. 
Astrangia princeps, Reuss (allied). 
Solenastrea astroites, Goldfuss. 
Heliastrea Guettardi, Defr. 
Porites polymorpha, Abich. 


EcHINODERMATA : 
Clypeaster turritus, Phil. 
Scille, Desm. 
Echinolampas complanatus, Abich. 


LAMELLIBRANCHIATA : 
Venus Aglaure, Brongn. 
Pecten suburmiensis, n. sp. 
convexocostatus, Abich. 
lychnulus, Fontannes. 


* Buanrorp, W. T.—‘“‘On the Nature and probable Origin of the Superficial 
Deposits in the Valleys and Deserts of Central Persia,” Quart. Journ. Geol. Soc. 
1873, vol. xxix. pp. 493-503. 

+ Buanrorp, W. T.—‘ Eastern Persia, an Account of the Journeys of the 
Persian Boundary Commission, 1870-1872’: 1876, vol. ii. p. 439-506. 

{ Asicu, H.-—‘ Geologie des Armenischen Hochlandes.’ 1882. 


MIOCENE MOLLUSCA OF LAKE URMI. 437 


Pecten Tournali, Serres. 
subopercularis, n. sp. 
— WMalvine (allied), Dubois. 
GASTEROPODA: . 
Turritella Archimedis, Brongn. 
turris, Orbigny. 
rotifera, Desh. (allied to). 
gradata, Menke, var. 
margarita, D. sp. 


These species are additional to those cited in Abich’s work 
of 1858, which were mostly obtained from the islands on Lake 
Urmi; with the following alterations in nomenclature: Pecten 
maximus to be P. Tournali, Serres; P. simplex 10 be P. Hol- 
gert, Geinitz. 

Dr. Rodler * in 1888 reported the occurrence of Jurassic fossils 
at the following localities off the eastern side of Lake Urmi:— 
Aktahu-dere, Tazeh-kend, Iditsch, Guschaisch, and the Karangu 
Valley. The genus Harpoceras, which was found in the Karangu 
Valley, he regarded as indicative of Middle or Upper Lias; 
whereas Perisphinctes from the other localities might belong to 
either the P. polyplocus-zone of Kimeridge age or the P. curvi- 
costa-zone, which is Callovian. Rodler’s researches also includes 
a chapter on the origin of Lake Urmi. 

This was followed in 1890 by a memoir from Dr. K. Anton 
Weithofer ¢ on the Jura and Cretaceous rocks of North-western 
Persia, in which the following Mollusca are described and par- 
tially figured, as having been collected to the east of Lake 
Urmi (Guschaisch and Tazeh-kend) in the neighbourhood of 
Maragha:— 

Upper Lias: 

Harpoceras cf. radians, Reinecke. 
cf. kurrianum, Oppel. 
Belemnites sp. radet. 

Pecten ef. disciformis, Schibler. 
Pleuromya sp. indet. 


* Ropusr, Dr. Aurrep.—< Hinige Bemerkungen zur Geologie Nordpersiens,” 
Sitzunsgb. K. Akad. Wiss. (Wien), 1888, vol. xevii. pt. i. pp. 203-212. 

+ Werrnorrr, Dr. K. Ant.—“ Ueber Jura und Kreide aus dem nordwest- 
lichen Persien,” Sitzungsb. K. Akad. Wiss. (Wien), 1890, vol. xcviii. pt. i. pp. 756— 
778, pls. i. & ii. 


438 MR. R. B. NEWTON ON THE 


Upper Ootire (Lower Kimeridge) : 
Perisphinctes Lothart, Oppel. 

cf. polyplocus, Reinecke. 
Belemnites persicus, ni. sp. 
Goniomya Rodleri, n. sp. 

Cretaceous (Neocomian): 
Olcostephanus tetrameres, D. Sp- 
-—— Straussi, n. sp. 

new form indet. 

—— (Hoplites?) cf. narbonensis, Pictet. 


Dr. Georg von dem Borne* in 1891 contributed a valuable 
monograph on the fossils from the Eastern side of Lake Urmi 
(Guschaisch, Chanajan, Aktahu-dere, Scurgan and Tazeh-kend), 
which were described and figured from the following horizons :— 


(1) Kelloway (anceps-zone). 
(2) Kelloway ? (macrocephalus-zone). 
(3) Upper Lias (Jurensis-zone). 


His identifications may be thus tabulated :— 


CEPHALOPODA: 

Belemnites cf. calloviensis, Oppel. 
Persicus, Weithofer. 
sp. (=B. acuwarius-group). 
Ludwigia nodosa, Quenst., sp. 
krakoviensis, Neumayr, sp. 
gigas, Quenst., sp. 
lunula, Zieten, sp. 
ef. punctata, Stahl, sp. 
spp- indet. 
Macrocephalites sp. indet. 
Stephanoceras stenostoma, n.sp. 
Perisphinctes curvicosta, Oppel. 
paneaticus, Notling. 
cyrus, D. Sp. 
tetrameres, Weithofer, sp. 
poculum, Leckenby, sp. 
Aerxes, Di. Sp. 
spp. indet. 
balinensis, Neumayr. 


* Bornu, Grora von peM.— Der Jura am Ostufer des Urmiasees.’  [Dis- 
sertation Thesis.] Halle, 1891; with several plates of fossils. 


MIOCENE MOLLUSCA OF LAKE URMI. 439 


Reineckia Straussi, Weithofer, sp. 
sp. 
Harpoceras Atropatenes, 0. sp. 
kapautense, 0. 8p. 
medi@, DQ. Sp. 
GASTEROPODA : 
Spinigera, n. sp. indet. 
Pleurotomaria sp. 
LAMELLIBRANCHIATA : 
Pecten ctf. disciformis, Schibler. 
Mytilus Matianus, n. sp. 
Trigonia Roxane, n. sp. 
Sp. 
Pleuromya urmiensis, 0. sp. 
like arata, Brauns. 
Gresslya sp. 
ANNELIDA : 
Serpula sp. 


DESCRIPTION OF THE SPECIES. 


The synonymy here employed is not intended to be exhaustive ; 
only the principal works are enumerated, and these must be 
referred to for more complete details. For the sake of brevity, 
the following memoirs, quoted in the synonymy, are mentioned 
under the name of the “Serial” in which they originally 
appeared :— 

Axsicu, H.—Ueber das Steinsaiz und seine geologische Stell- 
ung im Russischen Armenien: J/ém. Ac. Imp. Sci. St. 
Pétersbourg, 1858, ser. 6, vol. vii. 

Basrrrot, B. de.—Description Géologique du Bassin Tertiaire 
du Sud-ouest de la France (Mollusques Fossiles): Mfém. 
Soc. Hist. Nat. Paris, 1825, vol. 11. pt. 1. 

Fucus, T.—Ueber die von Dr. Tietze aus Persien mitgebrachten 
Tertiarvesteinerungen: Denkschr. K. Akad. Wiss. (Wien), 
vol. xli. pt. 2. 

Fucus, T.—Beitrige zur Kenntniss der Miocinfauna Aegyp- 
tens und der Libyschen Wiiste: Palgontographica, 1883, 
vol. xxx. 

Horners, M.—Die fossilen Mollusken der Tertiir-Beckens von 
Wien (Univalven und Bivalven); <Abhandl. k.-k. Geol. 
Reichs., 1856-1870, vols. 11. & iv. 


4AM) MR. R. B. NEWTON ON THE 


LAMELLIBRANCHIATA. 

OsTREA PSEUDODIGITALINA, Huchs. 

Ostrea pseudodigitalina, Fuchs, Denkschr. K. Akad. Wiss. 1879, vol. xli. 
pt. 2, pl. 3. figs. 4-6, p. 107. 

This species is of elongate shape, pointed at the summit 
and rounded basally ; it is related to O. digitata of Kichwald, 
but appears to differ in possessing cost of considerably less 
digitiform character. The upper valve is flat and ornamented 
with prominent growth-lines. Other closely allied species are 
O. Rholfsi of Fuchs, from the Siokuh mountains in Persia, and 
O. digitalina var. Rholfsi of the same author, from the Miocene 
beds of Egypt. The specimens are fragmentary ; representing 
four lower valves and three upper valves. 

Fuchs first recorded this species from the Miocene (=Schio- 
Schichten) of the Siokuh mountains in Persia. 

Locality. In a grey marly matrix from the beach of the 
island of Koyun Daghi. 

Formation. Miocene (Helvetian ?). 


OsTREA LAMELLOSA, Broccht. , 

Ostrea lamellosa, Brocchi, Conch. Foss. Subapennina, 1814, vol. ii. 
p. 564; Abich, Mém. Ac. Imp. Sci. St. Pétersbourg, 1858, ser. 6, vol. vii. 
pl. 5. fig. 3, p. 126 (68); Hornes, Abhandl. k.-k. Geol. Reichs, 1870, 
yol. iv. pl. 71. figs. 1-4, pl. 72. figs. 1, 2, p. 444. 

A single fragmentary upper valve is all that represents this 
shell; and it seems to agree remarkably well with Abich’s 
interpretation of the species as illustrated in his “ Steinsalz ” 
monograph. It is slightly convex near the summit, and the 
external surface exhibits the regular concentric and lamellose 
striations characteristic of this species. It has been previously 
recorded from Italy (Brocchi); Urmi (Abich); Vienna Basin 
(Hornes); Asia Minor (P. Fischer in Tchihatcheff); Greece 
(Gaudry), &c. 

Locality. In agrey marly rock from the beach of the island of 
Koyun Daghi. 

Formation. Miocene (Tortonian or Helvetian). 

The species is distributed through the Miocene and Pliocene 
periods, and still survives in the Mediterranean. 


ALEctRYONIA VIRLETI, Deshayes. 
Ostrea Virleti, O. excavata, O. pseudoedulis, Deshayes, Expéd. Sci. 
Morée, 1883, vol. iii. (Zool.), pl. 21. figs. 1-6, pp. 122-124. 


MIOCENE MOLLUSCA OF LAKE URMI. 441. 


Ostrea Virleti, O. excavata, Abich, Mém., Ac. Imp. Sci. St. Pétersbourg, 
1858, ser. 6, vol. vii. pl. 2. figs. 1, 2; pl. 3. figs. 1, 2; pl. 5. figs, 1, 2; 
pp. 124 (66), 125 (67). 

Ostrea Virlett, Fuchs, Denkschr. K. Akad. Wiss. 1879, vol. xli. pt. 2. 
pl. 4, p. 106; and Paleontographica, 1883, vol. xxx. pl. 9. (4), pl. 10. (5), 
p. 43 (25). 

Ostrea (Alectryonia) Virleti, R. B. Newton, Geol. Mag. 1899, p. 205. 

This is a most abundant species in Persia, occupying the 
topmost bed of the Miocene section at Koyun Daghi island, as 
given by Dr. Abich in his 1882 work. It is a form of some- 
what variable shape, though exhibiting external plications on both 
valves, which places it in the genus Alectryonia. These plications 
may sometimes be more or less obsolete according to age and 
wear ; the valves are mostly depressed, though the eacavata form 
has a very convex lower valve. Mr. Ginther’s specimens are 
fairly typical of the species and correspond with Dr. Abich’s 
figures of the same. 

The species has been previously recorded from the Miocene 
rocks of Morea (Deshayes); Siokuh Mountains, Persia (Fuchs) ; 
Egypt (Fuchs); Azores &c. (Mayer-Eymar); Crete (Raulin); 
Cyprus (Gaudry); Malta (Wright); and a variety of this species 
is recorded from the Plocene deposits of Altavilla, Italy, by 
Gregorio (Boll. Soc. Mal. Italia, 1884, vol. x.). 

Localities. Several specimens were obtained from the beach of 
the island of _Koyun Daghi; and one fragmentary valve came 
from the cliff-section of the same island. 

Formation. Miocene, probably about the age of the Leithakalk, 
which is Tortonian. 


Prcren Beupantt, Basterot. (Plate 29. fig. 2.) 

Pecten Beudantt, Basterot, Mém. Soc. Hist. Nat. Paris, 1825, vol. ii. 
part 1, pl. 5. fig. 1, p. 74; Hornes, Abhandl. k.-k. Geol. Reichs. 1867, 
vol. iv. pl. 59. figs. 1-3, p. 399 ; T. Fuchs, Denkschr. K. Akad. Wiss. 1879, 
vol. xli. pt. 2. p. 105; R. B. Newton, Geol. Mag. 1899, p. 207. 

A large convex lower valve, contained in this collection, appears 
to be referable to P. Beudanti. The fine, striated sculpture of 
the surface, however, through erosion by water or weathering, so 
conspicuous a feature of this species, is almost obliterated, 
although it can be obscurely traced in places. In every other 
respect it agrees with Hornes’ interpretation of Basterot’s shell. 
It is very transverse (dimensions in millimetres=height 87, 
length 110, depth 35), and bears about 16 wide plano-conver 

LINN. JOURN.—ZOOLOGY, VOL. XXVII. 33 


4.42 MR. R. B. NEWTON ON THE 


cost separated by grooves of rather less width. Sacco has 
recorded this species from the Helvetian Beds of Piedmont (Boll. 
Soc. Geol. Italiana, 1889, vol. viii. p. 331). 

The species has been previously known from Bordeaux 
(Basterot) ; Portugal (J. de C. Sowerby); Constantine (Coquand) ; 
Sardinia (Meneghini) ; Piedment (Sacco) ; Persia (Fuchs); Egypt 
(Newton). 

Locality. Tn a matrix of compact, cream-coloured foraminiferal 
limestone ; from the neighbourhood of Urmi. 

Formation. Miocene (Helvetian). 


PECTEN CONVEXO-cosTaTuS, Abich. 

Pecten convexo-costatus, Abich, Mém. Ac. Imp. Sci. St. Pétershourg, 
1858, vol. vii. pl. 1. figs. 40, 4c, p. 188; Geol. Armenischen Hochlandes, 
1882, pl. 1. figs. 6, 6a, pl. 8. fig. 2, p. 276; Fuchs, Paleeontographica, 
1883, vol. xxx. pl. 21 (xv.). figs. 1, 2, p. 40 (58). 

Two lower valves of this species are attached to a piece of 
yellowish-white limestone. They are much weathered so that the 
fine concentric strie are almost obsolete, but the characteristic 
convex and narrow coste numbering from 20-22 are prominently 
shown. This is a fairly distinct species, and not to be confused 
with Hichwald’s P. aduneus of 1830, from Poland, which has 
wider, fewer, and more depressed ribs, and which Hornes regarded 
asisynonymous with P. convexo-costatus, uniting both under the 
older name. 

Abich’s original specimens were obtained from the neighbour- 
hood of Urmi; the species is also recorded from Egypt by 
Fuchs. 

Locality. Guverchin Kala, north end of Lake Urmi. 

Formation. Miocene (Helvetian). 


PEcTEN SUBURMIENSIS, Abich. 

Pecten suburmuensis, Abich, Geol, Armenischen Hochlandes, 1882, pl. 8. 
figs. 1-8a, p. 276. 

This species is represented by a number of lower valves, which 
although rather fragmentary show the characteristic wide, 
curved, and depressed coste crossed by obscurely fine striations ; 
the umbonal region is rounded and incurved. The original 
specimens were described by Abich as from the Urmi Plateau. 

Localities. Two examples are in a drab-coloured limestone, 
and were found at Guverchin Kala, northern end of Lake Urmi; 
two others are in a greyer rock localised as from the same place ; 


MIOCENE MOLLUSCA OF LAKE URMI. 443 


the remaining specimens are considerably water-worn, having 
been obtained from the beach at the island of Koyun Daghi. 
Formation. Miocene (Helvetian). 


Prcren (AmussiopEcren) like BurpreaLensts, Lamarck. 

Pecten Burdigalensis, Lamarck, Annales du Muséum, 1809, vol. viii. 
p. 355 ; Goldfuss, Petrefacta Germaniz, 1835, vol. ii. pl. 96. fig. 9, pp. 66, 
80; Abich, Mém. Ac. Imp. Sci. St. Pétersbourg, 1858, vol. vii. p.{120 
(62); Hornes, Abhandl. k.-k. Geol. Reichs. 1867, vol. iv. pl. 65, p. 418. 

Amussiopecten Burdigalensis, Sacco, Moil. Terz. Piemonte, &c., 1897, 
pt. 24, pl. 15. figs. 1-7, p. 53. 

Pecten (Amussiopecten) Burdigalensis, R. B. Newton, Geol. Mag. 1899, 
p. 209. 

This form is represented by a very much worn specimen, with 
rather imperfect margins. It shows wide and slightly raised coste ; 
and the surface is covered with numerous obsolete concentric 
striations. Dr. Fuchs’ Persian shell from the Siokuh mountain, 
P. placenta, appears to be closely related to it (Denkschr. K. 
Akad. Wiss. 1879, vol. xli. pl. 2, figs. 1,2, p. 104). 

The true Burdigalensis is a well-known Miocene species, being 
found in both the Helvetian and Burdigalian divisions of that 
period, and it forms the type of Sacco’s sub-genus Amussiopecten. 

Locality. From the beach of the island of Koyun Daghi. 

Formation. Miocene (Helvetian ?). ; 


Prcren (OoPpECTEN) RoTUNDATUS, Lamarck. (Plate 29. fig. 1.) 

Pecten rotundatus, Lamarck, Hist. Nat. Anim. sans Vert. 1819, vol. vi. 
pt. 1, p. 179; T. Fuchs, Denkschr. K. Akad. Wiss. 1879, vol. xli. pt. 2, 
pl. 2. figs. 1, 2, p. 104; Fontannes, Hist. Periode Tert. Rhone, 1880, 
vol. vi. pl. 1, fig. 1, p. 161. 

Oopecten rotundatus, Sacco, Moll. Terz. Piemonte, &c., 1897, pt. 24, 
pl. 15. figs. 14, 15, p. 54. 

Original specific description :—“ P. testa suborbiculart, utrin- 
que convexa; radiis 14 ad 16 distinctis, convexis, versus limbum 
planulatis. 

“ Largeur 75 millimetres. Il est moins bombé que le P. muléi- 
radiatus et que le P. rugosus.” (Lamarck.) 

A small block of a light brown calcareous sandstone obtained 
from a conglomerate contains remains of this species; the best 
preserved valve having a height of 60 millimetres and a length of 
rather more than 70 millim. 

It agrees with the original diagnosis as given above, and 
corresponds with Dr. T. Fuchs’ later interpretation of the same, 


444, MR. R. B. NEWTON ON THE 


this author having identified it among some Persian fossils 
obtained from the Siokuh mountains. Lamarck’s original speci- 
men is localised as from Vence, near Grasse in France, which 
Fuchs considers as belonging to the “Horner Schichten ” 
division of the Miocene; it has been well figured by Fontannes, 
who tabulates it as of Lower Helvetian age. This species forms 
the type of Sacco’s sub-genus Oopecten, and its range in Italy, 
according to the same author, is from the top of the Aquitanian 
into the Helvetian. 

Locality. Seir Hill, 6-8 miles west of Urmi. 

Formation. Miocene (Helvetian or Burdigalian ?). 


PECTEN (FLABELLIPECTEN) sp. indet. 

This determination refers to a fragmentary pectinoid shell 
exhibiting a very slightly convex lower valve, with about 16 
rounded radial costee separated by very wide interspaces, some 
obscure concentric striations on the ribs and grooves being dis- 
cernible in places. 

It is probably an example of P. Besser’, Andrejowski, from the 
Polish Miocene (Bull. Soc. Nat. Moscou, 1830, vol. 1. pl. 6. fig. 1, 
p- 108), but the intercostal grooves are mostly filled with matrix, 
and the specimen is otherwise so poorly preserved that a more 
definite identification is not possible. 

There is no doubt as to its belonging to Sacco’s sub-genus 
Flabellipecten, of which the type is P. flabelliformis, Brocchi, sp. 

Locality. From the beach of the island of Koyun Daghi. 

Formation. Miocene (Helvetian). 


Cuiamys (Aiguiprcten) Matvinm, Dubois de Montpéreu 
affin., H. Abich. (Plate 29. fig. 4.) 

Pecten Malvine, affin., Abich, Geol. Armenischen Hochlandes, 1882, 
pl. 8. fig. 7, p. 278. 

The collection contains five fragments of a Pectinoid shell which 
appear to be the equivalent of that form regarded by Dr. Abich 
as allied to P. Malvine of Dubois de Montpereux (Conch. Foss. 
Wolhyni-Podolien, 1881, pl. 8. figs. 2, 3, p. 71). The most com- 
plete example has from 18-20 narrow, rounded coste, extremely 
fine at the umbone, but afterwards widening, and separated by 
deep grooves; the lateral ribs having a tendency to curve. The 
entire surface is covered by fine, equidistant, transverse, imbri- 
cating or squamose striations. The shell is probably a oun 
form, as there is no indication of the ribs dichotomizing. 


MIOCENE MOLLUSCA OF LAKE URMI. 445 


The true P. Malvine possesses a greater number of ribs, which 
radiate in straight lines from the umbone without any apparent 
curvature on the lateral areas. The present Persian specimens 
are probably more closely allied to P. opercularis of Linneus, 
from which they are most difficult to separate after comparison 
with the Pliocene representatives of this species contained in the 
“Searles Wood” collection in the British Museum. ‘This par- 
ticular form of Pecten belongs to P. Fischer’s sub-genus Aqui- 
pecten, founded in 1886 on P. opercularis, Linneus, sp. 

Locality. Beach specimens from the island of Koyun Daghi; 
Dr. Abich’s specimens were obtained from the “ Urmia Plateau.” 

Formation. Miocene (Helvetian). 


CHLAMYS sp. indet. 

Two specimens have been determined as above. One consists 
of an impression of a valve in a small block of limestone. It 
possesses numerous radial, almost contiguous coste, which are 
divided up into twos and threes such as are typical of P. gloria- 
maris of Dubois from the Polish Miocene, but without the 
echinations on the ribs distinguishing the European form. It is 
of ovately oblong shape, and from the character of the costze 
clearly belongs to the genus Chlamys. 'The cther specimen has 
similar costal details, and was obtained from the island of 
Koyun Daghi. 

Locality. In a cream-coloured limestone with foraminifera and 
other small organisms, found on the road from Sujbulak to the 
Plain of Solduz; and from the cliff of the island of Koyun 
Daghi. 

Formation. Miocene (Helvetian). 


CuHLAMyYS sp. indet. (Plate 29. fig. 3.) 

This specimen has about 16 rounded costs divided by deep 
and wide grooves which are feebly marked by extremely fine 
oblique striations ; the costz show obscure indications of dividing 
besides being ornamented with numerous slightly elevated scabre 
or spines. There is only one example of this form in the collec- 
tion; it has both valves attached and is considerably worn so 
that the sculpture is difficult to define; its dimensions in milli- 
metres=height 42, length 38, and depth 18. The specimen is 
probably related either to P. gloria-maris, Dubois, P. opercu- 
laris, Linneus, sp., or to P. pusio, Linneus, all of which are 


446 MR. R. B. NEWTON ON THE 


included under the genus Ohlamys of Boltem, 1798 (type Ostrea 
varia, Linneus), by Sacco and other authorities. 

Locality. Neighbourhood of Urmi. 

Formation. Miocene (? Helvetian). 


Carpium sp. indet. (Plate 29. fig. 5.) 

This is a sandstone-cast of a valve belonging to the genus 
Cardium, measuring the same both in its height and length, viz. 
19 millimetres, and in depth 8 mm. Where preserved, the 
vertical striations on the surface are numerous and very close 
together; in the ventral area obscure lines of growth can be 
traced. 

Without more material it is difficult to speak of the nearest 
ally of a specimen of this description. It slightly resembles a 
Lower Tertiary form from Asia Minor figured by d’Archiac in 
Tchihatcheff’s ‘ Asie Mineure,’ 1866-69, pl. xi. figs. 6,:7, p. 162, 
under the name of Cardium indet., which however is longer than 
high and scarcely so convex as the Persian shell. 

Locality. Found in the same brown calcareous sandstone 
accompanying P. rotundatus at Seir, 6 to 8 miles west of Urmi. 

Hormation. Miocene (Helvetian or Burdigalian). 


Venus AcLAuRa, Brongniart. (Plate 30. figs. 1, 2.) 

Corbis? Aglaure, Brongniart, Mém. Séd. sup. Calc.-Trapp. Vicentin, 
1823, pl. 5. fig. 5, p. 80. 

Venus Aglaure, Hornes, Abhandl. k.-k. Geol. Reichs. 1861, vol. iv. 
pl. 14. figs. 1-4, p.122; P. Fischer, “ Faune Tert. Moyen,” in Tchihatcheft’s 
‘Asie Mineure,’ 1866-69, p. 290; Abich, Geol. Armenischen Hochlandes, 
1882, pl. 4. fig. 5, p. 278. 

Original diagnosis.—Transversim fere elliptica, ventricosa, 
cancellata, lamellis transversis crebris ad latera plicato-crispis, 
serratis. (Brongniart.) 

This well-known species is represented by a frais specimen 
with closed valves somewhat imperfect and worn. The original 
sculpture of the shell can be fairly seen in places, exhibiting the 
close characteristic concentric lamellose structure crossed by fine 
longitudinal striations. The specimen is of typical shape, being 
transversely elliptical, having a short rounded anterior region 
and an obliquely curved postero-dorsal margin; the lunule and 
escutcheon are a good deal obscured by matrix. 

Dimensions (in millimetres). Height=about 60; length=68; 
depth =38. 


MIOCENE MOLLUSCA OF LAKE URMI. 4AT 


Brongniart’s original example of this species was obtained from 
the Oligocene (Tongrian) rocks of Castel Gomberto of North 
Italy (this correlation being according to Lapparent, ‘ Traité 
de Géologie,’ 1893, p. 1292). Under the name of V. granosa it 
has been recorded from the Miocene (Gaj series) deposits of 
Cutch, India (J. de C. Sowerby) ; it occurs in the Vienna Basin 
at Gauderndorf &c. (Hornes), in the Lower Miocene or Burdiga- 
lian stage ; “‘ Urmia Plateau’ (Abich); Asia Minor (P. Fischer); 
Saucats; Turin, &c. 

Locality. Guverjin Kala, north end of Lake Urmi. Specimen 
having a reddish-brown colour externally; the matrix within 
appearing to be a grey, marly, calcareous rock. (Collected by 
the Rev. C. Labaree.) 


Formation. Miocene (Burdigalian or Aquitanian). 


MERETRIX PERSIENSIS, n. sp. (Plate 30. figs. 3-5.) 


Shell transversely ovate, equilateral, anterior end truncate 
and short, posterior extremity rounded and produced; valves 
convex, dorsal margin rounded and long, surface ornamented 
with concentric sulcations ; beaks very anterior, incurved, having 
beneath a small, fairly deep, cordiform lunule; escutcheon 
lanceolate. 

Dimensions (in millimetres). Height=28 ; length=45; depth 
= 28), 

This species is represented by one specimen having both its 
valves united. It appears to differ from other forms of this 
genus in its transverse shape, the very anterior position of the 
umbones (which come beyond the shell-margins below), and 
the very convex rounded valves, which slope obliquely to the 
posterior and ventral margins. 

The specimen is quite free from external matrix, its lithological 
aspect being similar to that observed in the forms of Venus 
Aglaure and Meretria like inerassata. 

Locality. Guverchin Kala. (Collected by the Rev. C. Labaree.) 

Formation. Miocene (Burdigalian or Aquitanian). 


Mererrix allied to Incrassata, J. Sowerby. (Plate 30. 
figs. 6 & 7.) 

Venus incrassata, J. Sowerby, Mineral Conchology, 1817, vol. ii. pl, 155. 
figs. 1, 2. 

Cytherea merassata, Deshayes, Dese. Cog. Foss. Paris, 1825, vol. i. 


A448 MR. R. B. NEWTON ON THE 


pl. 22. figs. 1-3, p. 186; Abich, Geol. Armenischen Hochlandes, 1882, 
pl. 2. fig. 8, p. 289. 

Meretrix incrassata, R. B. Newton, Syst. List British Oligocene and 
Eocene Mollusca, 1891, p. 64. 

A single specimen, consisting of a cast showing both valves, 
appears to have the form and thickness of Meretrix inerassata, as 
figured by Deshayes from the Paris Basin Oligocene; and, in 
addition, it is very similar to Abich’s figure of this species, which 
depicts a specimen obtained from the Upper Hocene of Achalzik, 
in the northern part of the Armenian highlands. Our cast also 
shows obscure indications of a deep pallial sinus on the left valve. 

Its dimensions are:—Height=35 millimetres; length=29 ; 
depth=23. 

Locality. Guverchin Kala; of the same colour and aspect 
lithologically as observed in the Venws Aglaure and Meretri« 
Persiensis. (Collected by the Rev. C. Labaree.) 

Formation. Miocene (Burdigalian or Aquitanian). 


CarpiTa sp. indet. (Plate 29. figs. 6, 7.) 

This determination refers to a single specimen having both 
valves attached. It is inequilateral, somewhat depressed, and 
possesses about 20 radial, rather arched cost, which are orna- 
mented with a bead-like, squamulose structure, the prominences 
of which have been eroded by aqueous action. The anterior 
margin is short and rounded, whilst posteriorly it is oblique and 
produced ; lunule small and cordate. 

The specimen is in bad preservation, with the umbonal surfaces 
fractured. It appears to represent a new species, but without 
more material the present reference will suffice, although it may 
be said to bear a curious resemblance to the figure of Oardita 
mutabilis of d’Archiac & Haime (‘ Descr. Anim. Foss. Nummu- 
litique de l’Inde,’ 1854, vol. ii. pl. 21. figs. 83-6, p. 256), from the 
Eocene of Subathoo, North-west India. 

Dimensions (in millimetres). Height=16; length=20; depth 
=10. 

Locality. Beach of the island of Koyun Daghi. 

Formation. Miocene (Helvetian). 


GASTEROPODA. 


Stromevs like Bonen, Brongniart. 
This specimen consists of a fragmentary water-worn cast which 
appears to be referable to this species. Only the two latest 


MIOCENE MOLLUSCA OF LAKE URMI. 449 


whorls are present, the spire being absent. In the upper part 
of the last whorl there are undoubted indications of the thick 
tubercles which ornament that region in this species. 

8. Bonelli was originally described by Brongniart from the 
Miocene (Helvetian) beds of Turin (Mém. Terr. Séd. Cale.-Trapp. 
Vicentin, 1823, pl. 6. fig. 6, sp. 74). 

Locality. In a grey marly rock from the beach of the island of 
Koyun Daghi. 

Formation. Miocene (Helvetian). 


Conus sp. indet. 

Two casts of this genus are present in the collection, but 
they are much broken, water-worn examples, and therefore not 
determinable. 

Locality. In a grey marly rock from the beach of the island 
of Koyun Daghi. 

Formation. Miocene (Helvetian). 


PYRULA CINGULATA (Bronn MS.), Hornes. (Plate 30. figs. 8, 9.) 
Pyrula clathrata, Lamarck, Hist. Nat. Anim. sans Vert. 1822, vol. vii. 
. 141. 
fi Pyrula reticulata, Hornes, Abhandl. k.-k. Geol. Reichs. 1853, vol. iii. 
pl. 28. figs. 1-3, p. 268, non Lamarck, 1822. 

Pyrula cingulata, Hornes, Abhandl. k.-k. Geol. Reichs. 1856, vol. iii. 
p- 676. 

Represented by a brown-coloured natural cast exhibiting the 
shape and sculpture of this species. It has a long oval aperture, 
an acute and outwardly-curved labrum, a moderately excavated 
columella, and a very short .convex spire. The large body- 
whorl is ornamented with broad and rather distant transverse 
bands, these being crossed obscurely by finer lines. The ante- 
rior canal is absent, most probably through fracture though the 
end is now rounded from wear. The broad spiral bands seem to 
separate this species from P. condita, Brongniart. 

Dimensions (in millimetres). Length=50; diameter=34. 

This species, under the name of P. clathrata, Lamarck, was 
first known from the Miocene of the Touraine district of France, 
although mistaken by that author for a Paris Basin form. It 
has been recorded also from Italy (Bronn); Poland (Hichwald) ; 
Rhone Basin (Matheron); Portugal (G. B. Sowerby); Vienna 
Basin (Hornes), &e. 

Locality. Guverchin Kala. (Collected by the Rev. C. Labaree.) 

Formation. Miocene (Burdigalian or Aquitanian). 

LINN. JOURN.—ZOOLOGY, VOL. XXVII. 34 


450 MR. R. B. NEWTON ON THE 


Cassis sp. indet. (Plate 30. figs. 10, 11.) 

This specimen is an imperfect cast with a general resemblance 
to O. subharpeformis, dArchiac & Haime (Descr. Anim. Foss. 
Numm. 1’Inde, 1854, vol. 11. pl. 31. fig. 6, p. 317), from the Eocene 
rocks of India; but differing from that form in-being much 
shorter and broader, in the non-bifureate character of the vertical 
ribs, and the presence of some obscure concentric markings on 
the principal whorl. The aperture is narrowly elliptical- and 
oblique ; spire only slightly elevated. , 

Dimensions (in millimetres). Length=27; diameter=27. 

Locality. Guverchin Kala. Of similar lithological appearance 
to the specimen of Pyrula cingulata. (Collected by the Rey. C. 
Labaree.) 

Formation. Miocene (Burdigalian or Aquitanian). 


LatIRrus cRISPUS, Borson. (Plate 29. fig. 8.) 

Fusus crispus, Borson, Mem. R. Accad. Sci. Torino, 1821, vol. xxvi. 
p. 817; Deshayes, Appendix to Lyell’s ‘ Principles of Geology,’ 1838, 
pl. 1. fig. 8, p. 80; Michelotti, Desc. Foss. Mioe. l’Italie Sept., 1847, pl. 9. 
figs. 17, 18, p. 272; Hornes, Abhandl. k.-k. Geol. Reichs. 1853, vol. iii. 
pl. 32. fig. 3, p. 291. 

Original diagnosis.— Testa costata transversim sulcata ; plicis 
longitudinalibus fornicatis ; labio intus sulcato. (Borson.) 

A somewhat water-worn example is all that represents this 
species. It shows the characteristic longitudinal, elevated and 
rounded costulations, of which there are about eight on the 
principal whorl, separated by deep corresponding sulcations ; the 
transverse sculpture consisting of numerous regular striations. 
The anterior canal is broken and the spire is incomplete ; other- 
wise it preserves the fusoid and elongate shape of this form. 

Dimensions (in millimetres). Length=27 ; diameter=15. 

Borson’s original example is localised as Piedmont, and belongs 
to the Tortonian stage of the Miocene, Hornes’ specimen from 
the Vienna Basin (Mollersdorf) being of similar horizon. 

Locality. From the beach of Koyun Daghi island. 

Formation. Miocene (Tortonian) The specimen has a reddish 
tinge similar to what is present in some of the examples of 
Alectryonia Virleti, and probably came from the same bed. 


TurRRITELLA ARCHIMEDIS, Brongniart. (Plate 29. fig. 11.) 
Turritella Archimedis, Brongniart, Mém. Terr. Séd. sup. Cale.-Trapp. 
Vicentin, 1823, pl. 2. fig. 8, p. 55; Hornes, Abhandl. k.-k. Geol. Reichs. 


Linn. Soc. Journ. Zoon. Vou. XXVIII Pu. 29. 


Bullen Newton. 


S.Imp. 


Mantern Bro 


NOLOKCIE NUE. GIB0e IS ERO LARS) TUNEUNTI 


J. Green del et lth. 


Linn. Soc.Journn. Zoou .Vor.XXVI.Pu.30. 


Bullen Newton . 


Mintern Bros.imp. 


J.Green del.et lith. 


WOOCE INI, Slave ib, S NEO IvANAT) URN 


MIOCENE MOLLUSCA OF LAKE URMI. 451 


1855, vol. iii. pl. 43. figs. 13, 14, p. 424; Abich, Geol. Armenischen 
Hochlandes, 1882, pl. 8. fig. 14, p. 279. 

This species is represented by a fragment exhibiting three 
whorls of the upper part of the spire, with depressed, spirally- 
striated surfaces and margined by prominent bicarinations. 

The species occurs in Northern Italy (Brongniart); Vienna 
Basin (Hornes); Egypt (Fuchs); “‘Urmia Plateau,” north- 
western Persia (Abich), &e. 

Locality. Koyun Daghi (beach specimen). 

Formation. Miocene (Helvetian). 


TURRITELLA GRADATA, Hornes. Var., Abich. (Plate 29. fig. 9.) 

Turritella gradata, var., Abich, Geol. Armenischen Hochlandes, 1882, 
pl. 8. fig. 12, p. 280. 

This consists of a water-worn fragment showing three whorls 
with depressed surfaces, the spiral sculpture being nearly obso- 
lete, but having a prominent elevated keel at the base of each. 
Abich regarded his shell as a variety of Hornes’ 7. gradata from 
the Vienna Miocene (Abhandl. k.-k. Geol. Reichs. 1855, vol. ii. 
pl. 48. fig. 3, p. 420), with which it seems to be closely related ; 
his specimen was from the “ Urmia Plateau.” 

Locality. Beach of the island of Koyun Daghi. 

Formation. Miocene (Helvetian). 


TURRITELLA ROTIFREA, Deshayes, affin. Abich. (Plate 29. 
fig. 10.) 

Turritella rotifera aftin., Abich, Geol. Armenischen Hochlandes, 1882, 
pl. 8. fig. 11, p. 279. 

This is a fragmentary specimen which uudoubtedly belongs to 
the form determined by Abich as above. On account of the 
turreted whorls, their spiral sculpture and the equidistant pro- 
minent keels, Abich regarded this form as related to 7. rotifera 
of Deshayes (Desc. Coq. Foss. Env. Paris, 1832, vol. ii. pl. 40. 
figs. 20, 21, p. 275), from the Paris Basin Eocene. Abich’s speci- 
men was found on the “ Urmia Plateau.” 

Locality. Krom the beach of the island of Koyun Daghi. 

Formation. Miocene (Helvetian), 


EXPLANATION OF THE PLATES. 
Puate 29. 


Fig. 1. Pecten (Oopecten) rotundatus. Seir Hill. 
2. Pecten Beudanti. Neighbourhood of Urmi. 
3. Chlamys sp. indet. Neighbourhood of Urmi. 


452 MR. R. B. NEWTON ON A 


Fig. 4. Chlamys (Aiquipecten) Malvine aff. Koyun Daghi. 
5. Cardium sp. indet. Seir Hill. 
6, 7. Cardita sp. indet. Koyun Daghi. 
8. Latirus crispus. Koyun Dagh1. 
9. Turritella gradata, var. Koyun Daghi. 
10. Turritella rotifera affin. Koyun Daghi. 
Il. Turritella Archimedis. Koyun Daghi. 


Puate 30. 


Figs. 1,2. Venus Aglaure. (Fig. 2=sculptur emagnified.) Guverchin Kala. 
3,4,5. Meretri« persiensis, n. sp. Guverchin Kala. 
6,7. Meretriz allied to inerassata. Guverchin Kala. 
8,9. Pyrula cingulata. Guverchin Kala. 
10,11. Cassis sp. Guverchin Kala. 


The figures on both Plates are drawn natural size, with the exception of fig. 2 
on Plate 30. 


NOTE ON A PALAOZOIC LIMESTONE. 


By R. Buitzew Newroy, F.G-S8., of the British Museum 
(Natural History). 


Amone Mr. R. T. Giinther’s geological specimens from North- 
western Persia is a fragment of dark slate-coloured limestone, 
highly crystalline, and intersected with numerous calcite veins, 
which was obtained from the island of Shazalan on Lake Urmi. 

A microscopical examination of this rock proves that its 
structure is largely made up of foraminiferal remains which, with 
the kind assistance of Mr. Frederick Chapman, have been deter- 
mined as Hndothyra Bowmannt, Phillips, Valvulina bulloides, 
Brady, and Nodosaria radicula, Linneus. 

Generically, the most interesting of these forms is Endothyra, 
being essentially Carboniferous, the other genera having a far 
wider range in geological time. 

The nearest habitat to Shazalan which has hitherto yielded 
E. Bowmanni appears to be Sloboda, in the province of Toula, 
European Russia, reported some years ago by V. von Moller *. 

So far as can be ascertained, there is only one species of 
Paleozoic foraminifera yet recorded from this region, viz., 


* Mouurr, V. v.—‘ Die spiral-gewundenen Foraminiferen des Russischen 
Kohlenkalks,” Mém. Ac. Imp. Sci. St. Pétersbourg, 1878, ser. 7, vol. xxv. no. 9, 
p- 96, pl. iv. fig. 3, & pl. xii. fig. 2. 


PALEOZOIC LIMESTONE FROM LAKE URMI. 453 
Fusulina spherica, described and figured as a new form by 
Abich * in 1859 from the Bergkalk (= Carboniferous) formation 
of the Armenian and Persian plateaux, which shows resemblances 
to Fusulina cylindrica of Fischer, a characteristic species of 
European Russia. 

In reference to the distribution of the older rocks in this area, 
attention may be called to the geological plan of the islands on 
Lake Urmi published by Abich + in 1882, which is of considerable 
value. By this we can trace the extension of these ancient 
deposits through Koyun Daghi and Isbir to the western shores 
of Lake Urmi, where in each case they form the fundamental 
structure on which repose the various beds of the Miocene 
system. Mr. Gunther states, in some manuscript notes on this 
area, that a similar limestone as that occurring at Shazalan is to 
be seen at the northern end of Koyun Daghi, and possibly also 
on the neighbouring island of Arzu. 

There appear to be no references in literature to the geology 
of this small island of Shazalan ; so that the example of limestone 
brought home from that place by Mr. Giinther is of great 
interest, not only in adding to the list of Paleozoic Foraminifera 
from this neighbourhood and in establishing as Carboniferous 
the age of the rock in which they occur, but in extending farther 
northwards the distribution of this Paleozoic formation, so 
important an addition to the geological history of Lake Urmi. 


* Axicu, H.—“ Vergleichende geol. Grundziige der Kaukasusischen, Arme- 
nischen und Nordpersischen Gebirge,” Mém. Ac. Imp. Sci. St. Pétersbourg, 
1859, vol. vii. pl. 3. fig. 13, p. 489. 

t ‘Geologie des Armenischen Hochlandes,’ 1882, p. 275. 


LINN. JOURN.—ZOOLOGY, VOL. XXVitl. 35 


454 DR. W. G. RIDEWOOD ON THE HYOBRANCHIAL 


On the Hyobranchial Skeleton and Larynx of the new Aglossal 
Toad, Hymenochirus Boettgert. By W. G. RipEwoop, 
D.Se., F.L.S., Lecturer on Biology at St. Mary’s Hospital 
Medical School, London. 


[Read 2nd November, 1899. ] 
(Pave 31.) 


XeEvorus and Pipa have for many years been to herpetologists a 
fruitful source of discussion, by reason of the structural peculi- 
arities which mark them off so sharply from the other Anura, and 
also on account of the great differences which exist between them- 
selves. They are isolated types which fail to show any marked 
affinity with any of the Phaneroglossal Anura, and the relatively 
few features in which they resemble one another some writers 
would explain away as due to convergence and adaptation to 
similar conditions of life. The discovery of a third genus of 
Aglossal Toads marks the beginning of a new era in the history 
of this little suborder; and it is to Hymenochirus that attention 
will now be turned with the object of ascertaining how far the 
Anura Aglossa constitute a natural assemblage. 

A single specimen of this new form was discovered in 1896 at 
Ituri, in German East Africa, and was described by Tornier as 
Xenopus Boettgert (7. p. 163). Several specimens have since 
been collected from the Benito river, in the French Congo; and 
the skeletal and other characters detailed by Boulenger in the 
present year (2) go to show that this writer was fully justified in 
the claim which he put forward in 1896 (1) that Tornier’s new 
species of Xenopus was entitled to generic rank. 

Having already made an exhaustive study of the hy obranchial 
skeleton and larynx of Xenopus and Pipa (4), I applied to 
Mr. Boulenger for permission to examine these parts in one of 
the specimens of Hymenochirus belonging to the Natural History 
Museum. The request was graciously acceded to, and a male and 
a female specimen were placed at my disposal. For these my 
thanks are hereby gratefully tendered. 

There are certain features in the hyobranchial and laryngeal 
skeleton of Xenopus and Pipa which may be regarded, as dis- 
tinetive, occurring in both of these genera and not in any of the 
Phaneroglossal Anura. The features in question are :— 


SKELETON AND LARYNX OF HYMENOCHIRUS BOETTGERI. 455 


The presence of a hyoglossal foramen, transmitting the reduced 
hyoglossal muscle, and originating by a secondary union of 
the hyoidean cornua. 

The presence of a pair of large backwardly directed carti- 
laginous wings, developed from the branchial skeleton of 
the larva. 

The large size and complexity of the larynx, and the incor- 
poration of the thyrohyal bones into the laryngeal apparatus. 

The absence of vocal cords. 

Tt is a singular fact that all these five characters are exhibited 
by Hymenochirus. 

Tt does not follow, however, that the hyobranchial skeleton of 
Hymenochirus bears, on the whole, any close resemblance either 
to that of Xenopus or that of Pipa. In certain features the 
hyobranchial skeleton of Hymenochirus is unique. The hyoidean 
cornua, for instance, are very large and strongly ossified. Ossi- 
fication of the hyoidean cornua, though not uncommon in the 
Urodela, has hitherto been unknown to occurin the Anura. That 
Boulenger (2), describing a dried skeleton in which the larynx 
had not been preserved, should have mistaken these bones of 
Hymenochirus for the thyrohyals is thus quite excusable, seeing 
that in all other forms of Anura the only strongly ossified parts 
of the hyobranchial skeleton are the thyrohyals. 

The hyoid bones (Pl. 81. fig. 1, ch) are thinnest at about one- 
third of their length from the posterior end, and are flattened in 
the horizontal plane anteriorly. The hinder part slopes outward 
and upward; and the swollen extremity, which is not tipped 
with cartilage, is bound by a short, strong ligament to the 
inferior surface of the large otic capsule. Projecting hori- 
zontally from the mesial surface of each bone is a thin lamella of 
cartilage, with a convex free border, which is evidently the 
counterpart of the similarly placed lamella of the unossified 
hyoidean cornua of Xenopus (see 4. pl. 8. fig. 1). Anteriorly the 
flattened hyoid bones terminate in epiphysial plates which are 
cartilaginous in the female, but exhibit an irregular endochondral 
ossification in the male. 

Lying antero-internally to these latter is a median element, 
strongly ossified in both sexes, which is unique among adult 
Anura (fig. 1, ca). It is roughly peutagonal in shape, and from 
its relations to the surrounding parts might be regarded as the 
equivalent of the basihyal of Fishes. It is most important to 

35* 


456 DR. W. G. RIDEWOOD ON THE HYOBRANCHIAL 


note that it is not homologous with the median cartilage which 
is lodged between the two hyoidean cornua in the larval Xenopus 
(4. pl. 11. figs. 1 and 2, 6h); for the latter, by coalescence with 
surrounding parts, develops into a plate of cartilage which is 
behind the hyoglossal foramen. The median bone of Hymeno- 
chirus would correspond rather with the anterior of the two 
cartilages which occupy the median line in the hyobranchial 
skeleton of the larval <Alytes (see 6. figs. 1 and 2, ca). It is 
possible that this anterior copula of the Discoglossid larva may 
later prove to be the true basihyal element; but, for reasons 
already stated (5. p. 583), it is preferable, for the present, to 
apply the term basihyal to the posterior copula, or to avoid the 
use of the word altogether, until further research has satis- 
factorily demonstrated the homologies of the constituent parts 
of the hyoid apparatus of the Anura. 

The hyoidean skeleton is completed in front by two plates of 
cartilage, each exhibiting a small area of calcification (pa). These 
plates are doubtless the equivalents of the processus anteriores of 
the common Frog (see 5. fig. 11, pa), and represent also the thin 
lamelle of cartilage which project from the anterior edges of the 
hyoidean cornua in Xenopus (see 4. pl. 8. fig. 1). 

The front part of the hyobranchial skeleton is quite detached 
from the hinder part, and the relative positions of the two parts 
to one another and to the mandible and larynx are preserved in 
fig. 1, Plate 31. The front part undoubtedly belongs to the 
hyoidean arch exclusively, but it is possible that in the hinder 
part there is also some cartilage of hyoidean origin. This pro- 
position is based upon the fact that in Xenopus and Pipa 
the hyoglossal foramen is formed by the coalescence of the 
right and left hyoidean cornua in front of the hyoglossal sinus (4). 
The hyoglossal foramen of Hymenochirus (figs. 1 and 3, h) is 
small in size, but its identity is unmistakable, since it transmits 
a reduced hyoglossal muscle, running from the ventral surface 
of the larynx to the pharyngeal mucous membrane in front of 
the glottis. The tract of cartilage which lies anteriorly to this 
foramen (ch') may, therefore, be considered as a hyoid derivative. 
It corresponds with the pointed rod of cartilage which in Pipa 
projects in front of the hyoglossal foramen (4. pl. 9. fig. 1, ch’). 

In the presence of hyoidean cornua Hymenochirus differs from 
Pipa and resembles Xenopus. In fact, one may say that no 
porticn of the front half of the hyobranchial skeleton of Hymeno- 


SKELETON AND LARYNX OF HYMENOCHIRUS BOETTGERI. 457 


chirus is represented in the adult Pipa. The other half, how- 
ever, will bear a detailed comparison with the entire hyobranchial 
skeleton of the adult Pipa. The great alary cartilages (a), so 
characteristic of the Aglossa, slope outward and backward, and 
terminate in inwardly directed expansions, which partially 
overlie the thyrohyals. They extend some distance behind the 
posterior epiphyses of the thyrohyals in the female, but not 
in the male (cf. figs. 1 and 3). The external geniohyoid muscle 
is inserted about midway between the hyoglossal foramen and 
the postero-lateral extremity of the alary cartilage (fig. 1, ge), 
whereas in Xenopus it is inserted much nearer the foramen, and 
in Pipa quite close to the postero-lateral border of the cartilage. 
Lying antero-laterally to the insertion of this muscle is a thin 
lamellar extension of the cartilage, which finds its exact counter- 
part in Xenopus, but not in Pipa. The antero-lateral processes 
of the basal plate of Xenopus (4. pl. 8, fig. 1, ap) are not repre- 
sented, unless they are included in the semicircular tract of 
cartilage which lies in front of the hyoglossal foramen. The alary 
portion of the hyobranchial skeleton differs somewhat in shape 
in the two specimens examined, but the paucity of material 
precludes one from deciding whether these are normal sexual 
differences. 

In both sexes of Hymenochirus the thyrohyal bones have the 
form of tapering rods, as they have in the female Xenopus; and 
they are brought into intimate relation with the cricoid cartilage 
of the larynx, as is characteristic of both Xenopus and Pipa. 
The posterior ends terminate in large epiphysial cartilages ; the 
anterior ends converge, and are connected with one another by a 
small tract of cartilage, which in the female is continuous with 
the transverse bar of cartilage running behind the byoglossal 
foramen, but which is bound to the latter by a short stout lga- 
ment in the male. These last relations are exactly those obtaining 
in the two sexes of Pipa (see 4. pl. 9, fig. 1, 2, and fig. 5, 2’). 

-The larynx is considerably smaller in the female than in the 
male, and the thyrohyal bones are shorter, more slender, and set 
at a wider angle. The floor or ventral wall of the larynx of the 
female (fig. 3) is largely composed of membrane. It is supported 
by a horizontal ring of cartilage, from which there project 
postero-laterally a pair of slender cartilaginous bars. These 
expand at their extremities into the bronchial cartilages (br), 
and are confluent with the posterior epiphyses of the thyrohyals. 


458 DR. W. G. RIDEWOOD ON THE HYOBRANCHIAL 


The ring of cartilage is produced into a blunt point in front, 
while from the sides there rise vertically upwards, internal to the 
thyrohyal bones, but quite free from them, a pair of cartilagmous 
bars, which end at the sides of the arytenoid cartilages on the 
upper surface of the larynx (fig. 4, 67). The dorsal extremities 
of these cartilages appear to be the equivalents of the “ Scheu- 
klappenartige Fortsitze ” described by Henle (8) in the female 
Xenopus. 

As seen from above, the larynx bears a striking resemblance 
to that of the young Pipa (see 4. pl. 11. fig. 11). The dorsal 
part of the cricoid cartilage has the form of an arched band, 
running transversely between the posterior epiphyses of the thyro- 
hyals, and fused with them at its extremities. The arytenoids 
are simple, concavo-convex cartilages, constructed upon the type 
normal for the Phaneroglossal Anura. They each exhibit a small 
centre of ossification at the point where the tendinous extremity 
of the laryngeal dilator is inserted. 

There are no vocal cords in either sex, neither are there 
bronchial tubes. The wall of the lung is quite smooth, and 
exhibits no sacculation. It is as strongly vascular in the parts 
connected with the larynx as over the general surface; and it is 
worthy of notice that the bronchial cartilages of Hymenochirus 
are not more extensively developed than in such a phanero- 
glossal form as Bombinator (see 4. pl. 11. fig. 18, br). As in Pipa 
and Xenopus, the lung is bound to the abdominal wall by a 
stout pleural fold, which does not quite reach to the posterior 
extremity. 

In the male, the floor or ventral wall of the larynx is composed, 
entirely of cartilage, but the cartilaginous lamelle projecting 
from the inner borders of the thyrohyal bones (fig. 1, ¢’) are not 
confluert with the median cricoid. The latter terminates 
anteriorly in a sharp point, concealed in a ventral view by the 
meeting of the thyrohyal lamelle just mentioned. The vertical 
bars of the cricoid cartilage, which in the female lie internal to 
the thyrohyal bones, are also present in the male, and are similarly 
placed; but they can only be seen by cutting open the larynx. 
Their dorsal extremities are not free, but are fused with the 
rocfing part of the ericoid cartilage. The cricoid cartilage 
extends much farther forward on the upper surface of the larynx 
than it does in the female. It conceals the anterior part of the 
thyrohyals, and is confluent in front with the tract of cartilage 


SKELETON AND LARYNX OF HYMENOCHIRUS BOETTGERI. 459 


which has been regarded above as the united anterior epiphyses 
of the thyrohyals. The actual roof, z. e. the part of the cricoid 
behind the arytenoids, is saddle-shaped, and differs from that 
of the female in being longer, than broad, and in the greater 
thickness of its cartilage. The arytenoids are completely ossified, 
no cartilage remaining; and the glottis appears in the prepared 
skeleton as a rectangular hole. The sides of the glottis are 
formed by thick folds of mucous membrance occupying the 
right and left halves of this rectangle. The arytenoids are 
not confluent with one another as they are in the male Xenopus, 
nor are they backwardly produced into the interior of the larynx 
as in the male Pipa. 

The larynx of both sexes is thus of simpler construction than 
those of the two sexes of Pipa and Xenopus; but in its hyo- 
branchial skeleton Hymenochirus exhibits a most conflicting set 
of features. It is quite impossible to conclude, from evidence 
derived from this source, whether Hymenochirus is more 
primitive than the two previously known aglossal toads, or with 
which of these forms it is the more closely allied. Inconclusive, 
however, as the results are for this purpose, they constitute a 
strong argument in support of the view that the Aglossa are a 
natural group, and that the three genera now composing it have 
had a common ancestry. The discovery of Hymenochirus binds 
Pipa and Xenopus more closely together than before. 


PAPERS QUOTED. 


1. Bounrnerr, G. A.—“ A new Genus of Aclossal Batrachians.” 
Ann. & Mag. Nat. Hist. [6] xviii. 1896, p. 420. 

2. Boutenerr, G. A.—“ On Hymenochirus, a new Type of 
Aglossal Batrachians.” Ann. & Mag. Nat. Hist. [7] iv. 
1899, pp. 122-125. 

3. Hentz, D. J.—Beschreibung des Kehlkopfs. Leipzig, 1839. 

4. RipEwoop, W. G.—“ On the Structure and Development of 
the Hyobranchial Skeleton and Larynx in Xenopus and 
Pipa.” Linn. Soc. Journ., Zool. xxvi. 1897, pp. 53-128. 

5. RrpEwoop, W. G.—‘‘ On the Structure and Development of 
the Hyobranchial Skeleton of the Parsley-Frog (Pelodytes 
punctatus). Proce. Zool. Soc. 1897, pp. 577-595. 

6. Rrp—Ewoop, W. G.—“On the Development of the Hyo- 
branchial Skeleton of the Midwife-Toad (Alytes obstetri- 
cans).” Proc. Zool. Soc. 1898, pp. 4-12. 

7. TorniterR, G.—Kriechthiere Deutsch-Ost-Afrikas. Berlin, 
1897 (1896). 


460 DR. W. G. RIDEWOOD ON THE 


EXPLANATION OF PLATE 31. 


Fig. 1. Hymenochirus Boettgeri, male. Hyobranchial, laryngeal, and man- 
dibular skeleton, ventral view. (x 6.) 
. Laryngeal skeleton of same, dorsal view. 
3. Hymenochirus Boettgeri, female. Laryngeal skeleton, and hinder part 
of hyobranchial skeleton, ventral view. (x 6.) 
4. Laryngeal skeleton of same, dorsal view. 
5. Carpal skeleton of same, dorsal view. (xX 12; see page 462.) 


i) 


Reference Letters. 


So far as has been possible, the lettering used in Plates 8 and 9, Linn. Soc- 
Journ., Zool., xxvi. 1897, has been adopted in the present instance. 


a. Ala, or great wing of the hyobranchial skeleton. 
ar, Arytenoid. 
as, Angulosplenial bone. 
bl. Dorsal extremity of vertical cricoid rod. 
br. Bronchial cartilage. 
ce. Dorsal or roofing portion of the cricoid cartilage. 
ce’, Antero-ventral portion of cricoid. 
e''. Postero-ventral portion of cricoid. 
ca. Ossified copula (cf. 6. pl. 2. figs. 1 & 2, ca). 
ch, Ceratohyal or hyoidean cornu. 
ch'. Median cartilage formed by the secondary union of the byoidean cornua. 
d. Dentary bone. 
ep. Posterior epiphysis of the thyrohyal. 
ge. Area of insertion of the m. geniohyoideus externus. 
gt. Position of the glottis. 
h. Hyoglossal foramen. 
z. Isthmus between the anterior and posterior portions of the branchial 
skeleton. 
. Ligament equivalent to the cartilaginous isthmus of the female. 
pa. Anterior plate of the hyoidean skeleton (cf. 5. pl. 35. fig. 11, pa). 
pm. Outline of anterior part of lung. 
t. Thyrohyal bone. 
t'. Lamella of cartilage projecting from the thyrohyal bone in the male. 


Note on the Carpus of the new Aglossal Toad, Hymenochirus 
Boettgert. By W. G. Rroewoop, D.Sc., F.L.S. 


(PiatE 31. fig. 5.) 


WueEreEas the carpus of Xenopus is comparatively normal, and 
does not differ in any very essential respect from that of Disco- 
glossus or Bombinator, the carpus of Pipa is remarkable from 


CARPUS OF HYMENOCHIRUS BOETTGERI. 461 


the fact that the ulna is separated from the fifth metacarpal by 
a single bone, and not two bones as in all other Anurous 
Amphibians. 

While investigating the hyobranchial skeleton of Hymeno- 
chirus, an account of which appears in the pages immediately 
preceding, it occurred to me that it would be of considerable 
interest to determine, while the material was still in my hands, 
whether the carpus of this new genus could throw any light upon 
the question of the affinities of this form with the two previously 
known Aglossal Toads. 

An examination of the carpus of Hymenochirus disclosed the 
fact that in this genus, as in Pipa, a single bone intervenes 
between the uJna and the fifth metacarpal. In viewof the great 
variation to which the carpus of the Anura is subject, it is perhaps 
unwise to attach much importance to this fact; but it is certainly 
remarkable that the Hymenochirus of Africa should thus depart 
from the generalized type of carpus found in Xenopus (African), 
and should approach the hitherto unique variety exemplified by 
the American Pipa. 

The carpus of Hymenochirus (fig. 5, Pl. 31) consists of five 
elements, not including the radial sesamoid (s). This last occurs 
also in both Pipa and Xenopus. Ina joint paper on the Anuran 
Carpus and Tarsus by Prof. G. B. Howes and myself, the ventral 
surface of the carpus of Xenopus and Pipa is unfortunately 
figured as the dorsal surface, and the radial sesamoid is stated 
as occurring ventrally to the lunatum *—nmistakes which were 
pointed out by Jungersen t in 1891. The sesamoid in question 
is dorsal in position in all three genera, and is lenticular in shape. 
In Hymenochirus it occupies a more proximal position than in 
Pipa and Xenopus, and lies over the epiphysis of the radius,— 
a fact somewhat destructive to the theory put forward by Emery ¢ 
that this element, in Pipa, is an “intermedium carpi.” Since the 
dorsal radial sesamoid has been shown by Zwick § to occur also 
m the Frog and Toad, it cannot be regarded as distinctive of the 
Aglossa. 

The largest bone of the carpus is that which extends from the 
ulna to the fifth metacarpal (pk). It represents the pyramidale 


* Proc. Zool. Soc. Lond. 1888, pl. vii. figs. 1, 2 & 4, and p. 162. 
t+ Ann. Mag. Nat. Hist. [6] viii. pp. 193-206. 

t Ricerche Lab. Anat. Norm. Roma, iv. 1894, p. 10. 

§ Zeitschr. fiir wiss. Zool. lxiii. 1898, p. 102. 


462 CARPUS OF HYMENOCHIRUS BOETTGERI. 


or ulnare, confluent with the post-axial centrale, as in Pipa. The 
distal carpal of the fifth digit is probably absent and not included 
in this bone. The fourth carpal is free. Seen from above, it 
articulates with a part of the third metacarpal as well as the 
fourth, but an examination of the palmar surface of the carpus 
suffices to show that this element belongs exclusively to the 
fourth digit. 

In Ppa the third carpal is free, while that of the second digit 
is (presumably) fused with the naviculare or pre-axial centrale, 
but there is no distinct carpal to either the second or third digit 
in Hymenochirus. Whether the carpals of these digits have 
dwindJed away and left no traces, or whether they have fused 
with the naviculare, it is impossible to say. The appearances 
rather suggest that the latter is the more correct interpretation. 

The lunatum (J) is normal, and calls for no comment. 

The pollex is represented by a single lenticular bone, probably 
earpale 1, which lies on the ventral surface of the naviculare. It 
is concealed by the latter in a dorsal view, but its position is 
indicated in the figure by a dotted line (jo. 1). 

The terminology used in this note is the same as that employed 
by Prof. Howes and myself in 1888. The arguments propounded 
by Perrin, Emery, and Zwick do not appear to be of sufficient 
weight to warrant the relinquishing of the opinions we then 
held as to the morphological value of the parts of the Anuran 
manus. 


EXPLANATION or Fie. 5, Pratz 31. 
Carpal skeleton of Hymenochirus Boettgeri, dorsal view (x 12). 


7. Lunatum. 
n. 2,3. Naviculare confluent with the carpals of the second and third 
digits. 
pk. Pyramidale confluent with the post-axial centrale. 
po. 1. Carpal of the pollex. 

r, Radius. 
s. Radial sesamoid. 
uw. Ulna. 
4, Carpal of the fourth digit. 

', 3', 4', 5'. Metacarpals of the second, third, fourth, and fifth Hs digits 


Ridevrood. Linn. Soc. Journ. Zoor Vou XXVII. Pu. 3). 


W.G.R. del. J.Green lith. ~ Mintern Bros imp. 


HYMENOCHIRUS BOETTGERI. 
Hyobranchial Skeleton and Carpus. 


re ny 


EYE-SPOT IN EUGLENA VIRIDIS. 463 


On the Eye-spot and Flagellum in Luglena viridis. 
By Harotp Waaer, F.LS. 


[Read 2nd November, 1899.] 


(PiatE 32.) 


Everena viripis is commonly found in stagnant water which 
contains a considerable amount of organic matter. It is often 
found abundantly in the liquid which runs from manure heaps, 
forming at certain periods a dense green scum on its surface. 
It is found sometimes in water troughs, and on mud in roadside 
gutters in places where refuse water from houses is thrown; 
and it is often very abundant on sewage farms. 

Under ordinary conditions it is a free swimming organism, 
capable of moving very rapidly through the water by means of a 
cilium, or flagellum, which is placed at the anterior end of the 
body; but it can exist for a long time in a non-motile condi- 
tion in an encysted state, surrounded and protected by a very 
thick cell-membrane. It is, however, capable of passing again 
into the motile condition as soon as the environment becomes 
favourable. 


General Structure. 


An excellent account of the general structure and life-history 
of the genus Euglena is given by Klebs *, whose memoir records 
the literature on the subject up to the date of its publication. 
A large amount of information concerning the structure of the 
various species of Huglena and allied genera is given by Stein T, 
Butscblif, Saville Kent$; and, more recently, a good general 
account of the group will be found in the first volume of the 


* “ Ueber die Organisation einiger Flagellaten-Gruppen und ihre Beziehungen 
Zu Algen und Infusorien.” Unters. aus dem bot. Inst. zu Tubingen, Bad. I., 
1881-85. 
t Der Organismus der Infusionsthiere, III. Der Organismus der Flagellaten. 
Leipzig, 1878. 
{ Bronn’s Classen und Ordnungen des ‘Thierreichs, I. Protozoa, 1883-87. 
§ A Manual of the Infusoria. London, 1880-82. 


464 MR. HAROLD WAGER ON THE 


‘Traité de Zoologie Concrete’ by Yves Delage and Edgard 
Hérouard *. 

A memoir by Khawkine f also contains valuable information 
on the structure and physiology of Huglena viridis. For further 
information concerning the numerous memoirs in which the 
structure of Huglena is dealt with, the reader is referred to the 
papers quoted above, which contain complete bibliographical lists 
of the literature on the subject. 

Without attempting to give a complete account of the structure 
of Euglena viridis, it may be useful to briefly summarize what is 
contained on the subject in the memoirs already published, 
which my own observations enable me to confirm. 

In the free swimming condition, the animal is elongate and 
cylindrical in shape, slightly larger in the middle than at the 
ends; the anterior end being truncated, the posterior usually 
pointed. It is a unicellular organism, protected on the outside 
by a thin skin or layer of modified protoplasm which is striated 
obliquely by slightly elevated ridges. These can be made visible 
by crushing the cell and squeezing out the protoplasmic contents 
under a cover-glass. Under certain conditions, of which one 
appears to be malnutrition, I find that the cells, while still 
retaining their power of movement, become curiously distorted 
and deformed, and might easily be mistaken for distinct species. 
This apparently accompanies a process of slow disintegration. 

The protoplasm contains numerous chlorophyll-bodies, some- 
times scattered all over the cell, with the exception of a shert 
space at the anterior end which always remains colourless, but 
more often radiating from the centre, leaving both anterior and 
posterior ends free. In many cases, especially in cells freshly 
collected, it is not easy to distinguish the separate chlorophyll- 
bodies ; and this led Saville Kent (doc. cit.) to regard the chloro- 
phyll as diffused through the protoplasm ; but it is only necessary 
to keep such cells in obscurity for a short time, in ordinary tap- 
water, or to examine them under a high power of the microscope, 
to see that the chlorophyll-bodies are really definite organs of 
the cell $. 


* Vol. L. La Cellule et les Protozoaires. Paris, 1896. 

+ “ Recherches biologiques sur |’ Astasia ocellata n.s. et Euglena viridis.” 
Ann. des Sci. Nat. 7th series, vol. i., 1886. 

t Klebs, 7. c. p. 264. See also Jessie A. Sallitt, “On the Chlorophyll 
Corpuscles of some Infusoria,” Q.J.M.S. 1884. 


EYE-SPOT AND FLAGELLUM IN EUGLENA VIRIDIS. 465 


I find that the chlorophyll-bodies are capable of assuming 
different shapes—spherical, oval, elongate, and ribbon-like ; and 
to this may be due the diferene views as to their nature taken 
by various observers. 

The cell usually contains a large number of grains of a starch- 
like composition (paramylum), which are not coloured blue by 
iodine, and are therefore not pure starch. They possess a very 
definite and characteristic structure, which is correctly figured 
by Klebs *, presenting some resemblance to the structure of a 
typical starch grain in being composed of lamine, but differing 
from the starch grain in this, that the lamine are not concentric, 
and present the appearance of flat plates laid one upon the 
other. 

These paramylum grains may be found in any part of the cell, 
but under normal conditions are generally more numerous in 
the region of the chlorophyll-bodies. In cells kept in the 
dark, in a good nutrient solution such as a dilute solution of 
potato-starch T, the chlorophyll-bodies become pushed to the 
posterior end of the cell, whilst the anterior two-thirds of the 
cell become filled with a mass of paramylum grains. This 
results in the production of the so-called “white” or “ colour- 
less” forms. 

The nucleus is usually found at the posterior end of the cell, 
but it may occur in the middle, or very rarely at the anterior 
end; in the “ white” forms just described, it is found between 
the chlorophyll-bodies and the paramylum grains. In normal 
cells the nucleus is posterior to the chlorophyll-bodies; in the 
“white” forms itis anterior to them. It can be very easily seen 
in the living cell, and with care its structure can be made out. 
In osmic acid, and other hardening and preserving fluids, the 
structure is brought out very clearly. It consists of a central 
spherical nucleolus surrounded by a granular network. 

The anterior end of the cell has a depression leading into a 
narrow tube or gullet, out of which the flagellum passes. On 
the dorsal side of the gullet isa distinct eye-spot, and near it is 
a large clear space, called the principal vacuole; and on one 
side of this a single, pulsating vacuole. 


* Loc. cit. See plate ii. figs. 8a, 8b, 8c. 
f Khawkine, /. c. 


466 MR. HAROLD WAGER ON THE 


The Vacuole System and Gullet. 


At the anterior, colourless end of all species of Huglena and 
allied genera there is a sharply defined space, which was known 
to Ehrenberg, and has been described by Klebs * as the “ prin- 
cipal vacuole,’ into which open one or more pulsating vacuoles T. 
In Euglena viridis there is only one pulsating vacuole, but in 
other species there may be two or more, and in some cases the 
single pulsating vacuole is formed by the fusion of several 
smaller ones ¢. The principal vacuole possesses the power of 
slowly contracting, by which the liquid contained in it is dis- 
charged; but it is not to be regarded as a true contractile 
vacuole, but rather as a reservoir for liquid; and as a part of 
this liquid is poured into it by the true pulsating vacuoles, it 
would probably be better to describe it as an “excretory reservoir.” 
The gullet is a conical or tube-like depression in the blunt 
anterior end of the cell, and it is usually described and figured 
as terminating in the neighbourhood of the excretory reservoir, 
from which it is separated by a thin layer of protoplasm only. 

According to the view at present held, the excretory reservoir 
discharges its contents into the gullet through an opening which 
is made, at the moment when the contraction begins, by the 
rupture of the thin layer of protoplasm which separates the 
reservoir from the gullet §. Klebs ||, however, states that no 
opening can be observed in the excretory reservoir; it always 
remains sharply defined, and its slow contraction shows that such 
an opening is improbable. 

I found, however, upon a careful examination of Huglena 
viridis, that, contrary to the observations described above, there 
is a permanent communication between the excretory reservoir 
and the gullet. This is not easy to observe in the living cells, 
but is very clearly seen when they are placed in a half to one 
per cent. solution of osmic acid for several hours, or even days, 
then mounted in dilute glycerine and observed under a ;'yth inch 
oil-immersion objective, with the aid of a good sub-stage 
condenser. 

* Loe. cit. p. 246. 

+ Carter, “ Additional Notes on the Freshwater Infusoria in the Teen of 
Bombay,” Ann. & Mag. Nat. Hist. ser. 2, vol. xx. p. 34. Stein, 1. ¢. 

t Klebs, 7. c. 

§ Traité Zool. Concrete, i. p. 346. 

|| Loc, cit. p. 248. 


EYE-SPOT AND FLAGELLUM IN EUGLENA VIRIDIS. 4.67 


_ It is then’seen that the external opening of the depression at 
the anterior end of the cell is slightly funnel-shaped, that it 
leads into the gullet, a narrow tube-like passage, which is curved 
towards the dorsal side of the animal, and opens at its distal 
end into the so-called principal vacuole or excretory reservoir 
(Pl. 32. figs. 2-8). 

The cavity of the excretory reservoir is thus in communication 
with the exterior ; and this affords a more satisfactory explanation 
of the discharge of the liquid poured into it by the pulsating 
vacuole than that given by previous observers. 

This connection between the excretory reservoir and the 
gullet does not appear to have been recognized by previous 
observers; although Stein (J. c.) figures what appears to me to 
be a distinct connection between themin Euglena deses, Ehrbg., 
E. Spirogyra and Colactum calviwm, Stein. 

As to the function of this anterior cavity we know very little. 
The flagellum arises in it, and it is often stated that the ingestion 
of solid food material can take place through it, small particles 
of solid matter being forced down into the protoplasm at the 
base of the opening by the continual movement of the flagellum, 
where they become digested in the same way as in Ameba*, 

But it is doubtful whether this can be definitely proved, and 
the evidence at present brought forward in favour of it is not 
altogether satisfactory. I have constantly repeated the experi- 
ment, which is commonly taken as a demonstration that Euglena 
possesses the power of absorbing solid food-material, of putting 
finely powdered carmine in water containing Huglena-cells in 
active movement; and have observed that the motion of the 
flagellum causes a rapid and violent movement of the carmine 
grains, often sweeping them up to and past the mouth-opening 
in large numbers; but I have never been able to satisfy myself 
that any of them ever get into the gullet. Dangeard fT states 
that the nutrition in the whole group is distinctly vegetal in 
character; and that no solid particles penetrate the interior of 
the body. The gullet, according to him, certainly does not serve 
for the introduction of solid matter. 

Khawkine (J. ¢.) suggests that lquid nutriment may be 


* See, for example, Parker’s ‘ Elements of Biology,’ p. 40. 
+ “Recherches sur les Cryptomonadine et les Euglene.’” Le Botaniste, i. 
1889, p. 1. 


468 MR. HAROLD WAGER ON THE 


absorbed through the gullet, and he brings forward the following 
observation in support of this view. When Huglene are kept 
in dilute potato-starch solution in the dark, paramylum grains 
are produced in great abundance, gradually filling up the whole 
cell, and pushing the chlorophyll granules to the posterior end 
of the cell. The smallest paramylum grains are found at the 
anterior end of the cell in the neighbourhood of the gullet; and 
they gradually increase in size towards the middle of the cell, 
where they are much crowded together and nearly all of the 
same size. Khawkine’s explanation of this is that the small size 
of the grains near the gullet indicates that they begin to form 
in this region, and that it is probably through the gullet that 
the substances are absorbed which are necessary for their 
production. 

This is a very interesting observation; but it is obvious that 
we require further evidence before we can say definitely that 
liquid nutriment is absorbed wholly, or even partly, through the 
gullet. 

So far, then, all we can say concerning the function of this 
anterior cavity in the cell of Euglena is that it serves for the 
excretion of liquid from the pulsating vacuole ; but that it serves 
for the ingestion of either solid or liquid food has not been proved. 
Tn a later part of this paper I shall refer to its connection with 
the flagellum. 


Structure of the Eye-spot. 


It is well kuown that many motile cells of both animals and 
plants possess a red pigment-spot which is sharply defined 
from the protoplasm; and, from its apparent resemblance to 
the eye of Cyclops and Rotifers, was called by Ehrenberg an 
eye-spot. 

Our knowledge of its structure depends upon the researches 
of Leydig, Kunstler, Klebs (@. ¢.), Schilling, Franzé *, Overton fT, 
Pouchet, Johnson {, and others, for an account of which reference 


* Franzé, “Zur Morpholegie und Physiologie der Stigmata der Mastigo- 
phoren.” Zeit. f. Wiss. Zool. 1893, vol. lvi. p. 138. 

+ Overton, “Beitrag zur Kentniss der Gattung Volvox.” Bot. Centralbl. 
vol. xxxix. p. 65, 1889. 

+ Jobnson, “Observations on the Zoospores of Draparnaldia.” Bot. Gar. 
vol. xvili. p. 204, 1893. 


EYE-SPOT AND FLAGELLUM IN EUGLENA VIRIDIS. 469 


may be made to Zimmermann™*, and to the memoirs of Klebs 
and Franzé. 

The eye-spot was at first supposed to be a homogeneous red 
spot or disc; but Leydig 7 in 1856 described it in Euglena and 
some other forms as consisting of scarcely measurable, strongly 
refractive granules ; and this has since been confirmed by many 
other observers. 

According to Klebs (/. ¢.) the eye-spot of Euglena is a well- 
characterized body, both in form, inner structure, and chemical 
behaviour. Its form varies with the species, but is very 
constant for one and the same species. Its structure is appa- 
rently the same in all species, consisting of two substances—a 
plasmatic ground-mass forming a fine network, and a pigment 
which occurs in the form of drop-like bodies embedded in it. 
The presence of a plasmatic substance can only be inferred from 
the fact that the eye-spot can be caused to expand under pressure 
or by the action of swelling reagents, which results in the 
separation of the pigment-drops from one another. 

Franzé (/. c.) states that, m addition to the pigment, the eye- 
spot contains one or few to many strongly refractive bodies 
which are, in Huglena, composed of paramylum, grouped in a 
more or less regular manner, consisting of a central or excentric 
“crystal body,’ with smaller and always more numerous “ lens 
bodies’ surrounding it. Both the crystal body and the lens. 
bodies serve to concentrate the light, and the pigment not only 
absorbs light but s also sensitive to light. 

I have been quite unable to confirm Franzé’s observations 
either as to the crystal body or the lens bodies, although it is 
very easy to observe the granular structure described by other 
observers. I have examined several species of Huglena and 
Phacus; I have had Euglena viridis under observation in all its 
stages and under many different conditions of its existence, but 
I have never seen anything of the nature of paramylum bodies 
in or on the eye-spot. Nevertheless I have sometimes noticed, 
when examining the eye-spot with a magnifying-power of 500 
or 600 diameters, an appearance something like that figured by 
Franzé, which seems to be due, however, to the irregular outline 
of the eye-spot (it is very rarely as regular in outline as Franzé’s 


* Zimmermann, “Sammel. Referate &c., 10. Der Augenfleck (Stigma).” 
Beihefte zum Bot, Centralbl. vol. iv. p. 160, 1894. 

t Leydig, Lehrbuch der Histologie. (See Franzé, /. c.) 
LINN. JOURN.—ZOCLOGY, VOL. XXVII. 36 


470 MR. HAROLD WAGER ON THE 


figures would indicate), and to the fact that it is curved around 
the gullet and is sometimes sharply turned in here and there at 
the edge. The eye-spot has thus not only an irregular contour, 
but its surface is very uneven, and under a low power with poor 
definition it may sometimes present an appearance of colourless 
granules embedded in the pigment. Under a high magnifying- 
power with good definition, this appearance vanishes completely, 
and the eye-spot is resolved into the simple granular structure 
to which reference has already been made. 

The pigment granules are brightly refractive and have a very 
distinct outline. They form a single layer, and in Huglena 
viridis are easily separated from each other, especially when 
the cells are in the enyested condition. In some eye-spots the 
grauules are spherical and all of the same size, but in others they 
are more irregular in shape and of different sizes, and in such cases 
the eye-spot is more homogeneous in appearance, especially near 
the middle. 

By the action of strong putash solution the eye-spot swells up, 
and the pigment granules become separated from one another *. 
The arrangement of the granules is not very definite, but now and 
then they were found to be grouped in rows, sometimes radiating 
from the centre (fig. 1,a). The number of granules present 
varies, but not to any great extent: in a fairly large number of 
cases which I counted, I found that between 30 and 40 granules 
were the most frequent. If the action of the potash is continued 
for some time, the eye-spot disintegrates more or less completely 
into a number of granules (Pl. 32. fig. 1,d). Even in the living 
condition, the eye-spot sometimes breaks up into a number of 
separate granules which become distributed through the proto- 
plasm. This is frequently found to be the case in cells which 
have become encysted and surrounded by a thick wall. I have 
never seen it in elongate motile cells, although I have often 
noticed in the surrounding protoplasm a number of granules of 
the same size and colour as those in the eye-spot, and looking as 
if they had been separated from it. They can be easily dis- 
tinguished from the rusty-red granules, which appear in the 
protoplasm as a result of the disintegration of the chlorophyll 
grains, by their bright red or orange colour and greater refractive 
power. The nature of this colouring-matter has not been fully 


: Klebs, 7. ¢. 


EYE-SPOT AND FLAGELLUM IN EUGLENA VIRIDIS. 471. 


investigated, but the few observations which have been made 
seem to show that it is a chlorophyll derivative. According to 
Cohn, with whom Klebs agrees, the red pigment is hemato- 
chrome, and stands in genetic relationship to chlorophyll ; 
whilst Rostafinski* regards it as reduced chlorophyll. The 
rusty-red granules and the pigment of the eye-spot are similar 
in their behaviour towards alcohol, which in both cases, imme- 
diately it comes into contact with them, causes the granules 
to run together to form a homogeneous red mass or drop of oily 
substance, from which two colouring-matters at once become 
separated out—an orange-coloured substance which forms the 
main mass of the drop, and one or two small bright red droplets in 
the centre of it. The red colour soon disappears entirely, and the 
orange colour changes to yellowish green, then to green, and at 
the same time becomes smaller and smaller until it suddenly 
disappears, and in its place a small vacuolar-like body is left, 
surrounded by an irregular ring of some refractive substance. 

This reaction, although not conclusive by any means, indicates 
that the rusty-red granules which are derived from the chloro- 
phyll and the red pigment of the eye-spot have something in 
common, and supports Cohn’s statement that the latter is 
genetically connected with chlorophyll. It has been suggested 
that the pigment of the eye-spot is identical with the red 
colouring-matter, carotin, which occurs in the roots of Daucus 
Carota and in the orange or red chromatophores of many fruits 
and flowers, from the fact that it shows the characteristic blue 
colour with sulphuric acid rf. 

Guignard t has shown that in the Fucacee the orange- 
coloured chromatophores possess the same chemical reactions as 
the orange-coloured eye-spot; and he further shows that the 
eye-spot is formed from a colourless chromatophore found on one 
side of the nucleus, which at first becomes rapidly coloured yellow, 
then orange. 

In Euglena the eye-spot is found both in the motile and in the 
resting cells, and new eye-spots arise by division, as Klebs has 
also shown. Whether there is any formation of eye-spots de 
novo at any stage in the life-history of Huglena, such as occurs in 


* Klebs, /. ¢. 
t+ See Zimmermann, Botanical Micro-technique. 
t Revue Gén. de Bot. i. 1889. 
36% 


472 MR. HAROLD WAGER ON THE 


the motile cells of Fucacee* and Alege +, is doubtful. The 
breaking-up of the eye-spot which has been observed in old 
encysted cells may simply be a part of the general disintegration 
of the cell; if not, it would appear that a new formation of 
the eye-spot must take place on the resumption of the motile 
condition. This, however, requires further investigation. 

Without entering into any further discussion of these facts, 
it is clear, I think, that the eye-spot is a definite organ of the 
cell possessing a characteristic, but simple structure ; that it is 
probably a derivative of chlorophyll; that it is present in the 
motile cells only of those Algz in which it has been described and 
is formed de novo when they are produced; and that even in Ew- 
glena, although it is present both in the motile and in the resting 
condition, it disappears, or becomes broken up, in cells which 
remain long in the encysted condition. Weare probably justified, 
therefore, in concluding that the eye-spot is an organ of the 
motile stage of both animal and plant cells, specially connected 
with their power of movement. 


The Flagellum. 


The flagellum is an organ of the cell especially connected with 
its rapid movement through the water. If it is absent, the 
Euglena is only capable of a slow creeping movement which is 
caused by a peculiar contraction of the body, characteristic of 
the organism. This contraction appears to originate in the 
protoplasm, but the different forms assumed are probably to 
some extent dependent upon the elasticity of the limiting layer 
or cell-wall; and Khawkine (J. c.) suggests that contractile fibres 
are present—longitudinal ones over the whole of the cell, and 
transverse ones in the anterior region only. These, however, 
have not yet been seen, the only indication of any such contractile 
fibres being the oblique striz found on the cell-wall; and so far 
we have no evidence that these play any part in causing the 
contraction of the cell. 

The free movement of the cell through the water is entirely 
due to the flagellum, which by its rapid motion draws the cell 
after it, causing it at the same time to rotate around its longi- 
tudinal axis. Very little is known of the actual mechanical 
means by which this is accomplished, but an interesting discussion 


* Guignard, /. ¢. + Overton, Z. ¢., and Johnson, /. c. 


EYE-SPOT AND FLAGELLUM IN EUGLENA VIRIDIS. 473 


of the question will be found in the first volume of the ‘ Traité 
de Zoologie Concrete’ * by Delage and Hérouard. 

It is usually stated that the flagellum arises either on the dorsal 
wall of the gullet or from the protoplasm at its base. This, 
however, is not correct. The flagellum not only does not ter- 
minate in the gullet, but actually passes into the excretory 
reservoir, with which, as I have shown, the gullet is in open 
communication, and it becomes attached to its dorsal wall, or occa- 
sionally to its ventral, by a biturcate base (Pl. 32. figs. 2 to 8). 
This bifurcation extends as far as the anterior edge of the eye- 
spot, at which point the single whip-like portion of the flagellum 
begins (fig. 2). This mode of attachment of the flagellum has 
not, so far as I am aware, been observed before; it probably 
serves to give it a much firmer support, and may be of mechanical 
advantage to it in its movement. 

In addition to this bifurcation, we find another interesting 
structure connected with the flagellum. On one of the branches 
of the bifurcate base there occurs an oval swelling or enlargement, 
immediately in front of the eye-spot, and just below the point 
where the bifurcation begins (Pl. 32. figs. 2,8). It is in close 
contact with the eye-spot when the cell is in the elongate motile 
condition ; but it is not actually in organic connection with it, as 
shown (Pl. 32. figs. 7, 8), and in numerous cases which were 
observed of Euglena-cells in the resting condition, it was com- 
pletely separated from it (fig. 9). In some cases, this enlarge- 
ment appears in the position indicated in figs. 3 to 6, just above, 
or at, the point of bifurcation of the flagellum ; but from a careful 
examination of numerous specimens in which this appearance 
was seen, it seems to me that it is an effect due to the second 
branch of the bifurcation passing below the enlargement, and so 
becoming hidden from view (compare figs. 7 and 8). 

Although both the bifurcation and the enlargement are visible 
under favourable conditions, in the living cell, it is necessary in 
order to see them clearly to resort to the use of reagents. The 
best reagent for the purpose is a 1 per cent solution of osmic acid. 
This kills motile Huglena-cells at once, and leaves them in an 
expanded condition with their flagella distinctly visible. They 
should remain in this solution from three to forty-eight hours 
or more, and may be examined in dilute glycerine. In order to 
preserve them for future investigation, they may be kept in a 


* La Cellule et les Protozoaires, p. 305. 


47 4. MR. HAROLD WAGER ON THE 


50 per cent. solution of alcohol. Dilute glycerine is the best 
mounting medium, but fairly satisfactory preparations may be 
made either in glycerine-jelly or canada balsm. 

In good preparations the contents of the cell appear black, the 
eye-spot black and sharply defined, and the flagellum with its 
enlargement dark grey. 

I have occasionally succeeded in staining the flagellum and 
enlargement light green in a mixture of methyl-green and 
fuchsin. Fig. 2 is from a preparation made in this way and 
mounted in dilute glycerine. 


The Effect of Light on Buglena. 


In common with many other unicellular motile organisms 
which contain chlorophyll, Euglena is extremely sensitive to 
light. Our knowledge of the phenomena is mainly due to 
the investigations of Stahl *, Strasburger tT, and Engelmann f. 
They find that in general the cells are attracted by a light of 
moderate intensity and repelled by an intense light; but that 
the degree of sensitiveness which they exhibit varies considerably 
even in individuals of the same species. Further, both Klebs 
and Strasburger have shown that they do not lose their sensi- 
tiveness to light either in the dark or at a higher temperature; 
and Englemann has also shown that this power is independent 
to a high degree of a variation in the oxygen pressure. 

The effects produced by the action of light upon the motile 
cells of Euglena are very striking. They are strongly attracted 
by a bright light such as that of a gas-flame or incandescent 
burner focussed, by means of a substage condenser, upon a 
microscope-slide on which a drop of water containing Huglene 
has been placed. Such a spot of light will attract, in the space 
of about one minute, the majority of the cells in the field of the 
microscope, as seen by a one-inch objective; and in two minutes 
only very few will be found outside the light area. If a large 
number of cells are present, they will form a seething mass in 


* “ Ueber den Einfluss von Richtung und Starke der Beleuchtung auf einige 
Bewegunegserscheinungen im Pflanzenreiche.” Bot. Zeitung, 1883. 

+ “ Wirkung des Lichtes und der Warme auf Schwarmsporen.” Jena. 
Zeitschr. xii., 1878. 

t “Ueber Licht und Farbenperception niederster Organismen.”  Pifliiger’s 
Archiv, Bd. 29, 1882 (see J. R. M. 8. 1883, p. 81). 


EYE-SPOT AND FLAGELLUM IN EUGLENA YIRIDIS. 475 


the light space, perhaps two or three layers deep, all of them 
with the anterior end pointing downwards towards the source of 
light. On turning down the condenser so as to get the light 
spread evenly over the whole field of the microscope, they 
begin at once to move away rapidly in all directions, and in one 
or two minutes are found in all parts of the field. If the light 
is again focussed upon a small portion of the field, the Huglene 
turn round suddenly and make for it ; and a steady stream of 
them will be seen moving up to the light from all parts of the 
field. When the light space is shifted from place to place in 
the field of the microscope, the Huglene follow it; and if the 
mirror be moved backwards and forwards in such a manner 
that the light space is made to pass continually from one side of 
the field to the other, the Euglene arrange themselves all across 
the field in the area marked out by the passage to and fro of the 
light space. The rapidity with which they move up to the light 
space is much greater when there is no diaphragm over the con- 
denser than when one is present. ‘The diffuse light outside the 
bright spot apparently guides them. 

It is interesting to watch the movements of the Huglene in 
the light space. The majority of them move about freely, but 
appear to be unable to get out ; for directly the anterior end of 
an individual passes into the shade, it turns round completely 
and goes back again into the light. Some of them, however, may 
pass completely into the shade, and may even swim for some 
distance into the dark part of the field before turning round ; 
others again do not turn round completely at once, but move 
round the edge of the light space in a tangential direction for 
some distance before passing in again. 

Some of the cells which are strongly attracted to the light 
space move straight across it, and pass out again on the opposite 
side into the dark part of the field. After traversing this for 
some distance, they turn round and repeat the performance; 
and they may do this three or four times in succession before 
they come to remain in the light space. 

If a drop of water containing motile Huglene be placed on a 
glass slip and exposed to bright sunlight, the Huglene move to 
that side which is farthest from the sun. If the slip is turned 
round, they at once begin to move towards the opposite side, 
that which is now farthest from the sun; and in the space of 
three minutes a large number of Huglene are able to pass from 


476 MR. HAROLD WAGER ON THE 


one side to the other, across a drop of water a quarter of an inch 
in diameter. If in their passage across, the sun becomes obscured 
by clouds, or if a sheet of note-paper is placed between the sun 
and the drop of water, they stop at once, and either remain 
moving about in the middle of the drop or go back again to the 
side nearest the source of light. 

If they are kept in bright sunlight for any length of time, they 
come to rest and round themselves off ; and if they remain in a 
good light for some days, the cells gradually become encysted. 
The effect of a strong light is always to produce encystment. 

In a light of moderate intensity, the cells may remain motile for 
a very long time; but they always become rounded off at night, 
and may then lose their flagella and undergo division into two. 
Until they lose their flagella, however, they may be always 
brought into the motile condition again by exposing them to 
the light of a lamp or gas. This shows, as Klebs* has pointed 
out, that they do not lose their sensitiveness even in the dark so 
long as they can move. 

These experiments are sufficient to show that Euglena possesses 
a very definite light-perception. Engelmann has also shown 
that it is the colourless anterior end of the cell which is sensitive 
to light, and it is only when this comes into contact with light 
or shadow, that the cell reacts to the light by altering the 
direction of its movements. If the shadow falls upon the 
posterior chlorophyll-containing end of the cell, there is no 
reaction. He has further shown that Euglena prefers the blue 
portion of the spectrum, the following being the percentage dis- 
tribution of a typical case :— 


ieditoy Oranlgelereeeeee cree 1:4 per cent. 
Oranse ito Greeny ver eeace 0) eae, 
Greenies ee ee Near ib a 
Green tor Bluen sa. c.ueec Oe eee 
JUS HO LOCH) sag ncoosassnsoc LO;S ees 
Indigo to Vaolet <.02...5.... | Waa 


It has also been shown that swarmspores are more sensitive 
to the blue rays of the spectrum than to other parts. 

According to Franzé (J. ¢.) and Wildeman t motile Huglene are 
sensitive to heat, but they do not respond to it either so quickly 
or so intensely as to light. 

* Loe. cit. p. 263. 

t “Sur le thermotaxisme des Euglénes,” Bull. Soc. Micros. Belg. 1894. 


EYE-SPOT AND FLAGELLUM IN EUGLENA VIRIDIS. 477 


To determine the effect of heat, Franzé placed Huglene in a 
thin glass tube which was closed at both ends, and kept warm 
at one end by allowing a stream of hot water to flow over it. 
The tube was kept in the dark. At a temperature of 55° C. the 
Euglene became immobile from the effects of heat, but at a 
lower temperature, 30°-40° C., they mostly approached the source 
of heat. 

Wildeman obtained somewhat similar results. He placed 
Huglene in a tube with wet sand, in order to avoid convection- 
currents. It was then placed in the dark in a horizontal 
position, and warmed at one end, and he found that the Euglene 
accumulated at the warmer end of the tube at a temperature of 
30° C. On exposing tubes to light as well as heat, he found a 
considerable modification in the effects produced. If they were 
placed at right angles to the rays of light, the Huglene avoided 
the warm end of the tube. If they were placed in the same 
direction as the rays of light they moved towards the light, even 
when the opposite end of the tube was heated. 

My own experiments confirm in a general way the results 
obtained by these observers; but the Huglene are so slightly 
sensitive to heat as compared with light, that the subject requires 
further investigation before any very definite conclusions can be 
arrived at. 


The Function of the Eye-spot. 


Jt is commonly stated that the eye-spot is a light-perceiving 
organ. This does not imply that it possesses an actual visual 
function, but simply that it is connected in some way with those 
changes in the direction of the movements of the cell which are 
due to light. There is no direct proof of this, but we have a 
sufficient amount of indirect evidence to show that the statement 
is probably a correct one. 

Ehrenberg regarded the pigment-spot in Huglena as a light- 
perceiving organ on account of its general resemblance to the 
eyes of Rotifers and Cyclops; and it has since been shown 
that it resembles them in some respects both in structure and in 
its behaviour towards solvents and other reagents, such as iodine 
and sulphuric acid *. 

Further, in all those chlorophyll-containing unicellular or- 
ganisms which are very sensitive to light and capable, by means 


* Klebs, doc. cat. 


478 MR. HAROLD WAGER ON THE 


of their flagella or cilia, of responding quickly to changes in its 
direction or intensity, an eye-spot is present. Those motile cells 
which do not possess an eye-spot are either not sensitive to light 
at all, or only to a slight extent. Even in the zoospores of 
Chytridium, which according to Strasburger* are sensitive to 
light, it is interesting to note that at the base of the cilium there 
is a conspicuous orange-coloured oil-globule, which may act in 
the same way as an eye-spot. 

Again, Engelmann has shown for Euglena and Strasburger for 
swarmspores of Algw, that the rays of light which are most active 
in their influence upon the movements of these organisms are 
found in the region of the blue portion of the spectrum, and 
these are just the rays which, as indicated by the colour of the 
eye-spot, are absorbed by it. We conclude from this, therefore, 
that it is the light absorbed by the eye-spot, and not that trans- 
mitted through it, which is concerned in these movemeuts. 

Finally, it has been shown by Engelmann that it is the colourless 
anterior end of Euglena that is sensitive to light. ‘ Hence, in 
this case, a certain part of the body functions to a certain extent 
as an eye”. In this colourless anterior end of the cell, both the 
eye-spot and the apparatus which directly causes the movement 
of the cell—the flagellum—are placed. 

Having thus briefly stated the evidence we possess in favour 
of the conclusion that the pigment-spot of Huglena is a definite 
light-perceiving organ, we must now attempt some explanation 
of the way in which the light acts. 

It is obvious that, whatever may be the action of light, the 
movements of the cell, as well as any change in direction of its 
movements, are dependent directly upon the flagellum. Without 
it, as we have seen, the cell is only capable of a very siow con- 
tractile movement, of its body from place to place. It is evident 
therefore that those rays of light which are capable of exerting a 
material influence upon the movements of the cell can only do 
so by controlling or modifying in some way the mechanism by 
means of which the flagellum is caused to move. Now we have 
already seen that the flagellum and eye-spot are closely related 
to each other. The flagellum arises near the eye-spot and, on its 


* Jena. Zeitschr. xii. 1878, p. 568. 
+ Hertwig, ‘ The Cell,’ p. 100. 


EYE-SPOT AND FLAGELLUM IN EUGLENA VIRIDIS. AT 


way to the exterior, passes quite close to it. Moreover it possesses 
an enlargement on one of its basal supports which is placed im- 
mediately in front of, and in close contact with, the inner concave 
surface of the eye-spot. 

The presence of this enlargement of the flagellum in such a 
position at once suggests the simple explanation that the light- 
rays which are absorbed by the eye-spot cause a stimulation 
of the enlargement in some way ; and this stimulation reacting 
upon the flagellum causes its movements to become modified, and 
so results in a change in the direction of the movement of the 
cell. 

We have thus a combination of two structures, which may, 
if my explanation be correct, be regarded as an extremely simple 
form of eye, consisting of a specialized portion of the cell 
(protoplasm ?) possessing great sensitiveness to light-rays of a 
particular kind, and a pigment-spot, as a light-absorbing organ, 
in close contact with it. 

Whether it is the light absorbed by the pigment-spot which 
actually effects the change in the movements of the flagellum; 
or whether the pigment-spot simply prevents these rays from 
reaching one side of the enlargement, whilst the other side is 
left freely exposed to them, thus producing a difference of 
intensity on the two sides of the enlargement, it is at present 
impossible to say. We know that protoplasm itself is sensitive 
to light, and responds to it without the intervention of any such 
structure as a pigment-spot. It may be therefore that the 
arrangement just described in Euglena is one by which the 
active rays are, under certain conditions, cut off on one side of 
the enlargement, while the other is left exposed to them. This 
would result in a definite unequal illumination of the sensitive 
portion of the cell; and consequently the organism would tend 
to move into such a position that it would be more or less equally 
illuminated all round, and would be brought ultimately into the 
light-ray. 

I put forward this explanation tentatively, as the only one 
- which appears to me at present to afford an adequate inter- | 
pretation of the facts observed. It’ is evident, however, that 
it is a question which demands a more extended and laborious 
investigation than I have so far been able to undertake, before 
any very definite conclusions can be formulated. 


A80 MR. HAROLD WAGER ON THE 


Summary. 


(1) The structure of the eye-spot in Huglena viridis, HE. Ehren- 
bergii, and Phacus pyriforme is not as Franzé has described it. 
The so-called “crystal body ” and ‘lens bodies ” were not found 
in any of the individuals of the species examined by me. As 
earlier observers have stated, the eye-spot consists simply of a 
mass of pigment granules arranged in a single layer, and probably 
embedded in a protoplasmic framework, but the latter could 
not be seen, and its presence was only inferred from the fact 
that the pigment granules must be held together in some way, 
that the eye-spot as a whole is capable of expansion, and that the 
pigment granules can be separated from each other. 

(2) The gullet at the anterior end of the body does not end 
blindly, as was formerly supposed, near the principal vacuole, 
but actually opens into it, thus forming one continuous cavity 
open to the exterior. 

(8) The eye-spot is in close contact with the gullet at the 
point where it opens into the excretory reservoir, and curves 
around it slightly. 

(4) The flagellum is not attached to the wall of the gullet, 
but arises from near the posterior or basal side of the excretory 
reservoir, to which it is attached by a bifurcate base. The bifur- 
cation begins at about the level of the eye-spot, and in this region 
an enlargement of one of the basal supports takes place. This, 
which is oval in shape, is in close contact with the eye-spot, on 
its inner concave side. It is in the same position as that 
described by Franzé for his “ crystal body,” and may have been 
mistaken for this by him. 

(5) An explanation of this structure suggested by the author 
is that the light which is absorbed by the eye-spot in some way 
stimulates this enlargement, which in turn reacts upon the 
flagellum and causes its movements to become modified. This is 
supported by the facts which are stated in the three following 
paragraphs. 

(6) The rays of light which are absorbed by the eye-spot— 
those in the region of the blue of the spectrum—are the same as 
those which are active in modifying the direction of movement 
of the Zuglena-cell. 

(7) These rays can only influence the movements of the cell 
by acting upon‘or controlling in some way the mechanism by 
which the flagellum moves. 


Linn Soc. Journ. Zoon. Vou. XXVII Pi.32. 


Wager 


et 
ps 


Geo. West & Sons imp 


H. Wager del. 
MP. Parker lith. 


HUGLENA VIRIDIS. 


EYE-SPOT AND FLAGELLUM IN EUGLENA VIRIDIS. 481: 


(8) All those chlorophyll-containing motile cells which are 
very sensitive to light contain an eye-spot. Many other organisms 
which do not possess an eye-spot are sensitive to light, but not to 
so high a degree. 

(9) If the explanation given in par. 5 is correct, we have 
in Huglena an extremely simple form of light-organ, consisting 
of a specialized sensitive portion of the cell—the enlargement on 
the flagellum—and a light-absorbing pigment-spot in close con- 
tact with it. 


EXPLANATION OF PLATE 32. 


The figures have been drawn, unless otherwise stated, with the aid of the 
Camera lucida, and the ,,th oil-immersion of Leitz or the 2 mm. 14 apert. 
apochromatic objective of Zeiss and ocular 18. 


Fig. 1. Hye-spots from different individuals, showing the arrangement of the 
pigment-granules. 

Fig. 2. Side view of the anterior end of a motile cell, showing the flagellum and 
its enlargement in front of the eye-spot. This specimen was stained. 
in methyl-green and fuchsin for two hours, then washed in water and 
mounted in dilute glycerine. The whole of the red colour was washed 
out, the flagellum and its enlargement were stained light green; the 
eye-spot was left brown. 

Fig. 3. This shows the attachment of the bifurcate base of the flagellum to the 
dorsal wall of the excretory reservoir. The protoplasm formed a 
slight projection at the point of attachment, and was slightly more 
hyaline or refractive than the rest. 

Fig. 4. The flagellum is attached to the excretory reservoir nearer the base than 
in fig. 3, and each of its arms is slightly expanded at the base. The 
enlargement appears just at the place where the bifurcation begins. 

Fig. 5. A view of the anterior end of a cell as seen from below. 

Fig. 6. A slightly oblique view of the anterior end of a cell seen from above. 

Figs, 7 & 8. Two views of the same cell, showing the enlargement at some dis- 
tance from the eye-spot. 

Fig. 9. A freehand sketch of a portion of a cell in the resting condition. The 
excretory reservoir, the eye-spot, and the bifurcate base of the flagellum 
and its enlargement are shown. 


A482 MR. G. M. THOMSON ON 


On some New Zealand Schizopoda. 
By Geo. M. Txomsoy, F.L.S. 


[ Read 7th December, 1899.] 
(Pirates 33 & 34.) 


THouGH so many species of Crustacea have been described 
from New Zealand from time to time, yet the amount of col- 
lecting which has been done in the Colony, and especially in the 
North Isiand, is really very small. The result is that only a few 
individuals, representative of whole families, have in many cases 
been recorded. 

Trawling is now becoming common, and the opportunity of 
securing material in this manner will no doubt be more or less 
utilized. But up to the present, and excluding the fragmentary 
collections made during the ‘ Challenger’ Expedition, nearly ali 
the material hitherto studied has been obtained by shore collectors. 
The specimens described in this paper come under the same 
heading: they are the only Schizopod crustaceans which have 
yet been found in New Zealand waters, but probably the group 
will be found to be well represented. 


ScHIZOPODA. 
Fam. Mysip2. 


Genns SrrreLia, Dana. 


SIRIELLA DENTICULATA. (PI. 33. figs. 1-5.) 
Mysis denticulata, Thomson, Ann. & Mag. Nat. Hist. ser. 5, vol. vi. 
p- 1,§1880; Trans. N. Z. Inst. vol. xiii. p. 205, pl. vii. fig. 6, 1881. 

This species appears to be intermediate in most of its characters 
between S. Thompsoni, M.-Edwards, and 8. gracilis, Dana. I had 
provisionally included it in the former ; but Prof. G. O. Sars, to 
whom I sent specimens, considers it to be a new species. 

It resembles S. Thompsoni in the acutely pointed frontal pro- 
jection, in the large eyes with greatly expanded cornea, in the 
general form of the telson, and in size. 

On the other hand, its affinities with S. gracilis are seen in 
(1) the slender form of the body, (2) in having the last joint of 
the antennular peduncle rather short and stout, as in fig. 1, 
(3) in the form of the antennal scale, (4) in the divided propodal 


SOME NEW-ZEALAND SCHIZOPODA. 483 


joints of the legs, (5) in having the last caudal segment as long 
as the two preceding taken together, and (6) in the form of the 
apex of the telson. 

The adult specimens attain a length of fully 10mm. The 
antennal scale (fig. 2) is about three times as long as broad, the 
apex is obliquely truncate, and the outer margin is produced into 
an acute spine or denticle. The /egs (fig. 3) have the propodal 
joint distinctly divided into two articulations, the proximal being 
the shorter. The ¢elson (fig. 4) is rather slender, and tapers con- 
siderably towards the apex; the latter is very narrow and 
obtusely truncate, and bears three short spines between the 
rather long terminal spines of the margin, in this respect 
somewhat resembling S. gracilis. The uropods (fig. 5) project 
considerabiy beyond the telson. The inner plate is narrow- 
lanceolate, with the auditory apparatus very fully developed and 
the inner edge spinose. The outer plate is somewhat longer 
than the inner, and has the outer edge of the basal joint armed 
with spines, which in large specimens are. from ten to twelve in 
number, and which increase in size posteriorly. 

Hab. Ihave only met with this species in Otago Harbour, 
though it is probably common; but Mr. Suter has sent me one 
(mutilated) specimen from Lyttelton Harbour. In the Report 
on the Schizopoda of the ‘ Challenger’ Expedition, Prof. G. O. 
Sars states that both species of Siriella (S. Thompsoni and S. 
gracilis) were taken at the surface of the sea. The former has a 
very wide range, having been taken in the Atlantic and Pacific 
Oceans, and in the Tasman Sea; the latter occurs in the seas of 
the Indian Archipelago and in the Pacific Ocean. My specimens 
were taken in the dredge in 6 fathoms of water. I have only 
met with the species once. 


Genus TENAGOMYSIS, nov. gen. 


Generic characters.—Body slender, asin Mysis. Dorsal shield 
short, not covering more than half of the cephalothorax ; front 
obtusely pointed. Antennary scale narrow-lanceolate, setose on 
both margins. Mandibles with well defined molar tubercle 
palp with a much dilated second joint (Pl. 34. figs. 9 & 10) 
Maxille and maxillipedes as in Wysis. Feet rather slender and 
weak. 

Marsupial pouch in the female formed of three pairs of laminz 


484. MR. G. M. THOMSON ON 


increasing in size posteriorly, the first very small. Pleopoda 
in the female small and narrow, very feebly developed; in 
the male they are well-developed and two-branched ; in the first 
pair the inner ramus is very small and one-jointed, the outer 
of the usual natatory form; the remaining pairs have both rami 
long and subequal. Telson short, deeply cleft at the apex. 

Uropods narrow ; inner plate much shorter than outer, both 
densely furnished with marginal sete. Auditory apparatus well- 
developed. 

This genus appears to be intermediate between Leptomysis 
and Heteromysis, resembling the former in general structure and 
especially in the pleopoda and limbs. It differs from it, however, 
in the form of the front of the carapace, in the telson, and ina 
less marked degree in having the antennary scale one-jointed, in- 
stead of distinctly two-jomted. The resemblance to Heteromysis 
lies particularly in the dilated second joint of the mandibular 
palp. 

The genus is formed to contain a species which is not un- 
common on the coasts of New Zealand. 


TENAGOMYSIS NOVH-ZEALANDIA, n. sp. (Pl. 38. figs 6-8; 
Pl. 34. figs 9-17.) 

Form of the body linear, tapering slightly to the telson; 
length about six times the greatest breadth. 

Dorsal shield short, obtusely pointed in front in the median 
line ; evenly rounded on the posterior margin, only slightly pro- 
duced backwards on the lateral portions. 

Hyes \arge, cornea hemispherical, peduncles short and stout. 

Pedunele of the superior antenne with the basal joint subequal 
in length with the 2nd and 3rd together ; the appendage in the 
male is produced into a rounded process, furnished with a dense 
bunch of hairs at its base; inner flagellum less than half as long 
as the outer, very slender, destitute of hairs, and rather densely 
pigmented in great portion of its length (figs. 6 & 7). 

Scale of the znferior antenne obliquely lanceolate, more than 
twice as long as the peduncle, maximum width about one fifth of 
the length, densely furnished with plumose sete, which are very 
long on the curved inner margin; both the scale and the 
peduncle of the antenne are more or less ornamented with black 
stellate or frondose markings (fig. 8). 

Maxille of the first pair are normal. The second pair of 


SOME NEW-ZEALAND SCHIZOPODA. 485 


maxille have the terminal joint broadly obovate, and furnished 
with numerous strong setw on the outer margin. The mawilli- 
pedes and gnathopods present no very distinctive features. 

The legs are rather slender in structure, the propodos ending 
in a tuft of fine sete ; the exopodites are all well-developed, with 
a broad basal joint, bearing a ten- or eleven-jointed setose branch 
(fig. 12). 

The marsupial pouch in the female is formed of three pairs of 
lamelle, the anterior of which are very small, and the posterior 
very large. 

The pleopoda are rudimentary in the females. In the males 
the first pair has the inner ramus very short and one-jointed, 
obtuse at its apex, and with a lateral process produced across the 
long outer ramus; the latter is normally six-jointed (figs. 13 and 
14). The succeeding pairs have both rami well-developed, 
subequal in length and natatory, but the inner is always furnished 
with a process near its base which is produced more or less across 
the outer branch (fig. 15). elson short, only about half as long 
as broad, slightly narrowing at the sides, which are furnished 
with about twelve spines including the terminal one; the apex 
is deeply cleft, the sides of the notch being furnished with fine 
close-set pectinate spines and the centre defined by two long 
plumose sete (fig. 17). 

The uropods have the outer plate long and slender, somewhat 
obliquely truncate at the apex; the inner is about two-thirds as 
long as the outer, broadly lanceolate in form, and tapering to an 
obtuse apex, its inner margin has numerous spines between the 
sete ; both plates are densely setose on both margins (fig. 16). 

Length of the adult females, 10-16 mm. 

Hab. In the Kaikorai lagoon (brackish water), estuary of the 
Waikouaiti River, and rock-pools at Brighton, —all near Dunedin. 
Also dredged in the Bay of Islands from a depth of 8 fathoms. 


Fam. EUPHAUSIID#. 


Genus NyctrpHanss, G. O. Sars. 


NYCTIPHANES AUSTRALIS, G. O. Sars. 

Nyctiphanes australis, G. O. Sars, Report on the ‘Challenger ’ 
Schizopoda, p, 115, pls. xx. & xxi. figs. 1-7. 

Numerous specimens of this species were gathered by Dr. 
Benham and Mr. A. Hamilton at Port Chalmers. 

LINN. JOURN.—ZOOLOGY, VOL. XXVIL. 37 


A486 ON SOME NEW-ZEALAND SCHIZOPODA. 


The specimens originally described by Sars were all taken on 
the south and east coasts of Australia, “in the surface-net, and 
in most instances at night.” 


EXPLANATION OF THE PLATES. 


Puatse 33. 


Siviella denticulata x 43. 
Fig. 1. Eye and peduncle of antennule. 
2. Antennal scale, denuded of most of its setze. 
3. Leg of the 2nd pair. 
4. Telson. 
5. Uropod, denuded of sete. 


Tenagomysis nove-zealandiea. 


Fig. 6. Peduncle of antennule of 3, from below x43. 


fig ” »” of 2 ’ ” ~* 56. 
8. Antennal scale, x 43. 


Prats 34. 


Tenagomysis nove-zealandia. 


10. Mandibular palp. 

11. Second maxilla. 

12. Leg of the third pair of 3, x56. 
13. Ist pleopod, ¢, x 56. 

14. External ramus of same, x 125. 
15. 5th pleopod, 3, x 1286. 

16. Uropod, x 43. 

17. Telson, x 56. 


Fig. 9. Mandible. 
x< 125. 


dies 


ALS Huth,Lith. 


_ Linn. Soc.Jouaw Zoor.Von.XXVILP1. 33 


ees 


THOMSON. 


IES, SURE EIN [DIE NING WLANs 
6-8, TENAGOMYSIS NOVA_-ZEALANDIEA. 


A.S.Huth, Lith, 


Linn. Soc. Journ Zoor Vou.XXVII Pl. 34 


THOMSON. 


TENAGOMYSIS NOVA ZEALANDIE. 


ON THE STRUCTURE OF PORITES. 487 


On the Structure of Porites, with Preliminary Notes on the 
Soft Parts. By Henry M. Bernarp, M.A. Cantab., F.L.S. 


[Read 7th December, 1899.] 
(Puate 35.) 


In a paper lately read before this Society*, I endeavoured to 
ascertain the position of Porites in the Madreporarian system. 
A brief sketch of the structure was then given, sufficient to make 
the rest intelligible. The conclusion arrived at was that Porites 
resulted from a tendency towards very early budding, already 
noted in the Madreporide. This tendency, pushed still further, 
has produced a genus in which the budding takes place while 
the skeleton is still immature. In this way, the small size and 
the shallowness of the calicle, the perforation of the septa, 
and the reticular nature of the whole skeleton, which may be 
regarded as retrograde characters, can be reconciled with the 
presence of a flattened epitheca which, as elsewhere + explained, 
is characteristic of the highest Madreporarian specialization. 

In the present paper I propose to give an account of the 
structure of Porites in greater detail. During the year which 
has elapsed, my specimens have been examined and re-examined, 
and new structural details have come to light. One of them, 
namely, the discovery of the directive plane, and the bilateral 
symmetry of the calicle, will largely help to rescue the genus 
from the obscurity to which the smallness of its calicles and the 
complexity of its reticular skeleton have necessarily condemned it. 

I have also been able to cut sections of the polyps of a 
West-Indian form, one of the many from that region with low 
thick knobbed stems, alive only for a few centimetres at the top. 
A few brief notes on the first results of the microscopical 
examination of these are here appended. 


The Skeleton. 


Wall and Caenenchyma.—tThe distinction often drawn between 
these two ¢ has been due to the absence of any clear conception 
as to what the wall really is. I have already explained in my 
last paper how, in the Madreporide, the primitive epithecal 

* This vol., p. 127. t Journ. Linn. Soe., Zool. xxvi. p. 495. 

{ Compare, for example, Klunzinger’s statement that, in Porites, the calicles 
are united by the walls and not by.a ccenenchyma (Corallenthiere, ii. p. 39). 
The same is repeated by Martin Duncan, Linn. Soc. Journ., Zool. xviii., 1884. 


37 * 


488 MR. H. M. BERNARD ON 


wall became flattened out, and a secondary internal septal wall 
took its place, the living tissues clothing the whole of this septal 
wall down to the remains of the epitheca. This wall was 
primitively built up of the radially arranged Jaminate septa and 
their synapticular junctions. Secondary modifications occur, 
and the stiff radial arrangement of plates joined concentrically 
together dissolve down and change into a sponge-work in which 
the primitive elements are only just traceable. We then have 
a reticular wall which may be thick or thin. When acoral with 
such a septal wall buds, to form a stock, the reticular walls of 
parent and bud flow together. These combined walls form the 
ceenenchyma. The ccenenchyma can only be said to be absent 
when it is reduced to a minimum, 7.¢., when the calicles are 
separated by a single perforated plate. But, in reality, no 
sharp line can be drawn between the many degrees of thickness 
resulting from the fusion of the walls. 

It is true that an apparent distinction exists between the 
wall and the coonenchyma in certain cases, but in none will it 
bear examination. In Madrepora, the upper parts of the calicles 
(when young) project above the ccenenchyma, that is, above the 
fused basal parts of their walls. But as these calicles get older, 
the fusion usually rises, till, in the basal or older parts of most 
Madrepores, the calicles are quite submerged, and the fused walls 
and the coenenchyma are one and the same structure *. 


* Tt will be noticed that this description tends to limit the meaning of the 
word ‘ ccenenchyma’ to the fused outer or costal surfaces of purely septate thece. 
Its component elements, therefore, are septal and synapticular,—one might have 
said costal and synapticular. But, in these porous thece, division of radial 
structures into septal and costal portions can only be artificial. And it seems 
to me that the word ‘ costa’ had better be reserved for external ribs which are 
somewhat more naturally separate from the septa. 

If the term ‘ coenenchyma,’ is so limited, it not only excludes such tissues as 
that in which the calicles of Galaxea are embedded, which is epithecate in 
origin, but also all the stray proliferations of the skeleton which are frequently 
met with in the Madreporide and elsewhere. For instance, in Alveopora 
there can be no coste at all, the walls between adjoining calicles being 
morphologically equivalent to interlacing septal spines (Journ. Linn. Soc., Zool. 
xxyi. p. 495). And yet, in rare cases, on the undersides of stocks, a curious 
proliferation of the walls sometimes takes place, so that the calicles may be 
separated by a coarse reticulum almost like a normal coenenchyma. All such 
adventitious proliferations of skeleton I propose to call a ‘ pseudo-ccenenchyma.’ 
They are mostly found in the basal parts of stocks, where normally, as is well 
known, the basal skeleton merely thickens without forming any additional 
framework. 


THE STRUCTURE OF PORITES. A489 


In Montipora and in Porites, the calicle-walls fuse for their 
whole height, and the fosse are merely sunk into the common 
reticular skeleton. In Montipora itis impossible to assign definite 
portions of this reticulum to the individual calicles; but 
in many Porites this can be done, the surface being marked 
off into polygonal areas, each area being in close contact with 
those adjoining it, and with its surface often sloping inwards 
towards the fossa in the centre. Hence it has been stated that 
Montipora has a ccenenchyma, but Porites little or no ccenen- 
chyma (Milne-Edwards and Haime*; see also quotation from 
Klunzinger, ante, p. 487, footnote). An extended survey of 
Porites shows that this distinction is quite artificial. On many 
of the forms with these thick-walled calicles marked off into poly- 
gonal areas, these areas become gradually invisible in the older 
parts of the stock; while forms in which no areas are traceable 
at all, and in which the calicles are sunk straight into a reticular 
skeleton as they are in Montipora, are quite common. Milne- 
Edwards suggested that perhaps these latter should be placed in 
anew genus. This suggestion was carried out by Verrill 7, and 
further emphasized by Klunzinger, who placed Synarea, Verr., at 
the very end of the Poritide, because it alone had a ccenenchyma ; 
whereas, as above stated, calicles marked off into areas and 
calicles sunk in a level ccenenchyma can frequently be found on 
one aud the same stock ¢. 

Passing in review the various walls found in Porites, we shall 
see again how impossible it is to separate a group as a new 
geuus merely on account of the great thickness and level tops of 
their walls. 

Thin Membranous Walls—These walls have very different 
appearances according as the intrathecal skeleton rises to the 
level of the wall or sinks down below that level. In the former 
case we have a surface like that shown in fig. 5, Pl. 35; 
only here the skeletal elements are somewhat thickened. In 
these cases it is common to find the edge of the wall incom- 
plete, the calicles communicating freely one with the other. 
These communications doubtless become perforations as the 
stock thickens. 

In these thin-walled forms the wall itself is composed of a 

* Ann. Sci. Nat. 3rd ser., xvi. 1851, p. 24. 

+ Bulletin Mus. Comp. Zool. i. 1864, p. 42. 

{ For another supposed generic distinction between Porites and Synarea 

ee p. 494 footnote. 


490 MR. H. M. BERNARD ON 


single ring of synapticular bars or plates, and these may be 
arranged either zigzag round the calicle or in straight lines. 
In the latter case, the lines form the sides of polygons. These 
straight walls are, I expect, secondary; for, regarding the reticular 
wall as primitive, the zigzag wall would be the more natural 
derivative of such a reticulum. The zigzag line is formed by 
single synapticular bars joining the alternating costal edges of 
adjacent calicles. All stages of the straightening of this line 
can be found. It is the straight wall which rises above the 
surface as a thin membrane-like edge, and, in extreme forms, 
may give the whole surface an alveolate appearance. The 
whole intrathecal skeleton may remain deep down in the base 
of the membranous pits, or septal strie may slightly thicken 
the walls and serrate their edges. 

Reticular Walls—Of these there is an immense variety, 
and again the appearance is very. different according as the 
intrathecal skeleton rises to the level of the wall or is sunk down 
below that level. They are thick and thin, round-topped or with 
a sharp median ridge; or again, if the calicle-depressions are 
cylindrical, the intervening angles may be thick and reticular, 
while at the pomts where the calicles touch one another, the 
wall may be reduced to a single lattice-work. Of the thick walls, 
we have already referred to the variety in which straight median 
ridges mark off in polygonal lines the areas belonging to each 
calicle. These areas may sink inwards funnel-shaped towards 
their central fosse. Or, again, they may be quite level. In 
this case, the median ridge frequently disappears, and we have 
the conenchymatous group (formerly Synarea, Verrill). The 
most interesting section of these is formed by those in which 
the thick walls rise up secondarily into ridges or papille (fig. 6, 
Pl. 35), very similar to those found in Montepora. ‘This is an 
interesting case of similar specialization arising under similar 
conditions. But, in Montipora, the thick reticular walls are 
characteristic of the genus: hence these coenenchymatous 
developments on the tops of the walls are far richer and more 
varied than they are in Porites, in which the thick reticular 
wall is confined to a group only. I have so far found no traces 
of the ‘ tuberculate’ specialization * of the cceenenchyma which is 
so widely developed in Montipora. 

* Of. Brit. Mus. Madrep. iii., Introd. p. 9: also for figures, Ann. & Mag. 
Nat. Hist. xx. (1897) p. 117, pl. ii. 


THE STRUCTURE OF PORITES. 491 


These secondary ccenenchymatous developments have given 
rise to some confusion, their morphology not being always clear, 
especially when the calicles are crowded and the ridges appear 
as mere upward extensions of the walls. In the ‘ Challenger’ 
Report on the Reef Corals (xvi., 1886), two new forms, P. crassa 
and P. latistellata, Quelch, are described ; but, according to the 
classification there adopted, they should have been placed in the 
genus Synarea, which was established to contain all the Poritid 
forms with ridges or papille rising from the walls between the 
calicles. P. latistellata was so named because the individual 
calicles were measured as if these secondary ccenenchymatous 
ridges were the tops of the walls; and in an ailied form, where 
the ridges ran so as to separate the calicles into short linear 
series in the bottoms of narrow valleys, such series were thought 
to be the result of intracalicinal gemmation. This led Mr. Quelch 
to found a new genus, Napopora. Recent acquisitions by the 
British Museum have supplied us with links sufficient to connect 
the types of the new genus specifically with P. latistellata. 

A few of these different types of wall will be seen in Plate 35. 
Representatives of the two extremes can be seen and compared 
in figs.5 and6. How impossible it is to make generic distinctions 
between them, may be gathered from the fact that specimens 
occur in which part of the stock has walls even thinner than 
those shown in fig. 5; while another part of the same stock has 
walls as thick as those shown in fig. 6, although without the 
special coenenchymatous papille. 

The finer texture of the walls can be best discussed in con- 
nection with the septa. 

The Septa.--The number usually assigned to Porites is twelve, 
and it seems quite possible to separate Porites from Goniopora 
mainly on this point, Gontopora having typically 24. Dr. Verrill 
ascribed “‘12, sometimes 12 to 20, rarely 24” septa to Porites ; 
but I have found it better to class every Poritid with more than 
12 septa in the genus Goniopora, as Dana proposed. Mr. Quelch, 
again, claimed 24 septa for Porites (Chall. Rep. xvi.), on account 
of his species P. mirabilis, in which calicles occur with 3 cycles 
of septa. These, however, are obviously the large double calicles, 
one or two of which can be found on almost any stock and 
must be regarded as abnormalities. One is seen in fig. 5, 
P]. 35. ae 

With regard to the development of the septa in Porites 


492 MR. H. M. BERNARD ON 


no comprehensive survey has ever yet been attempted. Milne- 
Edwards and Haime described them as usually hardly distinct 
from the pali.* I give, in the accompanying diagrams, a series 
of their more important variations. These are found so in- 
timately linked together, more than one being observable on 
the same stock, that no generic distinctions can possibly be 
built upon them. 

1. Twelve distinct septa end freely and separately round the fossa. 


Diagrams showing the principal variations in the septal and palic 
formule in Porites. 


Slight swellings of their inner edges may or may not indicate a 
disposition to form pali. They may or may not be distinguishable 
into two cycles. Forms with such septa, of which the best known 
example is P. astreoides, Lamarck, were grouped into a separate 
genust Neoporites by Duchassaing and Michelotti. But the 
disposition to form pali and also to pass into the condition shown 
in fig. 2 renders it impossible to admit any such distinction. 


* Ann. Sci. Nat. 3rd ser., xvi. 1851, p. 25. 
t Or ‘subgenus,’ Pourtalés, Bull. Mus. Comp. Zool. iv. 1871, p. 85. 


THE STRUCTURE OF PORITES. 493 


2. The septa meet and fuse, and always in the way shown, two 
pairs on each side of a line passing through two opposite septa, 
the significance of which will be seen presently. 

3. The fusion of septa goes still further, and, in addition to 
the two pairs, a triplet is formed (fig. 3). Pl. 35. figs. 5 and 
6 show these fusions fairly clearly, and probably careful study 
might make them out on some of the other figures, but they 
ean naturally be best seen in forms in which the upper edges 
of the septa are nearly on a level with the walls. I have never 
found any other fusions of septa but these. And here we 
may note that we obviously have the directive plane in the line 
passing through this triplet and the columellar tubercle. The 
calicle is divided into two symmetrical halves; and whether the 
two eycles of septa are distinguishable or not in size and 
development, we can always now ascertain which are the 
primaries and which the secondaries. 

4 & 5. The septa which are best developed im fig. 1 become 
usually poorly developed in figs. 4 and 5 (diagr.), and their upper 
edges are interrupted. A portion, frequently only a granule, 
shows near the wall, and another portion appears as a palus at 
the tip of each septum. I propose to cali the peripheral portions 
of the septal edges, the septal granules. Their variations are 
found to supply new and valuable taxonomic characters. 

In these figures, the spaces between the septal granules and 
the pali are exaggerated, the figures not being intended to 
be more than diagrams. 

We now come to the pali, which are so very characteristic 
of the genus, but on the arrangements of which no light has 
hitherto been shed. 

The Pali.—These structures, though well shown in one of the 
earliest figures of Porites *, attracted no attention till Dana 
described them in his ‘ Zoophytes’+. He speaks of an inner and 
an outer ring of points, and adds that sometimes one of the 
inner unites with two of the outer to form a \-shaped palus. 
Since that time, the pali have always been treated as features 
of taxonomic importance, but nothing could be said about them 
than that their numbers were 5, 6 or more, and that they 
were large and prominent or the opposite, and occasionally 
V-shaped. 

* Hillis & Solander, ‘ Zoophytes,’ 1786, pl. 47. fig. 2. 
Tt P. 550. 


494 MR. H. M. BERNARD ON 


Reference to the digrams shows that they occur in a definite 
order, and that, in their development, they are closely associated 
with the fusions of the septa. 

Passing over the cases of those Porites with septa arranged 
as in fig. 1 (p. 492), and each with a slight paliform swelling, and 
therefore with traces of 12 pali, the number is limited to eight as 
shown in fig. 4. Of these eight, the four which arise at the 
points where the septa meet in pairs are usually much larger 
than the rest (Pl. 35. fig. 1). They may be very large indeed 
as compared with those arising from single septa. When the 
large palus is formed by the fusion of the three septa, as shown 
in fig. 5 (diagr.), we have, with the four large ones at the points 
of fusion of the pairs, five large pali. Whenever we have a ring 
of five large pali, we know where and how they arise, viz., at 
the five points in which the septa fuse (see many of the rings 
of pali, Plate 35. figs. 3 & 6). Among these will be seen 
many in which there is a small extra palus, making five principal 
pali and one minute palus. The last is that on the directive 
septum, which remains single as shown in fig. 5 (diagram). 

In Plate 35. fig. 6, it is easy to see the large pali arising from 
the points of fusion of the septa, and here and there having the 
V-shape described by Dana. But this arises not from the fusion 
of granules, as the great American naturalist believed, but from 
the fact that the fusing septa are slightly exsert *. 

Ag the pali will have to figure in all future specific descriptions, 
it is necessary to name them. I propose therefore to call those 
which arise at the points of fusion of the four pairs, the four 
‘Principals. The directive palus which often takes part in the 
triple fusion may be called the ‘ Directive principal. The rest 
may be called Supplementaries, and we have a directive supple- 
mentary and two lateral supplemeutaries ; one or both of these 
Jast-named may fuse with the directive principal. 

It should be noted that no palic formula seems to be constant 
throughout all the calices of a stock. When describing the 
formula of any species of Porites, all that is meant is that there 
is apparent in the colony a tendency to produce that particular 
formula. 

Having now described the septa and the pali, we may return 

* This prominence of the pali and septa in many of the Porites with 


developed coenenchymas was thought to be another generic distinction of 
Synarea, but the point is an unimportant one. 


THE STRUCTURE OF PORITES. 495 


to the finer texture of the walls which are built out of these 
septa, with their synapticular junctions. 

It is quite correct to say of Porites that the stiff radial and 
concentric symmetry of septa and synapticule which characterizes 
the typical Madrepore is here melted down into a reticulum ; 
a reticulum more loose because of the perforate and incomplete 
character of the septa. 

We find, however, interesting variations in this respect. Not 
a few Porttes still show traces of the stiff radial septa radiating 
outwards on the top of the walls as so many short ridges, e. 7., 
slightly seen in fig. 4, Pl. 35. These sometimes run from calicle 
to calicle, but are always slight and never approach the fine 
systems of parallel striz which are so well developed over the 
whole ccenenchyma in Madrepora and Turbinaria,as unmistakable 
evidence of the part which the laminate septa (or cost) play in 
the construction of the walls. 

The first stage of dissolution is probably that in which the 
vertical elements persist as trabecule or upright threads which 
end above the surface as granules. This is the most common 
condition of the surface of Porites. The connection between the 
walls can then be seen from the fact that the wall granules are 
only a repetition of those seen within the calicle—septal granules 
and pall. 

Not infrequently the horizontal elements become flattened 
flakes, so that the vertical section shows tiers of floors supported 
by short pillars. 

The last stage is that in which both vertical and horizontal 
elements melt down entirely into either a sponge-work, or into a 
system of flakes mostly lying horizontally. In these cases we 
do not usually find the surface covered with granules, they 
are present only so far as the vertical elements, the tips of 
which constitute these surface-granules, continue to be developed 
as pillars. 

The Columellar Tangle and Tubercle.—The base of the calicle in 
Porites always fills up sooner or later with a mass of reticular 
tissue, as emphasized by Dana*. This tissue may be conveniently 
called the Columeilar Tangle. The most symmetrical manner in 
which this can develop is as a regular ring joining the septa and 
pali. It rises to various heights. It may be very simpie and 


* Zoophytes, 1848, p. 117. 


496 MR. H. M. BERNARD ON 


open, or else, especially in cases where the horizontal elements 
are flaky, it may appear solid. I have never seen the columellar 
tangle as such protuberant or convex in Pordtes. 

And here it is worth noting that there appears to be some 
trace of dimorphism in the calicles of a stock, while in some the 
centre of the columellar tangle fills up with cross bars, in others 
the latter remains a deep open pit. These two forms on one and 
the same stock are so frequent, that they suggest some definite 
physiological significance (cf. infra). 

The tubercle which rises from the columellar tangle but is 
absent when the centre is hollow, is, as above noted, often found 
flattened in the directive plane and here and there joined to one 
of the directive pali. It is the equivalent of the directive keel 
seen bisecting the columellar tangles in many Turbinaria. It 
appears as if, in the earliest stages of budding, the directive septa 
met straight across the calicle, and that this columellar tubercle 
is the remains of the connecting link. This conclusion commends 
itself from the fact that a close study of columellar tangles, as 
in Turbinaria, where they are specially well developed, shows 
that they are primitively built up of the usually curled lower 
edges of the septa. 

These comparisons help to confirm my view of the relationship 
of Porites to the Madreporids. In the case of Madrepora, the 
directive plane in the buds falls in with the radial symmetry of 
the parent, and might almost be regarded as coincident with that 
of one of its cost. In Turbinaria although, in large stocks, the 
directive keels point almost all ways, yet, near the margins, 
they are often seen all pointing towards the growing edge. I 
have been fortunate enough to see this phenomenon in a thin 
explanate Porites (fig. 4, Pl. 35). In this figure the great 
majority of the directives point nearly uniformly up and down, 
that is, in the line of growth. If we refer to the diagrams, the 
direction of growth runs out between the four principal pali as 
indicated by the arrow (diag. 4 & 5, p. 492). 

Whether these directive planes, when found twisted all ways 
on a stock, indicate special relationships between adjacent indi- 
viduals of the colony it is impossible to say. I should be inclined 
to think that the individual as such is submerged except in 
Madrepora, and that the buds come from the common ccenosare. 

Before leaving this columellar tubercle, I would draw attention 
to the fact that I have inserted it in all the diagrams as a per- 


THE STRUCTURE OF PORITES. 497 


manent feature of the skeleton. While I believe it to have been 
a primitive element of importance, it is not now universally 
developed. I have already mentioned those scattered calicles 
which occur on most stocks in which the fossa is a deep pit, 
but, apart from these, we find in very deep calicles that the 
pali and the columellar tubercle are sometimes wanting. There 
can be little doubt that there is some correlation between the 
depth of the calicle and the development of pali. The same 
phenomenon occurs also in Goniopora. Stocks occur in which 
ereat variation exists in the depths of the calicles; for instance 
fig. 2, Pl. 35, is taken from one side of a stock in which the 
calicles on the opposite side are much deeper and show only the 
faintest traces of pali. 

It is not surprising, then, that it is just in the deeper calicles 
with all septa free, as is shown in diagram 1, that the columellar 
tubercle is most frequently absent, and the pali often only slightly 
traceable as faint swellings. But cases are not wanting in which, 
while the septa are free and the pali hardly traceable, the colu- 
mellar tubercle is well developed: I have found this in some West 
Indian species. The combination is interesting, because a Porites 
without pali but with a columellar tubercle exists in the Berlin 
Museum, and was named by Ehrenberg P. punctata. Milne- 
Edwards & Haime first suggested a new generic name for it, and 
called it Stylarea Mullert. But these authors suppressed this 
name the same year and reverted to Ehrenberg’s P. punctata. In 
recent years, however, Dr. Klunzinger* has again revived the 
genus, re-naming the original specimen Stylarea punctata, and 
placing it next to Porites. I find it necessary to add this genus 
Stylarea to the list of apparently needless genera given in my 
previous paper. For, not ouly is there nothing specially startling 
in the absence of pali and the presence of the columellar tubercle, 
but, among the many other variations presented by Porites, we 
actually have specimens in which the columellar tubercle is well 
developed while the pali are here and there only faintly traceable. 
As against the advisability of making a new genus on the original 
specimen of Ehrenberg, I should like to point out that it is so 
small that, as Ehrenberg suggested, it might easily be a young 
form in which adult conditions are not yet fully developed. I 
have frequently observed that young, and perhaps very rapidly 


* Corallenthiere, ii. 1879, which see for other references. 


498 MR. H. M. BERNARD ON 


growing stocks have characters very unlike those of the larger 
stocks close to which they are growing, and from which it is 
probable, though not always certain, that they have been derived. 

Growth-forms.—V ery young colonies consisting of a small mass 
of reticulum fillmg up an epithecal saucer are frequently met with. 
I have, however, never met with one in which the parent calicle 
was still recognizable. Such, however, must of course be postu- 
lated. The appearance, in cases in which the wall is reticular, is 
almost indistinguishable from that shown by young Montipores 
(see figs. 1 & 2, Ann. & Mag. N. H. (6) xx. pl. ii.). I have not 
found any young colonies of forms with membranous walls. <A 
knowledge of the early colonies would be very desirable: con- 
sidering the ease with which, in a large stock, one type of calicle 
passes into a very different type, we might expect them to vary 
considerably from the adult. 

The budding of shallow, saucer-like calicles from the sides of 
other shallow calicles is not likely to be very plastic. But among 
other causes of form-variation, we may note the local thickening 
of the walls. When this is very irregular and confined to small 
groups of calicles, it leads to the formation of bosses and knobs 
from which branches are easily developed. In this way purely 
branching forms many have been evolved, but the branches are 
for the most part thick and coarse ; elegantly branching forms 
are rare. 

Considerable variation occurs as to the depth to which the 
colony descends in branching forms. We find all extremes: 
merely the tips fora centimetre or two may be alive, or the living 
layer may extend right down to the base of the stock, twenty 
centimetres or more. 

As reef-builders, this genus of Stony Corals has long been 
famous. It is a conspicuous component of the outermost edge 
of the reef where the surf is most violent. The unfavourable 
conditions of existence at such spots may supply us with a 
clue to the dwarfing of the polyps, this having resulted in the 
building-up of almost solid coral-masses. Hence again, though 
branching forms are fairly numerous, they are insignificant as 
compared with the rounded masses of almost solid coral-rock 
(often many feet in diameter) which are most frequently met with. 
In the West Indies, we read of thick tangles of branching Porites 
spreading over the surface, but the branches are coarse, thick and 
matted together. : 


THE STRUCTURE OF PORITES. 499 


Dana thought that there were no foliate forms, the nearest being 
the sublamellate growths due to the fusion of the branches. 
There are, however, a few thin explanate forms which approach 
the foliate condition. But there is certainly no rich foliation 
such as we find in the foliate group of the Turbinarie and, only 
in lesser degree, in the Montipores. 

The various growth-forms pass so gradually into each other 
that it is only possible to group them in series. I place the ex- 
planate forms, which admit of being described separately, as the 
first division. The second consists of those explanate and en- 
crusting forms which throw up lobes and columns; these vary 
imperceptibly in two directions: (a) into columns, and (0) into 
branching forms, both without encrusting bases. The third group 
consists of glomerate forms beginning (a) with those with edges 
expanding while the centre thickens, and may either, by con- 
tinuous growth or by fresh relays, form great hemispherical 
masses, and ending (0) with those in which the stock develops at 
once as a rounded mass. ‘These again are often difficult to 
distinguish from one another: and hard and fast distinctions are 
impossible. 


Some Preliminary Notes on the Soft Parts. 


In the diagram of Porites which I sketched in my former 
paper™, in order to compare the structure of the skeleton upon 
its flattened epitheca with that of a Madreporid, the porous septa 
were purposely drawn very low. The diagram represents only 
an ideal parent calicle of a Porites. As a matter of fact, while, 
in very many forms, the calicle-depression is like that there shown, 
the underlying reticular skeleton occupied by the soft tissues is 
very much deeper. 

In sections of dried Porites, the staining of the living tissues 
penetrates some 3-4 mm. beneath the surface, and, if a piece be 
decalcified, the soft parts are left as a fleshy reticulum of nearly 
even thickness (3-4 mm.), from the surface of which the polyps 
project. The enteric cavities of the compound stock are simply 
a network of fine canals. 

On decalcifying a fragment of P. recta (?) from Jamaica, ' 
beneath the fleshy rind was found a cloudy mass of hyphe, 
threads of which had run up through the skeletal reticulum into 


* This vol., p. 135, fig. 1. 


500 MR. H. M. BERNARD ON 


the septa and pali. These threads kept the hyphal mass 
attached to the rind after the skeleton was dissolved away. 
Several different kinds of fungoid growths can be distinguished, 
many presenting appearances so interesting that the whole will 
be submitted to specialists in that branch of study, in the hope 
that some new biological facts will be forthcoming. 

Further, in the clear spaces left by the decalcified skeleton, 
are sections of an organism which is almost certainly a ciliate 
Infusorian. These are essentially like those figured by Moore 
of Spirostomum (Journ. Linn. Soe., Zool. xxiv. pl. 27),—an open 
angular network with staining granules at the nodes, the cilia 
not staining and passing out through a thin, deeply staining 
membrane which, in sections, is broken up by the cilia into a row 
of dots. How Infusoria live in the apparently solid coral I do 
not know. It is, however, worth recording that one not infre- 
quently finds the thinnest skeletal framework completely hollowed 
out by a system of thin-walled tubes, so that it is in reality not 
solid. JI have never hitherto found any clue to this phenomenon. 
The discovery of a large Infusorian in the spaces of a Porites- 
section which are, in life, occupied by skeletal bars, adds another 
to the organisms, Sponges, and Fungi, to whose agency the 
excavations above mentioned might possibly be due. 

There seems to be great variation in the shapes of the polyps, 
both contracted and protruded, judging from the published 
figures. But it is very doubtful how far any value can be 
attached to these variations. 

Nevertheless some of these differences should be noted. 
Lesueur *, who first figured the polyps of Porites, shows three 
different kinds in a species called by him “ P. astreoides,” pos- 
sessing a large disc, and short round tentacles each with a distal 
black point. Duchassaing? also figures a Porites without name, 
with large convex dise and short, slightly knobbed tentacles each 
with a black distal poimt. This so far agrees with the above- 
mentioned figure of Lesueur. Agassiz{, on the other hand, 
gives five figures of polyps of “P. astreoides,” viz., a young 
one, in which no tentacles are yet seen, and four adults in which 
the arrangement of the parts are not at all clear. The ring of 
lobes drawn round the mouth can hardly be the tentacles, while 


* Mém. Mus. Paris, vi., 1820. 
t Ooraliaires des Antilles, Suppl. 1864, pl. viii. fig. 2. 
t Florida Reef, 1880, pl. xvi, 


THE STRUCTURE OF PORITES. 501 


the peripheral fringe looks very like the mesenterial furrows 
which run down the sides of most Porites. The escape of a 
planula is shown. 

The other two polyps feared by Lesueur, P. recta and “ P. 
clavaria” (non Lamarck), have much smaller discs and short 
tentacles inclined to be pointed. Dana figures the polyps of 
P. levis as contracted down flush with the surface, the external 
mesenterial furrows radiating like spokes round the ring of knobs 
representing the contracted tentacles. The disc is fairly large 
and only furrowed by six mesenteries, whereas in most of the 
figures referred to, at least in which any furrowing of the disc is 
at all marked, the disc is furrowed by the full number of mesen- 
teries. This is the case also in the figures of the long polyps of 
“ P, furcata” given by Agassiz, in which the tentacles are thin, 
fusiform, and pointed. Saville Kent, in his ‘ Great Barrier Reef,’ 
figures the polyps of three Australian species, with thin, cylin- 
drical tentacles about as long as the diameter of the disc, each 
tentacle ending in a distinct spherical knob. The only specimen 
examined by myself which I would provisionally classify with 
P. recta, Lesueur, has rather a narrow column which suddenly 
enlarges to carry the twelve tentacles, which stand erect and 
short, stout and round-topped; the disc was rather small. 

Agassiz (J. c.) first figured the nematocysts, or rather the coiled 
threads of the nematocysts. On my sections these occur chiefly 
in small groups raised into hemispherical batteries. One large 
battery occurs at the tip of each tentacle, and a row of smaller 
batteries runs downits inner face. The stinging-threads were all 
I could see, and they were mostly coiled in more or less conical 
spirals, the cones pointing inwards, and each beneath what appears 
to be a small round aperture on the covering membrane of the 
battery. Between these batteries the ectoderm was largely com- 
posed of slime-cells. 

In addition to these small ectodermal stinging-threads, the 
cavity of the polyp contains great numbers of long, membranous 
sacs some 40 pin length, each with a long coiled thread; the coil is 
never a regular spiral, and the membranous sac is often collapsed 
upon the thread. No nucleus or communication with the exterior 
could be found, the bodies being loosely attached to the endoderm 
in great numbers in the tentacles, but also, though in smaller 
numbers, on the mesenteries. Examined with avery high power, 
the thread, which was as thick as the whole coil of the ordinary 

LINN. JOURN.—ZOOLOGY, VOL. XXVII. 33 


502 ON THE STRUCTURE OF PORITES. 


nematocyst (7. e. about 2 yx), appeared to have spiral lines running 
round it, and, when cut across, I saw it on more than one occa- 
sion hexagonal with the sides inwardly curved. Whileit is fairly 
safe to assume that these are defensive weapons *, more extended 
researches with better preserved material would be necessary to 
throw light on their structure and to discover their origin and 
the nature of their contents and method of discharge. * 

The internal tissues were much disorganized (1) by the sym- 
biotic alge and (2) by the great quantities of slime. The former 
were large, very numerous, and often found dividing. They 
occurred in greatest numbers in the extensible or projecting parts 
of the polyp, that is, doubtless, where the light can reach them, 
although they also occur scattered among the ccenosarcal canals. 

The slime seemed to have filled the internal cavity with a net- 
work of darkly staining strands, quite different from the bright 
carmine of the ectoderm-cells of the gullet. 

Among the fragments of the ccenosare which appear in the » 
sections separated by the clear spaces left by the decalcified 
skeleton, the interseptal loculi can be made out by their radial 
arrangement, and by the presence of the mesenterics, which are 
here correspondingly narrowed. Different conditions would 
doubtless be found in other forms in which the intrathecal 
skeleton did not rise to the level of the walls. 

Small as this contribution to the subject is, my work on the 
sections having been unexpectedly interrupted, it is enough to 
show the desirability of an extended study of the soft parts of 
different species of the genus. 


EXPLANATION OF PLATE 35. 

The six photographs here reproduced are from negatives kindly lent, for the 
illustration of this paper only, by the Trustees of the Natural History 
Museum, All are enlarged five times. 

Fig. 1. Porites with full number of pali typically arranged ; the directives 

point in all directions. 


* At the suggestion of my friend Prof. Howes, I have compared these organs 
with figures of large cells with thick, coiled threads given by Weymouth 
Reid (Phil. Trans. 1894, B) for the skin of the Hel, and Goodrich (Q. J. M. Sci. 
xxxix.) for the coelomic corpuscle of the Oligochxte Enchytreus. As Prof. 
Howes points out, these latter seem to be curious modifications of ordinary 
slime secretions. While I can trace only slight structural resemblance between 
these cells and those described above and Porites, the fact that here again 
they are associated with immense numbers of slime-vells suggests a line of 
enquiry which might be followed up. 


LINN. Soc. JOURN. ZOO, WOkL, XOKWIN Pit, BSD: 


BERNARD. 


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WEAVE, STRUCTURE OK IPORININES. 


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ON THE ATR-BLADDER IN NOTOPTERUS. 203 


Fig. 2. The form referred to on p. 488; on one side of the specimen the pali 
are developed, and on the other the calicles are deep and the pali quite 
obscured as a definite system. 

Fig. 3. A form with the more usual ring of five pali. Traces of the others and 
of the columellar tubercle can be seen. 

Fig. 4. A leaf-like specimen of P. exilis, Gardiner, showing the directives all 
pointing in the direction of growth (towards the top edge of the 
fioure). 

Fig. 5. A form with thin wavy walls and showing a double calicle; the colu- 
mellar tangle rises to the surface, and here and there unites the pali in 
a ring. 

Fig. 6. A ceenenchymatous form, the ceenenchyma rising into rounded ridges 
and papille ; the pali are slightly exsert and are often slightly V-shaped. 
In a few cases the directive principal and its adjacent lateral supple- 
mentaries can be seen forming a very blunt broad arrow (the triple 
fusion mentioned on p. 490). 


The Air-bladder and its Connection with the Auditory Organ 
in Notopterus borneensis. By Prof. T. W. Bripae, Sc.D., 
E.L.S., Mason University College, Birmingham. 


[Read 21st December, 1899. ] 
(PLates 36 & 37.) ; 


CONTENTS. 
Page 
Tis, JIMWTROUNCEWON “hon saonacocéodcodcosucee aaa OCH e NEE RED REM CE ter cruearacadoccadonad 503 
il. General Structure and Relations of the Air-bladder ..................006 504 
il. Internal Structure, and the Relations and Attachments of the Air- 
loladderitoxtheySkeletonimreeeresseereeeae ence renee cee eee eee eee 507 
iy. Relations of the Air-bladder to the Skull and Auditory Organs _...... 513 


yv. Cranial Fontanelles and their Relation to the Auditory Organ...... .«- O16 


vig “ae ANtchiuomny7 Oran, ih scensenqdagnacdadcoseedepeonenedeneedeonsoscuoooosodabos 518 
vii. Comparison with Notopterws Pallasti .........cccccccesceceeceeeee AnagauOGoC 519 
Wii, (ClommpeREOMn YAN Oiluee WENOSE — coccoapocosodusoon de seosocabsocooeDoscNDOCS 522 
ix. Remarks on the supposed Auditory Function of the Air-bladder ...... 531 
say JBMMbKoTER OOK? GooadendgacdosoohodasuosdodcokedoLcaEpEraedcce seesonadsoddeacdecsoue 53k 
xi. Explanation of the Plates, Reference-letters .............ss-eceseeseceenes 539 


I. Inrropvcrion. 
Waite recently dissecting the air-bladder and associated struc- 
tures in a specimen of Notopterus borneensis, Bleeker, certain 
features were noticed in which this species differed from Woto- 
pterus Pallasi, C. & V.,as described by Cuvier and Valenciennes 
(4. pp. 189-141). How far the differences observed are due to 
ashe 


504 PROF. T. W. BRIDGE ON THE 


imperfections in the account given by the two distinguished 
French zoologists whose names have been mentioned, or to the 
existence of genuine variations in which WV. borneensis deviates 
from WV. Pallasii, [am not in a position to decide. Nevertheless, 
and apart from the question of specific variations in the structure 
of a particular organ, there are other features in the air-bladder 
of WV. borneensis, and more especially its connection with the 
auditory organ, which, perhaps, are net without importance to 
those interested in the study of the obscure but fascinating 
problem of subaqueous audition. For these reasons I venture 
to give a brief account of the air-bladder of WV. borneensis, and, 
as the organ has not bitherto been figured in any species of the 
family, to supplement the description by suitable figures. 

I desire to express my grateful thanks to the Council of the 
Royal Society for a grant from the Research Fund in aid of this 
and other kindred investigations. 


(J. Tur Generar Srructtre aNnD RELATIONS OF THE 
ATIR-BLADDER. 


The specimen examined was 38 cm. long, and to this length 
the postanal portion of the body or tail contributed 28 em. 

For convenience in description, the air-bladder may be said to 
consist of three well-defined portions, which, from their regional 
disposition, may be termed (1) the anterior or pre-ccelomic, 
(2) the abdominal or ceelomic, and (3) the caudal or post-ccelomie. 


Se 


ud 


Lateral view of the air-bladder of Nofopterus borneensis. About two-thirds 
nat. size. a@b.p, abdominal portion ; /.c.c, left caudal cecum; d.p, ductus 
pneumaticus; fc, filiform ceca ; /.a.c, left auditory caecum ; @s, esophagus ; 
sb.s, subspherical sac; ¢.p, its tubular prolongation; v.d, ventral diver- 
ticulum. 


The abdominal portion (see fig., ab.p.) is more or less cylin- 
drical in shape, but much deeper behind than in front, and 


ATR-BLADDER IN NOTOPTERUS BORNEENSIS. 505 


occupies the usual position beneath the vertebral column, being 
separated from the latter by the kidney. The ventral surface 
only is invested by the peritoneum, the line of reflection of this 
membrane on to the inner surface of the abdominal wall being 
coincident with the junction of the lateral with the ventral wall 
of the organ. Posteriorly, the peritoneum is reflected down- 
wards on to the hemal arch and spine of the first caudal vertebra 
and the first radial element (‘“interspinous bone” or “ pterygio- 
phore”’) of the anal fin, and therefore does not accompany the 
caudal prolongations of the air-bladder. 

At its binder extremity, immediately anterior to the first 
radial element of the anal fin, the abdominal portion of the 
bladder divides into two caudal cxea (J.c.c.), each of which at its 
commencement, if not so wide, is nearly twice the vertical dimen- 
sion of the former. The two ceca extend backwards on opposite 
sides of the tail, and, gradually contracting, finally terminate in 
pointed and almost filiform extremities about 13 cm. behind the 
anus, or, approximately, about the middle of the length of the 
tail. In its course along the tail each cecum is situated wholly 
internal to the caudal musculature of its side, and in immediate 
contact with the hemal spines of the caudal vertebre and the 
supporting radial elements of the extensive anal fin. The dorsal 
and external walls of each cecum are invested by a strong 
aponeurotic membrane, the outer surface of which receives the 
lusertions of the inner margins of the fibrous septa separating 
the caudal myotomes. 

Along nearly the whole length of each of the caudal prolonga- 
tions of the air-bladder, the ventral margin gives off a fringe of 
numerous short but relatively wide diverticula (v.d.), which form 
pairs with their fellows of the opposite side of the tail. With 
the exception of the first two of the series, each diverticulum 
eventually terminates by subdividing into an anterior and a 
posterior slender, filiform cecum (f-c.), which extend ventrally 
towards the base of the anal fin, and are intercaiated between, 
and partially covered by, the superficial flexor muscles of the 
coutiguous dermal fin-rays, but are external to the corresponding 
deep flexors. The second diverticulum has three filiform ceca, 
while the first remains simple aud undivided. The series of 
ventral diverticula and their filiform prolongations gradually 
diminish in size from before backwards, and eventually cease at 
some little distance anterior to the tapering terminal portions of 
the caudal extensions of the bladder. 


506 PROF. T. W. BRIDGE ON THE 


The anterior or pre-ccelomic section of the air-bladder consists 
posteriorly of a somewhat subspherical sae (sd.s.), separated 
externally from the abdominal portion of the organ by a shallow 
transverse or obliquely disposed groove. In this groove the 
subvertebral portion of the mesonephros curves downwards on 
each side to fuse with the large anterior and unpaired portion 
of that organ, or “ head-kidney,” which is situated dorsad to the 
heart and gills, but ventrad to the subspherical sac. Anteriorly, 
the sac abruptly contracts to form a somewhat tubular or slightly 
inflated, median prolongation (¢.p.), which extends forwards 
immediately above the dorsal extremities of the hinder branchial 
arches, in contact with the ventral surfaces of the centrum of 
the first vertebra and the basioccipital, and by each of its lateral 
surfaces is in relation with the branchial branches of the 
corresponding pneumogastric nerve. A. constriction separates 
the subspherical sac from its tubular prolongation, and at the 
same time transmits on the left side the cceliac branch of the 
dorsal aorta. Eventually the tubular prolongation divides into 
two somewhat narrower, cecal, auditory cornua, each of which 
(l.a.c.) diverges somewhat from its fellow and passes upwards and 
outwards, as well as forwards, in order to reach the outer surface 
of the auditory capsule of its side, where, as Cuvier and Valen- 
ciennes (op. cit. p. 140) have described in the case of Notopterus 
Pallas, it becomes intimately associated with the enclosed mem- 
branous labyriath of the organ of hearing. Between the origins 
of the two auditory ceca, and on the dorsal side, the dorsal aorta 
may be seen passing backwards to reach the bony aortic groove, 
to which further reference will subsequently be made. 

The whole of the anterior section of the air-bladder is situated 
in front of the abdominal cavity, and consequently none of its 
various divisions or chambers are invested externally by the 
peritoneum. 

It is perhaps worthy of note that anteriorly and dorsally the 
branchial cavity is prolonged forwards on each side of tne skull 
for some distance, parallel to the lateral surface of the auditory 
capsule, and directly external to the corresponding auditory 
cornua of the air-bladder. This singular extension of the 
branchial cavity practically takes the form of a cecal diverticulum 
(Pl. 36. fig. 1, a.b.c.), ending blindly in front by reason of the 
gradual contraction of its walls, but communicating behind with 
the general branchial cavity. Into eavh cecal diverticulum the 


ATR-BLADDER IN NOTOPTERUS BORNEENSIS. 507 


dorsal portions of the first and second branchial arches (6r.", br.”), 
with their branchial lamellae, extend for some distance; the 
remaining arches, however, lying posterior to the opening of the 
diverticulum into the dorsal portion of the branchial cavity. 
The relatively thin inner wall of the branchial diverticulum is 
coextensive with, and directly and closely invests the outer wall 
of, the corresponding auditory cecum (i.a.c.), and hence in this 
region the air-b!adder and its gaseous contents are more intimately 
related to the external medium in which the Fish lives than 
is the case at any other point. 

For a Teleost the ductus pneumaticus (text-fig., and Pl. 36. 
fig. 3, d.p.) is remarkably short and unusually wide, its length 
not exceeding 3 mm., while its lumen, even in a spirit-preserved 
specimen, is approximately the same in diameter. The cesophageal 
opening of the ductus is in the mid-dorsal line, and about 8 mm. 
behind the last branchial cleft. The aperture by which the 
ductus communicates with the air-bladder is situated a little to 
the left of the median ventral line, and directly posterior to the 
oblique groove (0.g.) separating the subspherical sac from the 
abdominal portion of the organ. The ductus is surrounded at its 
cesophageal extremity bya strong sphincter muscle, but no valvular 
mechanism in connection with either of its apertures could be 
detected. 

No gas-secreting or gas-absorbing ‘‘red-glands”’ or “ red- 
bodies” were to be found in the air-bladder. 


III. Tue Inrernat Srrucrore, aND THE RELATIONS AND 
ATTACHMENTS OF THE ATR-BLADDER TO THE SKELETON. 


In the condition of their inner or mesial walls, and in the 
relation of these structures to the skeletal elements of the tail, 
the caudal prolongations of the air-bladder present ‘several 
interesting features. . 

With regard to the character and disposition of the caudal 
skeletal elements, it may be mentioned that the hemal spines of 
the suprajacent vertebre are relatively short (Pl. 36. fig. 2, h.s.). 
The radial elements of the anal fin (7.¢.), on the contrary, are of 
considerable length in the anterior part of the tail, although 
they gradually become much shorter as they are traced backwards 
towards the caudal fin; and as two of them, forming a pair, are 
associated dorsally with each hemal spine, it follows that the 
radial elements are twice as numerous as the caudal vertebre, at 


508 PROF. T. W. BRIDGE ON THE 


any rate in the region of the air-bladder. At the commencement 
of the tail only the ventral third of the radial elements serve for 
the origin of the flexor muscles of the anal fin-rays, the dorsal 
two-thirds, and the short hemal spines with which the former 
interdigitate, being quite free from muscular attachment. With 
the gradual shortening of the radial elements towards the middle 
portion of the tail, where the air-bladder terminates, nearly the 
whole length of each radial element becomes invaded by the origin 
of the flexor muscles. The area over which the various radial 
elements are wholly devoid of muscular attachment is coextensive 
with that of the inner wall of each of the two closely-related 
caudal ceca. The series of radial elements and hemal spines are 
connected together by a thin, but tough, sheet of fibrous tissue, 
which extends from one to the other and fills up the intervals 
between them, and, with the caudal skeletal elements above 
mentioned, form the only separation between the caudal cxca of 
opposite sides of the tail. It may be added, that the series 
of ventral diverticula occupy the intervals between the successive 
pairs of radial elements. 

The dorsal and outer walls of each caudal cecum are of 
moderate thickness, as also are the external walls of the ventral 
diverticula, while the filiform ceca have much thinner walls. In 
each case, nevertheless, a relatively thick outer fibrous stratum 
or tunica externa, and an extremely thin lining or tunica interna, 
consisting of an internal epithelial stratum, supported externally 
by a thin layer of connective tissue, can readily be recognized. 
The inner wall of each caudal cecum, including also that of each 
of its primary ventral diverticula, on the contrary, is of extreme 
tenuity, consisting only of the tunica interna, and, moreover, is 
closely adherent to the outer surfaces of the radial elements and 
heemal spines, and also to the fibrous sheet which stretches between 
them. In fact, in the intervals between these skeletal elements, 
the median fibrous sheet and the attenuated inner walls of the 
caudal ceca are all that separate the cavities of the cxca of 
opposite sides of the tail; and so thin are the inner walls of 
these that, when the cavity of either of them is exposed by the 
removal of its outer wall, the various skeletal elements (A.s., r.e.) 
appear as if completely bare of any investing tissue and to 
project freely into the lumen of the bladder. 

In addition to the investment of the cuter surfaces of the 
caudal skeletal elements by the tunica interna of the inner wall, 


AIR-BLADDER IN NOTOPTERUS BORNEENSIS. 509 


the dorsal and ventral walls are also firmly attached to the 
skeleton. Along its inner or mesial dorso-lateral margin, where 
the relatively thick dorsal wall becomes continuous with the 
attenuated inner wall, the tunica externa becomes somewhat 
thickened and terminates by becoming firmly attached to the 
bases of the hzemal spines and also to the dorsal extremities of 
the two radial elements with which each spine is associated. 
Ventrally also the tunica externa thickens and forms a series of 
strong transversely disposed ridges projecting from the floor of 
the cecum, and separating the orifices leading into the ventral 
diverticula. Traced towards the inner wall of the cecum, the 
ridges cease by becoming inserted into the adjacent sides of two 
contiguous radial elements and to the connecting fibrous mem- 
brane which extends between them. On the other hand, if traced 
ventrally, the fibres of each ridge split to form the posterior wall 
of one diverticulum and the anterior wall of the next succeeding 
diverticulum, both walls nevertheless retaining their mesial 
attachments to the contiguous radial elements between which 
the diverticulum is situated. 

Near the dorsal wall of each caudal cecum, the fibrous sheet 
between the radial elements presented a series of oval vacuities 
(d.v.), which were disposed in regular order between the succes- 
sive pairs of radial elements which are attached to the vertebral 
hemal spines. Over most of these vacuities the thin inner wall 
of the caudal cecum appears wanting, and hence the cavity 
of each cecum seems to communicate with that of its fellow 
at these points. In one or two instances, however, it is 
clear that no such communication exists, for, notwithstanding 
the vacuity in the fibrous sheet, the opposed inner walls of the 
two ceca remain intact over the area of the vacuity. It is 
difficult, therefore, to be quite certain that these vacuities are 
associated with normal perforations in the opposed walls of the 
cea, or that the cavities of the latter really intercommunicate 
during life, or in perfectly fresh specimens, more especially as, 
owing to its thinness and fragile character, the tunica interna is ex- 
tremely likely to break down and disintegrate where unsupported 
by the much stronger and more resistant fibrous sheet, unless 
more than ordinary care is taken with the preservation of the Fish. 

A similar series of small, oval, or rounded vacuities (v.v.) in 
the interradial fibrous membrane was also present towards the 
ventral side of each caudal cecum, where the menbrane in 


510 PROF. T. W. BRIDGE ON THE 


question separates the closely related inner walls of the two series 
of ventral diverticula, and, as regards their relations to the caudal 
skeleton elements, these correspond in position with the dorsal 
series. Wherever these ventral vacuities existed, corresponding 
perforations in the opposed inner walls of the diverticula were 
also present, and each of the latter seemed to communicate with 
its fellow of the opposite side of the tail. In some instances it 
was possible to determine the continuity of the inner wall of a 
diverticulum with the corresponding wall of its fellow at the 
edges of the vacuity ; and for this reason I am inclined to 
believe that the caudal ceca are really in communication with each 
other ventrally, through the fusion and subsequent perforation 
of the inner walls of their ventral diverticula. Nevertheless, it 
is perhaps desirable that the caution needful in the case of the 
apparent dorsal perforations should also be observed here, at 
all events until fresh or well-preserved specimens of Notopterus 
have been submitted to examination. 

A somewhat interesting numerical and regional correspondence 
is to be observed in the anterior half of the tail between the 
various caudal skeletal elements and the arrangement of the 
cecal outgrowths from the caudal divisions of the air-bladder. 

Thus, the ventral diverticula are situated exactly opposite the 
intervals between the series of hemal spines, and, at the same 
time, occupy the interspaces between successive pairs of radial 
elements ; that is, each diverticulum fills up the interval between 
two contiguous pairs of radial elements which, dorsaily, are 
attached to the hemal spines of two successive caudal vertebre. 
Hence, therefore, the diverticula closely agree in number with 
the pairs of radial elements, and in position may be said to be 
intervertebral. Again, each of the series of fibrous ridges 
which separate the orifices of communication between the 
ventral diverticula and the caudal ceca, is attached to the 
opposed surfaces of a pair of radial elements as well as to the 
fibrous membrane between them; and as each ridge coincides 
dorsally with the hemal spine to which the two radial elements 
belong, it is vertebral in its relations to the axial skeleton. 
La-tly, the filiform ceca agree in number with the radial 
elements, the posterior cecum of one diverticulum, and the 
anterior cecum of the next succeeding diverticulum, being 
situated immediately external to the corresponding factors of 
two contiguvus pairs of radial elements. 


AIR-BLADDER IN NOTOPTERUS BORNEENSIS. onal 


The dorso-lateral margins of the abdominal portion of the air- 
bladder (Pl. 36. fiy. 3) are firmly attached to the transverse 
processes of certain of the trunk vertebre and to the proximal 
portions of their costal elements, the latter being so intimately 
related to the bladder as to produce a series of faint trans- 
versely-disposed grooves in the lateral walls of that organ. As 
previously stated, the peritoneum invests the ventral surface of 
this portion of the air-bladder; and it may be added, that in this 
region it assumes the condition of an exceptionally tough fibrous 
membrane which is firmly adherent to the veutral wall of the 
bladder. Posteriorly, the peritoneum is reflected downwards on 
to the hemal arch and spine of the first caudal vertebra and the 
first radial element of tbe anal fin, and is, moreover, firmly 
attached to these skeletal structures. Hence it follows, that the 
ventral wall of the abdominal section of the air-bladder is firmly 
attached posteriorly to the anterior caudal skeletal elements. 
The walls of this portion of the bladder are of moderate and 
equal thickness throughout. Internally, the cavity of the 
bladder is subdivided into two lateral compartments by a vertical 
longitudinal septum, which is continuous dorsally and ventrally 
with the corresponding walls of the bladder (fig. 3, J.s.). Poste- 
riorly, the septum increases in height with the increasing 
vertical dimension of the bladder, and, at the point where the 
latter subdivides into the two caudal ceca, the binder margin of 
the septum is inserted into the anterior face of the hemal arch 
and spine of the first caudal vertebra and the proximal portion of 
the first radial element of the anal fin. Anteriorly, the longi- 
tudinal septum deviates from the median plane towards the lett 
side, and, so far as its ventral portion is concerned, the septum 
ceases immediately behind and a little to the right side of the 
internal aperture of the ductus pneumaticus (/.s., d.p.). At this 
point the dorsal portion of the septum, which, it may be men- 
tioned, extends forwards into the subspherical sac, is connected 
by a uarrow obliquely-transverse septum (¢.s.) with the left 
lateral wall of the bladder along the line of the external oblique 
groove (0.g.) separating the sac from the abdominal portion of 
the bladder*. The effect of this singular unsymmetrical dis- 
position of the longitudinal septum, and the presence of an oblique 
transverse septum on the left side only, combined with the 


* In fig. 3 that portion of the left lateral wall of the bladder which is 
traversed by the oblique groove is indicate as a slender strip. 


512 PROF. T. W. BRIDGE ON THE 


position cf the aperture of the ductus pneumaticus slightly to 
the left of the median line, is (1) that the left lateral eompart- 
ment is smaller than the right; (2) that the orifice of communi- 
cation between the subspherical sac and the left lateral chamber 
is reduced to the condition of a relatively small aperture, which 
is in striking contrast to the widely open, direct communication 
between the sac and the lateral compartment of the right side; 
and (3) that while the ductus pneumaticus opens directly into 
the left compartment, its connection with the right chamber is 
somewhat indirect aud takes place through the cavity of the 
subspherical sac, round the free anterior margin of the ventral 
portion of the longitudinal septum (fig. 3). 

Like the abdominal portion of the bladder, the subspherical 
sac (Pl. 36. fig. 3, sb.s.) is firmly attached along its dorso-lateral 
margins to the transverse costiferous processes of certain of the 
trunk vertebree, viz., the second to the fifth, inclusive. Its 
dorsal wall is extremely thin, and, in fact, is represented by 
the tunica interna alone; the latter stratum, in the absence of 
the mesonephros in this region, being closely adherent to the 
Jateral and ventral surfaces of the anterior vertebral centra. 
Internally, the sac is partially subdivided by an extremely thin 
median longitudinal septum (/.s.’), which projects downwards 
from its dorsal wall, and is a direct continuation of the dorsal 
half of the longitudinal septum previously mentioned as unsym- 
metrically dividing the cavity of the abdominal portion of the 
air-bladder. The septum is best developed behind, and gradually 
diminishes in height towards the anterior limit of the sac. Its 
free ventra] margin is remarkably thick and forms a strong, arch- 
hike band, the fibres of which diverge behind to strengthen and 
stiffen the inner and outer lips of the orifice by which the sac 
communicates with the left lateral compartment of the preceding 
portion of the bladder. Anteriorly also the band divides, but 
into three fibrous bundles, the more laterally-situated of which 
diverge downwards into the Jateral walls of the sae and strengthen 
the outer lips of the aperture through which the sac com- 
municates with its anterior tubular prolongation, while the mesial 
fibres continue their forward course. 


AIR-BLADDER IN NOTOPTERUS BORNEENSIS. 513 


TY. Renartions oF THE ATR-BLADDER TO THE SKULL AND 
Avpirory ORGANS. 


The structure of the tubular prolongation of the subspherical 
sac and of the auditory ceca, and the relations of both to the 
skull and the auditory organ, will be better understood after a 
brief description of certain structural features in connection 
with the hinder part of the skull (Pl. 37. fig. 4). 

The basioccipital (4.0.) is somewhat hourglass-shaped, being 
slighily constricted at the junction of the hinder third with the 
anterior two-thirds of its length, and expanded towards either 
extremity, but much more so in front than behind. The hinder 
half of the bone is produced ventrally into a laterally-compressed 
bony keel, which is traversed by a moderately deep, median, longi- 
tudinal groove, bounded by prominent lateral ridges, and trans- 
mitting the initial section of the dorsal aorta. 

Traversing the lateral surface of the basioccipital, immediately 
ventrad to the sutural articulation of this bone with the ex- 
occipital and opisthotic, is a well-marked oblique ridge, which, 
on each side, extends backward and downward and finally 
terminates in a free projecting process (a.p.), closely applied to 
the ventro-lateral surface of the centrum of the first vertebra 
after the fashion of an accessory articular process. Between 
the aortic groove ventrally and the oblique ridge dorsally, the 
lateral surface of the basioccipital is traversed by a fairly deep 
groove (b.g.), the direction of which is obliquely upward and 
forward towards the outer surface of the auditory capsule of 
the same side. Anteriorly to the commencement of the aortic 
groove the basioccipital ceases to be laterally compressed, and, 
instead, becomes greatly swollen laterally, assuming in fact a 
_ distinctly bullate appearance. The lateral surface of this portion 
of the bone is also traversed by a well-defined groove, which, 
however, is merely an extension forwards of that mentioned 
above. Ventrally, the anterior half of the basioccipital is in 
relation with the hinder portion of the parasphenoid (ps.), and in 
front the bone articulates by means of an irregular squamous 
suture with the hinder margin of the prootie (pro.). 

There is a complete series of periotic bones, which, in the 
main, exhibit the usual relations one to another and to the 
adjacent cranial bones. The opisthotic (op.) forms the postero- 
lateral portion of the auditory capsule, articulating with the 


514 PROF. T. W. BRIDGE ON THE 


exoccipital (¢o.) behind, the pterotic (pt.) above, with the supero- 
Jateral margin of the basioccipital (60.) below, and in front with 
the prootic {(pro.). The relatively large prootic (pro.) forms the 
anterior portion of the periotic capsule, and articulates above 
with the sphenotic (spe.) and pterotic (pt.), below with the para- 
sphenoid (ps.), and posteriorly and dorsally with the opisthotie 
(op.), while the posterior portion of the bone forms a thin 
squamous lamina which extends backwards, overlapping the 
lateral surface of the basioccipital (40.), and at the same time 
closing in the anterior portion of the basioccipital groove. In 
addition, the ventral portion of each prootic sends inwards a strong 
horizontal process (Pl. 37. fig. 5, pro.), which unites with its 
fellow in the floor of the cranial cavity behind the pituitary fossa. 
The pterotic (Pl. 37. fig. 4, pt.) constitutes the superior lateral 
margin of the auditory capsule, and also contributes the usual 
articular surface for the proximal extremity of the hyoman- 
dibular ; anteriorly, the bone is in articular relation with the 
upper surface of the prootic and with the sphenotic, and behind 
overlaps the dorsal margin of the opisthotic. The sphenotie 
(spo.) is a small nodular ossicle, wedged in between the upper 
margin of the prootic and the overlapping anterior extremity of 
the pterotie. 

As already indicated, the anterior termination of each lateral 
basioccipital groove is overlapped by the squamous posterior 
extension of the prootic, and thus becomes converted into a 
short, but relatively spacious, bony cul-de-sac (c.s.), continuous 
behind with the open portion of the groove, but terminating 
blindly in front. The inner or cranial wall of the cul-de-sac is 
coincident with a somewhat considerable fontanelle (Pl. 37. 
fig. 5, a.f.), which is encircled above, below, and in front by the 
prootie (pro.), and behind is limited by the anterior margin of 
the opisthotic (op.) and by the antero-superior border of the 
basioccipital (40.), and in the dried skull places the cul-de-sac in 
free communication with the cavity of the auditory capsule. The 
formation of this auditory fontanelle is apparently due to the 
widening of the normal sutures which separate the surrounding 
periotic elements one from another and from the basioccipital. 
An extremely thin fibrous membrane, which in texture, colour, and 
appearance closely resembles the membrane closing the supero- 
lateral cranial fontanelle, extends between the margins of the 
auditory fontanelle, and constitutes the only separation between 


AIR-BLADDER IN NOYTOPTERUS BORNEENSIS. 515 


the lumen of the bony cul-de-gac and the perilymph-containing 
cavity of the auditory capsule. 

Returning now to the air-bladder (Pl. 36. fig. 3), it may be 
stated that the dorsal surface of the tubular prolongation (¢.p.) 
of the subspherical sac is closely moulded to the ventral surface 
of the centrum of the first vertebra, and also to the ventral and 
lateral surfaces of the hinder part of the basioccipital. From 
its laterally-compressed shape, the ventral portion of the latter 
bone forms a median keel projecting downwards and pushing 
before it the dorsal wall of the subjacent part of the air-bladder, 
which, consequently, appears as if partially subdivided internally 
by an incomplete, but very thick, longitudinal partition (J.p.), 
while dorso-laterally the bladder fills up the grooves on the 
lateral surfaces of the basioccipital. Laterally and ventrally, 
this portion of the air-bladder is free from any special relations 
or attachments to the skeleton, and hence its moderately thick 
walls consist of both tunica externa and tunica interna: elsewhere, 
however, the walls are intimately related to the skeleton, and 
then are either firmly attached thereto or become greatly reduced 
in thickness. Thus, dorso-laterally, on each side, the tunica 
externa ceases by becoming inserted into the oblique bony ridge 
on the lateral surface of the basioccipital. In the mid-dorsal 
line, the tunica externa is not only considerably thickened by 
mesial fibres derived from the arch-like band which forms the 
free ventral margin of the longitudinal septum of the sub- 
spherical sac, but is also attached to the lips of the bony aortic 
groove in such a way as to convert the groove into a canal. 
Between the mid-dorsal and the dorso-lateral skeletal attach- 
ments, however, the tunica externa is wanting, and all that 
represents the proper wall of this part of the bladder is the thin 
tunica interna investing and lining the hinder portion of the 
grooves on the lateral surfaces of the basioccipital, into which 
the lateral portions of the bladder are received. The diameter 
of the tubular portion of the bladder is about 6 mm. 

The two auditory ceca (J.a.c., r.a.c.) communicate with the 
preceding part of the bladder by relatively wide orifices; and at 
this point the simple non-septate lumen of each cecum is about 
3°5 mm. in diameter. From their origins the two cweca diverge 
obliquely upward and forward towards the outer surfaces of 
their respective auditory capsules. In its forward extension, 
each auditory cecum traverses the anterior portion of the oblique 


O16 PROF. T. W. BRIDGE ON THE 


groove on the lateral surface of the basioccipital, and, with only 
a slight reduction in calibre, appears to terminate at the sutural 
junction of the latter bone with the prootic. The outer lateral 
wall of each cecum is complete, but both dorsally and ventrally 
its tunica externa ceases by becoming firmly inserted into the 
corresponding lips of the basioccipital groove in which the 
cecum is lodged. Towards the anterior termination of the 
cecum even the tunica externa of the outer wall disappears 
through its continuity with the hinder margin of the prootic 
bony lamina, which, anteriorly, converts the groove into a cul- 
de-sac. The inner wall, on the contrary, is throughout formed by 
the tunica interna alone, and, moreover, is closely adherent £0 
the sides of the basioccipital groove. 

Reduced to a simple wall of tunica interna of extreme tenuity, 
the terminal portion of the auditory cecum now enters the bony 
cul-de-sac (Pl. 37. fig. 4, c.s.), and, on its mner or eranial side, 
becomes closely applied to the outer surface of the membrane 
closing the auditory fontanelle (fig. 5, a.f-). 


V. Tue CrantaL FonTANELLES AND THEIR RELATION TO THE 
AUDITORY ORGAN. 


In their account of the bones of the skull in Notopterus 
Pallasii, Cuvier and Valenciennes (4. p. 143) describe two in- 
teresting vacuities in the postero-superior aspect of the cranial 
roof. “Ces mastcidiens ont en;avant une tres-profonde 
échancrure, qui cerne prés des deux tiers du giand trou pari¢to- 
mastoidien, dont les cétés du crane sont percés, une échancrure 
du frontal postérieur contribue aussi 4 former le cercle de ce 
trou. Ce grand trou, analogue 4 celui que nous avons observé 
dans ’ Alose et dans plusieurs autres Clupées, mais beaucoup plus 
semblable encore 4 ce que existe dans le Mormyre, est bouché 
par une couche peu épaisse de cette mucosité graisseuse, qui 
remplit les cavernes du crane et sur laquelle passe la peau mince, 
nue et sans écailles de la téte. Par ce trou on pénétre largement 
dans Vintérieur de la cavité du crane, et l’on voit presque sans 
dissection, aprés avoir toutefois enlevé toutes ses parties externes, 
les canaux semicirculaires supérieurs, leur ampoule commune et 
une portion du sae qui contient l’otolithe.” 

This description is in the main an accurate one, but may never- 
theless be supplemented in certain details. 


AILR-BLADDER IN NOTOPTERUS BORNEENSIS. 517 


Two longitudinal grooves traverse the lateral portions of the 
cranial roof, and are bounded for the anterior part of their 
extent by prominent bony ridges on the external surface of the 
frontals, and more posteriorly by similar parallel ridges on the 
upper surface of the pterotic (Pl. 37. figs. 4 & 6, 7). Externally, 
the grooves are closed in and converted into complete canals by 
the thin, scaleless, superficial skin of the dorsal surface of the 
head. In these grooves or canals are lodged the main lateral 
sensory canals of the head and their respective supra-orbital 
prolongations. At its hinder extremity each groove suddenly 
deepens into 2 shallow basin-shaped, or funnel-like, oval de- 
pression, at the bottom of which is the somewhat smaller, 
but also oval, cranial fontanelle (Pl. 37. fig. 6, ef), which in 
the dried skull communicates internally with the cavity of the 
auditory capsule, precisely as described by Cuvier and Valen= 
ciennes. The margins and sides of the funnel are smooth and 
rounded, and are formed anteriorly, and also to a large extent 
on each side, by the deeply concave posterior border of the 
pterotic (pé.), and completed laterally and behind by che epiotic 
(ep.), the opisthotic (op.), and the exoccipital (eo.). The long 
axis of the mouth of the funnel measured 10 mm., and its trans- 
verse dimension 6 mm. The fontanelle is smaller, the cor- 
responding dimensions being 6 mm. and 4 mm. respectively. 
Across the fontanelle, and firmly attached to its margins, is 
stretched a thin, fibrous, drum-head-like membrane, which 
is in relation internally with the somewhat fatty perilymphatic 
tissue of the interior of the auditory capsule. 

There is, however, one point to which Cuvier and Valenciennes 
make no reference, viz.—the relations of the main sensory 
canal of the head to the cranial fontanelle and the mem- 
brane closing it. As it passes directly dorsad to the proximal 
or supra-clavicular element of the pectoral girdle, the sensory 
canal traverses the axis of a somewhat cylindrical bone, and 
then enters the lateral longitudinal groove or canal to which 
reference has just been made. At this point the sensory canal 
expands considerably, and assumes a singular cavernous or sinus- 
like appearance, practically filling the bony groove in which it is 
lodged. The lateral and inner walls of the sensory canal are 
here strengthened by two longitudinally-arranged, thin, demi- 
cylindrical bones, or sensory canal ossicles, the convex inner 
surfaces of which are in close relation with the cranial fontanelle 

LINN. JOURN.—ZOOLOGY, VOL. XXVII. 39 


518 PROF. T. W. BRIDGE ON THE 


and the whole extent of the outer surface of its drum-head 
membrane, while the outer wall of the sensory canal is in contact 
with the external scaleless skin of this part of the head. 

Practically, therefore, the cavernous sensory canal and its 
investing ossicles completely separate the cranial fontanelle and 
its membrane from the superficial skin. Anteriorly to the 
fontanelle the sensory canal is still of considerable width, but, 
as it passes dorsad to the orbital cavity, gradually contracts to 
more normal dimensions. 


VI. Tar Avuprrory Or@aN. 


The utriculus and the sacculus of each side oceupy a spacious 
common recess excavated in the substance of the corresponding 
lateral half of the large basioccipital (Pl. 87. fig. 5, w.s.r.). So 
large is the recess, that the outer portion of nearly the anterior 
two-thirds of the bone is reduced to the condition of a thin 
partially transparent shell, and corresponds to the externally 
bullate portion to which reference has already been made; while 
a thin, vertical, bony partition in the axis of the basioccipital is all 
that separates each recess from its fellow of the opposite side of 
the skull. The two recesses are partially roofed in by the mesial 
union of horizontal ingrowths from the two opisthotic bones (op.), 
in a fashion which recalls the method by which the similarly- 
situated saccular recesses are roofed by the exoccipitals in the 
Siluroid Fishes. The utriculus occupies the anterior two-thirds 
of the utriculo-saccular recess, and is relatively of large size. 
The much smaller sacculus fills up the hinder third of the recess, 
and is connected with the utriculus by a very short but obvious 
ductus sacculo-utricularis. Anteriorly and dorsally, the utriculus 
gives oft a conical, forwardly-directed diverticulum, which occupies 
a recess of corresponding shape excavated in the prootic*. Near 
the origin of the diverticulum, the ampullary extremities of the 
horizontal and the anterior vertical semicircular canals communi- 
cate with the utriculus by opening into a small recessus utriculi. 
The great utricular otolith, or “ sagitta,”’ is nearly as large as 
the cavity of the utriculus itself; and it is interesting to note 
that, anteriorly and dorsally, the otolith is produced into a con- 
spicuous, tapering, conical process, which extends into the 


* Cf. Ridewood’s account of the auditory organ in the Clupeide (9). 


ATR-BLADDER IN NOTOPTERUS BORNEENSTS. 519 


utricular diverticulum inthe prootic. The membrane which closes 
each auditory fontanelle (a.f-), and is invested externally by the 
tunica interna of the auditory cecum of the air-bladder, is almost 
in contact internally with the outer wall of the utricular diver- 
ticulum. The semicircular canals have the normal arrangement, 
both as regards their relations to the utriculus and to the various 
periotic bones. It may be mentioned, however, that each 
supero-lateral cranial fontanelle (¢.f) is almost completely en- 
circled by the three semicircular canals—the horizontal canal in 
the pterotic (p?.) curving round the lower or outer border, the 
posterior vertical canal in the epiotic (ep.) and opisthotic (op.), 
and the anterior vertical canal in relation with the inner surface 
of the prootic (pro.), being similarly situated with regard to the 
anterior and hinder margins respectively ; while the vertically- 
disposed common stem, or sinus utriculi superior, by which the 
two latter canals join the utriculus, crosses the fontanelle at a 
short distance internal to the membrane which closes it. 

Careful examination failed to reveal the existence of any con- 
nection between the sacculi of opposite sides by means of the 
supra-cerebral or sub-cerebral union of their endolymphatic ducts, 
such as has been described by Weber (12) for Silurus glanis, and 
in the case of other Siluroids by Bridge and Haddon (2) and 
Ramsay Wright (13), and by Hasse (6) for the Herring (Clupea 
harengus) ; or of any communication between the two utriculi, 
similar to the sub-cerebral connection which has been described 
in the last-mentioned Teleost by Weber (12), and in the Shad 
(Clupea alosa) by Breschet (1). It must be admitted, however, 
that the single specimen of Wotopterus examined by me was 
not sufficiently well-preserved to quite justify purely negative 
conclusions on these points. 


VII. Comparison witn Nororrervs Pazzasrr. 


As regards the general structure and relations of the various 
divisions of the air-bladder, the account above given agrees 
generally with that of 1. Pallasii by Cuvier and Valenciennes. 
In certain details, however, there are important discrepancies, 
which may be due either to the existence of structural variations 
in the two species, or possibly to errors in the description of the 
air-bladder of V. Pallasiz. The more important of these differences 
will now be considered. 

39* 


520 PROF. T. W. BRIDGE ON THE 


In their account of the air-bladder of NV. Pallasii, Cuvier and 
Valenciennes (4. pp. 189-141) make no mention of the existence 
of aseries of branched diverticula similar to those which fringe 
the ventral margins of the caudal prolongations of the air-bladder 
in LV. dorneensis; neither do these authors refer to the extreme 
tenuity of the inner walls of these portions of the bladder, or to 
their attachments and relations to the caudal skeletal elements, 
nor to the existence of vacuities in the closely related inner walls: 
of the caudal cca, by which the cavities of the two ceca freely 
interecommunicate. 

In their description of the caudal portion of the bladder in 
N. Pallasii, Cuvier and Valenciennes (op. cit. p. 189) remark :— 
“Tl existe sur la surface externe des cornes un organe singulier, 
comme glanduleux, divisé par un nombre considérable de filets 
blanchatres, anastomosés entre eux en petits lobules, que l’on ne 
pourrait séparer par la dissection qu’avec beaucoup de peine. 
Cet organe, qui couvre presque tout le bas de la corne, ne dépasse 
guére la moitié de sa longueur.” 

No trace of any such gland-like structure could be detected in 
N. borneensis ; and as no organ of a similar character is known to: 
be associated with the external surface of the air-bladder in any 
other Fishes, it is impossible to hazard even a conjecture as to its 
nature in WV. Pallasi. 

Tn their description of the abdominal and anterior portions of 
the air-bladder, Cuvier and Valenciennes apparently failed to 
note the skeletal attachments of the organ, or the general 
tendency of the bladder to lose its external fibrous coat wherever 
its walls enter into intimate relations with the cranial or vertebral 
elements of the skeleton. The unsymmetrical subdivision of the 
abdominal section of the bladder seems also to have escaped their 
notice. 

The account given by Cuvier and Valenciennes of the mode of 
termination of the auditory ceca and their relations to the mem- 
branous labyrinth in Motopterus Pallasii is so different from 
my own observations on Notopterus borneensis, that I venture to 
quote their remarks 2n extenso. 

After describing the extension of the air-bladder towards the 
cranium, it is stated : “ De la elle donne deux cornes qui s’engagent 
dans lintérieur de la boite cérébrale sous les mastoidiens, en 
passant entre l’os et le sac de Voreille. Ces cornes s’avancent 


AIR-BLADDER IN NOTOPTERUS BORNEENSIS. 521 


dans Vintérieur de la boite cérébrale jusque sur la grande aile du 
sphénoid, et atteignent la hauteur de la scissure qui sépare 
le second tubercule, ou le tubercule optique du cerveau, du 
troisiéme, derriére lequel existe le cervelet. En pénétrant dans 
la boite cérébrale la vessie perd ses tuniques fibreuses, ou plutot 
eest la seule tunique propre ou membraneuse de la vessie qui 
s’avance ainsi dans la cavité du crane. On voit en dedans de la 
corne le sac qui contient la pierre de l’oreille. Tl y a done ici 
communication médiate entre la vessie et l’organe de louie; 
e’est le seul exemple que je connaisse d'une communication 
aussi intime entre la vessie et l’organe de l’oule” (op. cit. p. 140). 

It is quite certain that the auditory cornua or ceca do not 
enter the cranial cavity in JV. borneensis, but, on the contrary, 
are wholly extra-cranial. The statement quoted above that “ en 
pénétrant dans la boite cérébrale la vessie perd ses tuniques 
fibreuses, ou plutot c’est la seule tunique propre ou membra- 
neuse de la vessie qui s’avance ainsi dans la cavité du crane,” 
correctly describes the behaviour of the auditory ceca on entering 
the bony culs-de-sac in which their terminal extremities are 
lodged; and it is therefore possible that Cuvier and Valenciennes 
have mistaken this canal for a portion of the cranial cavity. It 
hag been shown, however, that this canal has no communication 
with the eranial cavity, inasmuch as it remains separated there- 
from either by the thin membrane which closes the auditory 
fontunelle, or by the various periotic bones surrounding it. 
Hence it follows that the subsequent statement, “Il y a done 
ici communication médiate entre la vessie et l’organe de l’ouie,” 
is obviously erroneous as regards Notopterus borneensis, whether 
it implies an open communication between the two organs, or the 
existence of a direct connection by the simple apposition of their 
limiting walls; and I entertain little doubt that the statement is 
equally inapplicable to Notopterus Pallasi. 

It is also stated by Cuvier and Valenciennes that Nofopterus 
affords the only instance known to them “ d’une communication 
aussi intime entre la vessie et ’organe de l’ouie; car je n’hésite 
pas a répéter ici que celle qui avait été annoncée dans |’Alose ou 
dans le Hareng, et dans plusieurs autres poissons, u’existe 
réellement pas” (op. cit. p. 140). 

The reference to the Allis Shad (Clupea alosa) and the Herring 
(CO. harengus), as Ridewood (9. p. 40) has pointed out, is evidently 


522 PROF. T. W. BRIDGE ON THE 


based on the misconception that previous writers had described 
an open communication between the air-bladder and the 
auditory organ, such as Valenciennes apparently believed to exist 
only in Wotopterus. It is now scarcely necessary to add that no 
such open communication exists, or has even been affirmed to 
exist, in any Fish except Wotopterus Pallasii; and, for the 
reasons given above, it is extremely improbable that the latter 
species offers any exception to the general rule, or, so far 
as this point is concerned, differs in any way from its congener, 
Notopterus borneensis. 


VIII. Comparison with oTHER TELEOSTS. 

Perhaps the most interesting point in the air-bladder of 
Notopterus is the combination which it exhibits of structural 
features, some of which are unique, while others are individually 
characteristic of widely different genera or species of Teleosts. 

The extension of the air-bladder from its normal position in 
the abdominal region into the tail is by no means of infrequent 
occurrence in Teleosts, although it may take place in various 
ways. In some species (e. g., species of Hwocetus) the organ is 
prolonged backwards without undergoing subdivision into the 
expanded hemal canal of the anterior part of the tail (11. p. 222). 
In others the caudal extension takes the form of an unsym- 
metrical prolongation of the entire organ along the left side of 
the tail, as in Ophiocephalus, or along the right side, as in the 
Characinoid Alestes Hasselquistii, C. & V. (A. dentex, Mull. & 
Trosch.) [11. p. 222] *, and in the Siluroids Cryptopterus micro- 
nema, Blkr., and C. micropogon, Blkr. (2. pp. 202-3). More 
frequently, perhaps, the air-bladder subdivides anteriorly to the 
first caudal hemal arch, and in the form of two bilaterally- 
arranged cecal prolongations extends for a variable distance on 
either side of the tail and internal to the lateral caudal muscu- 
lature, as in some Sparide, Scombride, and Carangide (11. 
pp- 221-2), and also as in Notopterus. But in none of the 
Teleosts above mentioned, except Notopterus, or in any others with 
which I am acquainted, do the caudal prolongations of the air- 
bladder exhibit the slightest tendency to branch, or to develop 
structures in any way comparable to the singular fringe of 
bifurcate ventral diverticula which are so characteristic of 
Notopterus borneensis; and very rarely in anv Teleost are the 


* On the authority of Cuv. & Val. 


AIR-BLADDER IN NOTOPTERUS BORNEENSIS. 523 


inner walls of the caudal cxca reduced to so attenuated a condition 
through the intimacy of their relations and attachments to the 
caudal skeleton, as is the case in the last-mentioned species. Even 
in the two species of Oryptopterus, where the caudal portion of 
the air-bladder is in contact with the subvertebral hemal arches 
and spines, the walls of the organ are of uniform thickness and 
are quite free from any special connection or attachment to the 
skeleton. 

Perhaps, on the whole, the air-bladder of certain species of 
Sparide (e. g., species of Box) approaches more nearly to that of 
Noptopterus than does the bladder of any other Teleosts. In 
Box vulgaris, C. & V., not only are caudal ceca present, but the 
inner or mesial walls of these structures are devoid of an outer 
fibrous coat, or tunica externa, and the tunica interna, which 
alone remains, closely invests the opposite sides of the hemal 
arches and spines of the caudal vertebra. The resemblance is 
further heightened by the fact that in Sparus salpa, L. (Box 
salpa, CO. & V.), as Weber (12. p. 71 e¢ seg.) pointed out, auditory 
ceca are also present, although in the location of their connection 
with the auditory fontanelles and in some minor details the 
latter Sparoid does not precisely agree with Notopterus. On the 
other hand, Bos has no bifurecate ventral diverticula in connection 
with the caudal ceca, and the air-bladder is wholly destitute of 
internal septa, and of a ductus pneumaticus in the adult. 

The extension of the air-bladder into the tail in Notopterus, 
as no doubt is also the case in many other Teleosts, is to be 
associated with the extreme shortness and laterally-compressed 
shape of the abdominal portion of the body, which, if the bladder 
is to acquire its normal degree of development as a hydrostatic 
organ, necessitates its prolongation into the caudal region *. 

In the disposition of the internal septa, and especially in the 
development of a principal longitudinal septum which, anteriorly, 
meets an incomplete transverse septum, the ccelomic or abdominal 
portion of the air-bladder of Mofopterus presents some approxi- 
mation to the characteristic T-shaped arrangement of the 
primary septa in the bladder of a considerable number of 
Siluroids ; and the resemblance is rendered still more marked by 
the fact that in both the carinate shape of the suprajacent axial 
skeleton involves a partial subdivision of the anterior portion of 
the bladder through the inpushing of its dorsal wall in the 


* See reference to remarks by Ginther (7. footnote to p. 491). 


524 PROF. T. W. BRIDGE ON THE 


median line. On the other hand, however, Notopterus differs 
from the more typical Siluroids in the absence of the transverse 
septum on the right side, the unsymmetrical division of the air- 
bladder by the longitudinal septum, and also in the fact that it 
is the basioccipital which is carinate ventrally and not, as in 
Siluroids, the confluent centra of the “ complex” vertebra. 

An additional resemblance to many Siluroids is also apparent 
in the tendency of the outer fibrous coat of the air-bladder to 
become invaded by ossific deposit whenever it becomes attached 
to, or inserted into, adjacent portions of the axial skeleton. The 
bony ridges which bound the basioccipital grooves for the 
reception of the auditory czeca, and the squamous hinder portions 
of the prootizs which form the outer walls of the bony culs-de-sac 
wherein these ceca terminate, almost certainly owe their existence 
to the ossification of the tunica externa at the points where it is 
attached to these cranial bones. 

By no means the least noteworthy of the many interesting 
structural features in connection with the air-bladder of Woto- 
pterus borneensis is the extreme shortness and relatively wide 
calibre of the ductus pneumaticus, and the position of its ceso- 
phageal aperture in close proximity to the last pair of branchial 
clefts. Such a combination of features is eminently charac- 
teristic of the Acipenseroid, Crossopterygian, Amioid and 
Lepidosteoid Teleostomi and of the Dipnoi, but is rarely to be 
found in Teleosts, although an approximation thereto may be 
noted in such genera as Arapaima, Heterotis, and Gymnarchus, 
at any rate to the extent that in these Teleostei the ductus is 
botk short and wide. 

In the absence of gas-secreting or gas-absorbing “red glands” 
and “ red bodies,” Notopterus agrees with the generality of those 
Teleosts in which an open ductus pneumaticus is retained 
throughout life. 

From a physiological pomt of view, the most noteworthy 
feature in the air-bladder of Notopterus is its intimate relation 
with the auditory organ. 

The presence of antero-lateral cecal outgrowths from the air- 
bladder is by no means uncommon in Teleostean Fishes, especi- 
ally in the Sparide, Sciznide, Cottide, and Gadide ; and such 
outgrowths may even extend so far forwards as to become more 
or less closely related to the skull, but it is only in a compara- 
tively limited number of genera that the air-bladder acquires any 


AIR-BLADDER IN NOTOPTERUS BORNEENSIS. 525 


special connection or physiological relationship with the auditory 
organ. As regards the precise nature of the connection between 
the two organs in different Fishes, three principal methods may be 
distinguished :— 

A. Auditory ceca are present, and the anterior extremity of 
each is closely applied to a fontanelle in the outer wall of the 
auditory capsule, the utricular portion of the membranous 
labyrinth and the surrounding perilymph being in relation with 
the inner surface of the fibrous membrane by which the fontanelle 
is closed. In no part of their course are the auditory ceca 
enclosed within bony canals or grooves, and no connection 
between the auditory organs of opposite sides of the head has so 
far been described. 

According to Stannius (11. p. 171) this method of connection 
is characteristic of Priacanthus* macrophthalmus, C. & V. 
(P. arenatus, C. & V., or P. cruentatus, C. & V.t), among the 
Serranide ; of certain species of Berycide pertaining to the 
genera Myripristis and Holocentrum; and possibly of Hyodon 
claudulus (H. tergisus, Les.), the solitary representative of the 
North-American freshwater family of the Hyodontide. Jeffery 
Parker (8) bas recorded an essentially similar arrangement in the 
New Zealand Gadoid, Lotella (Pseudophycis) bacchus; and 
Weber (12. pp. 71-72) for such Sparide as Sparus salpa, Linn. 
(Box salpa, C. & V.), and S. sargus, Linn. (Sargus Rondeletz, 
©. & V.). 

B. Instead of being closed by a fibrous membrane, the auditory 
fontanelles are open, and through each of them passes a cecal 
diverticulum from the corresponding utriculus, which thus 
becomes directly and closely applied to the anterior extremity 
of an auditory cecum. 

Those Teleosts which afford examples of this method of 
connection by direct apposition are also characterized by certain 
other noteworthy modifications. Thus, for the greater part of 
its forward course each slender auditory cecum is enclosed in 
bone, first traversing a groove and subsequently a canal in the ex- 
occipital bone, and finally terminates by dividing into two distinet 
vesicular enlargements, of which one lies in a chamber excavated 


* The generic name “ Triacanthus” given by Stannius (/oc. cit.) is apparently 
a misprint for Priacanthus. 

t For synonyms of P. macrophthalmus, vide Brit. Mus. Cat. Fishes, 2nd ed. 
vol. i. p.. 353 & p. 356. 


526 PROF. T. W. BRIDGE ON THE 


in the pterotic bone and has no special relations with the auditory 
organ. ‘he other vesicle occupies a globose chamber in the 
prootic, and there becomes closely applied to the wall of the 
corresponding utricular diverticulum, which enters the chamber 
through an auditory fontanelle in the prootic. In some 
instances there appears also to exist a connection between the two 
utriculi in the form of a transverse sub-cerebral canal, which, 
however, is not to be regarded as homologous with the similarly 
situated ductus endolymphaticus of the Cyprinoid and Siluroid 
Teleosts. Such a utricular connection was first discovered in 
Clupea harengus by Weber (12. p. 77), and subsequently 
by Breschet (1) in C. alosa. On the other hand, the evidence 
as to the existence of a supra-cerebral connection between 
the two sacculi, as affirmed by Hasse (6), or between the two 
utriculi, as stated by Breschet (1. p. 17), is too conflicting to 
admit of any definite conclusion being drawn *. 

The preceding arrangement, which appears to be restricted to 
the physostome family of the Clupeide, was first described by 
Weber in the Herring (Clupea harengus). Recently, Ridewood 
(9. p. 26) has contributed an excellent revision of the anatomical 
relationships of the two organs in the six British species of 
Clupeide, viz.:—Clupea harengus, Linn., the Pilchard (C. pil- 
chardus, Walb.), the Sprat (C. sprattus, L.), the Allis Shad 
(C. alosa, I..), the Thwaite (C. jfinta, Cuv.), and the Anchovy 
(Engraulis enchrasicholus, Cuv.). These species apparently 
include all the Clupeide in which a connection between the 
auditory organ and the air-bladder has so far been described. 

C. In a third and last series of Teleostean fishes, viz., the 
physostome families of the Cyprinide, Characinide, Siluridee, and 
Gymuotide, the connection between the air-bladder and the 
auditory organ attains its maximum complexity and physio- 
logical importance, and is effected, not by auditory ceca, but by 
means of a chain of movable Weberian ossicles. The auditory 
organs of opposite sides of the head are connected together by the 
mesial union of the two endolymphatic ducts, one from each 
sacculus, and the consequent formation of a transverse sub- 
cerebral connection between the two sacculi. From the point of 
union of the two ducts a median sinus endolymphaticus, enclosed 
in a similar median extension of the perilymphatic spaces of the 


* For a discussion of this point, vide Ridewood (9. pp. 38-89). 


AIR-BLADDER IN NOTOPTERUS BORNEENSIS. 527 


two auditory capsules (sinus impar), is prolonged backwards to 
its physiological connection with the most anterior of the series 
of Weberian ossicles *. 

Whether Hyodon claudulus is rightly to be regarded as an 
example of the first method, as stated above, or not, is by no 
means quite clear. The description of the connection between 
the auditory organ and air-bladder in this Teleost, as given by 
Stannius (op. cit. p. 171), is as follows: — 

“ Bei Hyodon claudulus—und ganz analog verhilt sich Nofo- 
pterus—communicirt das vordere Ende desSchwimmblasenkorpers 
durch enge Oeffnungen mit zwei sphiirischen dickwandigen 
Blasen. Jede derselben legt sich in eine Vertiefung der Knochen 
der Hinterhauptsgegend ihrer Seite und haftet eng an den 
letzteren. Dem vorderstea Theile jeder dieser Blasen entspricht 
eine Oeffnung in den Knochen, die inwendig von einem Theile 
des Vestibulum, auswendig aber von der innersten Haut dieser 
Blase bekleidet ist, indem die weisse Faserhaut derselben im 
Umkreise der tiusseren Gehorsoffnung aufhort und nicht tiber 
letztere selbst sich fortsetzt.” 

From this account it would seem that, as in the Clupeide, the 
auditory fontanelles in Hyodon are not closed by fibrous mem- 
branes, and, consequently, the auditory cwxca, after losing their 
outer fibrous coat, are either closely related to the vestibular 
walls, or in actual contact therewith, or, at all events, are not 
separated by any intervening fibrous membrane. If, however, 
the description given by Stannius is correct, then the statement 
“und ganz analog verhalt sich Notopterus”’ is scarcely applicable 
to that Teleost, inasmuch as there is no doubt as to the existence 
of a separating membrane in Wotopterus. 

It would be interesting to ascertain if any tubular communi- 
cation between the two vestibuli, similar to that present in the 
Clupeide, exists also in Hyodon. So far as I am aware, no 
observations on this point have yet been made. 

Comparison of Wotopterus with the various Teleosts mentioned 
above, proves that it furnisbes an additional example of the first 
of the three methods by which the air-bladder and auditory organ 
are brought into physiological relationship, although in one or 
two minor features an approach to the second type is indicated. 

Thus in Notopterus there is no direct contact between the 


* For references vide Bridge & Haddon (2. p. 65 et seq.). 


528 PROF. T. W. BRIDGE ON THE 


walls of the utriculi and those of the auditory cwca, the two 
structures being separated by the membranes which close the 
auditory fontanelles; neither is there any open tubular commu- 
nication between the utricular and saccular portions of the 
auditory organs of opposite sides of the head by means either of 
sub-cerebral or supra-cerebral connections. In these features 
Notopterus closely agrees with those Teleosts in which the first 
method has been adopted. On the other hand, it is equally 
evident that to some extent Notopterws approaches the second 
type and resembles the Clupeide in the fact that, for a part of 
their course, the auditory ceca are enclosed in bony grooves and 
for the terminal portion of their extent occupy the interior of 
bony culs-de-sac, and also in the origin of the ceca from an 
anterior tubular portion of the air-bladder. 

It is perhaps worth remarking that some little variation in 
minor details exists amongst those Teleosts which offer examples 
of the first and second methods, more especially with regard to 
the precise position of the auditory fontanelles and the nature of 
the cranial or periotic bones which form their boundaries, and 
also in the degree of tenuity of the closely related walls of the 
auditory prolongations of the air-bladder. 

For example, in Sparus (Boa) the fontanelle is represented by 
‘Weber (12. tab. vii. fig. 62) as being situated posteriorly to the 
foramen for the exit of the Vagus uerve. In Pseudophycis 
bacchus the auditory fontanelle is described by Parker (loc. cit.) 
as situated between the basioccipital and the opisthotic, and 
immediately beneath the Vagus foramen. In AHolocentrum 
spiniferum, Gthr., the funtanelle is bounded by the exoccipital, 
opisthotic, and prootic bones, and is anterior to the Vagus fora- 
men; and in Wotopterus, as we have seen, the prootic, opisthotie, 
and basioccipital encircle the fontanelle. Finally, in the Clupeide 
the fontanelle is a simple perforation in the prootic alone, and 
hence is more anteriorly placed than in any other Teleosts. 
There can be no doubt, I imagine, that the precise location of 
the fontanelle and the nature of its limiting bones, are due to the 
varying extent to which the auditory ceca are prolonged for- 
ward, and perhaps also to variations in the size of the fontanelle 
itself. 

‘Of much more importance is the degree of tenuity of the 
walls of the auditory ceca at their point of contact or closest 
relationship with the auditory organs ; for upon this will depend 


AIR-BLADDER IN NOTOPTERUS BORNEENSIS. 529 


the degree of intimacy and perfection of the physiological 
connection between the two organs. 

On this point it may be mentioned that in Pseudophycis 
bacchus, as described by Parker (op. cit.), the anterior extremity 
of each auditory cecum forms a thickened pad which is closely 
applied to the membranous sheet closing the corresponding 
auditory fontanelle. From this account it may be legitimately 
inferred that both the outer fibrous coat of the cecum and 
the tunica interna are applied to the fontanelle and its mem- 
brane. In Sparus (Weber, op. cit.), on the contrary, the tunica 
externa of the auditory caecum ceases at the margins of the 
fontanelle, and only the extremely thin tunica interna is applied 
to the membrane which closes the aperture. So far as this point 
is concerned Notopterus closely resembles Sparus; but in the 
Clupeidz, and possibly in Hyodon, the connection of the two 
organs becomes even more intimate, inasmuch as it is effected 
by the actual apposition of the limiting walls of auditory ceca: 
and utricular outgrowths. 

The derivation of the second and more intimate type of con- 
nection between the air-bladder and the auditory organ from the 
first method is easy to imagine, as the process simply involves 
the atrophy of the fibrous membranes closing the auditory 
fontanelles, so as to admit of the actual contact of the two 
organs by the direct apposition of outgrowths from each. A 
much more difficult problem is the genesis of the third method 
by means of Weberian ossicles, and so far no satisfactory solution 
has yet been offered. There is every probability, however, that 
the Weberian mechanism has been independently evolved, but 
that the initial stages of its evolution have not yet been dis- 
covered, if they exist, in any Fishes at present living. 

In his account of the connection between the air-bladder 
and auditory organ in Psewdophycis bacchus, Parker (op. cit.) 
remarks : “ The anterior end of the air-bladder fits closely against 
the hinder end of the skull and is produced outwards into paired 
pouches which are in contact with the thin skin beneath the oper- 
culum and in front of the shoulder-girdle.” To this arrangement 
Notopterus offers a parallel in the close relation of the auditory 
cea to the cecal diverticula of the branchial cavity, and, there- 
fore, to the external medium in which the fish lives. . The 
Siluroids (2) also afford examples of a somewhat similar 
modification. In the majority of these Teleosts the lateral walls 


530 PROF. T. W. BRIDGE ON THE 


of the anterior chamber of the air-bladder are in contact, often 
over a considerable area, with the superficial skin behind the 
pectoral girdle. 

Notopterus is by no means the ouly Teleost in which the 
existence of cranial fontanelles, more or less closely associated 
with the auditory organ, has been recorded. Nevertheless, the 
position of these vacuities, and their precise relation to the 
membranous labyrinth, exhibit a wide range of variation in 
different Teleosts. 

In Notopterus, as we have seen, the fontanelles are situated on 
the upper surface of the hinder part of the skull and immediately 
dorsad to the auditory capsules. 

According to Stannius (11. p. 170) cranial fontanelles essen- 
tially similar to those of Notopterus exist also in Hyodon 
claudulus. 

In the Herring (Clupea harengus, L.) “the ventro-external 
surface of the sacculus lies over a membranous fenestra of the 
skull-wall, situated between the exoccipital, basioccipital, and 
prootic bones, and described by several authors as a foramen 
ovale; the sacculus-wall is here separated from the mucous 
membrane of the mouth by this membrane only ” (Ridewood, op. 
cit. p. 38). 

In referring to certain of the Macruride, Stannius (op. cit. 
p. 170)* states :—‘‘ Bei Lepidoleprus trachyrhynchus t findet 
sich seitlich am Hinterkopfe tuber dem oberen Ende der Kiemen- 
spalte eine trichterformige von dinner Haut gesschlossene Grube, 
welche in den zur Aufnahme des Gehororganes bestimmten 
Theil der Schedelhohle hineinragt. Zwischen der Innenfliche 
ihrer Haut und dem Labyrinthe liegt eine faserig-gallertartige 
Substanz.” Similar fontanelles are said to be present in Z. celo- 
rhynchus, but not in L. norvegicus. 

The skin-covered fontanelles on the postero-superior surface 
of the skull of Mormyrus somewhat resemble those of Lepido- 
leprus, except that immediately beneath the external skin there 
is a perfectly free and very thin lamina of bone which partially 
covers each fontanelle (Stannius, op. cit. p.170¢). Fontanelles, 


* On the authority of Otto, Tiedemann u. Treviranus Zeitschrift f. Physiol., 
Bd. ii. 8. 86 

t =Macrurus trachyrhynchus, Risso. 

{ Apparently on the authority of 8. Heusinger, Meckel’s Archiv f. Anat. u. 
Physiol. 1827, Bd. i. S. 324. 


ATR-BLADDER IN NOTOPTERUS BORNEENSIS. 531 


similar to those of Mormyrus, are also said to be present in 
Gymnarchus niloticus (Stannius, J. c. *). 

Mention may also be made of the prevalence of median 
fontanelles in the cranial roof in many Siluroids, although in 
these Fishes the structures in question have no special relation 
to the auditory capsules f. 

Tt is certainly worthy of remark that the existence of cranial 
fontanelles, as a possible avenue for the transmission of sound 
vibrations from the water to the auditory apparatus, has been 
recorded not only for those Fishes in which the air-bladder has 
no connection with the membranous labyrinth (e. g., Mormyrus, 
Macrurus), but also, and apparently quite as frequently, even in 
those Teleosts in which such a connection is known to exist 
(e. g., Clupea, Hyodon, and Notopterus). 


IX. REMARKS ON THE suPPOSED AuDIToRY FUNCTION 
OF THE AIR-BLADDER. 

The physiological value of the connection between the air- 
bladder and the auditory organ is still one of the unsolved 
problems of comparative physiology, and this statement seems 
to be equally applicable to any of the three principal methods 
by which such a connection is established. The reason for this 
is, no doubt, that practically nearly all attempts to solve the 
problem have been based upon anatomical evidence alone ; 
and, as must be admitted, morphological data are often unre- 
liable, and even misleading, when employed as the sole basis for 
physiological deductions, and not infrequently appear to support 
with remarkable impartiality conclusions of a widely different 
character. When the problem has received the attention of the 
experimental physiologist, then, and then only, may a final and 
satisfactory solution be expected. 

The question has been most frequently discussed in connection 
with the Ostariophysez {, and, so far, comparatively little atten- 
tion has been devoted to those Teleosts (e. g., Sparus, Notopterus, 
Hyodon, Clupea, &c.) in which the connection of the air-bladder 
and auditory organ is effected by other means than the presence 
of Weberian ossicles. In venturing to discuss certain physio- 


* According to Hrdl, see Stannius, /. c., footnote to p. 170. 

+ Professor Howes has reminded me of the large lateral occipital fontanelles 
in the Cyprinide (c.f. Sagemehl, Morph. Jhb. Bd. xvii. p. 495). 

{ The physostomous families of the Cyprinids, Siluride, Characinide, 
and Gymnotide, in which a Weberian mechanism is present. 


5382 PROF. T. W. BRIDGE ON THE 


logical points in connection with the latter Fishes, I must again 
emphasize my conviction that physiological conclusions drawn 
solely from anatomical data must be regarded as little more than 
tentative and provisional suggestions. 

At the conclusion of his account of the mode of connection 
between the air-bladder and the auditory organ in certain species 
of Sparus, Weber (op. cit. p. 72) states his views as to the 
physiological value of the connection in the following words :— 

“Prima utilitas hee est, ut tremoros soniferi aque toti 
piscium corpori communicati, a vesica natatoria elastica recipi- 
anter, in se colliquantur, et in membranam, fenestram vestibuli 
ossei obducentem, transferantur, cuius tremoros liquorem, cavum 
cranii replentem, movent itaque vestibulem membranaceum 
ipsum, ab aqua cranii circumdatum, afficiunt ad quam quidem 
utilitatem vesica natatoria, quippe que singulis costis affixa est, 
aptissima videtur.” 

On this theory the air-bladder, by reason of its connection 
with the auditory organ, becomes a physiological accessory to 
audition, in addition to its usual and normal function as a hydro- 
static organ. 

In discussing Weber’s view of the auditory function of the air- 
bladder and Weberian ossicles in the Ostariophysex, it was 
pointed out by Bridge and Haddon (2. p. 276) that, assuming 
sound-waves to be transmitted from the air-bladder to the 
auditory organ by these ossicles, the fact that such stimuli 
would, in the first instance, affect certain median and unpaired 
structures common to both auditory organs (sinus impar and 
sinus endolymphaticus) involved the difficulty that the mem- 
branous labyrinths of opposite sides of the head would be affected 
with equal intensity and simultaneously, and, consequently, as 
no differential stimulation took place, the Fish would be incapable 
of appreciating the direction of the sound. 

Sérensen (10. pp. 185-186), in attempting to mitigate the force 
of this criticism, remarks :—“ As far as I can see this is the only 
real objection that can be made against the theory of Weber. 
But I do not judge it to be of great importance. If this 
objection were absolutely valuable, no human being would be 
able to decide if a sound arises before or behind him when the 
direction of the sound coincides with the symmetrical plane of the 
body ; and this, however, we are able to decide. And, would 
not this objection be just as valuable if you presume the sound 
not to be transmitted to the ear through the air-bladder? Are 


AIR-BLADDER IN NOTOPTERUS BORNEENSIS. 53a 


not the Fishes deprived of the means by which the higher 
Vertebrata are able to judge the direction of the sound, viz., by 
turning the head (or external ears) to the right or to the left? 
It must also be kept in mind that it is always very difficult 
thoroughly to understand that beings differently conditioned 
know how to use the powers with which they are endowed, 
especially when these powers are inferior to those bestowed 
upon us.” 

Sérenson’s attempt to lessen the importance of what he terms 
the only “real objection” to Weber’s theory does not impress 
one as being very successful. ‘The capacity for appreciating the 
direction of sounds, whether employed as a means of securing 
prey, or escaping from enemies, or, as in the case of gregarious 
Fishes, as a means of keeping together in shoals for breeding or 
other purposes, must be of primary importance in any modifica- 
tion of the auditory organ in the direction of giving to its 
possessor exceptional powers of hearing. Fishes may have no 
power of rotating or inclining the head, but it must not be for- 
gotten that a slight deflection of the long axis of the body to the 
right or left will at once enable the differential action of the two 
auditory organs to come into play, and the Fish would be in a 
position to appreciate the direction from which the sound is 
travelling. An Ostariophysean, in so far as those parts of the 
auditory apparatus which it possesses in common with all other 
Fishes are concerned, is in much the same position as regards 
the sense of direction. It is quite true that the two auditory 
organs are in open communication by means of a sub-cerebral 
transverse ductus endolymphaticus ; but it is also obvious, I 
think, that the auditory organ, right or left, turned towards 
the direction of the sound, will be stimulated appreciably sooner, 
or it may be more forcibly, than its fellow, and hence the cog- 
nizance of direction. Again, is it not possible, or even probable, 
that sound-waves reaching the auditory organs simultaneously 
by two distinct ebannels will have their effect nullified, and 
the sense of direction seriously interfered with? In fact it 
seems extremely probable that sound stimuli which, according 
to Weber’s hypothesis, are received by the auditory organs 
through the air-bladder and Weberian ossicles, would have 
the effect of confusing any sense of direction based upon 
similar stimuli, generated by the same cause and at the same 
moment, but pursuing, as in the generality of Fishes, the usual 

LINN. JOURN.—ZOOLOGY, VOL. XXVII. 40 


584 PROF. T. W. BRIDGE ON THE 


direct path through the bones and other structures of the head 
to the membranous labyrinths. It may be also urged that this 
objection loses none of its force when it is considered that the 
cranial bones are far more pervious to sound-waves travelling in 
water than, as is the case with terrestrial animals, in air. 

If this argument has any force, does it not suggest that the 
Weberian mechanism, cousidered as an accessory to audition, 
may prove a positive disadvantage to the Fish in so far as the 
sense of direction is concerned ? Finally, if the Weberian 
mechanism is a means of increasing the acuteness of the sense of 
hearing, is it not a little remarkable that sounds heard through 
this agency convey no idea of direction, especially when the 
latter is of so much importance to the animal, and that the 
sense of direction should depend upon the perception of sounds 
of an obviously more limited range of intensity which reach the 
auditory organs directly through the head ? 

Other objections which may be urged against Weber’s theory 
as applied to the Ostariopbysex are scarcely pertinent to the 
present discussion, which is: designedly restricted to those Fishes 
in which there is no Weberian apparatus and the auditory organ 
and air-bladder are connected by other methods. 

It may at once be affirmed that the same “ real objection ” is 
equally applicable to such Fishes as Wotepterus, Sargus, Sparus, 
Hyodon, Clupea. 

In some of these Fishes (e. g., Sparus, Sargus, and Clupea) 
there is no longitudinal bipartition of the air-bladder, which, 
therefore, for the greater. part of its extent encloses a simple 
undivided cavity. Sound-waves in the gases of the air-bladder 
travelling along the auditory ceca will ultimately affect the 
auditory organs to an equal extent and simultaneously. Hence, 
these Fishes will derive no sense of direction from auditory 
stimuli reaching the membranous labyrinths by such channels. 
Whatever sense of direction they possess will be derived from 
those stimuli which reach the auditory organs by the usual path 
through the skull; but here, as in the Ostariophysex, we are con- 
fronted with the probability that such sense of direction will be 
interfered with by the stimuli received through the air-bladder. 

From this point of view the condition of the air-bladder 
in Notopterus is especially significant. For more than three- 
fourths of its extent the organ consists of two lateral chambers ; 
and if, by an anterior extension of the longitudinal septum, these 


AIR-BLADDER IN NOTOPTERUS BORNEENSIS. 535 


compartments had become entirely separated from each other, 
and each had remained continuous with the auditory cecum of 
its side, it is at least conjecturable that each chamber would be 
somewhat differently affected by sound-waves impinging on its 
own side of the body and eventually propagated to the auditory 
organ of the same side. In this way,it seems possible that 
stimuli received by the auditory organ through the intervention 
of the air-bladder might be competent to give rise to a sense 
of direction. So far, however, from this being the case, the 
longitudinal septum ceases at the very point where its forward 
prolongation seems to be most desirable; and, in consequence, the 
auditory ceca arise from an anterior, median, tubular portion of 
the bladder, which is only slightly divided longitudinally by the 
carinate growth of the suprajacent axial skeleton. Hence, as in 
non-septate air-bladders, sound-waves are propagated to the 
auditory organs simultaneously and with equal intensity. 

There is also another point which deserves consideration in 
discussing the possible auditory function of the air-bladder in 
these Fishes. 

Well-marked cranial fontanelies are present in several of 
them, and these structures are not only paired but each is in 
immediate relation with the auditory organ of its side of the 
head. Do not these skin-closed fontanelles afford a better 
channel for the transmission of sound-vibrations to the mem- 
branous labyrinths than the air-bladder ?, and is not one of these 
structures somewhat superfluous if we regard both of them as 
sound-transmitting organs? Moreover, so far from interfering 
with the sense of direction, cranial fontanelles would probably 
facilitate the differential action of the two auditory organs. 

If the connection of the air-bladder with the auditory organ is 
not subservient to the sense of hearing, can any other function 
be assigned to it? 

In the case of the Ostariophysee, Ramsay Wright (13) and 
Bridge and Haddon (2) have supported the view that the 
connection of the two organs was possibly for the purpose of 
enabling these Fishes to appreciate the varying degrees of tension 
of the gases in the air-bladder, resulting from corresponding 
variations of hydrostatic pressure produced by locomotor move- 
ments involving differences of depth. Subsequent reflex or 
voluntary impulses, it was suggested, might find expression in 
the exercise of some form of regulatory control over the liberation 

40* 


536 PROF. T. W. BRIDGE ON THE 


of gas through the ductus pneumaticus, so that only so much gas 
will be eliminated as will suffice to maintain the Fish in a plane 
of least effort. 

In attempting to extend this theory to such Teleosts as 
Sargus, Hyodon, Priacanthus, Notopterus, and Clupea, it must 
be at once admitted that there are grave difficulties in the way. 
It is easy to conceive how distension or contraction of the air- 
bladder, produced by variations in the superincumbent column of 
water, would be competent to give rise to stimuli affecting the 
auditory organ. A tendency to over-distension may be conceived 
to produce such a bulging of the anterior extremities of the 
auditory ceca as would modify the condition of the perilymph 
surrounding the auditory organs, and impart a stimulus to the 
sensory epithelium of those organs. The objection to this view 
is, however, that it is very difficult to see in what way the 
contingent efferent impulses will find expression. In Fishes like 
Notopterus and Clupea, where a ductus pneumaticus is present, 
the existence of some kind of regulatory control over the libera- 
tior. of gas from the air-bladder is possible; but this suggestion 
is obviously inapplicable to such Fishes as, for example, the 
species of Sparus and Sargus in which the ductus atropbies in 
the adult. In the latter genera, variations in the amount of gas 
present in tke air-bladder must depend upon the relatively slow 
proccsses of gaseous secretion or absorption; and it 1s at least 
within the bounds of conjecture that the connection of the air- 
bladder and auditory organ forms part of a reflex mechanism by 
which the varying tensions of the gases of the air-bladder cou- 
stitute a stimulus to the auditory organ and central nervous 
system, and, ultimately, by reflex action lead to such a modifica- 
tion of the rate of secretion or absorption as will vary the amount 
of gas in the bladder in accordance with the requirements of tlie 
Fish. The special advantage to the Fish may be that secretion 
and absorption will take place more rapidly than is the case 
where pressure-stimuli have no special means of affecting a 
sensory organ, and adjustment to varying hydrostatic pressures 
effected with greater promptitude. Nevertheless, it must be 
acknowledged that there is at present but very little evidence, 
either physiological or anatomical, which can be adduced in 
support of these suggestions. 

Admitting, however, the want, or rather the paucity, of 
evidence for such tentative suggestions, is it not possible that the 
connection of the auditory organ and the air-bladder may have 


AIR-BLADDER IN NOTOPTERUS RORNEENSIS. 537 


to do with a simpler physiological réle? The danger which a 
Fish incurs from over-distension of the air-bladder as the result 
of a too rapid rise in the water to the reduced pressure of a 
higher level has been emphasized elsewhere (2); and, without 
involving any alteration in the rate of gaseous absorption, the 
afferent impulses communicated to the auditory organ may, 
through appropriate efferent channels, lead to such modifications 
ot the locomotor movements as will enable the Fish to guard 
against over-distension, and return to its normal plane of 
equilibrium or “ least effort”? at a greater depth. 

It is a significant illustration of the difficulties of the problem 
that even Weber, who was primarily responsible for the theory 
of the auditory function cf the air-bladder in those Fishes in 
which that organ is connected with the membranous labyrinth, 
was sufficiently far-seeing to admit the possibility of a second 
function. Immediately following the quotation with reference 
to Sparus, which has been previously given, Weber proceeds 
(12. pp. 72-73) :— 

“ Altera vesice natatoriz utilitas hee est, ut aére vesice com- 
pressione alvi in appendices superiores impulso, membranaque 
fenestram utramque vestibuli ossei obducente tensa et introrsum 
pressa, aqua cranii cavum replens adeoque vestibulum membra- 
naceum, ad hac aqua circumdatum ipsum prematus. 

“Quam ob rem, si cranio aperto et labyrintho rmembranaceo 
conspicuo vesiza natatoria manu comprimatur, facile hic vesice 
natatorie contracte effectus oculis percipitur. Per vesicam enim 
manu compressam liquor cranium replens non solum propellitur, 
sed vestibulum membranaceum ipsum quoque motu liquoris 
cranil commovetur.”’ 

The two paragraphs just quoted suggest that Weber certainly 
entertained the idea of the air-bladder being the means of con- 
veying to the auditory organ stimuli due to its varying degrees 
of distension, such as, we may presume, are naturally brought 
about by variations of depth and, therefore, of pressure; and,. 
further, prompt the remark that the term “ Weberian theory ” 
may be as reasonably applied to this view as to the “ auditory 
theory ” with which the name is more generally associated. 

In conclusion, it may be affirmed that there are obvious objec- 
tions both to the auditory and hydrostatic views of the physio- 
logical raison d’étre of the conuection between the air-bladder 
and the auditory organs, which, in the present state of our 
kuowledge of the individual functions of those organs, cannot 


538 PROF. T. W. BRIDGE ON THE 


easily be explained away in accordance with either theory. With 
a more precise and definite acquaintance with the physiology of 
the organs in question, and a more extended knowledge of the 
habits and mode of life of the Fishes concerned, it is possible 
that some definite conclusions may be arrived at; but the final 
solution of the problem must rest with the experimental 
physiologist. 

Whatever the significance of the inter-reJationship of the air- 
bladder and the auditory organ, there can be no question as to 
its physiological importance. Such a connection presents varying 
degrees of perfection and specialization in different Teleosts, and 
its independent evolution in widely different families seems to 
suggest the possibility that it is correlated with the requirements 
of special or local conditions of life, and, while attaining its 
maximum development in the dominant families of freshwater 
Teleosts *, is nevertheless present in simpler forms in a limited 
number of both freshwater and marine Fishes. 


X. BIBLIOGRAPHY. 


1. BrescuEet.—Recherches sur lorgane de l’oule des Poissons. 
Paris, 1838. 

2. Briper, T. W., & Happon, A. C.—‘‘ Contributions to the 
Anatomy of Fishes. II. The Air-bladder and Weberian 
Ossicles in the Siluroid Fishes.” Phil. Trans. Roy. Soc. 
vol. 184, 1893, B. pp. 65-333. 

3. Bripar, T. W., & Happon, A. C.—‘‘ The Air-bladder and 
Weberian Ossicles in the Siluroid Fishes.” Proc. Roy. 
Soe. vol. li. 1892, pp. 189-157. 

4. Cuvier & Vatenctennes.—Histoire Naturelle des Poissons. 
Tome xxi., 1848. 

5. Cuvier & VaLencrennes.—Histvoire Naturelle des Poissons. 
Tome xx., 1847. 

6. Hasse, C.—‘* Beobachtungen tiber die Schwimmblase der 
Fische.” Anatomische Studien, Bd. i. 1873, pp. 583-610. 

7. Owzn, R.—On the Anatomyjof Vertebrates, vol. i. Londen, 
1866. 

8. Parker, T. J.—‘‘ On the Connection of the Air-bladder and 
the Auditory Organ in the Red Cod (Lotella bacchus).” 
Trans. New Zealand Institute, vol. xv. 1882, p. 284. 
(Abstract.) 


* 3. p. 153. 


AIR-BLADDER IN NOTOPTERUS BORNEENSIS. 539 


9. RripEwoop, W. G.—‘‘ The Air-bladder and Ear of British 
Clupeoid Fishes.” Journ. Anat. & Phys. vol. xxvi. 1892, 
pp. 26-42. 

10. Sédrensen, W.—“ Are the Extrinsic Muscles of the Air- 
bladder in some Siluride, and the Elastic Spring- 
apparatus of others, subordinate to Voluntary production 
of Sounds?” ‘“ What is, according to our present know- 
ledge, the Function of the Weberian Ossicles?” Journ. 
Anat. & Phys. vol. xxix., 1894-95. 

11. Srannrus, H.—Handbuch der Zootomie der Wirbelthiere. 
Die Fische. Berlin, 1854. 

12. Weser, E. H.—“‘ De aure et auditu Hominis et Animalium. 
Pars i., De aure Animalium Aquatilium.” Lipsiz, 1820. 

13. Wrieut, R. Ramsay, McMurricn, J. P., Macurium, 
A. B., & McKenztz, T.—“ Contributions to the Anatomy 
of Amiurus.”’ Toronto, 1884. Reprinted from the Proc. 
Canadian Institute (n. s.), vol. 11. 1884, p. 251. 


XI. EXPLANATION OF THE PLATES. 
PuatEe 36. 


Fig.1. Lateral view of the left branchial cavity, to show the relation of its 

anterior cecal extension to the left auditory cornu of the air-bladder. 
The operculum and pectoral girdle have been removed. (Nat. size.) 

Fig. 2. View of the interior of a portion of the left caudal prolongation of the 
air-bladder. The outer wall of the bladder has been partly removed 
and partly reflected above and below. A portion of the outer wall of 
a ventral diverticulum has also been cut away in order to show one 
of the ventral series of vacuities. (Nat. size.) 

Fig. 8. Lateral view of the anterior and abdominal portions of the air-bladder. 
The outer wall of the left half of the bladder, except that portion of it 
which is traversed by the oblique transverse groove between the sub- 
spherical sac and the abdominal part of the organ, which has been left 
as a narrow oblique strip, has been removed, so as to show the longi- 
tudinal septum throvghout its entire ength. The outer wall of the 
left auditory czxcum has been removed, and the esophagus and ductus 
pneumaticus laid open. (X 2.) 


Puate 37. 
Fig. 4. Lateral view of the posterior portion of the right side of the skull. 


Fig. 5. oe the interior of the hinder part of the right half of the cranial 
cavity, as seen in a vertical longitudinal section of the skull. (x 2.) 

Fig. 6. Supero-lateral view of the posterior portion of the right side of the 
skull. (x 2.) 


540 


AIR-BLADDER IN NOTOPTERUS BORNEENSIS. 


Reference letters. 


a.o.c. Anterior diverticulum of the left branchial cavity. 


ad.p. 
af. 


als. 
ap. 


Abdominal portion of the air-bladder. 

Auditory fontanelle. 

Alisphenoid. 

Accessory articular process for the centrum of the first vertebra. 


. Basioccipital groove. 
. Basioccipital. 
. First and second branchial arches. 


Cranial fontanelle. 


. Entrance to bony cul-de-sac. 

. Ductus pneumaticus, 

. Dorsal vacuity. 

- Dorsal wall of the air-bladder. 
. Exoccipital. 

. Epiotic. 


Frontal. 


. Kiliform ceca. 
. Lateral groove for the sensory canal of the head. 
. Heemal spine. 


Left auditory cecum. 


. Left caudal ceecum of the air-bladder. 

. The inpushed dorsal wall of the tubular portion of the bladder. 

. Longitudinal septum in the abdominal part of the bladder. 

. Longitudinal septum in the subspherical sac. 

. Cisophagus. 

. Oblique groove separating the subspherical sac from the abdominal 


part of the bladder. 


. Opisthotic. 

. Parietal. 

. Prootic. 

. Parasphenoid. 

. Pterotic. 

. Aperture leading to right auditory csecum. 
. Radial or interspinous element of anal fin. 
. Subspherical sac. 

. Sphenotic. 

. Supraoccipital. 

. Tubular portion of the air-bladder. 

. Transverse septum. 

. Utriculo-saccular recess in the basioccipital. 
. First vertebra. 

. Ventral diverticulum. 

. Ventral vacuity. 


Foramen for the exit of the Vagus nerve. 


Bridége. | Lins Soc. Journ. Doon Vou. XXVII Pr.36. 


Ga nat. size. 


3 nat. Size. 


FW. Crispe del. ad nat. 
ME Parker Tithe Geo, West & Sons imp. 


NOTOPTERUS BORNEENSIS. 


Linw Soc. Journ. Zoou. Vou. XXVII Pi.37 


| Bridge. 


iss 
x< 


EE ESOS 


ee 


SOas= 


KR 


Geo, West & Sons imp. 


E.J.Partridge del.ad nat. 
M.P Parker lith. 


NOTOPTERUS BORNEENSIS. 


ONS 
ani 


Kemet eh 
Quang 
mayen os 


ON NASAL SECRETORY SACS IN TELEOSTEI. 541 


On the Presence of Nasal Secretory Sacs and a Naso-pharyngeal 
Communication in Teleostei, with especial reference 
to Cynoglossus semilevis, Gthr. By H. M. Kyzz, M.A. 
(Communicated by Prof. G. B. Howes, Sec. Linn. Soc.) 


[Read 18th January, 1900-] 
(PuaTE 38.) 


THE observations embodied in this paper have been made 
during the course of a prolonged research into the anatomy 
of the Flat-fishes (Heterosomata). It is considered advisable to 
publish them separately because, though the research is far from 
completed, the facts to be described have a certain importance 
apart from the main series of results. In order to carry on my 
work successfully I spent some time at the British Museum of 
Natural History, and I desire to acknowledge my indebtedness to 
Mr. G. A. Boulenger, F.R.S., of the Zoological Department of that 
Institution. Every facility was given to me for my work, and his 
advice, generously offered, has aided mein many directions. I am 
proud to acknowledge also the friendly counsel and masterly 
criticism of Prof. G. B. Howes, F.R.S., to whom indeed the 
appearance of this paper in its present form is due. 

It is generally believed and taught that Fishes possess no 
secretory apparatus in connection with their olfactory organs, and 
that in the Teleostei these organs have no direct communication 
with the mouth. These characters, and more especially the 
latter, have been considered as almost exclusively distinctive of 
the air-breathing Vertebrates—so much so that Huxley, in his 
famous paper on Ceratodus Forsteri*, discussing the communica- 
tion present in the Dipnoi, considered it necessary to raise and 
answer the question—of what use are “‘ internal nares ” to purely 
branchiate animals? Internal nares in water-breathing verte- 
brates seem indeed unnecessary, because the respiratory apparatus 
is in immediate communication with the mouth, so that where 
they do occur a special explanation has to be sought. As 
with the Dipnoi and Myzxine, so with the case to be herein 
described, the function of the naso-pharyngeal communication 
seems quite clear, though its presence may run counter to our 
preconceived notions, 


* Proe. Zool. Soc. 1876, cf. pp. 24 & 180. 


542 MR. H. M. KYLE ON NASAL 


In order to obviate a confusion of terms which might readily 
arise in dealing with the subject of this paper, I shall refer to 
the organ of smell as the olfactory organ; to certain accessory 
derivatives of that organ as nasal sacs; to the one or two external 
apertures of the olfactory organ of the Teleostei as anterior and 
posterior nostrils; and to the apertures and passages of com- 
munication between the nose and the month as the posterzor or 
internal nares. 

The existence of nasal sacs in connection with the olfactory 
organs of Teleostei has been noted by Owen, who in ‘The Anatomy 
of Vertebrates’ (p. 329) mentions their occurrence in the Mack- 
erel (Scomber scombrus L.) and in the Wolf-fish (Anarrhichas 
lupus, L.). In the latter he states that the “reservoir passes 
backward (expanding) as far as the back part of the palate, 
where it ends blindly ; ” and adds, “ the prolongation of the single 
nasal cavity in the Lamprey is analogous to this.” Although 
Owen* mentions the Mackerel as possessing these reservoirs, 
they are comparatively slightly developed. Solgerr has more 
recently discovered a similar organ in the Stickleback. 

These are not the only species, however, where such “ reser- 
voirs”” occur. They are met with in those forms which lead 
a semi-sedentary life—in the Blenniide, and (more largely 
developed) in the Labride and Scorpenide. Sometimes there 
are two “sacs” in connection with each olfactory organ— 
Scorpenide ; sometimes only one—Blennide and Labride. 

These sacs are simple continuations of the nasal cavity and 
possess no secretory function. They are “ reservoirs” as Owen 
called them, associated with the retaining, circulation, and 
changing of the water which passes through the nostrils. Their 
walls are not muscular, and their capacity is determined merely 
by the movements of the ascending processes of the premaxille 
and the maxille and palatines. Solger suggests that they may 
also furnish a habitat and breeding-place for Infusoria ! 

A further specialization of the nasal sacs is found in certain 
groups of flat-fishes, and here also the number present for 
each olfactory organ varies—in the Halibut, Plaice, and Turbot 


* Milne-Edwards also states that the nasal organ in the Mackerel possessed 
a ‘cul-de-sac.’ Leg. sur la Physiologie et l’Anat. Comp. tom. xi., 1877, p. 4/6. 

+ Solger, B.: ‘ Notiz tiber die Nebenhohle des Geruchsorgans von Gastrosteus 
aculeatus, L.” Zeit. fiir Wiss. Zool., Bd. 57, 1894, p. 186. 


SECRETORY SACS IN THLEOSTEI. 543 


there are two, in the Sole tribe only one, in the aberrant Sail- 
fluke (Lepidorhombus whiff) none at all. In the Halibut, Plaice, 
and Turbot tribes, when the sacs are developed they lie anterior 
to the ethmoid, and are closely connected with the pterygo- 
palatine and maxillary bones (Pl. 38. fig. 1, m.s.,-n.s.,). In the 
genus Solea that of the right or eyed side also lies anterior to the 
ethmoid, whilst that of the left or blind side extends posteriorly 
over the roof of the mouth to as far back as the posterior third 
of the parasphenoid (fig. 2, U.n.s.). In the above-mentioned 
groups the function of these sacs is entirely secretory, their 
blind ends resting on a layer of fatty tissue immediately over 
the integument lining the roof of the mouth (fig. 1, f¢.). In 
the genus Solea the larger, posteriorly directed sac acts for the | 
most part as a reservoir just as in the cases cited. A small 
quantity of mucus is certainly always found in it, but mixed with 
a large percentage of sea-water. 

In order to render fully clear the significance of these facts, it 
is necessary to state the correlated differences in form and struc- 
ture of the several species of flat-fishes. This would lead too far 
away, however, from the subject at present in hand, and it need 
only be said that the Soles are better adapted for more sand- 
loving habits than are the Halibut, Plaice, and Turbot. 

Leaving the physiological significance of these sacs for dis- 
cussion later, we may turn now toa remarkable form first de- 
scribed by Dr. Giinther *, a native of the China Seas, and called 
by him Cynoglossus semilevis. This species is included under a 
sub-family of the Sole group, but is well marked off from the 
true Soles, and is probably of separate origin. In European 
waters it has a near ally in the small Ammopleurops (Plagusia 
of the French ichthyologists) of the Mediterranean. The charac- 
teristics of the true Soles—the curved snout projecting in front of 
the mouth, the small eyes and largely developed olfactory organs, 
the slender and slightly developed opereular bones with contours 
completely hidden by the skin, the comparatively small branchial 
openings, and various other internal peculiarities of skull and 
skeleton—are accentuated in Cynoglossus, and indicate a greater 
adaption to sand-loving habits than is found even among the 
true Soles. The degeneration of the fins evidenced in these 
latter reaches its extreme. The pectoral fins have entirely 


* Ann. & Mag. Nat. Hist. ser. 4, xii. 1873, p. 379, and ser. 7, i. 1898, p. 261. 


544 MR. Il, M. KYLE ON NASAL 

disappeared and one ventral only remains, the dorsal and anal are 
continuous round the tail, and the tail itself, carrying further 
the change begun in Solea, has become “ pseudo-diphycercal.” 

These characteristics give the impression not merely of sand- 
loving habits, but also that the animals bury their comparatively 
heavy jaws and snout in the sand or gravel, whilst their long, 
slender, and flexible tails move freely in the water. And in 
this position they are more dependent upon their sense of 
smell than upon their powers of sight for the detection of their 
food. 

Correlated with their mode of life in some mysterious way is 
the peculiar development of their lateral-line system. On the 
right or blind side there is only one lateral line, on the left or 
eyed side there are three of them—one median, the other two near 
the bases of the dorsal and anal fins. The median and dorsal 
lines are continued on to the head, over the occipital region of 
which a connecting branch joins the two. The median line gives 
off the usual branch over the preoperculum to the mandible, and 
continues forward almost to the anterior border of the head, 
where it joins the dorsal line, which has followed the contour of 
the head and is continued round the border to the curved snout. 

Five specimens referred to this species, Cynoglossus semilevis, 
have been examined, one of them in detail, the other four only 
with regard to certain doubtful points *. Although these are 
classed by Dr. Gunther as representatives of one species, the 
individual examined in detail differs so markedly from the 
other four that some systematists would not hesitate to make it 
the type of a separate species or even genus, and the advisa- 
bility of this will be considered when the facts concerning it 
have been described. 

The characters already enumerated are shared by all five, and 
in external appearance there is only one point of difference 
between the divergent specimen and the others. In it, an acces- 
sory branch of the lateral line passes backward from the ring 
round the snout towards the posterior nostril between the eyes. 
In the others this is absent. Itis difficult to say what value can 
be put upon this character. In many nearly allied species, e. g. 
Synaptura, it 1s in exactly the same position ; and in many of the 
North American flat-fishes (especially on the Pacific coast) 


* As these were registered specimens, I was very fortunate in having so 
many to examine and in being permitted to handle them so fully. 


SECRETORY SACS IN TELEOSTEI. 545 


Jordan* describes an accessory branch of the lateral line as 
passing backward from the head along the back of the dorsal 
fin. It is connected with other peculiarities, and must have 
some relation to the mode of life of the different species; and 
in the case of Cynoglessus it is present along with certain 
characters which plainly show the greater adaptation of its 
possessor to more sand-loving habits than those which do not 
have it. 

In the case of Solea, it has been said that one of the nasal sacs 
extends backward over the roof of the mouth, whilst the other 
is quite separate and lies anterior to the ethmoid. In the four 
specimens of Cynoglossus, so far as could be ascertained by 
means of a seeker without actual dissection, there is a single 
larze sac lying over the roof of the mouth, resembling in position 
the large posteriorly directed sac of Solea. Into this both nasal 
canals open—one from each nasal cavity, The nasal organs, 
which are placed symmetrically on each side of the head, are thus 
in communication with a large “cul-de-sac ” which occupies the 
entire area overlying the median portion of the roof of the mouth. 

The step is not great from this combination to that found in 
the divergent specimen. The roof of the mouth in this jig 
pertorated by a large oval opening (PI. 38. figs. 3, 4, 5, ¢), around 
which the mucous membrane is thrown into a broad rim or fold, 
projecting inward and underlying a portion of the central 
chamber, which in position corresponds to that enclosed by the 
sac of the other four specimens. When this rim ig cut through 
anteriorly, two comparatively large openings are seen (fig. 5) 
—one on each side of the median line. These lead into 
the nasal canals which pass upwards and forwards—one on 
each side of the parasphenoid—internal to the palatines, until 
they reach positions anterior to and alongside of the ethmoid, 
and open at their upper extremity into the posterior portion of 
the nasal cavities (fig. 3,¢). On the eyed side of tke head the 
canal passes downward from the nasal cavity close to the 
membranous lining of the lower orbit (fig. 3, m.p.c.). These 
canals are tolerably large, and form an effective means of 
communication from the exterior, through the nasal cavities, 
to the mouth. 


* Jordan, D.S., & Goss, D. K.: “A Review of the Flounders and Soles 
(Pleuronectidz),” Rep. U.S. Comm. of Fish and Fisheries, 1886.—Jordan, D.S., 
& Hyermann, B. W.: ‘The Fiskes of Noith and Middle America,’ 1898, 


546 MR. H. M. KYLE ON NASAL 


The other structures in connection with the mouth vary little 
in the different specimens. In all, the jaws of the upper (eyed) 
side are quite bare, whilst those of the lower (blind) side display 
strong series of chisel-shaped teeth (fig. 3, t.m.). The mandibular 
and maxillary ‘“ breathing-valves”* are strongly developed 
(fig. 4, 6.v.), and the only appreciable difference is seen in the 
length of the gill-filaments. These are very long in all specimens, 
exceeding the usual length found in the Heterosomata, but are 
slightly longer in the divergent specimen than in the others. 
Although dissections of four of the specimens were not made, it 
is most probable that in other characters of the head and body 
all five are alike, since by analogy I find in the Plaice and 
Flounder, in which the internal structure differs little, that what 
important differences there are may be seen externally. 

The question then presents itself whether the single specimen 
of Cynoglossus is entitled to be taken as the type of a new 
species. It is possible, indeed, that such a species really exists, 
because if this peculiar combination of characters occurs in one 
individual, there is no reason why it should not occur in many 
others. On the other hand, it is possible that this individual 
may be quite unique, and that the accessory portion of the 
lateral line and the increased length of the gill-filaments may 
have arisen after the perforation of the roof of the mouth in the 
life-history of it alone; but this does not seem very probable. 
The lateral line in other forms is developed at a very early stage, 
and if an accessory branch is present, we should imagine that it 
would arise about the same time as the main portions. Hence 
the perforation of the roof of the mouth must have appeared in 
the earlier stages of life ; and, if so, is possibly inherited—that is 
to say, a distinet species may exist whose characters are fairly 
constant and reappear in the offspring. It is possible again that 
we have here only the beginning or the foreshadowing of a new 
species. If we reason from the principle of Natural Selection, 
we may conclude that the possession of a naso-pharyngeal 
communication would result in great advantage to a fish living 
in the sand. Or we may say, with Dohrn?, that the change in 


* Owen, R.: ‘Anatomy of Vertebrates,’ vol. i. p. 413.—Dahleren, W.: “The 
Maxillary and Mandibular Breathing-valves of Teleostean Fishes,” Zool. Bull. 
Boston, 1898. Smitt, F. A.: ‘Scandinavian Fishes,’ p. 263. 

t+ Dohrn, A.: ‘Das Princip des Functionswechsels.’ Leipzig, 1874. 


SECRETORY SACS IN TELEOSTEI. alir/ 


function, from secretory to water-retaining, being accompanied 
by changes in the tissues, may have led to a great change in 
structure, and that Natural Selection has been thereby confronted 
by variations which it will convert into specific differences. Or, 
discarding Natural Selection and questions of ‘‘advantage ” 
and ‘‘survival of the fittest,” it seems simpler and more natural 
to believe that the remaining longer than usual with the head 
buried im the sand, z.e.a slight change in habit*, may have 
brought about the perforation of the roof of the mouth, in 
which case the presence of the naso-pharyngeal communication, 
accompanied by the change of habit and habitat, would have a 
“ discriminative ” as distinguished from a “ selective’ value. 

Whichever way the matter be argued, we see how this im- 
portant ‘ modification ” gives us the possibility of a new species, 
although it is better perhaps to wait for additional evidence 
before recording it as such in the classification of the Hetero- 
somata. 

The discovery of a naso-pharyngeal communication in only one 
specimen so far does not, however, lessen the interest attaching 
to its presence. Such facts are rare in the class Pisces, so that, 
when they do occur, their general importance is so great as to 
render the question of their specific value a somewhat secondary 
matter. Theimportance and interest do not lie in the uniqueness 
of this single individual, but in the occurrence in the Teleostei of 
on organ hitherto unknown in them and considered as almost 
peculiar to the air-breathing Vertebrates. 

We may now turn our attention to more general consider- 
ations, and in the first place to those of function. The first 
stage, or most elementary condition, where “reservoirs” with 
water-retaining functicn are present among Fishes, is found in 
Labrus, Scorpena, Gastrosteus, and Anarrhichas, but it is absent 
in the Gadide so far as examined, as also in the Herring. These 
sacs are also absent from the Sail-fluke, a highly specialized 
Heterosomid, which lacks the “‘recessus orbitalis’’> and has 
departed from the sand-loving habits of the other flat-fish. These 
facts lead to the conclusion that the presence of nasal “sacs” is 


* In another species of the Sole-group, it was found that certain parasites 
(Lernea?) had made their way into the nasal sacs and had caused a perforation 
of the roof of the mouth ! 

t Cf. Holt, EH. W. L.: “Studies in Teleostean Morphology.” Proc. Zool. 
Soc. 1894, p. 422. 


548 MR. H. M. KYLE ON NASAL 


an adaptation to semi-sedentary, as opposed to migratory, habits 
of life. 

The function of these sacs in the first stage is moreover aptly 
described, as has been said, by the word “reservoir.” In the 
tolerably quiet life these animals lead, the water containing 
odoriferous particles will not pass so freely over the olfactory 
epithelium as in the case of the free-swimming migratory forms. 
These sacs are, however, distended and constricted by the 
movements of the premaxille and maxille, and are thus able 
to draw in water, the odoriferous particles in this way coming 
into contact with the sensory epithelium without necessitating 
any movement on the part of the animal as a whole. 

The second stage, where definite secretory sacs are present, 
has been found so far only in the family of the Heterosomata 
comprising the Halibut, Plaice, and Turbot groups. The secre- 
tion is forced from the sacs into the nasal cavity and over the 
sensory epithelium by the movements of the premaxille and 
maxille, in a fashion similar to the water in the previously 
described species. 

The function of these secretory sacs is not very evident. They 
are absent in other Teleostei so far as is known, and this might 
mean that the olfactory organ is of little importance or that 
the epithelium is maintained sufficiently sensitive by water 
alone. In tke air-breathing Vertebrates the nasal secretion is of 
importance in cleansing the organ, in keeping the sensory 
epithelium in a healthy sensitive condition, and in aiding towards 
bringing odoriferous materials into a state of solution favourable 
to their full appreciation. When the secretion 1s present in 
Fishes, therefore, it may signify that the olfactory organ is much 
used and of great importance. Such, indeed, would seem to 
be the case with the flat-fishes mentioned. In the life these 
animals lead as ground-feeders, searching for their food almost 
entirely by sense of smell, this secretion may be of as much 
service in cleansing the olfactory epithelium and maintaining it 
sensitive as the nasal secretion of the air-breathing Vertebrates. 

In the Sole group the secretory has given place for the most 
part to a water-retaining function. This change might at first 
sight appear strange, as a return to a previous condition, although 
the animals are more sand-loving in their habits than even the 
Plaice, Halibut, and Turbot, and in most cases have their 
olfactory organs as largely developed as these forms. But the 


SECRETORY SACS IN TELEOSTEI. 549 


tactile sense, as shown in the development of papille and 
filamentous outgrowths * of the integument, here aids and even 
replaces the olfactory organ to a great extent. There is probably 
not the same necessity, therefore, for the epithelium to be kept 
in a high degree of sensitiveness as is the case with the Halibut, 
Plaice, and Turbot. Further, it is well known that a large 
quantity of mucus is secreted from the external surface in the 
Soles ; and this, entering the nasal organ with the water-currents, 
may replace definite secretory sacs and effect the cleansing and 
preservation of the epithelium as efficiently as the needs and 
mode of life of the animals require. 

When we come to Cynoglossus, we find the last stage in the 
structural specialization, apparently the complete return of 
the nasal “sacs” to the earliest function of water-retaining. 
The sacs are not secretory, and, further, there are no tactile 
filaments round the head; and it may be that the mucus from the 
skin enters the nasal cavity and acts as a nasal secretion, just as 
has been suggested for Solea. 

In the divergent specimen of Cynoglossus, the perforation 
of the roof of the mouth brings into consideration a totally new 
function; but by the change that bas occurred, the raison d’étre 
of the former function of water-retaining is still as efficiently 
fulfilled. The odoriferous particles which the closed sacs induce 
to pass over the olfactory epithelium will be drawn through the 
nasal cavity during the process of respiration, by the movements 
of the mouth and gill-covers. The respiratory function, however, 
although it bas arisen secondarily, probably becomes the more 
important. 

The manner by which the Teleostean fishes respire has recently 
been carefully described by Dahlgren (/. ¢.), who shows what an 
important role is played by the maxillary and mandibular 
“ breathing-valves ” (fig. 4, 6.v.). These are well-developed in 
the divergent specimen of Cynoglossus; but it is probable that in 
the mode of life which these animals lead, the circular fold 
beneath the central sac which receives the internal nares has 
taken the place of, or at least may act in the same manner as, the 
breathing-valves. Both would function when the animal’s head 
was free in the water, but when the jaws were buried in the 
sand, the nasal respiratory canals and this “ respiratory-fold” 

* Raffaele, F.: ‘“Papille e organi di senso cutaneo nei Pleuronettidi del 
genere Solea: nota preliminare.” Naples, 1886. 

LINN. JOURN.—ZOOLOGY, VOL. XXVILI. 41 


550 MR. H. M. KYLE ON NASAL 


would be most in action, opening and shutting synchronously 
with the gill-covers. When the gill-covers rise, and so increase 
the cavity of the mouth, whilst the posterior edges of the 
branchiostegal membranes close the gill-openings, this fold 
will also rise, and water will enter the mouth from the nasal 
passages. Conversely, when the gill-covers fall, the fold will 
press on the internal nares and close them; whilst the water 
from the mouth, passing between the gill-arches, bathing the 
gills, will escape by the gill-openings *. 

The foregoing discussion of the functions of the nasal sacs and 
their specialization, leads on to a consideration of Huxley’s 
conclusions with respect to the use of the communication between 
the nose and mouth in the fishes with which he dealt. In his 
paper on Ceratodus (1. c.), after comparing the Dipnoi and Selachi 
with regard to the nasal organ, he raised the question—of what 
use are such nasal passages and internal nares to purely bran- 
chiate animals? In answering this, he considered that in all 
probability they are primarily connected with respiration when 
the mouth is closed; and, secondarily, that by their means a 
constant stream of water containing odoriferous particles would 
be brought into contact with the. Secncn epithelium of the 
olfactory organs. 

What has been advanced in the fenemetne pages is so far in 
complete accord with both of these conclusions, but Huxley went 
beyond this and, reasoning from the second, concluded that the 
posterior nostrils of the Teleostei, where they occur, have most 
probably a function similar to the internal nares, viz., to aid in 
ensuring the adequate passage of odoriferous particles over the 
sensory epithelium. 

This conclusion is, however, open to doubt. When internal 
nares are present, the pumping action which draws the water 
through the nasal passages is carried on by means of the gill- 
covers and floor of the mouth in common with that concerned in 
respiration. But in those Teleostei where internal nares and 
nasal sacs are absent and posterior nostrils are present, this 
pumping action during respiration cannot involve the olfactory 
organ. Hence the essential conditions which would render the 


* Howes has proposed to distinguish this buccal mechanism of respiration 
characteristic of all the Ichthyopsida as stomatophysous, and that characteristic 
of the Amniota, and taking place only in the presence of a costal sternum, as 
somatophysous. Of. Jour. Anat. & Phys. vol. xxiii. p. 272. 


SECRETORY SACS IN TELEOSTET. 551 


physiological significance of the posterior nostrils similar to that 
of the internal nares are absent. 

The walls of the posterior nostrils, again, act as valves whose 
function is to let water pass from within outwards. In the free- 
swimming forms, therefore, when both nostrils are present, it is 
probable that the movements of the fish through the water 
suffice to induce the passage of water through the nasal cavity ; 
and this flow is controlled by the anterior nostril, whose 
walls may be prolonged into a contractile tube or flap-like 
covering. Where only one external nostril is present, it is the 
anterior which must carry on the functions of the two of the 
other forms. And hence, if analogies are to be drawn, the 
anterior nostrils might be likened physiologically to the internal 
nares, and the posterior nostrils to the gill-openings. Similarly, 
when nasal sacs are present the displacement of the sacs ensures 
the circulation of water through the nasal cavity, just as the 
movements of the mouth and gill-covers cause the passage of 
water through the mouth, and the posterior nostrils are again 
only comparable to the gill-openings. 

We may proceed now to a consideration of the other water- 
breathing Vertebrates in which the function of respiration is in 
part carried on by a naso-pharyngeal communication. In the 
Cyclostomes we find an analogy to what is seen in Cynoglossus. 
In the Petromyzontide there is a prolongation from the nasal 
cavity backward to the roof of the mouth; in the Myxinide the 
latter is pierced, and the communication thus opened is supposed 
to fulfil a respiratory function. The origin and development of 
this so-called prolongation of the nasal organ have been described 
by Dohrn* for Petromyzon, and recently somewhat briefly by 
Dean + for Bdellostoma. 

Leaving aside the apparent remarkable differences alleged by 
Dean in the development of these two forms, it is clear that the 
internal nares of the Myxinide are formed by tne communication 
of the hypophysis with the gut. According to Dean the hypo- 
physis arises in Bdellostoma before the externa) openings of the 


* Dohrn, A.: “Studien zur Urgeschichte des Wirbelthierkorpers. III. Die 
Enstehung u. Bedeutung der Hypophysis bei Petromyzon Planert.” Mitt. Zool. 
Stat. zu Neapel, 1883. Vide Howes, G. B.: “ On the Affinities, Inter-relation- 
ships, and Systematic Position of the Marsipobranchii.” Tr. Biol. Soe. 
Liverpool, vol. vi., 1891, p. 122. 

+ Dean (Bashford): ‘On the Embryology of Petromyzon Stouti.’ Kupffer’s 
Festschrift, Jena, 1899, p. 269. 


552 MR. H. M. KYLE ON NASAL 


mouth and nose appear; and if this allegation should hold good 
also for Myxine, a special importance must be ascribed to such an 
early communication of the hypophysis with the gut, and it would 
furnish an argument in favour of Dean’s view that the Myxinide 
and the Petromyzontide stand to each other in somewhat the 
same relation as the recent Selachians to the recent Ganoids. 
But, however widely they may be separated, the hypophysis would 
appear to present a condition analogous in the one group to 
that of the closed nasal-sac state aforementioned, and in the 
other to that of the open nasal canals in Cynoglossus. 

The assertion of Dean that the hypophysis in Bdellostoma 
opens into the gut before the external opening of the nasal 
organ is formed, if correct, further leads to the conclusion that 
the so-called “internal nares” of the Myxinide, although a 
primary formation, is only secondarily connected with respiration. 
The reverse is the case in Cynoglossus, and thus there arises 
a good example of convergency in evolution, since a similar 
structure possessing a similar function, but having a totally 
different origin, would appear to have arisen in two separate 
groups of the animal kingdom. 

On the other hand, there is so much diversity of opinion with 
regard to the development of the internal nares in the higher 
Vertebrates, that reference to them is somewhat difficult. 
According to Balfour’s theory *, the nares arise from a single 
depression lying anterior to, and one on each side of, the mouth. 
This depression, as the embryo develops, takes the form of a 
longitudinal slit, and a little later passes through a stage similar 
to that of the adult Selachian. By the fusion of the adjacent 
tissues over the depression, this slit becomes tranformed into a 
canal with an opening at either end—these openings represent 
the external and internal nares. The later development differs in 
the different groups. In the Amniota, Balfour believed that the 
maxillary region of the face so develops in relation to the canal 
that the two openings become widely separated, the outer passing 
upwards and forming the external nares, the inner passing 
inwards and forming the internal nares. In the Amphibia it was 
believed that the origin of the internal nares is distinct from that 
above described, being “secondary,” through perforation of the 
roof of the mouth after the latter is developed ; and the develop- 
ment of the nares in the Dipnoi was considered to be similar to 


* Balfour, KF. M.: ‘Comparative Embryology,’ vol. ii. pp. 533-538. 


SECRETORY SACS IN TELEOSTEI. 558 


that in the Amniota, except that the upward rotation of the 
external nostril does not take place. In the Teleostei, lastly, it 
was believed that the homology still holds good, and that both 
nares are rotated outward and upward. Hence the posterior 
nostrils of the Teleostei would be homologous to the external 
nostrils of the higher Vertebrates, and the anterior to their 
internal nares. 

These generalizations have been in part confirmed and in part 
refuted by more recent workers. For the Teleostei, Sagemehl *, 
though believing that he was refuting Balfour’s view, in reality 
corroborated it. For the Dipnoi, Semon tT has shown that the 
nares develop as Balfour suggested; and he thus confirms 
Huxley’s conclusions (J. ¢.), drawn from a comparison of the 
adult conditions, that, as concerning their nostrils, the Dipnoi 
and cartilaginous fishes are closely related. An important. 
modification of Balfour’s view has, however, to be noted. In the 
Dipnoi the communication between the nose and mouth is naso- 
labial, not naso-pharyngeal, the posterior aperture being morpho- 
logically disposed external to the teeth which arise on the vomer 
and palatine bones. Further, there is no true “palate”; and 
the question whether the maxille of fishes are truly homologous 
with the maxille of the higher Vertebrates is left open (Semon, 
l.c., p. 45). As regards the internal nares, it is thus evident that 
no true homologue to that of Cynoglossus exists in the class 
Pisces. 

In the higher Vertebrates, however, if the internal nares arise 
secondarily in the Amphibia, as suggested by Balfour, there is 
the possibility of such an homology. Hochstetter {, however, has 
thrown doubt upon the whole of Balfour’s conclusions with 
regard to tbe origin of the internal nares in the Amniota. He 
shows that the ‘“ palate” is not formed by the maxillary bones, 
but by the fusion of the external and median primitive nasal 
processes, and that the internal nares then arise secondarily by 
the perforation of the palate. These observations were made 
upon certain mammalian forms ; and in the same forms as well as 


* Sagemehl, M.: “Das Cranium von Amia calva, L.” Morph. Jahrbuch, 
Bd. ix. p. 221. 


+ Semon, R.: “Die aussere Entwickelung das Ceratodus Forstert.” Denksch. 
der Med.-Naturwiss. Gesellsch., Jena, Bd. iv. 1893, pp. 44-45. 
+ Hochstetter, F.: ‘‘ Ueber die Bildung der inneren Nasengange oder 


primitiven Choanen.” Verh. d. Anat. Gesellsch. 1891 (Anat. Anz. Bd. vi 
Suppl.) p. 145, 


504 MR. H. M. KYLE ON NASAL 


in others Keibel * has more recently, in opposition to His tT, con- 
firmed those which concern the formation of the palate, though 
he is doubtful about the secondary origin of the internal nares. 

If the conclusions of Hochstetter had been the last word on 
the matter, it would have been possible to frame an homology 
between the internal nares of Cynoglossus and those of the higher 
Vertebrates. If the latter arise secondarily in ontogeny, then it 
is probable that at their first beginning they were also of 
secondary origin, and there‘ore distinct from the primitive 
internal nares of the Dipnoi. Hence we might have in Oyno- 
glossus a glimpse of what may have occurred at a remote period 
of time, when the air-breathing Vertebrates were but in process 
of evolution. 

But if Keibel’s suggestions (see footnote {) are well-founded, 
then we must accept, in a modified form, Balfour’s view con- 
cerning the origin of the internal nares as the true one, and upon 
this no comparison could be made between Cyxoglossus and the 
higher Vertebrates. Morphologically, Cynoglossus would then 
be perfectly unique, possessing not only the homologues of the 
internal nares in the morphological sense, but additional organs 
also which represent the internal nares in the physiological 
sense. The internal nares of Cynoglossus should then be called 
‘‘ pseudo-nares ” or ‘“ pseudo-choani,” which would emphasize 
their morphological distinction from the “ choani” of the higher 
Vertebrates, whilst implying their physiological similarity. 

It is of interest to note, in conclusion, that this discovery in 
the Teleostei of a distinctive peculiarity of the bigher Vertebrates, 
is not without a parallel. Warm blood has been found in the 


* Keibel, F.: ‘Zur Entwickelungsgeschichte und vergleichenden Anatomie 
der Nase u. des oberen Mundrandes (Oberlippe) bei Vertebraten.” Anat. Anz., 
Ed. viii. 1893, p. 473. 

t His, W.: “ Die Entwickelung der menschlichen und tierischen Physio- 
gnomieen.” Archiv f. Anat, u. Phys. (Anat. Abth.), 1892, p. 399. 

t In his short paper Keibel concerns himself mostly with the palate and 
upper lip of the Vertebrates, and it is only secondarily that he throws out 
suggestions as to the furmation of the internal nares. Hence his meaning is 
not very easily determinable, but it seems to be as follows:—The internal 
nares are homologous and primary structures wherever they appear in the 
Vertebrate kingdom. ‘They arise from the inner portion of the primitive nasal 
groove—as described by Balfour—but instead of the maxille growing in 
between the primitive nostril and nares, it is the ‘“ palate”—formed from the 
median and external primary nasal processes. The Selachii represent, there- 
fore, the most primitive condition, where the primary nasal groove persists. 


SECRETORY SACS IN TELEOSTEIL. 505 


Tunny, viviparity occurs in Zoarces and in other Teleosts; and 
in Anableps it is said that ‘the vascular yolk-sac is provided 
with villi which absorb nutriment”’ from the fluid secreted by 
the walls of the dilated ovary within which the embryo develops *. 
The Elasmobranchs offer numerous instances of this kind; and 
Professor Howes (who has aided me liberally with the literature 
throughout my work) has at the last moment drawn my attention 
to another case which compares in its ‘‘ uniqueness” with the 
divergent specimen of Cynoglossus. In the Anurous Ampbibia 
be has shown that the epiglottis—an accessory voice-organ 
peculiar to Mammals—is of frequent occurrence in an elementary 
form, liable to great individual variation. In Chiroleptes australis 
the epiglottis was developed in one specimen, a male; in another, 
also a male, it was insignificant ; whilst in a third, a female, the 
epiglottis was small and the accompanying “ epilaryngeal folds ” 
absent. He informs me that three more adults, which he has 
examined since his paper was written, were wholly destitute of 
the organs in question. So far, therefore, only one specimen of 
this species has been found with these organs well-developed. 
With the exceptioa perhaps of this last case, the foregoing are all 
examples of separate specializations in the respective modes of 
life of the animals, and show once more how plastic is the 
organism in the grasp of its environment. 


EXPLANATION OF PLATE 38. 


Fig. 1. Semidiagrammatic transverse section across the nasal region of a 


Pleuronectid (Plaice or Halibut), to show the nasal secretory sacs, 
2.8.4 —NeSo40 


The Dipnoi ocsupy the second stage, where a rudimentary true palate is repre- 
sented by the hinder portion of the upper lip anterior to the teeth, but where 
no true maxille are developed. The palate (?) is here formed by the nasal 
processes (¢f. Semon, J. ¢.), and the naso-pharyngeal communications arise from 
the primary nasal grooves. The Amniota show the third and last stage. The 
palate and the internal nares are formed as above, and the maxillz grow round 
anteriorly to complete the external boundaries of the mouth. 

* Haddon, A. C.: ‘The Study of Embryology,’ p. 98. (Cf. Wyman— 
Boston Journ. Nat. Hist. vol. vi. p. 432.) My best thanks are due to Mr. A. 
W. Kappel, the resourceful Librarian of the Linnean Society, for the pains 
with which he has determined this reference. 

t Howes, G. B.: “On an unrecognized feature in the Larynx of the 
Anurous Amphibia.” Proc. Zool. Soc. 1887, p. 497. 


NASAL SECRETORY SACS IN TELEOSTEI 


Fig. 2. Semidiagrammatic longitudinal section through the skull of Solea 
lascaris, to’ show the nasal sacs, 7.2.8. and /.7.s. 


a dissection from the left side, showing the left 


ventral view of the roof of the mouth, showing the 


3. Cynoglossus semilevis: 
olfactory organ (0./.), the mandibular tooth-mass of the right side 
(¢.m.), and indicating, diagrammatically in dotted lines, the course of 
the left naso-pharyngeal canal (z.p.c.), and the position of the 
aperture of the naso-pharyngeal sac (¢). 

4. The same: 
aperture of the naso-pharyngeal sac (c), and, diagrammatically in 
dotted lines, the course of the naso-pharyngeal canals (z.p.c.). The 
arrows point towards the internal openings. 

5. The same: 


a dissection to the level of the naso-pharyngeal sac, the 


left wall of which (¢,f.) has been turned outwards to show the course 
of the left naso-pharyngeal canal (7.p.c.), and the slit-like expansion 
of the naso-pharyngeal canal of the right side (m.c.) as it opens into 


the sac (¢). 


Reference Letters. 


4., basioccipital. 
b.v., breathing-valves. 
c., sac in roof of mouth into which 
the naso-pharyngeal canals 


open. 
c.f., cut edge of fold round sac (¢). 
e., ethmoid. 


e.l., left eye (displaced). 
¢.0., opening of naso-pharyngeal 
canal into nasal cavity. 
Ff, fold round naso-pharyngeal 
sac (¢). 
jr., frontal. 
f.t., fatty tissue. 
h., hyomandibular. 
i.m., premaxilla. 
Ln.s., left nasal sac. 
Lim., left premaxilla. 
J.m., left maxilla. 
m., maxilla. 
M., mouth. 


md., left mandible. 
mt., metapterygoid. 
n., nasal bone. 
n.c., nasal cavity. 
1.8.,—N.S.,, nasal sacs. 
.p.c.. naso-pharyngeal canal. 
o.t., left olfactory organ. 
op., operculum. 
p. palatine. 
p.o., preoperculum. 
pr, prootic. 
p.s., parasphenoid. 
gu., quadrate. 
r.i.m., right premaxilla. 
rw., vight maxilla. 
r0.c., opening of right naso-pharyn- 
geal canal into sac (c). 
s.0., supraoccipital. 
t.m., tooth-mass of right mandible. 
w., turbinal. 
v., vomer. 


linn SOC Momma Zoor Von XeQullaen OVS, 


H.M.K del. Nantern Bres.imp. 
J, Green lth. 


INAS AL, SiNGS WN VEIL OS Wan S| 


rant 
i 
id 


INDEX. 


[Synonyms and native names are printed in ¢adics. 


A star is added to names 


which appear to be here used for the first time.] 


Abramis, 389, 390. 
urmianus * , Giinther, 389, 391. 
Abretia affinis, Gray, 155. 
Acalephe, stages of, 301. 
Acanthias, 48, 49, 55. 
vulgaris, 47. 
Acanthochiton, 180. 
Acanthonotosoma, Boeck, 75. 
cristatum, Owen, 75, 117. 
Acanthonotus cristatus, Owen, 75. 
Acanthopleura spinigera, Low, 180. 
Acanthostepheia, Boeck, 74. 
Malmereni, Goés, 74, 117. 
Acanthyllis caudacuta, 250. 
collaris, 249. 
Acar acerea*, Melv. § Stand., 186; 
mentioned, 206. 
divaricata, Sow., 186. 
domingensis, Lam., 186. 
sculptilis, Reeve, 186. 
Acari of Lake Urmi, by A. D. Michael, 
407. 
Accipitres, wings of, 247. 
Aceros, Boeck, 73. 
phyllonyx, VM. Sars, 73, 117. 
Acridiidee of Lake Urmi, 417. 
Acridiodea, 417. 
Acronycta centralis, Staud., 411. 
Actzon solidulus, Lénn., 155. 
Actzonide, 155. 
Actinia, 303, 304. 
equina, mentioned, 275, 297, 298, 
303, 304. 
Actiniaria, 296. 
Actinozoa, 434, 485, 486. 
Adonis vernalis, 26. 
Adopea lineola, Ochsenh., 411. 
Adula lanigera, Dér., 184. 
/Kgina, Kroyer, 80. 
spinosissima, Stimps., 80, 117. 


/Epyornis, 251. 

AXquipecten Malvine, Duhois de 
Montpéreux, 444; mentioned, 431, 
452. 

Agaronia nebulosa, Lam., 157. 

Agelenidz from Corral, 16, 20. 

Aglossa pinguinalis, Linn., 414. 

Agrotis Christophi, Stawd., 411. 

ypsilon, Mozt., 411. 

Aiptasia annulata, mentioned, 3053. 

Air-bladder and its Connection with 
the Auditory Organ in Notopterus 
borneensis, by Prof. T. W. Bridge, 
503-540. 

Alaba pyrrhacme, Melv. § Stand., 170, 
ftnote 171. 

Alburnus, 354. 

Filippii, Kessler, 390. 

punctulatus, Kessler, 389, 390. 
Alcedinide, 234. 

Alcyonaria, filaments of, 296. 

Alectryon, mentioned, 160. 

fretorum*, Melv. g Standen, 

159. 

suturalis, Zam., 159. 

Alectryonia Virleti, Desh., 440; men- 
tioned, 431, 450. 

Alestes dentex, Miiller § Trosch., 

522. 

Hasselquistii, Cuv. § Val., 522. 
Algx, 472; swarmspores of, 478. 
Alibrotus, Milne-Edw., ftnote 70. 
Alicula cylindrica, Helbling, 155. 
Allium sibiricum ved senescens, 42. 
Alpheus polaris, Sabine, 63. 

Altai Mountains, Zoology and Botany 
of the, by H. J. Elwes, 23-46. 

Alteutha purpurocincta, Norm., 112. 

Alveopora, 131, 132, 136, 148, 144, 
145, 488. 


LINN. JOURN.—ZOOLOGY, VOL. XXVII. 42 


558 


Alytes, 456; hyobranchial skeleton of 
larva, 456. 
obstetricans, mentioned, 459. 
Amathilla, Spence Bate, 79. 
homari, Mabr., 79, 117. 
pinguis, Kroyer, 79, 117. 
Amaurobius Platei*, Pickard-Cam- 
bridge, 15, 18. 
Ameira, Boeck, 97. 
exigua, 7. Scott, 97, 118. 
longipes, Boeck, 97, 118. 
longiremis, 7. Scott, 97, 118. 
reflexa, 7. Scott, 97, 118. 
Amia, 57. 
calya, Linn., ftnote 553. 
Ammonite, Note on a Jurassic, of Lake 
Urmi, by G. CO. Crick, 418. 
Ammopleurops, mentioned, 548. 
Amphibia, Anurous, 5595. 
Amphibia and Reptilia of Lake Urmi, 
by G. A. Boulenger, 578-381. 
Amphilochus, Spence Bate, 71. 
oculatus, Hansen, 71, 117. 
Awphioxus, mentioned, 307. 
Amphipoda, 61, 69, 395. 
Amphiscopus continens, Rosen, 392. 
Amphithoé bicuspis, Kroyer, 74. 
fulvocincta, M. Sars, 77. 
leviusculus, Kroyer, 78. 
pulchella, Kroyer, 74. : 
Amphithonotus Malmgrent, Goés, 74.° 
Amphithopsis, Boeck, 78. 
glaber, Boeck, 75. 
glacialis, Hansen, 78, 117. 
Amussiopecten, 448. 
Burdigalensis, Lam., 443. 
Burdigalensis, Sacco, 443. 
Amussium pleuronectes, Linn., 183. 
Amymone, Claus, 96. 
spherica, Claus, 96, 118. 
Anableps, 555. 
Anadara antiquata, Linn., 186. 
clathrata, Reeve, 186. 
Anaitis calophylla, Hanley, 196. 
thiara, Dillwyn, 196. 
Anaitis plagiata, Linn., 413. 
usgentaria, Staud., 413. 
Anarrhichas, mentioned, 547. 
lupus, Linn., 542. 
Anas boscas, wing of, 243. 
, var. domestica, 244, 256. 
Anaspides, characters of, 343. 
tasmanie, G. M. Thomson, 342, 
- 844. 
Anceus elongatus, Kroyer, 66. 
Anemone patens, 26. 
Pulsatilla, 26. 
Angulus philippinarum, Hanley, 201. 
procrita*, Melv. & Stand., 201, 
206. 
vernalis, Hanley, 201. 


INDEX. 


Annelida of Lake Urmi, 439. 

Anomalurus, The Position of, as’ in- 
dicated by its Myology, by F. G. 
Parsons, 317-334. 

Anomiidz of Torres Straits, 181. 

Anonyx, Kroyer, 69. 

Kdwardsti, Kroyer, 71. 

littoralis, Kroyer, 79. 

minutus, Kroyer, 69. 

nugax, Phipps, 69, 70. 

Anoura Dendyi * , Lubbock, 836, 337. 
spinosa * , Lubbock, 338, fig. 337. 
Tasmaniz * , Lubbock, 335. 

Anseres, wings of, 247. 

Antaria mediterranea, Claus, 115. 

‘Anthea cereus, mentioned, 278. 

Antilope sp., 377. 

Antilopidze, Pliocene, of Maragha, 377. 

Anura, 454, 

Aglossa, 454-459. 

Phaneroglossa, 454-459. 
Apherusa Jurinei, mentioned, 79. 
Aplustridee of Torres Straits, 155. 
Aplustrum amplustre, Linn., 155. 
Apollon pusillum, Brod., 164. 
Apterona erenulella, Bruand, 413; 

mentioned, 367, 416. 
Apteryx, 251. 
Aquila chirysaétus, Zinn., mentioned, 
361. oo 
Aquilegia glandulosa, 42. 
eae (Simpulum) gemmatus, Reeve, 
63. 


(——) pileatus, Linn., 163. 

“ Aquintocubitalism,” Some Facts con- 
cerning the so-called, in the Bird’s 
Wing, by W. P. Pycraft, 236-256. 

Arachnactis, mentioned, 297, 298, 3U0, 
307 ; larva of, 310. 

Arachnida and Chilopoda of Lake 
Urmi, by R. I. Pocock, 399-406. 

Aranea Bruennichi, Scop., 400. 

eaxtensa, Linn., 400. 
13-guttata, Rosst, 400. 

Araneee of Lake Urmi, 400. 

Arapaima, 524. 

Arca navicularis, Brug., 185. 

zebra, Swains., 185. 
zebuensis, Reeve, 185. 

Arcidee of Torres Straits, 185. 

Arcinella spinosa, Brod., 193. 

Arcopagia pinguis, Hanley, 201. 

Savignyi, 4. Ad., 201. 
tessellata, Desh., 201. 
Arctiade of Lake Urmi, 411. 
Argas persicus, Fischer, 407; mentioned, 
366. 
reflexus, Fabr., 407. 
Argiope, Sav., 400. 
Bruennichi, Scop., 409. 
Argiopide of Juan Fernandez, 16, 18. 


INDEX. 


Argynnis, 39. 
freya, 43. 
frigga, 43. : 
Kefersteini, 43. 
maia, Cramer, 408. 
Ariadna maxima (Nic.), 15, 17. 
Aricia annulus, Zinn., 164. 
arabica, Linn., 164. 
Arpacticus nobilis, Baird, 111. 
Artemia, 398; mentioned, 357, 359. 
fertilis, mentioned, 398. 
gracilis, mentioned, 398. 
monica, mentioned, 398. 
Milhausenii,  Schmankewitsch, 
mentioned, 359, 396, 398. 
salina, Leach, mentioned, 356, 359, 
395, 397, 398. 
urmiana*, R. ZT. Giinther, 395; 
mentioned, 356, 359, 396, 398. 
utahensis, mentioned, 398. 
Artiodactyla, Pliocene, of Maragha, 377. 
Asaphis, mentioned, 198. 
coccinea, Mart., mentioned, 198. 
deflorata, Zinn., 198. 
dichotoma, Anton, mentioned, 198. 
Ascomyzontide, 113. 
Asellus, 366, 395. 
Asio accipitrinus, 256. 
Astacus homari, Fabr., 79. 
Astoma, 407. 
eryllaria, Le Baron, 407, 417. 
Astrea, 128. 
Defrancei, Milne-Hdw., 434, 435. 
Hllisiana, Defr., 434, 435. 
erandistella, Abich, 435. 
Guettardi, Defr., 434, 435. 
Astropora, ftnote 150. 
Astragalus, 42. 
Astralium petrosum, Mart., 175. 
Astrangia princeps, Hewss, 436. 
Astrea argus, Lam., 425. 
Defrancei, Milne-Edw., mentioned, 
425, 426. 
grandistella, Abich, mentioned, 429. 
Guettardi, Defr., mentioned, 425. 
irregularis, Defr., ftnote 428. 
turonensis, Michelin, ftnote 427. 
Atilia galaxias, Reeve, 161. 
Atrichia, 248. 
Atrina nigra, Chemn., 184. 
Atylus, Leach, 78. 
carinatus, Fabr., 78, 117. 
Atys debilis, Pease, 155. 
(Alicula) cylindrica, Helbling, 155. 
Auditory Organ in Notopterus borne- 
ensis (Prof. T. W. Bridge), 503- 
540. 
Augiades sylvanus?, Hsper, 411. 
Aulacomya hirsuitus, Lam., 184. 
Aulactinia, mentioned, 298, 308, 309; 
larvee, 276. 


559 


Aulactinia stelloides, MeMurrich, men- 
tioned, 269, 272. 

Aulica Ruckeri, Crosse, 157. 

rutila, Brod., 157. 
Sophiz, Gray, 157. 

Aurelia, mentioned, 304. 

Australasian Collembola, On some, by 
the Right Hon. Sir John Lubbock, 
334-838. 

Aves, Eutaxy and Diastataxy in Aves 
generally, 283. 

Avicula ala-corvi, Ohemn., 183. - 

aquatilis. Reeve, 183. 
crocea, Chemn., 183. 
malleoides, Reeve, 183. 

Aviculidz, 183. 

Azar coarctatus, Gmel., 198. 


Balanide, 116. 
Balanoglossus, ftnote 257. 
Balanophylha, mentioned, 
308. 
regia, mentioned, 310, 
Balanus, Lister, 116. 
erenatus, Bruguiére, 116, 118, 
poreatus, Da Costa, 116, 118. 
Barbatia decussata, Sow., 185. 
fusca, Brug., 185. 
imbricata, Brug., 185, 
lima, Reeve, 185. 
tenella, Reeve, 185. 
trapezina, Lam., 185. 
volucris, Reeve, 185. 
(Acar) aerexa*, Melv. & Stand., 
186, 206. 
(——) divaricata, Sow.. 186. 
(——) domingensis, Lam., 186. 
(——) sculptilis, Reeve, 186. 
(Venusta) lactea, Linn., 185. 
Barbus. 336. 
caucasicus, Kessler, 385; men- 
tioned, 364, 386. 
ciscaucasicus, 386. 
Goktschaicus, mentioned, 286. 
miliaris, de Filippi, 386. 
Bathymedon, G. O. Sars, 73. 
obtusifrons, Hansen, 73, 117. 
Bathynella, On the Characters of the 
Crustacean Genus, by W. T. Cal- 
man, 308-344, 
natans, Vejdousky, 338, 344. 
Bathyoscopus poceilus 7ead Bythoscopus 
pecilus, Herr.-Schaff:, 367, 416. 
Batrachia, 380. 
Bdellostoma, 551, 552. 
Beaver, gastric gland in, 1. 
Belemnites acuarius, 438. 
calloviensis, Oppel, 438. 
persicus, Weithofer, 438, 
sp., 437, 458. 


304, 


49% 


560 


Bernard, Henry M., On the Structure 
of Porites, with Preliminary Notes 
on the Soft Parts, 487-502. 

, Recent Poritids, and the Posi- 
tion of the Family in the Madre- 
porarian System, 127-149. 

Bird’s Wing, Aquintocubitalism in the 
(W. P. Pycraft), 236-256. 

, * Quintocubitalism ” in the 
(P. Chalmers Mitchell), 210-286. 

Bittium, mentioned, 168. 

torresiense*, Melv. § Stand., 
168, 206. 

xanthum, Watson, mentioned, 
168. 

Bivonia Quoyi, H. g A. Ad., 170. 

Blenniidx, 542. 

Bolivaria brachyptera, Padl., 417. 

Bombinator, 46; bronchial cartilage 
of, 458, 461. 

Botany and Zoology of the Altai 
Mountains, by H. J. Elwes, 23-46. 
Boulenger, G. A., Reptilia and Am- 

phibia of Lake Urmi, 378-381. 

Box, 528. 

salpa, Cuv. § Val., 525, 528. 
vulgaris, Cuv. & Val., 528. 
Brachyura, 394. 
Bradya, Boeck, 94. 
minor, 7. g A. Scott, 94, 95, 
117. 
typica, Boeck, 94, 117. 

Branchipus, mentioned, 359. 

Bridge, Prof. T. W., The Air-bladder 
and its Connection with the Auditory 
Organ in Notopterus borneensis, 
503-540. 

Bryophila perla, Schiffermiiller, 412. 

Bryozoa, 434, 435. 

Buccinide, 159. 

Bucerotes, 248. 

Bufo viridis, Lawr., 381. 

** Bulbus ventriculi,” 10. 

Buliminus (Amphiscopus) continens, 

Rosen, 392. 
(Chondrulus) Bayerni, Parr., 392. 


( ) didymodus, Bottger, 

( ) tetrodon, Mortillet, 392. 

( ) tridens, Miiller, 392. 

(——) , var. Bayerni, Parr., 
392. 

) ——, var. diffusus, Mouwss., 

392. 

(——) ——, var. major, Kryn., 
392. 


(Zebrinus) Hohenackeri, Krynicki, 
392 


Bulla Adamsi, Menke, 15h. 
punctulata, 4. Ad., 155. 
Bullide, 155. 


INDEX. 


Bunodes gemmacea, mentioned, 276. 
verrucosa, mentioned, 276. 

Burr, Malcolm, Orthoptera of Lake 
Urmi, 416-418. 

Buthus, Leach, 404. 

afghanus, Pocock, 404, 405. 

caucasicus, Hischer, 404; men- 
tioned, 406. 

caucasicus, Nord. = parthorum, 
Pocock, 406. 

, subsp. persicus*, Pocock, 

404; mentioned, 367. 

eupeus, C. Koch, 405. 

parthorum, Pocock, 405, 406. 

persicus, Pocock, 404, 405. 

thersites, C. Koch, 405. 

vesiculatus * , Pocock, 405; men- 
tioned, 406. 

Butler, A. G., Insecta (Lepidoptera 
Rhopalocera) of Lake Urmi, 408- 
411. 

Bythocythere flexuosa, G. S. Brady, 
89. 


Bythoscopus pecilus, Herr.-Schdff., 
307, 416. 


Calanide, 90. 

Calanus, Leach, 90. 
finmarchicus, Gunner, 

tioned, 91, 117. 

hyperboreus, Kroyer, 90, 117. 
longus, Lubbock, 91. 

Calliopius, Liiljed., 78. 
leviusculus, Kroyer, 78, 117. 

Calliostoma fragum, Phi/., mentioned, 


90; men- 


speciosum, A. Ad., 177. 
(Eutrochus) septenarium *, Melv. 
&§ Stand., 176, 206. 

Calman, W. T., On the Characters of 
the Crustacean Genus Bathynella, 
338-344. 

Caleenas, 226. 

Caloptenus italicus, Linn., 407, 417; 


infested with Astoma_ gryllaria, 

417. 

Calyptrea  equestris, Linn., var.?, 
171 


tortilis, Reeve, 171. 
Canarium dentatus, Lénn., 165. 
Cancer boreas, Phipps, 64. 

gammarus galba, Montagu, 69. 

locusta, Linn., 79. 

nugax, Phipps, 69. 

oculatus, O. Fabr., 64. 
Candona, Baird, 83. 

candida, 83. 

——, var. claviformis, 83. 

Harmsworthi*, Thos. Scott, 83, 

NG 
rostrata, mentioned, 82, 83. 


INDEX. 


Cantharidus torresi, #. A. Sinith, 175. 

Cantharus mollis, Gowld, 159. 

Canthocamptus, Westw., 100 ; 

tioned, 101, 339. 

parvus, Z. & A. Scott, 100, 118. 
Stromiz, Baird, 106. 

Capercaillie, 38. 

Capoéta, mentioned, 349, 382, 385. 
Gotschaica, ftnote 385. 
gracilis, Keyserl., 364, 383, 385. 
Sevanei, De Filippi, 383, 585. 
Steindachneri, Kess/er, 384. 

Capra sibirica, Pallas, 37. 

Caprella, Lam., 81. 
dubia, Hansen, 81, 117. 
microtuberculata, G. O. Sars, 81, 

MING 

, var. spinigera, 81. 
septentrionalis, Kroyer, 81, 117. 

Caprellidea, 80. 

Capreolus pygargus, Pallas, 37. 

Caprimulgi, 248, 

Capulide, 171. 

Carangide, 522. 

Carcharias laticauda, 47. 

“ Cardiadrisen region,” 2. 

Cardiidz, 190. 

Cardita cardioides, Reeve, 188. 
marmorea, [?eeve, 188. 


men- 


mutabilis, d’Archiac &  Haime, 
mentioned, 448. 
sp., 431, 448, 451. 
Carditide, 188. 
Cardium, 446. 
arenicolum, Reeve, mentioned, 
192. 
maculusum, Wood, mentioned, 
192. 
mauritianum, Desh., mentioned, 
192. 


obovale, Sow., mentioned, 192. 

sp., 446, 451. 

tenuicostatum, Lam., 190. 

(Leevicardium) Bechei, 
Reeve, 192. 

( ) biradiatum, Brug., 192. 

( ) lyratum, Sow., 192. 

(Papyridea) papyraceum, Chemn., 
192. 


Ad. § 


(Trachycardium) dianthinum * , 
Melv. & Stand., 190, 206. 
(——) elongatum, Brug, 190. 


(——) lacunosum, Reeve, 190. 
(——) maculosum, Wood, 191. 
( ) rubicundum, Reeve, 191. 


(-—) rugosum, Lam., 191. 

(——) serricostatum*, Melu. & 
Stand., 191, 206. 

) transcendens * , 
Stand., 191, 206. 

(——) variegatum, Sow., 192. 


Melu. & 


561 


Cariama, 253, 254, 

Carnivora, 377. 

Carpus, Note on the, of the new 
Aglossal Toad, Hymenochirus Boett- 
geri, by W. G. Ridewood, 461-463. 

Caryatis, Romer, mentioned, 194, 

regularis, Smith, 194. 

Caryophyllia cyathus, mentioned, 278, 
297, 298, 317. 

Casmaria vibex, Linn., 164. 

Cassididz, 164. 

Cassis sp., 4382, 450, 452. 

subharpeformis, dd Archiac & 
Haime, mentioned, 450. 

(Casmaria) vibex, Linn., 164. 

(Semicassis) torquata,  feeve, 
164. 

Cassowary, wing of, 251. 

Castellaria modesta, Reeve, 158. 

Castor, gastric gland in, 2. 

Cataclysme bilineata, Linn., 413. 

Catocala elocata, Esper, 412. 

neonympha, Esper, 412. 

Caudal Diplospondyly of Sharks, some 
observations on the, by W. G. Ride- 
wood, 46-59. 

Cavolinia longirostris, Lesson, 154. 

Cavoliniide, 154. 

Cellastrea irregularis, de Blainv., ftnote 
428. 

Cellepora gracilis, Miinst., 434, 435. 

Centropagidee, 91. 

Centropborus, 49. 

Cephalodiscus dodecalophus, McIntosh, 
on the Discovery and Development 
of Rhabdite-“ cells” in, by F. J. Cole, 
256 -268. 

Cephalopoda, 438. 

Ceratodus, mentioned, 550. 

Forsteri, 541, ftnote 553. 

Cerebratulus, 266. 

Cerianthus, mentioned, 303, 304. 

Ceriopora anomala, Abich, mentioned, 

434, 435. 
palmata, D’Orb., mentioned, 434, 
435. 
Cerithiidee, 166. 
Cerithium asper, Zinn., mentioned, 


citrinum, Sow., 167. 

echinatum, Lam., 167. 

morus, Lam., 167. 

novee-hiberniz, A. Ad., 167. 

petrosum, Wood, 167. 

pinguis, A. Ad., mentioned, 168. 

piperitum, Sow., 167. 

salebrosum, Sow., 167. 

tuberculatum, Linn., 
167. 

variegatum, Quoy, 167. 

zebrum, Kéener, 167. 


mentioned, 


562 INDEX. 


Cerithium (Colina) -contractum, Sow., ; Chione (Omphaloclathrum) _embri- 
: thes*, Mely. g& Stand., 195, 

(—— ) teniatum, Sow., 168. 206. 

(Lampania) zonale, Brug., 168. (——) Lamarckii, Gray, 196. 

(Vertagus) aluco, Linn., 167. (——) Listeri, Gray, 196. 

( ) lineatum, Zam., 167. ( ) marica, Linn., 196. 

(——) pulchrum, A. Ad., 168. ( ) subnodulosa, Hanley, 196. 

(——) Sowerbyi, Kiener, 168. ) toreuma, Gould, 196. 

(——) vertagus, Linn., 168. Chiroleptes australis, 555. 

Cervide, 29. Chironomus oceanicus, 416. 
Cervus, 29. sp., 416. 
asiaticus, Severtzoff, 36. Chiton pictus, Reeve, 180. 
canadensis asiaticus, Lydekker, Chitonidee, 180. 
38, 30. Chlamys, 431. 
—— ——, var. Luehdorfi, Bolau, cuneatus, Reeve, 183. 
37. sp., 445, 451. 
——, var. sibirica, Severtzoff, varia, Linn., 446. 
Bc (Aiquipecten) Malvine, 431, 452. 
,var.songarica, Severtzof, | Chondrulus Bayerni, Parr., 392. 
36. didymodus, Bottger, 392. 
canadensis Luchdorfi, Lydekker, tetrodon, Mortillet, 392. 
33, 37. tridens, Mii/ler, 392. 

elaphus, 29, 31, 34. , var. Bayerni, Parr., 392. 

elaphus, var., Radde, 37. ——.,, var. diffusus, Mouss., 392. 

eustephanus, Blanford, 29, 982, , var. major, Kryn., 302. 

33, 36. Chrysame peregra, Reeve, 157. 

isubra, Noack, 37. rubritineta, Reeve, 158. 

Luehdorfi, Bolau, 37. Chrysopa perla, larva of, 414. 

maral, 29, 33. Chrysophanus thersamon, 410. 

sibirica, Severtzoff, 33. , var. omphale, Klug, 410. 

songarica, 33. thetis, Klug, 410. 

Ceryle, 248. Chrysostoma paradoxum, Born, 175. 

aleyon, 249. Chytridium, zoospores of, 478. 

~ americana, 250. Cingulina spina, Cr. § Fischer, 173. 

maxima, 249, Circe castrensis, Linn., 194. 

rudis, 249. gibbia, Lam., mentioned, 194. 
torquata, 249. pectinata, Linn., 194. 

Cestracion, 49. rivularis, Born, 194. 

Philippi, 47. Cirripedia, 60, 61, 116. 

Chama divaricata, Reeve, 193. Citharoscelus Kochii, Pocock, 15, 16. 

fibula, Reeve, 193. Cladocopa, 61, 89. 

fimbriata, Reeve, 193. Clanculus atropurpureus, Gould, 175. 

pellis-phoce, Reeve, 193. unedo, A. Ad., 175. 

pulcheila, Reeve, 193. Clausia e/ongata, Boeck, 91. 

reflexa, Reeve, 193. Cleippides, Boeck, 78. 

(Arecinella) spinosa, Brod., 193. quadricuspis, Heller, 78, 117. 
Chamidez, 193. Clemmys caspia, mentioned, 363. 
Characinide, 526. leprosa, mentioned, 363. 
Charcharodus althese, Hiibner, 411. Cletodes, Brady, 104. 

Chelyconus radiatus, Gmel., 156. longicaudata, Brady § Roberés., 104, 
Chicoreus adustus, Lam., 161. 118. 

, var. fuscus, Dér., 161. similis, 7. Scott, 104, 118. 
axicornis, Lam., 161. tenuipes, 7: Scott, 104, 118. 
capucinus, Lam., 161. Clisiocampa castrensis, Linn., 413. 
cervicornis, Lam., 162. Ciubionide, 15, 17. 
ramosus, Linn., 162. Clupea, 531, 534, 536. 

Chilopoda and Arachnida of Lake alosa, Linn., 526; mentioned, 519, 
Urmi, by R. I. Pocock, 399-406. 621. 
Chione (Omphaloclathrum) Chemnitzii, finta, Cuv., 526. 
Hanley, 195. harengus, Weber, 526 ; mentioned, 


(——) costellifera, Ad., 195. 519, 521, 530. 


INDEX. 


Clupea pilchardus, Walb., 526. 

sprattus, Linn., 526. 
- Clupeide, 526. 

Clypeaster, 419, 424, 433. 
altus, Philippi, ftnote 421, 422. 
altus, Lam., 422, 434, 435. 

, var. turritus, Philippi, 422, 

435. 

crassicostatus, Agass., 424, 454. 

diversicostatus, Abich, 435. 

faloriensis, Dunc. g Sladen, men- 
tioned, 424. 

gibbosus, fisso, mentioned, 421, 
422. 

Guentheri *, Gregory, 420; men- 
tioned, 421, 422, 430. 

imperialis, Michelin, 
tioned, 420, 430. 

- Martini, Desmoulins, 422. 

melitensis, Mich., mentioned, 422, 
423. 

Michelotti, Ag., mentioned, 422, 
423. 

olisiponensis, Michelin, 419, 420. 

Scillx, Desm., 436. 

Scilla, Philippi, ftnote 421. 

suboblongus, Pomel, mentioned, 
422, 

turritus, Abich, 421. 

turritus, Philippi, 

436. 

Cobitis persa, Heckel, 391. 

Coccyges, 248. 

Codakia exasperata, Reeve, 200. 
fibula, Reeve, 200. 
interrupta, Lam., 200. 

Ceelenterata, 301. 

Ceelotes, 21. 

Coenonympha pamphilus, 408. 

Coenopsammia, 307. 

Colacium calvium, Stein, 467. 

Cole, EH. J.. On the Discovery and 

- Development of Rhabdite-‘‘cells” 
in Cephalodiscus dodecalophus, Mc- 
Intosh, 256-268. 

Colias croceus, 410. 

edusa, Fabr., 410. 
electra, 410. 
hyale, 410. 
mongola, 43. 

Colii, 248. 

Colina contractum, Sow., 168. 
pinguis, Ad., 168. 
teniatum, Sow., 168. 

Collembola, On some Australian, by 
the Right Hon. Sir John Lubbock, 
334-338. 

Columba, 216, 227, 228, 252. 
domestica, 256; wing of, 243. 
livia, 227. 

Columbe, 233, 234, 248. 


men- 


419; 


421, 


563 


Columbella varians, Sow., 160. 
(Atilla) galaxias, Reeve, 161. 
(Conoidea) flava, Brug., 161. 
ee semiconvexa, Lam., 
(Pygmza) fulgurans, Lam., 160. 
(—— , a. eufulgurans, 160. 

(——) —, p. punctata, Laim., 

160 


(—— ) Tyleri, Gray, 161. 
( ) versicolor, Sow., 161. 

Columbellide, 160. 

Columbula, 213, 248, 252, 253. 
livia, 219, 229. 
picui, mentioned, 212, 2138, 215, 

222, 236. 

Conide, 156. 

Conoclypeus, mentioned, 422. 

Conoidea flava, Brug., 161. 

Conomurex luhuanus, Linn., 166. 

Conus marmoreus, Linn., 156. 
sp., 431, 439. 

(Chelyconus) radiatus, Gmel., 156. 
(Coronaxis) minimus, Linn., 156. 
OTe Sa figulinus, Linn, 
6. 
(Hermes) nussatella, Linn., 156. 
(Lithoconus) litteratus, Zinn., 156. 
, var. millepunctatus, 
Lam., 156. 
(Nubecula) striatus, Linn., 156. 
(Bhizoconus) mustelinus, Awass, 
156. 
(-—) vitulinus, Hwass, 156. 
(Textile) canonicus, Hwass, 156. 

Copepoda, 61, 90, 395. 

Coraciz, 248. 

Corbis ? Aglaure, Brong., 446. 

Corbula crassa, Hinds, 199. 
Macgillivrayi, Smith, 199. 
taheitensis, Zam., 199. 
truncata, Hinds, 199. 

Cormorant, Note on the Hxternal 
Nares of the, by W. P. Pycraft, 
207-209. 

Coronaxis minimus, Linn., 156. 

Corvus, 227, 248, 

Corydalis, 42. 

Corynactis, mentioned, 303. 

Coryne, 258. 

Coscinarza, Milne-Edw. § Haime, 144; 

mentioned, 143. 
Botte, Milne-Edw. & Haime, 145. 
meandrina, Kéwnz., 145. 
monile, Forsk., 145. 

Cosmoporites, Duchass. § Michel., 143, 
147, 149. 

Cottide, 524. 

Coua, 251. 

Cranes, 248, 251. 

Craspedia marginepunctata, Goeze, 413, 


564 


Crassatella ziczae, Reeve, 188, 

Crassatellidz, 188. 

Crax, 248. 

Crenatula flammea, Reeve, 184. 

Cricetus frumentarius, 318. 

Crick, G. C., Note on a Jurassic 
Ammonite of Lake Urmi, 418 

Criotherium argalioides, Ma7., 377. 

Crista gibbia, Lam., mentioned, 194, 

pectinata, Linn., 194. 

Cronia amygdala, Kiener, 163. 

Crotophaga, 251. 

Crustacea of Lake Urmi, by R. T. 
Ginther, 394-398. 

, Report on the Marine and Fresh- 
water, from Franz-Josef Land, col- 
lected by Mr. William S. Bruce of 
the Jackson - Harmsworth Expedi- 
tion, by Thos. Scott, 60-126. 

Cryptoplax oculatus, Q. ¢ G., 181. 

striatus, Lam., 181. 

Cryptopterus, 523. 

micronema, Blkr., 522. 

micropogon, Blkr., 522. 
Ctenocardium fornicatum, Sow., 1938. 

fragum, Linn., 193. 

imbricatum, Sow., 193. 
Ctenoides fragilis, Chemn., 182. 

tenera, Chemn., 182. 

Cuckoo, 251. 

Cuculidz, 233. 

Cucullza concamerata, Chemn., 188. 

Cuma Rathkii, Kroyer, 65. 

Cumacea, 61, 65. 

Cupido ade var. ripartii, Preyer, 

409. 
aedon, Christoph., 409. 
agestis, Schiffermiiller, 409. 
bellargus, var. oceanus, Bergstr., 
409. 
dama, var. ?, Staud., 409. 
damon, Schiffermiiller, 409. 
endymion, Schiffermiiller, 409. 
icarus, Rott., 409. 

Cuspidaria latisulcata, Zen.-Woods, 
202. 

Cuspidariidz, 202. 

Cyathomorpha conglobata, Reuss, 436. 

gregaria, Catullo, 436. 

Cyclocypris, Brady & Norm., 82. 

globosa, G. O. Sars, 82, 117. 

Cyclopicera nzgripes, Brady & Roberts., 

113. 


Cyclopide, 92. 
Cyclopina, Claus, 93. 
gracilis, Claus, 93, 117. 
Core O. F. Miiller, 938; eyes of, 
aT 
bisetosus, Rehberg, 94. 
Brucei*, Zhos. Scott, 98; men- 
tioned, 94, 117. 


INDEX. 


Cyclops chelifer, O. F. Miller, 111. 
eae Baird, 112. 
Cyclostrema, Marryat, mentioned, 178, 
Cyclostrematide, 177. 
Cylichna arachis, Q. & G., 155. 
Cynodonta cornigera, Lum., 158. 
Cynoglossus semilevis, Giimther, On 
the Presence of Nasal Secretory Sacs 
and a Naso-pharyngeal Communi- 
cation in Teleostei, with especial 
reference to, by H. M. Kyle, 541- 
555. 
Cyprea, 150. 
Comptoni, mentioned, 152. 
felina, Gray, 164. 
helvola, Zinn., mentioned, 151. 
(Aricia) annulus, Linn., 164, 
( ) arabica, Linn., 164. 
(Luponia) errones, Linn., 164, 
(—) flaveola, Linn., 164. 
(——) lynx, Linn., 164. 
( ) ziczac, Linn., 164. 
Cypreide, 164. 
Cypria lacustris, Li//eb., 82. 
Cypridina brenda, Baird, 89. 
Cyprinide, 193, 526. 
Cyprinodon, 382. 
Cypris crassa, O. F. Miiller, 82. 
- globosa, G. O. Sars, 82. 
Cypselidze, 234, 250. 
Cythara chionea* , Melvill & Standen, 
156 


gracilis, Reeve, 156. 
pulchella, Reeve, 156. 
Cythere, Miiller, 84. 
cluthe, Brady, Crosskey, § Robert- 
son, 84, 117. 
contorta, Norm., 88. 
costata, G. 8. Brady, 85, 117. 
declivis, Norm., 86. 
dunelmensis, Norm., 85, 117. 
emarginata, G. O. Sars, 85, 117. 
globulifera, G. S. Brady, 84, 117. 
latissima, Norm., 87. 
limicola, Norm., 84, 117. 
marginata, Novm., 84, 117. 
mirabilis, G. S. Brady, 85, 117. 
septentrionalis, G. S. Brady, 85, 
117. 
tuberculata, G. O. Sars, 85, 117. 
variabilis, Baird, 88. 
Cytherea incrassata, Deshayes, 447. 
Cythereis dunelmensis, Norm., 85, 
emarginata, G. O. Sars, 85. 
tuberculata, G. O. Sars, 85. 
Cytheridea, Bosqguet, 86. 
papillosa, Bosquet, 86, 117. 
punctillata, G. S. Brady, 86, 
117. 
Sorbyana, Jones, 86, 117. 
Cytheropteron, G. O. Sars, 87. 


INDEX, 


Cytheropteron angulatum, Brady & 
Robertson, 88, 117. 
latissimum, Norm., 87, 117. 
punctatum, G. S. Brady, 87; 
mentioned, 88, 117. 
pyramidale, G. S. Brady, 87, 117. 
subcircinatum, G. O. Sars, 87; 
mentioned, 88, 117. 
Cytherura, G. O. Sars, 86. 
clathrata, G. O. Sars, 87, 117. 
fulva, Brady § Robertson, 87, 117. 
undata, G. O. Sars, 86, 117. 


Dacelo, 249. 
gigas, 249. 

Dactylopus, Claus, 101, 104. 
coronatus, 7. Scott, 105, 118. 
longirostris, Claws, mentioned, 101, 

105, 118. 
minutus, Claws, mentioned, 101. 
Stromii, Baird, 105, 118. 

, var. arcticus * , 7. Scott, 106. 

tenuiremis, Brady & Robertson, 
105 ; mentioned, 106, 118. 
tisboides, Claus, 104, 118. 

Dajus, Kroyer, 68. 
mysidis, Kroyer, 65, 68, 116. 

Daphnia sp., 395. 

Daucus Carota, carotin in roots of, 

471. 

Decapoda, 394. 

Decticidx, 418. 

Decticus, 371. 
albifrons, Fabr., 418. 
assimilis, F7%ed., 416, 418. 

Deiopeia pulchella, Linn., 411. 

Delavalia, Brady, 98. 
arctica * , 7, Scott, 98, 118. 
Giesbrechti, 7. Scott, mentioned, 


98. 
mimica, 7. Scott, 98, 118. 
palustris, Brady, mentioned, 98, 
99. 


reflexa, Brady & Roberts., 98, 118. 
robusta, Brady § fRobverts., 98; 
mentioned, 99, 118. 
Delphinula formosa, Reeve, 177. 
Delphinulide, 177. 
Dendrochelidon mystacea, 249. 
Dendroconus figulinus, Linn., 156. 
Dentaliide, 181. 
Dentalium javanum, Sow., 181. 
pseudo-sexagonum, Desh., 181. 
Dermatomyzon, Claus, 113. 
nigripes, Brady & Roberts., 113, 
118 


Diala albugo, Wats., 170. 

Diastopora gemmifera, 
mentioned, 435. 
Diastylis, Say, 65. 

Rathkei, Kroyer, 65, 116. 


Abich, 4384; 


565 


Dibranchia, 199. 
Dicholophus, 234, 248. 
Dicranura vinula, Linn., 412, 


‘Dictynide, 15, 18. 


Dione inflata, Sow., 193. 

Diosaccus, mentioned, 106. 

Diplodonta, mentioned, 197. 

ethima*, Melv. & Stand., 197, 
206. 

suberassa, Smith, 197. 

subglobosa, Smith, 197. 

Dipnoi, 541, 550. 

Diptera and Neuroptera (Hemerobiid) 
of Lake Urmi, by R. T. Ginther, 
414-416. 

Discoglossus, 461. 

Divaricella Macandresx, H. Ad., 199. 

ornata, Reeve, 200. 

Divaricellze, 200. 

Doliide, 164. 

Dolium perdix, Linn., 164. 

variegatum, Linn., 164. 
(Malea) pomum, Linn., 164. 

Donacilla rhomboides, Gmel., 201, 

semitorta, Sow., 201. 

virgulata, Hanley, 201. 
Dosinia czrulea, Reeve, 194, 

histrio, Gael., 194. 

Draba ochroleuca, 42. 

Dracocephalum grandiflorum, 42, 

Dreissena, mentioned, 353. 

Drillia torresiana, Smith, 156. 

Dryas octopetala, 43. 

Duck, wing of, 243. 

Duerden, J. E., The Edwardsia-stage of 
the Actinian Lebrunia,and the Forma- 
tion of the Gastro-ccelomic Cavity, 
269-316. 

Dulichia, Kroyer, 80. 

spinosissima, Kréyer, 80, 117. 

Dysdera crocota, C. Koch, 15. 

maxima, Niec., 17. 
Dysderide, 15, 17. 


Echinodermata, 179, 307, 434, 435, 
436. 
Echinoidea, Fossil, of Lake Urmi, by J. 
W. Gregory, 419-424. 
Echinolampas, 433. 
complanatus, Adbich, 434, 435, 
436. 
Hetinosoma, Boeck, 94; mentioned, 95, 


atlanticum, Brady §& Roberts., 96; 
mentioned, 95, 117. 

curticorne, Boeck, 95, 117. 

melaniceps, Boeck, 95, 117. 

Normani, 7. & A. Scott, 95; men- 
tioned, 96, 117. 

propinquum, 7. g& A, Scott, 95, 
117. 


066 


Ketinosoma pygmeum, 7. g A. Scott, 
95, 117. 
Sarsi, Boeck, 94; mentioned, 95, 
117. 
tenwe, Brady & Roberts., 96. 

Edentata, 377. 

Edriophthalma, 61. 

Hdwardsia-stage, The, of the Actinian 
Lebrunia, and the Formation of the 
Gastro-celomic Cavity, by J. EH. 
Duerden, 269-316. : 

Eelisia tricarinata, Ad. & Reeve, 172. 

Elephantide, 377. 

Elk, 37. 

Elwes, H. J., On the Zoology and 
Botany of the Altai Mountains, 
23-46. 

Enchytrus, mentioned, ftnote 502. 

Endothyra, mentioned; 351. 

Bowmanni, Phillips, 
452. 
Engina concinna, Reeve, 159. 
Reevei, Zryon, 159. 
Engraulis enchrasicholus, Cuv., 526. 
Enhydrosoma, Boeck, 104. 
curvatum, Brady g Roberts., 104, 
118.: 
Entomostraca, 60, 61, 81, 395. 
Ephydra halophila, 415, 416. 
riparia, mentioned, 556; larvee of, 


mentioned, 


salinaria, 415. 
urmiana*, R. T. Giinther, 415; 
larva of, 416. 
Epidromus antiquatus, Hinds, 163. 
Bednalli, Brazier, 163. = 
Epinephele hispulla, Esper, 408. 
lycaon, var. lupinus, Costa, 408. 
Episema glaucina, Esper, 412. 
Equide, 377. 
Equus Prejvalskii, 41. 
Erebia, 39. 
afer, Hsper, 408. 
ero, 43. 
lapponica, 43. 
maurisius, 43. 
tyndarus, 42. 
Eremia, 370. 
Ericacex, 43. 
Eristalis, larvee of, 356. 
Erycinide, 189. 
Eryx jaculus, Linn., 380. 
, var. miliaris, 380. 
Eucheeta Philippi, 91. 
norvegica, Boeck, 91, 117. 
Euchelus angulatus, Pease, 177. 
atratus, Gmel., 177. 
Euclidia mi, Clerch, 412. 
Eucythere, Brady, 86. 
declivis, Norm., 86, 117. 
Euglena deses, Hhrend., 467. 


“INDEX. 


Euglena Ehrenbergii, 480. 
Spirogyra, Szein, 467 ; cells in, 467. 
viridis, on the Hye-spot and Fla- 
gellum in, by Harold Wager, 
463-481. 
, vacuole system and gullet, 
466-468. 
—, structure of the eye-spot, 
468-472. 
—, flagellum, 472-474. 
——.,, effect of light on, 474-477. 
, function of the eye-spot, 
477-481. 
Eulima, 173. 
australasiaca *, Melv. 
173, 206. 
latipes, Watson, 173. 
(Leiostraca) acicula, Gould, 173. 
Eulimide, 173. 
EHumeces, 380. 
algeriensis, 380. 
Schneideri, Daud., 379 ; mentioned, 
361. 
Eupelte, Claus, 112. 
purpurocincta, Norm., 112, 118. 
Euphausiide, 485. 
Eupbyllia, 310, 311; mentioned, 297. 
rugosa, mentioned, 310. 
Hupithecia subumbrata, Schiffermiiller, 
413. 
Eupleura jucundum, A. Ad., 164, 
perca, Parry, 164. 
pulchra, Gray, 164. 
Kupolia, 266. 
Eurylemus, 248. 
javanicus, 250. 
EKurypelma sp., 16. 
Eurypyga, 248, 250, 254. 
Kusirus, Kréyer, 76. 
cuspidatus, Kréyer, 76, 117. 


g§ Stand., 


Kuthemisto libellula, Mandt, men- 
tioned, 69. 
Eutrochus septenarium*, Melv. ¢ 


Stand., 176. 

Exoceetus, 522. 

Hye-spot and Flagellum, On the, in 
Huglena viridis, by Harold Wager, 
463-481. 


Falco altaicus, 438. 
sacer, 43. 

Fasciolaria filamentosa, Zam., 158. 

Fasciolariids, 158. 

Fin-foot, 248. 

Fishes of Lake Urmi, by Dr. A. 

Ginther, 381-391. 

Fissurella corbicula, Sow., 179. 
lanceolata, Sow., mentioned, 179. 
(Glyphis) corbicula, Sow., 179. 
(——) Jukesi, Reeve, 179. 

Fissurellidx, 179. 


INDEX. 


Flabellipecten, 431. 


sp., 444. 
Flabellum, 132, 134. 
Flagellum, in Euglena viridis, by 


Harold Wager, 463-481. 
Flat-fishes, 541. 
Foraminifera, 435. 
Forficula auricularia, Zénn., 416. 
Forficularia, 416. 
Fossil Corals of Lake Urmi, by J. W. 
Gregory, 424-430. 
Fragum unedo, Linn., 192. 
Fucacee, 471, 472. 
“ Hundusdrisen,” 2. 
Fusulina cylindrica, Fischer, mentioned, 
. 453. 
spheerica, Aich, mentioned, 453. 
Fusus crispus, Borson, 450. 


Gadide, 547. 

Galaxea, mentioned, ftnote 488. 

Galeodes, Oliv., 361, 402. 
arabs, C. Koch, 402. 
citrinus, Pocock, 403, 404. 
eyrus, Pocock, 404. 
darius, Pocock, 403. 
truculentus * , Pocock, 402; men- 

tioned, 403, 404. 

-Galeommiidie, 189, 

Galeus, ftnote 46. 
canis, 47. 

Galidee, 524. 

Gallinula Campbelli, Gray, 165. 
canarium, Linn., 165. 
Isabella, Zam., 165. 

Sibbaldi, Sow., 165. 

urceus, Linn., 165. 

variabilis, Swains., 165. 
_ vittatus, Linn., 165. 

Gallus, 248. 
bankiva, var. domestica, 255, 

Gammaracanthus, Spence Bate, 79. 
loricatus, Sabine, 79, 117. 

Gammaridea, 69. 

Gammarus, Fadr., 79. 
carinatus, Fabr., 78. 
locusta, Linn., 69, 79, 117. 
loricatus, Sabine, 79. 
pinguis, Kroyer, 79. 
pulex, De Geer, 366, 395. 

Gampsonychidex, 343. 

Ganoris rape, var. ergane, Hiibner, 410. 

Gari anomala, Desh., 198. 
marmorea, Desh., 198, 
ornata, Desh., 198. 
prestans, Desh., 198. 
pulcherrima, Desh., 198. 

Gasteropoda, 154, 434, 435, 437, 439, 

448, 
Gastric Glands of Marsupialia, On the, 
by J. Johnstone, 1-14, 


567 — 


Gastrochena cuneiformis, Zam., 199. 
plicatilis, Desh., 199. 
Gastrochenide, 199. 


‘Gastrosteus, mentioned, 547. 


aculeatus, Linn., mentioned, ftnote 
542. 
Gayenna maculatipes, Keys., 15, 18. 
Gazella brevicornis, 377. 
deperdita, Gaudr., 377. 
subgutturosa, mentioned, 362. 
Gena striatula, A. Ad., 178. 
Gentiana altaica, 42. 
Geometridx, 4138. 
Geopelia cuneata, mentioned, 212, 213, 
215, 235. 
tranquilla, mentioned, 212, 218, 215, 
220, 222. 
Geophaps, 213, 215. 
plumifera, 212, 215, 220 
Geotrygon, 212, 213. 
montana, mentioned, 212, 236. 
Gigantostraca, 60. 
Giraflide, 377. 
Gitana, Boeck, 72. 
Sarsii, Boeck, 72, 117. 
Glyphis corbieula, Sow., 179. 
Jukesi, Reeve, 179. 
Glyphostoma rugosa, Migh., 156. 
Gnathia, Leach, 66. 
elongata, Kroyer, 66; mentioned, 
67, 116. 
Gnathostomata, 61. 
Gobio Kessleri, mentioned, 387. 
persa *, Giinther, 386, 391. 
uranoscopus, 387. 
Goniastrea, 136. 
Goniomya Rodleri, Weithofer, 438. 
Goniopora, 129, ftnote 130, 131, 137 
158, 140, 141, 142, 148, 144, 
148, 149, 491, 497. 
Stokesi, 148. 
Gorgonia, mentioned, 310. 
Goura, 226. 
Gregory, J. W., Fossil Corals of Lake 
Urmi, 424—430. 
, Fossil Kchinoidea of Lake Urmi, 
419-424, 
Gresslya sp., 439. 
‘Grues, 248. 
Gryllodea, 418. 
Gryllotalpa gryllotalpa, Zinn., 371,418. 
Gryllotalpide, 418. 
Giinther, Dr. A., The Wild Sheep of the 
Urmi Islands, 374-376. 
, Fishes of Lake Urmi, 381-391. 
Ginther, Robert T., Contributions to 
the Natural History of Lake Urmi, 
N.W. Persia, and its Neighbourhood, 
345-453. 
—, Crustacea of Lake Urmi, 394- 
398.. 


bo 
(SC) 
Or 


568 


Ginther, R. T.. Neuroptera (Hemero- 
biide) and Diptera of Lake Urmi, 
414-416. 

Guillemot, wing of the embryo, 242. 

Gymnarchus, 524. 

niloticus, mentioned, 531. 

Gymnotide, 526. 

Gyrineum (Apollon) pusillum, Brod., 

164. 


(Eupleura) jucundum, 4. Ad., 164. 
(Lampas) ranelloides, Reeve, 163. 


Haddon, Prof. A. O., First Expedition 
to the Torres Straits in 1888-89, 150- 
206. 

Hadena bimaculosa, Zinn., 411. 

Halcyon, 249. 

chloris, 249. 
saneta, 249. 
vagans, 249. 

Halimedon obtusifrons, Hansen, 73. 

Haliotids, 150, 179. 

Haliotis Philberti, Serves, 434, 435. 

varia, Linn., 179. 

(Padollus) ovina, Chemn., 179. 

(Teinotis) asinina, Lénn., 179. 
Halirages, Boeck, 77. 

fulvocinctus, M/. Sars, 77, 78, 117. 

Halmopota salinaria, 415. 

Halophilus, 416. 

Haminea brevis, Q. & G., 155. 

crocata, Paase, 155. 

Hamster, 318. 

Haploplax purus, Sykes, mentioned, 
180. 


Harpacticide, 94. 
Harpacticus, Milne-Hdw., 111. 
chelifer, Miidler, 111, 118. 
—, var. arcticus* , 7. Scott, 111, 
118. 
gracilis, Claus, mentioned, 111, 112. 
Harpoceras, 437. 
atropatenes, Borne, 439. 
kapautense, Borne, 439. 
kurrianum, Oppel, 437. 
medi, Borne, 439. 
radians, Reinecke, 437. 
Heliastrzea Defrancei, mentioned, 426. 
Guettardi, Defr., 486. 
Haimei, d’ Archiac, ftnote 427, 
Helicella acutistria, Bottger, 391. 
apicina, Lamk., mentioned, 392, 
parableta, Bottger, 391. 
pisiformis, Pfezffer, 392. 
sp., 392. 
Helicogena figulina, Parreyss, 391. 
Helicophora rotundicornis, Weith., 377. 
Heliornis, 248, 253, 254. 
Helix (Helicogena) figulina, Parreyss, 
391 


Hemerobius ?, larva of, 414, 416, 


INDEX. 


Hemicardium subretusum, Sow., 192. 
(Ctenocardium) fornicatum, Sow., 
193 


( ) fragum, Linn., 1938. 
( ) imbricatum, Sow., 193. 
(Fragum) unedo, Linn., 192. 

Hemilepistus, 394. 

Hemimactra aspersa, Sow., 199, 

Hemipodes, 248. 

Henopomus tricornis, Kroyer, 67. 

Heptadactylus lambis, Linn., 166. 

Heptanchus, 49, 51. 

Heracleum, 453. 

Hermes nussatella, Linn., 156. 

Herodiones, 247. 

Herpetocypris, Brady § Norm., 82. 
arctica *, Zhos. Scott, 83, 117. 
dubia *, Thos. Scott, 82, 117. 

Hesperiide, 411. 

Hesperornis, 251. 

Heterographis pyrethrella, Herr.- Schéff., 

413. 


Heteromysis, mentioned, 484, 

Heteropora, 129. 

Heterosomata, 541, 546, 547, 548. 

Heterotis, 524. 

Hexacorallia, 302. 

Hexactiniz, 303, 308. 

Hima plebecula, G/d., 160, 

Himantornis, 250. 

Hipparchia briseis, 408. 

, var. turanica, Stgr., 408. 

circe, Fabr., 408. 

pelopea, Klug, 408. 
Hipparion mediterraneum, Hensel, 377. 
Hippolyte Gatmardii, Milne-Edw., 63. 

Phippsti, Kroyer, 63. 
Holocentrum, 525. 

spiniferum, Giinther, 528. 
Hoplites narbonensis, Pictet, 488. 
Hoplonyx, G. O. Sars, 70. 

similis, G. O. Sars, 70, 116. 

Hoplophoria coralligens, H. V. Wilson, 
mentioned, 270, 271. 

Hyena eximia, Roth et Wagn., 377. 

Hydrozoa, 301. 

Hyla arborea, Linn., 381. 

, var. Savienyi, Aud., 381. 

Hymenochirus Boettgeri, Note on the 
Carpus of the new Aglossal Toad, 
461-463. 

, On the Hyobranchial Skeleton and 
Larynx of the new Aglossal Toad, 
454-460. 

Hyobranchial Skeleton and Larynx of 
the new Aglossal Toad, Hymeno- 
chirus Boettgeri, On the, by W. G. 
Ridewcod, 454-460. 

Hyodon, 527, 529, 531, 534, 536. 

claudulus, Les., 625, 527, 530. 
tergisus, Les., 525. 


INDEX. 


Hyodontide, 525. 
Hyperia, Latr., 69. 
galba, Montagu, 69, 116, 
oblivia, Kroyer, 69. 
Hyperiidea, 69. 
Hypsopygia costalis, Fabr., 414. 
Hystricomorpha, 320, 321, 331, 334. 


Tanthina fragilis, Linn., mentioned, 172. 
Smithiz, Reeve, 172. 
Tanthinide, 172. 
Ibex of the Altai, 37. 
Ichthyopsida, mentioned, ftnote 550. 
Ictitherium hipparionum, Gaudr., 377. 
Idya, Philippi. 112. 
furcata, Baird, 112; mentioned, 
113, 118. 
minor, 7. & A. Scott, 113, 118. 
Ino excelsior * , Melv. §& Stand., 116. 
Insecta (Lepidoptera Phaleenz) of Lake 
Urmi, by Sir G. F. Hampson, 411- 
414. 


(Lepidoptera Rhopalocera) of 
Lake Urmi, by A. G. Butler, 408- 
411. 

Tris ruthenica, 27. 

tigridia, Bunge, 42. 
Isanda, Ad., mentioned, 178. 

coronata, Ad., mentioned, 178. 
Ischnochiton (Haploplax) purus, Sykes, 

mentioned, 180. 

( ) sp., 180. 
Ischyrocerus, Kroyer, 80. 

anguipes, Kroyer, 80, 117. 
Tsopoda, 61, 65, 394. 


Janira, Leach, 67. 
tricornis, Kroyer, 67, 116. 
Johnstone, James, On the Gastric 
Glands of the Marsupialia, 1-14. 
Jonesiella, Brady, 97. 
spinulosa, Brady § Roberts., 97, 
118 


Julodis levicostatus, L. § G., 370. 


Kagu, 248. 

Katelysia scalarina, Lam., 196. 

Kellia, mentioned, 189. 

physema*, Melv. § Stand., 189, 
206. 

Koala, Anatomy of, 1. 

Kyle, H. M., On the Presence of Nasal 
Secretory Sacs and a Naso-pharyngeal 
Communication in Teleostei, with 
especial reference to Cynoglossus 
semileevis, Giinther, 541-555. 


Labride, 542. 

Labrus, mentioned, 547. 

Lacerta viridis, Laur., 378. 

, var. strigata, Hichw., 378. 


569 


Lacertilia, 378. 

Levicardium Bechei, Ad. §- Reeve, 192. 
biradiatum, Brug., 192. 
lyratum, Sow., 192. 

Lagomorpha, 320, 331. 

Lake Urmi, Contributions to the 
Natural History of, N.W. Persia, 
and its Neighbourhood, by R. T. 
Gunther, 345-453. 

Lamellibranchiata, 434-436, 439, 440. 

Lampania zonale, Brug., 168. 

Lampas ranelloides, Reeve, 1638. 

Lamprops, Sars, 65. 

fuscata, G. O. Sars, 65, 116. 
Lamprostoma maculatus, Lenn., 175. 
Laophonte, Philippi, 101. 
curticauda, Boeck, 102 ; 
tioned, 103, 118. 
depressa, 7. Scott, 102, 118. 
horrida, Norm., 101; mentioned, 
102, 118. 
intermedia, Z. Scoté, 102, 118. 
longicaudata, Boeck, 102, 118. 
perplexa *, 7. Scott, 103, 118. 
similis, Claus, 102; mentioned, 
103, 118. 
Laophontodes, 7. Sco?z, 103. 
typicus, 7. Scot¢, 103, 118. 

Lapwing, wing of the, 242. 

Larix sibirica, 43. 

Lasiocampide, 413. 

Lathrodectus, Walck., 400. 

tredecim-guttatus, Ross?, 400. 
, var. erebus, Sav., 400. 
Latirus crispus, Borson, 431, 450, 452. 
polygonus, Gmel., 158. 
, var. tessellatus, Kobelt, 158. 
(Peristernia) australiensis, Leeve, 
158. 

Lebrunia, the Edwardsia-stage of the 
Actinian, and the Formation of the 
Gastro-celomic Cavity, by J. H. 
Duerden, 269-316. 

Lebrunia coralligens, H. V. Wilson, 
mentioned, 271, 275, 300. 

Leda Darwini, Smith, 188. 

Leiostraca acicula, Gould, 173. 

thoracica, mentioned, 103. 

Lepidoleprus, mentioned, 530. 

collorhynchus. mentioned, 530. 

norvegicus, 530. 

trachyrhynchus, isso, mentioned, 
530 


men- 


Lepidoptera Phalenze of Lake Urmi 
(Sir G. F. Hampson), 411-414. 
Lepidoptera Rhopalocera of 
Urmi (A. G. Butler), 408-411. 
Lepidorhombus whiff, mentioned, 543. 
Leptognathia, G. O. Sars, 66. 
longiremis, Lilljeborg, 66, 116. 
Leptomysis, mentioned, 484. 


Lake 


570 


Leuciscus, mentioned, 349. 
aula, mentioned, 389. 
cephalus, Linn., 387. 
gaderanus*, Giimther, 364, 388, 
391. 
ulanus * , Giinther, 364, 387 ; men- 
tioned, 388, 391. 
Leucosarcia, 213, 215, 227. 
picata, 212, 215, 235. 
Leucothoé phyllonyx, M. Sars, 73. 
Leucotis, Swains., mentioned, 171. 
Libitina angulata, Lam., 193. 
Lima squamosa, Lam., 182. 
(Ctenvides) fragilis, Chemn., 182. 
( ) tenera, Chemn., 182. 
(Limatula) bullata, Born, 182. 
-——) torresiana, #. A. Smith, 
182, 206. 
(Mantellum) areuata, Sow., 182. 
( ) inflata, Zam., 182. 
Limatula bullata, Born, 182. 
torresiana, HL. A. Smith, 182, 206. 
Limestone, Note on a Paleozoic, from 
Lake Urmi, by R. Bullen Newton, 
452-453. 
Limicole, 247. 
Limide, 182. 
Limnea palustris, Miller, 393. 
stagnalis, Linn., 393. 
truncatula, Miiller, 393. 
Limopsis cancellata, Reeve, 188, 
Woodwardi, A. Ad., 188. 
Linnzea borealis, 43. 
Linum ecxruleum, +2. 
Lioconcha hebrea, Lam., 193. 
picta, Lam., 194. 
Sowerbyi, Desh., mentioned, 194. 
tigrina, Lam., mentioned, 194. 
Liotia varicosa, Reeve, 177, 
Litharea, 143. 
Lithobius, Leach, 399. 
forficatus, 399. 
mutabilis, ZL, Koch, 399. 
persicus * , Pocock, 399. 
Lithoconus litteratus, Linn., 156. 
, var. millepunctatus, Lazm., 
156. 
Lithophagus canaliferus, Hanley, 184. 
gracilis, Phil., 189. 
Hanleyanus, D/r., 185. 
teres, Phil., 185. 
Litiopide, 170. 
Littorina filosa, Sow., 170. 
(Melaraphe) mauritiana, Lam., 
170. 
Littorinide, 170. 
Lixus bardanex, £., 370. 
Lloydia serotina, 42. 
Locustodea, 418. 
Lomvia troile, 256 ; wing of, 242, 
Lophophaps plumifera, 215. 


INDEX. 


Loripes Haddoni*, Melvill & Standen, 
200, 206. 

Lotella bacchus, 525, 588. 

Lubbock, Sir John, on some Australasian 
Collembola, 334-338. 

Lucina (Codakia) exasperata, Leeve, 


( ) fibula, Reeve, 200. 

( ) interrupta, Zam., 200. 

(Divaricella) Macandree, H. Ad., 

199. 

( ) ornata, Reeve, 199. 

Lucinacea, 199. 

Lucinidex, 199. 

Ludwigia gigas, Quenst., 438. 
krakoviensis, Newmayr, 438. 
lunula, Zieten, 438. 
nodosa, Quenst., 438. 
punctata, Stahl, 438. 

Luponia errones, Linn., 164. 
flaveola, Linn., 164. 
lynx, Linn., 164. 
ziczac, Linn., 164. 

Lutraria arcuata, Desh., 199. 
rhyncheena, Jones, 199. 

Lyczna, 39. 
dama, Steud., 409. 
omphale, Klug, 410. 
orbitulus, 43. 
pheretes, 43. 
thetis, Klug, 410. 

Lyceenidee, 409. 

Lycosa, Latr., 370, 401. 
australis, H. Simon, 16, 22. 
Fernandezi * , Pickard-Cambridge, 

16, 21. 
ferox, Lucas, mentioned, 402. 
Guentheri * , Pocock, 401. 
implacida, Nic., 16, 21. 

Lycosidx, 16, 21. 

Lymantria dispar, Linn., 412. 

Lymantriadx, 412. 


“a 


Mabuia septemtzeniata, Reuss, 379. 
Macaria murinaria, Schiffermiiller, 413. 
Machetes pugnax, 240, 255. 
Machilis, mentioned, 407. 
pera ?, 416; Acarid upon, 416. 
Macrocephalites sp., 438. 
Macroglossa stellatarum, Linn., 412. 
Macrura, 61, 63, 394. 
Macruride, 530. 
Macrurus, 531. 
trachyrhynchus, Risso, mentioned, 
ftnote 530. 
Mactra achatina, Chemn., 199. 
apicina, Desh., 199. 
(Hemimactra) aspersa, Sow., 199. 
(Oxyperas) Coppingeri, Smith, 199. 
Mactridx, 149. 


INDEX. 


Madrepora, Hhrenbd., mentioned, 128, 
129, 296, 487, 488, 495, 496. 
Madrepora Astrzopora, 129. 
Phyllopora, 129. 
Porites, 129. 
Madreporaria, 296, 308. 
Madreporarian System, Recent Poritide, 
and the Position of the Family in the, 
by Henry M. Bernard, 127-149. 
Meandrarea, 148. 
Magadis* , Melv. § Stand., 174. 
eumerintha * , Melv. § Stand., 174, 
206. 
Malacostraca, 60, 61, 63, 394. 
Malea pomum, Linn., 164. 
Mamma aurantia, Lam., 172. 
Flemingiana, Récluz, 172. 
Mammalia, gastric glands in, 2. 
, Pliocene, of the bone-beds of 
Maragha (R. T. Giinther), 376. 
Manatus, stomach of, 2. 
australis, 3. 
Mangilia, mentioned, 157. 
(Cythara) chionea *, 
Stand., 156, 206. 
)g gracilis, Reeve, 156. 
) pulchella, Reeve, 156. 
(Glyphostoma) rugosa, M igh., 156. 
Manicina, mentioned, 995, 308. 
areolata, mentioned, 293. 
Manis, gastric gland in, 2. 
javanica, 3. 
Mantellum arcuata, Sow., 182. 
inflata, Zam., 182. 
Mantodea, 417. 
Maraenobiotus, Mrazek, 99; mentioned, 


Melv. § 


Vejdovskyi, Mrazek, 99, 118. 
Maragha, Bone-beds of, the Pliocene 
Maimalia of (R. T. Giinther), 376. 
ae rginele (Persicula) ovulum, Sow., 

157. 
Marginellide, 157. 
Marsupialia, Gastric Glands of, by 
J. Johnstone, 1-14. 
aR Pentelici, Gaud. et Lart., 377. 
be aril 
Mathilda eurytima, Melv. g§ Stand., 
170. 
Meandrina, 144. 
Megalatractus, 159. 
proboscidiferus, Lam., 158. 
Megapodes, 247. 
Melaraphe mauritiana, Lam., 170. 
Meleagrina anomioides, Reeve, 183. 
margaritifera, Linn., 184. 
tegulata, Reeve, 184. 
Meles canescens, mentioned, 362. 
maraghanus, Kitdl., 377. 
Melitzea aurinia, Rott., 408. 
Melo diadema, Zam., 157. 


571 


Melvill, James Cosmo, and Standen, 
Robert, Report on the Marine Mol- 
lusea obtained during the First Expe- 

‘ dition of Prof. A. C. Haddon to the 
Torres Straits, in 1888-89, 150-206. 

Membranipora fenestrata, Hichw., 434, 
435. 

Menura, 248. 

Meretrix incrassata, J. Sow., 447, 452. 
tncrassata, R. B. Newton, 448. 
persiensis *, &. B. Newton, 447; 
~ mentioned, 448, 431, 432, 452. 

Merops apiaster, mentioned, 363. 

Merria, Gray, mentioned, 171. 

Mesochra Brucei, Richard, mentioned, 
100. 

Mesodesma precisa, Desh., 198. 

Mesodesmatide, 198. 

Meta Menardi, 19. 
nigrohumeralis *, Pickard-Cam- 

; bridge, 16, 18. 

Metanastria trifolii, Schiffermitller, 413. 

, ab. terreni, Herr.-Schdéff, 413. 

Metopa, Boeck, 72. 
neglecta, Hansen, 72, 117. 
pusilla, G. O. Sars, 72, 117. 
sinuata, G. O. Sars, 72, 117. 

Metridia, Boeck, 91. 
armata, Boeck, mentioned, 91. 
hibernica, Brady, mentioned, 91, 

92. 
longa, Lubbock, 91, 117. 

Metridium, mentioned, 303, 304, 308. 

Michael, A. D., Acari of Lake Urmi, 
407. 


| Microsolena, 143. 


Microstella atlantica, Brady & Roberts., 
96. 

Mierctheca, 178. 

Acidalia *, Melv. §& Stand., 177; 
mentioned, 178, 206. 
erenellifera, Ad., mentioned, 177, 
178. 
Minolia glaphyrella, Melv. 
176. 
pudibunda, Fischer, 176. 
vitiliginea, Mke., 176. 
Misophria, Boeck, 92. 
pallida, Boeck, 92, 117. 

Misophriade, 92. 

Mitchell, P. Chalmers, On so-called 
“‘Quintocubitalism” in the wing of 
Birds; with special reference to the 
Columba, and Notes on Anatomy, 
210-236. 

Mitra episcopalis, Linn., 157. 

ferruginea, Lam., 158. 

Greeffei, Cr., 158. 

levizonata, Sow., 158. 

tricolor, Montr., 158. 
(Castellaria) modesta, Reeve, 158. 


§ Stand., 


572 


Mitra (Chrysame) peregra, Reeve, 157. 
(——) rubritincta, Reeve, 158. 
(Pusia) dichroa, Ad. § Reeve, 158. 
(Strigatella) decurtata, Reeve, 158. 
(Turricula) corrugata, Lam., 158. 
(Vulpecula) intermedia, Kéener, 

158. 


Mitrella semiconvexa, Lam., 161. 

Mitridx, 157. 

Moa, 251. 

Modiola arborescens, Chemn., 184. 

australis, Gray, 184. 
cinnamomea, Chemn., 184. 
lignea, Reeve, 184. 
philippinarum, Hanley, 184. 
(Adula) lanigera, Dkr., 184. 

Modiolaria Cumingiana, Dkr., 184. 

Modulidz, 169. 

Modulus obtusalis, Phi/., 169. 

Moerchiella spirata, Sow., 171. 

, var. artensis, Montr., 171. 

, var. deformis, Sow., 171. 

Mollusea, Land and Freshwater, of 
Lake Urmi, by Edgar A. Smith, 391- 
398. 

, Marine Tertiary (Miocene) of Lake 
Urmi, by R. Bullen Newton, 480- 
452. 

——, Report on the Marine, obtained 
during the First Expedition of Prof. 
A. C. Haddon to the Torres Straits, 
in 1888-89, by James Cosmo 
Melvill and Robert Standen, 150- 
206. 

Monia ione, Gray, 181. 

Monoculodes, Stimpson, 72. 

borealis, Boeck, 72, 117. 
latimanus, Goés, 73, 117. 
longicornis, Boeck, 73. 
Schneideri, G. O. Sars, 73, 117. 

Monoculopsis, G. O. Sars, 73. 

longicornis, Boeck, 73, 117. 
Monoculus finmarchicus, Gunner, 90. 
Monodactylus Lamarcki, Gray, 165. 

melanostomus, Swains., 165. 

Monodonta canaliferus, Lam., 175. 

Monoxenia, mentioned, 303. 

Montipora, 131, 135, 139, 144, 145, 
146, 147, 148, 489, 490. 

Montiporine, 143. 

Moraria, mentioned, 99. 

Mormyrus, mentioned, 530-531. 

Moths of Lake Urmi, 411-414. 

Munna, Kroyer, 67. 

Fabricii, Kroyer, 67, 118, 116. 

Kréyeri, Goodszr, 67, 116. 
Munnopsis, M/. Sars, 68. 

typica, M. Sars, 68, 116. 
Murex acanthopterus, Lam., 162. 

brevispina, Lam., 161 

eurypteron, Reeve, 162. 


INDEX, 


Murex triformis, Reeve, 162. 
(Chicoreus) adustus, Lam., 161. 


( ) , var. fuscus, Dhr., 
161. 

(——) axicornis, Zam., 161. 

( ) capucinus, Lam., 161. 

(——) cervicornis, Lam., 162. 

( ) ramosus, Linn., 162. 


(Ocinebra) salmoneus*, Melv. & 
Stand., 162, 206. 

(Pteronotus) saibaiensis*, Mel. g- 
Stand., 161, 206. 

Muricide, 161. 

Musk Deer, 37. 

Musophaga, 251. 

Mustelus, 48, 49, 54. 
vulgaris, 47. 

Mya vulsella, Linn., 184. 

Myide, 199. 

Myodocopa, 61, 89. 

Myology of Anomalurus (F.G. Parsons), 

317-334. 

Myomorpha, 320, 321, 331, 334. 

Myoxide, 11. 

Myoxus avellanarius, 3, 10, 11. 

Myripristis, 525. 

Myside, 482. 

Mysis, Latr., 64; mentioned, 483. 
denticulata, G. M. Thomson, 482. 
oculata, Fabr., 64, 116. 

Mytilicardia crassicostata, Lam., 188. 
muricata, Sow., 188. 
variegata, Brug., 188. 

Mytilide, 184. 

Mytilus horridus, Dkr.. 184. 
Matianus, Borne, 439. 
(Aulacomya) hirsutus, Zam., 184. 

Myurella subulata, Linn., 155. 

Myxine, 541. 

Myxinide, 551, 552. 

Myzopontius, Giesbr., 114. 
pungens, Gzesbr., 114, 118. 


Napopora, Quelch, 143, 144, 149, 491. 

Nares, Note on the External, of the 
Cormorant, by W. P. Pycraft, 207- 
209. 

Narica, Récluz, mentioned, 171. 

Nassa, 160. 

arcularia, Zinn., mentioned, 151. 

fretorum, mentioned, 160. 

monile, Kien., mentioned, 160. 

plebecula, Gow/d, mentioned, 160. 

(Alectryon) fretorum*, Melv. & 

Stand., 159, 206. 

( ) suturalis, Zam., 159. 
(Hima) plebecula, Gould, 160. 
(Niotha) albescens, Dkr., 159. 
(——) cremata, Hinds, 159. 

(——) multicostata, A. Ad., 160. 


INDEX. 


Nassa (Niotha) rotunda, Melv. ¢ Stand., 
160. 

Nassaria suturalis, A. Ad., 159. 

Nasside, 159. 

Natica areolata, Récluz, 172. 
Gaultieriana, Petit, 172. 
unifasciata, Lam., mentioned, 

151. 
(Mamma) aurantia, Lam., 172. 

) Flemingiana, Récluz, 172. 
(Neverita) bicolor, Phzl., 172. 

Naticidee, 172. 

Natural History, Contributions to the, 
of Lake Urmi, N.W. Persia, and its 
Neighbourhood, by R. T. Ginther, 

845-453. 

Nausithoé, mentioned, 305. 

Nebalia, mentioned, 341, 342. 

Nemachilus persa, 364, 391. 

Neoporites, 143, 147, 149, 492. 

Nerita polita, Linn., 173. 

, var. aurora, Dkr., 173. 

signata, Macleay, 174. 

undata, Linn., 174. 

(Peleronta) funiculata, 
174. 

( ) plicata, Linn., 174. 

(Thelicostyla) albicilla, Linn., 
174 


Reeve, 


Neritid, 1753. 

Neritina, mentioned, 353. 

Neuroptera (Hemerobiidz) and Diptera 
of Lake Urmi, by R. T. Ginther, 
414 416. 

Neverita bicolor, Phi/., 172. 

Newton, R. Bullen, Marine Tertiary 
(Miocene) Mollusca of Lake Urmi, 
430-452. 

, Note on a Palxozoic Limestone 
from Lake Urmi, 452-453. 

New Zealand Schizopoda, on some, by 
George M. Thomson, 484-486. 

Niotha albescens, Dér., 159. 

fretorum * , Melv. §& Stand., 159. 
rotunda, Melv. ¢ Stand., 160. 

Noctuide, 411. 

Nodosaria radicula, Zinn., mentioned, 
452. 

Nomophila noctuella, 
414. 

Notodontids, 412. 

Notopterus, 519. 

borneensis, Bleeker, 
508. 
Pallasii, C. ¢ V., mentioned, 503, 
504, 506, 516, 519-522. 
Nubecula striatus, Lznn., 156. 
Nucula obliqua, Lam., 188. 
simplex, 4. Ad., 188. 
Nuculide, 188. 
Nutcrackers, &8. 


Schiffermiiller, 


mentioned, 


LINN. JOURN.—ZOOLOGY, VOL. XXVII. 


573 


Nyctiphanes, G. O. Sars, 485. 
australis, G. O. Sars, 485. 
Nymphalide, 408. 


‘Obeliscus dolabrata, Linn., 173. 


terebelloides, 4. Ad., 173. 
Obtortis, Hedley, ftnote 171. 
pyrrhacme, ftnote 171. 
Ocinebra, mentioned, 162. 
salmoneus*, Melv. § Stand., 162. 
Ocydromus, 250-252. 
Cidalius nigrofasciatus, De Geer, 417. 
Cidiceros latimanus, Goés, 73. 
lynceus, M. Sars, 72. 
Cidipoda Schochii, Br., 417. 
Gidipodide, 371, 417. 
Gina, 212, 213. 
capensis, mentioned, 212. 
Mneis, 39. 
bore, 43. 
sculda, 43. 
Oidemia Stejnegeri, 38. 
Oithona, Baird, 92. 
helgolandica, Claus, mentioned, 
92 


plumifera, Baird, mentioned, 92. 
similis, Claus, 92, 117. 
spinifrons, Boeck, mentioned, 92. 
spinirostris, Claus, mentioned, 92. 
Oleostephanus Straussi, Weithofer, 438. 
tetrameres, Wetthofer, 438. 
(Hoplites?) narbonensis?, Pictet, 
438. 
ele (Strephona) episcopalis, Lam., 
Wc 
Olivancillaria nebulosa, 
Lam., 157. 
Olivide, 157. 
Ompheledsnnn Chemnitzii, Hanley, 
95 


(Agaronia) 


costellifera, Ad., 195. 
embrithes*, Melv. § Stand., 195, 
206. 

Lamarckii, Gray, 196. 

Listeri, Gray, 196. 

marica, Linn., 196. 

subnodulosa, Hanley, 196. 

toreuma, Gould, 196. 

Oneaa, Philippi, 115. 

mediterranea, Claws, 115, 118. 
Oncxade, 115. 

Onesimus, Boeck, 71. 

Edwardsii, Kroyer, 69, 71, 117. 
Oniscus aculeatus, Lepechin, 76. 
Oopecten, 445. 

rotundatus, Zam., 443. 

rotundatus, Sacco, 443. 
Operculata, 61. 

Ophidia, 380. 

Ophiocephalus, 522. 

Ophiops elegans, Ménétr., 361, 378. 
43 


574 


Opisthocomus, 223, 224, 233, 234, 248, 
251, 256. 
Orbicella, 425. 
Defrancei, Milne-Edw. § Haime, 
424, 425, 426, 427. 
Bllisi, Defr., mentioned, 428. 
Guentheri * , Gregory, 426, 
Guettardi, Defrance, mentioned, 
425. 
Haimei, d’ Archiac, 427. 
vesiculosus, Milne-Hdw. & Haime, 
mentioned, 426. 
Orchomenella, G. O. Sars, 69. 
minuta, Kroyer, 69, 116. 
Orneodes sp., 414. 
Orneodide, 414. 
Orthomesus nivosa, Reeve, 175. 
Orthoptera of Lake Urmi, by Malcolm 
Burr, 416-418. 
Orycteropus Gaudryi, Maj., 377. 
Ostariophysex, 534. 
Ostracoda, 61, 81. 
Ostrea, 429. 
digitalina, var. Rholfsi, 440. 
digitata, Hichw., mentioned, 440. 


excavata, Desh., 434, 435, 440, 
441, 

lamellosa, mentioned, 431, 434, 
435, 440. 


pseudodigitalina, mentioned, 431, 
440 


pseudoedulis, Desh., 440. 

Rholfsi, Fuchs, mentioned, 440. 

tubercularis, Lam., 181. 

varia, Linn., 446. 

Virleti, Desh, 
436. 

Virleti, Fuchs, 440, 441. 

(Alectryonia) Virdedi, R. B. Newton, 

441. 


360, 429, 434, 


Ostreide, 181. : 
Ostrich, wing of, 251. 
Ovide, 377. 
Ovis ammon, Linn., 29. 
cycloceros, 375. 
Gmelini, 375. 
ophion, 361, 374. 
, var. urmiana*, A. Giinther, 
374. 
orientalis, 374. 
, var. urmiana, 376. 
Poli, 29. 
steatopygus, mentioned, 362. 
Ovula (Radius) Angasi, A. Ad., 164. 
Owl, wing of, 243. 
Oxyperas Coppingeri, Smith, 199. 


Pachydrilus, 358. 
Pachytrachelus, 418. 

sp., 418. 

striolatus, Fieb., 418. 


INDEX. 


Padollus ovina, Chemn., 179. 
Palzemonetes varians, 358. 
Paleoryx Pallasii, Gaudr., 377. 
Paleozoic Limestone, Note on a, from 
Lake Urmi, by R. Bullen Newton, 
425-458. 
Palamedeide, 234. 
Pandora sp., 202. 
Pandoride, 202. 
Papaver alpinum, 42. 
Paphia glabrata, Desh., 196. 
mitis, Desh., 196. 
Papilio afer, Esper, 408. 
agestis, Schiffermiller, 409. 
althee, Hubner, 411. 
aurinia, Rott., 408. 
cardui, Linn., 408. 
circe, Fabr., 408. 
damon, Schiffermiuller, 409. 
daphnis, Schiffermiller, 409. 
daplidice, Linn., 410. 
edusa, Kabr., 410. 
endymion, Schiffermiller, 409. 
ergane, Hubner, 410. 
hispulla, Esper, 408. 
tcarus, Rott., 409. 
lineola, Ochsenh., 411. 
maia, Cramer, 408. 
oceanus, Bergstrasser, 409. 
pamphilus, Linn., 408. 
ripartii, Preyer, 409. 
sylvanus, Esper, 411. 
Papilionide, 410. 
Papilionine, 410. 
Papyridea papyraceum, Ohemn., 192. 
Paradoxostoma, Fischer, 88. 
flexuosum, G. S. Brady, 89, 
117. 
variabile, Baird, 88, 117. 
Parallelipipedum semitortum, Lam., 
Paramphithoé, Bruzelius, 74. 
bicuspis, Kroyer, 74, 117. 
fragilis, Goés, 77. 
monocuspis, G. O. Sars, '74, 75, 
117. 
pulchella, Kroyer, 74, 117. 
Paraphysa manicata, H. Stmon, 15, 
16. 
Parapleustes, Buchholz, 75. 
glaber, Boeck, 75, 117. 
Parathemisto, Boeck, 69. 
oblivia, Kroyer, 69, 116. 
Pardalisca, Kroyer, 76. 
cuspidata, Kréyer, 76, 117. 
Parembola litterata, Linn., 196. 
radiata, Chemn., 196. 
Parilimya, Melv. § Stand., 202. 
Haddoni*, Melv. & Stand., 202. 
Parnassius delius, 43. 
Eversmanni, 43. 


INDEX. 


Pareediceros, G. O. Sars, 72. 
lynceus, M/. Sars, 72, 117. 
Parsons, F. G., The Position of Anoma- 
lurus as indicated by its Myology, 
317-3934. 
Passeres, 248. 
Pecten, 433. 
aduncus, Hichw., mentioned, 442. 
benedictus, Lam., 434, 435. 
Besseri, Andrejowski, 444. 
Beudanti, Basterot, 441, 451. 
blandus, Reeve, 183. 
Burdigalensis, Lam., 


431, 435, 


convexocostatus, <Abich, 432, 485, | 


436, 442. 
crassicostatus, Sow., 183. 
Crouchi, Smith, 183. 
disciformis, Schiibler, 437, 439. 
flabelliformis, Brocchi, 434, 435; 
mentioned, 444. 
gloria-maris, Dubois, mentioned, 
445. 
Holgeri, Geinitz, 437. 
lemniscatus, Reeve, 183. 
lentiginosus, Reeve, 183. 
limatula, Reeve, 183. 
lychnulus, Fontannes, 436. 
madreporarum, Petit, 183. 
Malvine, Abich, 444, 445. 
Malvinz, Dubois, 437. 
maximus, Linn., 435. 
multiradiatus, Zam., mentioned, 


opercularis, Zin2., mentioned, 445. 

pallium, Zinn., 183. 

placenta, Fuchs, mentioned, 443. 

planicostatus, Abich, 435. 

pusio, Zinn., mentioned, 445. 

rugosus, Lam., mentioned, 443. 

senatorius, Gmel., 183. 

simplex, Michelotti, 434, 435, 437. 

subopercularis, Abich, 437. 

suburmiensis, Abich, 431, 432, 436, 
442. 

Tournali, Serres, 437. 

varius, Linn., 435. 

(Aiquipecten) Malvine, Dubois de 
Montpéreux, 444. 

(Amussiopecten) Burdigalensis, 
Lam., 443. 

(——_) Burdigalensis, R. B. Newton, 
443 


(Chlamys) cuneatus, Reeve, 183. 

(Flabellipecten) sp., 431, 444. 

(Oopecten) rotundatus, Lam., 448, 
451. 


(Pseudamussium) argenteus, Reeve, 
183, 


Pectinidz, 183. 
Pectunculus, mentioned, 187. 


575 


Pectunculus Hoylei *, Melv. § Stand., 
187, 206. 
vitreus, Lam., 187. 

Pedetes, 332, 333, 334. 

Pedicularis comosa, 42. 
foliosa, 42. 
verticillata, 42. 

Pelagia, mentioned, 303. 

Pelecypoda, 181. 

Peleronta funiculata, Reeve, 174. 
plicata, Linn., 174. 

Pelobates fuscus, Lawr., 381. 

Pelodytes punctatus, mentioned, 459. 

Pentatoma, 367. 
baccarum (?), 367, 416. 

Perdix barbata, Padlas, 38. 

Perisphinctes, 437. 
balinensis, Newmayr, 438. 
curvicosta, Oppel, 419, 437, 438. 
cyrus, Borne, 438. 

Lothari, Oppel, 438. 
paneaticus, Noetling, 438. 
poculum, Leckenby, 438. 
polyplocus, Reinecke, 437, 438. 
sp., 438. 

tetrameres, Weithofer, 438. 
Xerxes, Borne, 438. 

Perissodactyla, 377. 

Peristeridz, 215. 

Perna attenuata, Reeve, 184. 
australica, Reeve, 184. 
lentiginosa, Reeve, 184. 

Persicula ovulum, Sow., 157. 

Petalopus declivis, G. O. Sars, 65. 

Petalosarsia, Stebbing, 65. 
declivis, G. O. Sars, 65, 116. 

Petromyzon, 551. 

Planeri, ftnote 551. 
Stonti, ftnote 551. 

Petromyzontidx, 551, 552. 

Phacus, mentioned, 469. 
pyriforme, mentioned, 480. 

Phalacrocorax carbo, mentioned, 207. 

Phalene of Lake Urmi (Sir F. G. 

Hampson), 411-414. 

Phapide, 215. 

Phascolarctus, gastric gland of, 2, 7-8. 

Phascolomys, gastric gland in, 1, 5-7. 

Phasianella (Orthomesus) nivosa, Reeve, 

175. 

Phasianus, 248. 

Phenacolepas, Pi/sbry, mentioned, 179. 
lingua-viverree * , Melv. § Stand. 

179, 206. 

Philomedes, Lidjeb., 89. 
brenda, Baird, 89, 117. 

Phlogeenas, 213, 227. 
cruentata, 215, 220. 
luzonieca, 215. 

Pheenicophaés, 251. 

Pheenicopteres, 247. 


576 
Pholadomya, Sow., 202, mentioned, 153, 
203. 


candida, Sow., mentioned, 203, 204. 
caspica, Agass., mentioned, 203. 
Haddoni, Meek, mentioned, 205, 
206. 
Loveni, Jeffr., mentioned, 204, 205. 
Richardsii, mentioned, 205. 
(Parilimya) Haddoni*, Melv. & 
Stand., 202. 
Pholadomyidz, 202. 
Pholas, mentioned, 204. 
costata, Linn., mentioned, 204. 
Pholcidez, 16, 20. 
Pholcus americanus, Nic., 16, 20. 
Phos scalaroides, 4. Ad., 159. 
Phosinella clathrata, A. Ad., 171. 
Photis, Kroyer, 80. 
tenuicornis, G. O. Sars, 80, 117. 
Phrynocephalus, 370. 
helioscopus, Pall., 378. 
persicus, 378. 
, var. Horvathi, L. v. Méhely, 
378. 
Phryxotrichus roseus (Walck.) ?, 15, 
16 


Phyllangia alveolaris, Catudlo, 436. 
grandis, Reuss, 436. 

Phylloccenia, 429. 

Archiaci, Milne-Hdw. § Haime, 
428, 434, 435. 

irradians, Milne-Hdw. & Haime, 
mentioned, 429. 

Phyllopoda, 395. 

Pici, 248. 

Pickard-Cambridge, F. O., On some 
Spiders from Chili and Peru col- 
lected by Dr. Plate of Berlin, 15-22. 

Pierinz, 410. 

Pigeon, wing of, 243. 

Pigeons, 248. 

Anatomical Reasons for the suppo- 
sition that Hutaxic Pigeons are 
not Primitive, 215. 
Systematic Position of the Eutaxic 
Pigeons, 214. 
Pinna fumata, Hanley, 184. 
(Atrina) nigra, Chemn., 184. 
Pinus Cembra, 38, 44. 
sylvestris, 43, 45. 
Pipa, 454-462. 
Pitar, Romer, mentioned, 194. 
regularis, Smith, 194. 

Pitta, 248. 

Placuna lobata, Sow., 181. 

Placunanomia (Monia) ione, Gray, 
181. 

Plagusia, mentioned, 543. 

Planaxide, 169. 

Planaxis sulcatus, Born, 169. 

(Quoyia) decollatus, Quoy, 169. 


INDEX. 


Planorbis marginatus, Draparnaud, 
393. 
Pleopodophora, ftnote 339. 
Pleurocantha spinosissimum, G. O. 
Sars, 67. 
Pleurogonium, G. O. Sars, 67; men- 
tioned, 118. 
inerme, G. O. Sars, 67; mentioned, 
68, 116. 
spinosissimum, G. O. Sars, 67, 
11 


Pleuromya arata, Brauns, 439. 
sp., 437. 
urmiensis, Borne, 439. 
Pleuronectes limanda, mentioned, 95. 
Pleurotomaria sp., 439. 
Plicatula australis, Lam., 181. 
imbricata, Mke., 182. 

Pliocene Mammalia of the Bone-Beds of 
Maragha (R. T. Ginther), 376-878. 
Pocock, R. I., Chilopoda and Arach- 

nida of Lake Urmi, 399-406. 
Podascon, Giard § Bonnier, 68. 
Stebbingi, Giard & Bonnier, 68, 
116. 
Podocopa, 61, 82. 
Podophthalma, 61. 
Peecilostoma, 61. 
Polemonium pulchellum, 42. 
Polycope, G. O. Sars, 89. 
orbicularis, G. O. Sars, 89, 117. 
Polyplacophora, 180. 
Polystomella quaterpunctata, Adzch, 
435. 
Polytrema spongiosa, d@’ Ord., mentioned, 
434. 


spongiosa, Phi/., 435. 
Pontocypris, G. O. Sars, 83 ; mentioned, 
84 


hyperborea *, 7. Scott, 83, 117. 

Porastra, 141. 

Porites, On the Structure of, with Pre- 
liminary Notes on the Soft Parts, by 
Henry M. Bernard, 487-502. 

Porites, 143, 146-149, 429, 430. 

astreoides, Lam., 147, 492, 500. 

clavaria, Leswewr (non Lam.), men 
tioned, 501. 

crassa, Quelch, 491. 

dendroidea, Abich, mentioned, 434. 

exilis, Gardiner, 503. 

furcata, Agass., mentioned, 501. 

latistellata, Quelch, 491. 

leiophylla, Reuss, 430, 435. 

levis, Dana, mentioned, 501. 

mirabilis, Quelch, 491. 

polymorpha, <Adich, ftnote 429, 
435, 436. 

punctata, Hhrenb., 497. 

recta, Lesueur, mentioned, 499, 501. 

recta (?) from Jamaica, 499. 


INDEX. 


Poritidz, Recent, and the Position of 
the Family in the Madreporarian 


System, by Henry M. Bernard, 127- 
149. 


Poritidz, mentioned, 489. 
Poritine, 143. 
Potamides (Terebralia) sulcatus, Born, 
169. 
(Tympanotonus) palustris, Linn., 
169. 


( ) retiferus, Sow., 169. 
Potamilla, 259. 

Priacauthus, ftnote 525, 536. 
arenatus, Cuv. f Val., 525. 
cruentatus, Cuv. § Val., 525. 
macrophthalmus, Cuv. § Val., 

525. 

Primula nivalis, Pad/., 42. 

Parryi, A. Gray, 42. 
Prionastrea irregularis, Defrance, 428. 
Proboscidea, 377. 

Procardia, mentioned, 204. 

Prostrepsiceros (?) sp., 377. 

Protarza, 143. 

Protochordata, 257. 

Protoryx Gaudryi, Mqaj., 377. 
longiceps, Maj., 377. 

Psammobia rasilis *, Melv. § Stand., 

197, 206. 

(Gari) anomala, Desh., 198. 

( ) marmorea, Desh., 198. 

(——) ornata, Desh., 198. 

(——.) prestans, Desh., 198. 

( ) pulcherrima, Desh., 198. 
Psammobiids, 197. 

Psammocora, Dana, 145, 147. 

Pseudalibrotus, Della Valle, 70. 
littoralis, Kroyer, 70, 116. 

Pseudamussium argenteus, Reeve, 183. 

Pseudocalanus, Boeck, 91. 
elongatus, Boeck, 91. 

Pseudocythere, G. O. Sars, 88. 
caudata, G. O. Sars, 88, 117. 

Pseudophycis bacchus, 525, 528, 529 ; 

air-bladders and auditory organ in, 
529. 

Pseudotanais, G. O. Sars, 66. 
forcipatus, Lilijeb., 66, 116. 

Psittaci, 247, 250. 

Psophia, 234, 248, 251, 253, 254. 

Psychide, 413. 

Ptarmigan, 38. 

Pterocera (Heptadactylus) lambis, 

Innn., 166. 

Pterocletes, 247. 

Pteronotus, mentioned, 162. 
acanthopterus, Lam., mentioned, 
eurypteron, Reeve, 162. 
saibaiensis * , Melv. ¢ Stand., 161. 
triformis, Reeve, mentioned, 162. 


577 


Pteropoda, 154. 

Ptychoptera paludosa, larvee of, 356. 

Pupa granum, Draparnaud, 393. 

signata, Mousson, 393. 
Purpura alveolata, Reeve, 162. 
persica, Linn., 162. 
(Cronia) amygdala, Kiener, 163. 
(Stramonita) rustica, Lam., 163. 
(Thalessa) hippocastanum, Lam., 
Pusio dichroa, Ad. & Reeve, 158. 
luculenta, Reeve, 158. 

Pycraft, W. P., Note on the External 
Nares in the Cormorant, 207-209. 

, Some Facts concerning the so- 
ealled ‘“ Aquintocubitalism” in the 
Bird’s Wing, 236-256. 

Pygmiea fulgurans, Lam., 160. 

, a. eufulgurans, 160. 

, B. punctata, Lam., 160. 

Tyleri, Gray, 161. 
versicolor, Sow., 161. 

Pygopodes, 247. 

Pyralide, 413. 

Pyralis farinalis, Zinn., 414. 

Pyrameis cardui, Linn., 408. 

Pyramidellidx, 173. 

Pyrausta aurata, Scop., 414. 

cespitalis, Schiffermiiller, 414. 

Pyrethrum pulchellum, 42. 

Pyrgodera cristata, F. de W., 417. 

Pyrgomorpha grylloides, Latr., 417. 

-—., var. nov. Guentheri * , Burr, 
417. 
Pyrgomorphide, 417. 
Pyrula cingulata (Bronn MS.), Hornes, 
432, 449, 450, 452. 
clathrata, Lam., 449. 
condita, Brong., 449. 


Quail, 38. 

“ Quintocubitalism,” On so-called, in 
the Wing of Birds; with special 
reference to the Columbz, and Notes 
on Anatomy, by P. Chalmers 
Mitchell, 210-236. 

Quoyia decollatus, Quoy, 169. 


Radius Angasi, A. Ad., 164. 
Raeta Grayi, H. Adams, 199. 
Rajide, 52. 
Ralli, 247. 
Rallus, 250. 
Rana Camerani, Bouleng., 381. 
esculenta, Linn., 380. 
, var. ridibunda, Pall., 380. 
Red Deer, 29. 
Reindeer, 37. 
Reineckia sp., 439. 
Straussi, Weithofer, 439. 


578 


Reptilia and Amphibia of Lake Urmi, 
by G. A. Boulenger, 378-381. 

Rhabdite - “cells” in Cephalodiscus 
dodecalophus, McIntosh, On the 
Discovery aud Development of, by 
F. J. Cole, 256-268. 

Rhabdopleura, 258. 

Rhachotropis; S. Smith, 76, 77. 

aculeata, Lepechin, 76, 117. 

Rhea, wing of, 251. 

Rheum Rhaponticum, 43. 

Rhina, 48, 49. 

Rhinoceros sp., 377. 

Rhinocerotidx, 377. 

Rhinochetus, 284, 248, 254. 


Rhinolophus hipposideros, Bechst., 
370. 
Rhipicephalus sanguineus, Latr., 370, 
407. 


simus, C, LZ. Koch, 370, 407. 
Rhizoconos read Rhizoconus, 156. 
BRhizoconus mustelinus, Hwass, 156. 

vitulinus, Hwass, 156. 

Rhizothrix curvata, Brady & Roberts., 
104. 

Rhodactis, mentioned, 280, 295, 308. 

Sancti-Thomz, mentioned, 293, 

309. 

Rhodarea, 143, 144, 149. 

Rhodeus, 382. 

Rhodostrophia calabraria, Zeller, 413. 
inconspicua, Butler, 413. 

Ridewood, W. G., Some observations on 
the Caudal Diplospondyly of Sharks, 
46-59. 

——, On the Hyobranchial Skeleton 
and Larynx of the new Aglossal 
Toad, Hymenochirus Boettgeri, 454— 
460. 

——., Note on the Carpus of the new 
Aglossal Toad, Hymenochirus Boett- 
geri, 461-463. 

Rimula exquisita, 4. Ad., 179. 

Rissoide, 171. 

Rissoina scolopax, Sow., 171. 
thaumasia, Melv. § Stand., 171. 
triangularis, Wats., 171. 
(Moerchiella) spirata, Sow., 171. 


(——) ——, var. artensis, Montr., 
171. 

(—) , var. deformis, Sow., 
171. 

(Phosinella) clathrata, A. Ad., 
171. 


Robertsonia, Brady, 96. 
tenuis, Brady & Roberts., 96, 
118. 
Rodentia, stomach in, 2. 
Roe, 37. 
Rotifers, eyes of, 477. 
Rozinante, Stebbing, 77. 


INDEX. 


Rozinante fragilis, Goés, 77, 117. 
Rubrius, #. Simon, 21. 
annulatus*,  Pickard-Cambridge, 
16, 20. 
subfasciatus, Stm., 21. 
Rugosa, 302. 


Sabella, 259. 

Samotherium Boissieri, a7., 377. 

Sarcinula astroites, Goldj/., mentioned, 
428. 

Sargus, 534-537. 

Rondeletii, Cuv. §& Val., 525. 
Saturnia pyri, Schiffermiiller, 412. 
Saturniade, 412. 

Satyrine, 408. 

Satyrus lupinus, Costa, 408. 
pelopea, Klug, 408. 
turanica, Stgr., 408. 

Saxifraga oppositifolia, 42. 
umbrosa, 43. 

Scalaria lyra, Sow., 172. 

obliqua, Sow., 172. 

subauriculata, Sow., 172. 
Scalariidze, 172. 

Sealiola elata, Semper, 178. 

Scaphandride, 155. 

Scaphella Turneri, Gray, 157. 

Scaphiodon gracilis, Keyserl., 383. 

Sieboldii, Steind., 383, 385. 
Seaphopoda, 181. 

Schizochiton incisus, Sow., 181. 

Schizopoda, 61, 64. 

, On some New Zealand, by George 

M. Thomson, 482-486. 

Schizothorax, mentioned, 386. 

Sciznide, 524. 

Scincus cyprius, Cuwv., 379. 

Seintilla Alberti, Smith, 189. 

hyalina, Desh., 189. 
Sciuromorpha, 320, 331, 334. 
Sclerochilus, G. O. Sars, 88. 

contortus, Norm., 88, 117. 
Sclerocrangon boreas, Phipps, 64, 116. 
Scolopendra, Linn., 400. 

affinis, Newp., 400. 

canidens, Newp., 400. 

dalmatica, Koch, 400. 

oraniensis, Lucas, 400. 

Scomber scombrus, Linn., mentioned, 

542. 

Scombride, 522. 

Scoparia cembrae, Haworth, 414. 

Scopus, 250. 

Scorpena, mentioned, 547. 

Scorpenide, 542. 

Scorpiones, 404. 

Seoter Duck, 38. - 

Scott, Thos., Report on the Marine and 

Freshwater Crustacea from Frauz- 


INDEN. 


Josef Land, collected by Mr. William 
S. Bruce, of the Jackson-Harmsworth 
Expedition, 60-126. 

Scrobiculariide, 201. 

Scutella gibbosa, Hésso, ftnote 421. 

Scutellidium, Claus, 113. 
tisboides, Claus, 113, 118. 

Scutellina, Gray, mentioned, 179. 

Scutigera, Latr., 399. 
coleoptrata, Linn., 399. 

Scutus uneuis, Linn., 179. 

Scyllium, 47, 48, 49, 55. 
canicula, 47. 
catulus, 47, 49. 
stellare, 47. 

Scymnus, 48, 49, 54. 
lichia, 47, 48. 

Scyphistome, mentioned, 301. 

Scyphomedusz, mentioned, 301. 

Scyphostoma, mentioned, 301. 

Seyphozoa, larva of, 301. 

Scytodide, 15, 16. 

Selachii, 550. 

Semele duplicata, Sow., 201. 

Jukesi, A. Ad., 201. 
lamellosa, Sow., 201. 
Semicassis torquata, Reeve, 164. : 
Semifusus (Megalatractus) proboscidi- 
ferus, Lam., 158. 

Senectus argyrostomus, Linn., 175. 
chrysostomus, Linn., 175. 
foliaceus, Phil., 175. 
sparverius, Gmel., 175. _ 

Separatista Blainvilleana, Petiz, 169. 

Septifer nicobaricus, Chemn., 184. 

Seraphs terebellum, Linn., 166. 

Seriema, 248. 

Serpula, mentioned, 433. 

Sp-5 
Sharks, Some observations on the Caudal 
Diplospondyly of, by W. G. Ridewood, 

46-59 


Sheep, Wild, of the Urmi Islands, by 
Dr. A. Ginther, 374-376. 
Sicarius thomisoides, Waick., 15, 16. 
Siliquaria Cumingii, Morch, 170. 
ponderosa, Mérch, 170. 
Siluridse, 526. 
Siluroids, 522. 
Silurus, 382, 383. 
Chantrei, 383. 
glanis, Linn., 364, 365, 382, 383, 
519. 
triostegus, Heckel, 383. 
Simpulum gemmatus, Reeve, 163. 
pilearis, Linn., 163. 
Siphonaria sipho, Sow., 154. 
Siphonariide, 154. 
Siphonium maximus, Sow., 169. 
Siphonostoma, 61. 
iriella, Dana, 482. 


i 


579 


Siriella denticulata, G.M. Thomson, 482; 
mentioned, 486. 
gracilis, Dana, 482, 483. 
Thompsoni, Milne-Hdw., 482, 483. 
Sistrum arachnoides, Lam., 163. 
cavernosum, Reeve, 163. 
chrysostomum, Desh., 163. 
concatenatum, Lam., 163. 
elatum, Blainv., 163. 
fiscellum, Chemmn., 163. 
heptagonale, Reeve, 163. 
margariticolum, Brod., 163. 
ochrostoma, Blainv., 163. 

Smerinthus populi, Zinn., 412. 

Smith, Edgar A., Land and Freshwater 
Mollusca of Lake Urmi, 391-393. 

Solea, mentioned, 544, 545. 

Solenastrza astroites, Goldf., 436; men- 

tioned, 428. 

columnaris, Reuss, 428. 

turonensis, Michelin, 427; men- 
tioned, 428. 

Solenidze, 198. 

Solenocurtus (Azar) coarctatus, Gimel. 
198. 

Soletellina virescens, Desh., 198. 

Solifugze, 402. 

Sparassus bombilius *, Pickard-Cam- 
bridge, 15, 17. 

Sparide, 522, 523, 524. 

Sparus, 528, 529, 531-536. 

salpa, Linn., 523, 525. 
sargus, Linn., 525. 

Spheerium lacustre, Miller, 393. 

Sphingide, 412. 

Sphingonotus satrapes, Sawss. ?, 417. 

Spiders from Chili and Peru, On 
some, by F. O, Pickard-Cambridge, 
15-22. 

Spinax niger, 47. 

Spinigera, sp , 439. 

Spirontocaris Gaimardii, Milne-Edw., 

63, 116. : 
Phippsii, Kroyer, 63, 116. 
polaris, Sabine, 63, 116. 

Spirostomum, mentioned, 500. 

Spondylide, 181. 

Spondylus barbatus, Reeve, 182. 

bifrons, Goldf., 434, 435. 
foliaceus, Chemn., 182. 
nicobaricus, Chemn., 182. 
ocellatus, Reeve, 182. 
pacificus, Reeve, 182. 

Squalius turcicus, /%dippi, mentioned, 
387. 

Standen, Robert, and James Cosmo 
Melvill, Report on the Marine 
Mollusca obtained during the First 
Expedition of Prof. A. ©. Haddon 
to the Torres Straits, in 1888-89, 
150-206. 


580 


Starnoenas, 213. 
eyanocephala, 213, 215, 226, 
236. 
Steganopodes, 247. 
Stellio caucasicus, 370. 
Stenhelia, Boeck, 97. 
reflexa, 7. Scott, 97, 118. 
Stephanoceras stenostoma, 
438. 
Stomatella Mariei, Cr., 178. 
sulcifera, Lam., 178. 
Stomatiide, 178. 
Stramonita rustica, Zam., 163. 
Strephona episcopalis, Lam., 157. 
Strigatella decurtata, Reeve, 158. 
Striges, 248. 
Stringops, 251. 
‘Strombide, 165. 
Strombus, 431. 
Bonelli, Brong., 431, 448, 449. 
(Canarium) dentatus, Linn., 165. 
(Conomurex) luhuanus, Linn., 
166. 
(Gallinula) Campbelli, Gray, 165. 
) canarium, Linn., 165. 
(——) Isabella, Lam., 165. 
(——) Sibbaldi, Sow., 165. 
( ) ureeus, Linn., 165. 
(——) variabilis, Swazns., 165. 
(——) vittatus, Linn., 165. 
(Monodactylus) Lamarcki, Gray, 
165. 
(——) melanostomus, Swains., 


Borne, 


‘Stylareea, Milne-Edw. § Haime, 497. 
Miilleri, WMilne-Edw., 497. 
punctata, Klunzinger, mentioned, 

497: 

Stylochus, 266. 

Subula muscaria, Lam., 155. 

Suide:, 377. 

Sun-bittern, 248. 

Sus erymanthius, Roth. et Wagn., 

377. 
Synarea, Verrill, mentioned, 1438, 144, 
149, 487, 490, 491. 
crassa, Quelch, 491. 
latistellata, Quelch, 491. 
Synchloé daplidice, Linn, 410. 
Syncoryne, 258. 
Syntomide, 411. 
Syntomis persica, Kol/., 411. 
Syrian Names of Animals living in the 
Urmi Basin, 371-374. 
Syrnium aluco, wing of, 243. 
Syrrhoé, Goés, 75. 
erenulata, Goés, 75, 76, 117. 


‘Teniolate, 301. 
Tanais forcipatus, Lilljeb., 66. 


INDEX. 


Tanais longiremis, Lilljeb., 66. 
Tapes Deshayesii, Han/ey, 196. 
malabarica, Sow., 196. 


(Parembola) litterata, Linn., 
196. 
( ) radiata, Chemn., 196. 


(Textrix) sulcosa, Phil., 196. 

) textrix, Chemn., 196. 

Tarache luctuosa, Schiffermiiller, 412. 

sulphuralis, Zinn., 412. 

Tectarius malaccanus, Phil., 170. 

Tectus fenestratus, Guel., 175. 

Teinostoma, Ad., mentioned, 178. 

Teinotis asinina, Linn., 179. 

Teleostean Fishes, 526. 

Teleostei, On the Presence of Nasal 
Secretory Sacs and a Naso-pharyngeal 
Communication in, with especial 
reference to Cynoglossus semilzvis, 
Giinther, by H. M. Kyle, 541-555. 

Telestes leucoides, Fidéppi, mentioned, 
389. 

Tellimya ephippiolum*, Melv. & 

Stand., 189, 206. 
triangularis, Gould, 189. 
Tellina (Angulus) philippinarum, Han- 
ley, 201. 
(—— procrita*, Meiv. & Stand., 
201, 206. 
(——) vernalis, Hanley, 201. 
(Arcopagia) pinguis, Hanley, 201. 
) Savignyi, A. Ad., 201. 
( ) tessellata, Desh., 201. 
(Donacilla) rhomboides, 
201. 
( ) semitorta, Sow., 201. 
(——) virgulata, Hanley, 201. 
(Tellinella) asperrima, Hanley, 
200. 
( ) iridescens, Bens., 200. 
( ) staurella, Zam., 200. 
(—-) virgata, Linn., 200. 
(——) vulsella, Chemn., 201. 
(Tellinides) emarginata, Sow., 
201. 

Tellinella asperrima, Hanley, 200. 
iridescens, Bens., 200. 
staurella, Zam., 200. 
virgata, Linn., 200. 
vulsella, Chemn., 201. 

Tellinidse, 200. 

Tellinides emarginata, Soe., 201. 

Telphusa fluviatilis, 366, 374. 

Tenagomysis, G. M. Thomson * , 483. 

nove-zealandix, G. M. Thomson * , 
484 ; mentioned, 486. 

Terebellum subulatum, Zinz., 
tioned, 151. 

Terebra (Abretia) affinis, Gray, 155. 

(Myurella) subulata, Linn., 155. 
(Subula) muscaria, Zam., 155. 


Gimel., 


men- 


INDEX. 


Terebralia sulcatus, Born, 169. 
Terebride, 155. 
Teredinide, 199. 
Teredo nucivora, Speng., 199. 
Testudo ibera, mentioned, 363, 407. 
Tetracorallia, 302. 
Tetragnatha, Latr., 400. 
extensa, Linn., 400. 
Tetraogallus altaicus, 38. 
urogalloides, 38. 
Teutana grossa, C. Koch, 15, 16, 20. 
Textile canonicus, Hwass, 156. 
sulcosa, Phil., 196. 
textrix, Chemn., 196. 
Thais Cerisyi, Godart, 410. 
Thalessa hippocastanum, Lam., 162. 
Thalestris, Claus, 106 ; mentioned, 110. 
forficula, Claus, 108, 118. 
forficuloides, T. § A. Scott, men- 
tioned, 108. 
frigida *, ZT. Scott, 108, 118. 
helgolandica, Claws, 106, 118. 
hibernica, Brady § Roberts., men- 
tioned, 107, 108. 
Jacksoni*, Z. Scott, 109; men- 
tioned, 110, 118. 
mysis, Claus, mentioned, 107, 109, 
110. 
polaris * , Z. Scot, 106. 
rufoviolascens, mentioned, 110. 
Thamnarexa, Thurm. &§ Héallon, men- 
tioned, 429. 
polymorpha, Adich, 424, 429, 450. 
Thelicostyla albicilla, Linn., 174. 
Theraphosidee, 15, 16, 17. 
Theridiide, 16, 20. 
Theridion grossum, C. Koch, 20. 
tepidariorum, C. Koch, 16, 20. 
Thomisotdes terrosa, Nic., 16. 
Thomson, George M., On some New 
Zealand Schizopoda, 482-486. 
Thoracica, 61. 
Thorellia, Boeck, 93. 
brunnea, Boeck, 93, 117. 
Thylacodes unovee-hollandiz, 
170. 
Thyrostraca, 60, 61. 
Vhysanoessa, Brandt, 64. 
neglecta, Kroyer, 64, 116. 
Thysanopoda neglecta, Kroyer, 64. 
Vichopora, Quelch, 148, 144, 149. 
Tinamus, 24, 
Tineidee, 414. 
Tmeticus Defoei * , Pickard-Cambridge, 
NGy, 1), 
Platei *, 
1©). 
Toad, Aglossal, 454-462. 
Tonicia confossa, Gould, 180. 
fortilirata, Reeve ?, 180. 
Tornatina gracilis, 4. Ad., 155. 


Rouss., 


Pickard-Cambridge, 16, 


581 


Tornatinide, 155. 

Torres Straits, Report on the Marine 
Mollusca obtained during the First 
Expedition of Prof. A. 0. Haddon to 
the, in 1888-89, by Jas. Cosmo Mel- 
vill and Robert Standen, 150-206. 

Trachycardium dianthinum~* , Melv. § 

Stand., 190, 206. 
elongatum, Brug., 190. 
lacunosum, /ceve, 190. 
maculosum, Wood, 191. 
rubicundum, Reeve, 191. 
rugosum, Lam., 191. 
serricostatum *, Melv. & Stand., 
191, 206. 
transcendens*, Melv. 4 Stand., 
191, 206. 
variegatum, Sow., 192. 

Tragoceros amaltheus, Gaudr., 377. 

Treronidee, 215. 

Triacanthus, Stannius, apparently a 
misprint for Priacanthus, ftnote 525. 

Trichotropidide, 169. 

Tridacna elongata, Lam., 189. 

serrifera, Lam., 190. 
squamosa, Lam., 190. 
Tridacnide, 189. 
Triforis, mentioned, 166. 
gigas, Hinds, 166. 
(Ino) excelsior * , Melv. §& Stand., 
166, 206. 
Trigonia Roxane, Borne, 439. 
sp., 439, 
uniophora, Gray, 188. 

Trigoniide, 188. 

Tripoblastica, 306. 

Tritonide, 163. 

Tritropis, 77. 

““Triturationsorgan,” 3. 

Trivia oryza, Lam., 165. 

staphylea, Linn., 163. 

Trochide, 175. 

Trochosa, 22. 

Trochus (Lamprostoma) 

Linn., 175. 
(Tectus) fenestratus, Gmel., 175. 

Trogones, 248. 

Trombidiide, 407. 

Trombidium sericeum, 407. 

sericeum locustarum, Riley, 407. 

Tropidonotus anoscopus, 380 

cyclopium, 380. 
ferox, 380. 
tessellatus, Laur., 380. 

Trumpeter, 248. 

Truxalide, 417. 

Truxalis unguiculata, Ramb., 417. 

Tubinares, 247. 

Turacus, 251. 

Turbinaria, ftnote 130, 148, 495, 496, 
499. 


maculatus, 


LINN. JOURN.— ZOOLOGY, VOL. XXVIII. 4A, 


582 


Turbinellidze, 158. 

Turbinidee, 175. 

Turbo marmoratus, Linn., 175. 
petholatus, Linn., 175. 


(Senectus) argyrostomus, Linn., 
175. 
( ) chrysostomus, Linn., 175. 


(——) foliaceus, Phil., 178. 

( ) sparverius, Gmel., 175. 
Turnices, 248. 
Turricula corrugata, Lam., 158. 
Turritella Archimedis, Brong., men- 


tioned, 360, 4381, 457, 460, 
452. 

eradata, Menke, mentioned, 360, 
437. 


——, Hornes, 451, 452. 

——, var., Abich, 431, 451, 452. 

leptomita, Melv. & Sykes, men- 
tioned, 172. 

margarita, Abich, 437. 

multilirata, Ad. g Reeve, 170. 

rotifera, Desh., 481, 437, 451, 


452. 
turris, d@ Orb., mentioned, 360, 
431, 437. 


Turritellide, 170. 
Turtur, 226. 
chinensis, mentioned, 211, 236. 
Tympanotonus palustris, Linn., 169. 
retiferus, Sow., 169. 
Typhlotanais, G. O. Sars, 65. 
finmarchicus, G. O. Sars, 65, 116. 
Tyrannus, 248. 


Ulochlena hirta, Hiibner, 412. 
Umbonium vestiarium, Linn., 175. 
Ungulinide, 197. 
Unio batavus, 393. 

Sieversi, Drouet, 393. 

——, var. Kobelti, 393. 


Valvulina bulloides, Brady, mentioned, 
452. 
Vanellus cristatus, 255. 
——, wing of, 242. 

Vanikoride, 171. 

Vanikoro, Quoy, mentioned, 153, 174. 
cancellata, Lam., 171. 
Gueriniana, Récluz, 

174. 
Veneridz, 193. 
Venus Aglauree, Brong., 432, 436, 446, 
447, 448, 452. 
Aglaure, Hornes, 446. 
granosa, J. de C. Sowerb., men- 
tioned, 447. 
incrassata, J. Sowerb., 447. 


mentioned, 


INDEX. 


Venusta lactea, Linn., 185. 

Vermetide, 169. 

Vertagus aluco, Linn., 167. 
lineatum, Lam., 167. 
pulchrum, A. Ad., 168. 
Sowerbyi, Kiener, 168. 
vertagus, Linn., 168. — 
(Bivonia) Quoyi, 4. § A. Ad., 

70. 
(Siphonium) maximus, Sow., 169. 
(Thylacodes) novee-hollandiz, 
Reuter, 170. 
Vertebrates, air-breathing, 547. 
Viatrix globulifera, Duchass., men- 
tioned, 270. 

Voluta nivosa, Linn., mentioned, 157. 
Norrisi, Gray, mentioned, 157. 
piperita, Sow., mentioned, 157. 
Sophia, mentioned, 152. 

(Aulica) Ruckeri, Crosse, 157. 
(——) rutila, Brod., 157. 
(——) Sophia, Gray, 157. 
(Scaphella) Turneri, Gray, 157. 

Volutida, 157. 

Vulpecula intermedia, Kiener, 158. 

Vulsella lingulata, Lam., 184. 


Wager, Harold, On the Eye-spot and 
Flagellum in Huglena viridis, 463— 
481. 

Water-Rail, Common, 251. 

Westwoodia, Dana, 111. 

nobilis, Baird, 111, 118. 

Wild Sheep of the Urmi Islands, by 
Dr. A. Gunther, 374-376. 

Wing, embryo of, 238. 

probable origin of the Diastataxie, 
252. 
Wolf-fish, 542. 
Wombat, Anatomy of, 1. 


Xanthia ocellaris, Bork., 412. 
Xanthoptera triangularis, Warr., 412. 
Xenopus, 454-462. 
Boettgeri, 454. 
Xestoleberis, G. O. Sars, 86. 
depressa, G. O. Sars, 86, 117. 


Yponomeuta padellus, Linz., 
416 


367, 


Zaus, Goodsir, 112. 
spinatus, Goodsir, 112, 118. 
Zebrinus Hohenackeri, Krynicki, 392. 


INDEX. 083. 


Zoantharia, mentioned, 280, 301. | Zosima spinulosa, Brady & Roberts., 
larva of, 309, 310. 97. 
Zoanthidex, 296, 297. | Zosime, Boeck, 96. 
Zoarees, 555. he typica, Boeck, 96, 117. 
Zoology and Botany of the Altai | Zygena, 49. 
Mountains, on the, by H. J. Elwes, | Cuvieri, Boisd., 413. 
20-46. | Zygenide, 413. 


END OF THE TWENTY-SEVENTH VOLUME. 


PRINTED BY TAYLOR AND FRANCIS, RED LION COURT, FLEET STREET. 


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1639776 
APRIL 1. ass 


THE JOURNAL 


' OF 


THE LINNEAN SOCIETY. 


Vou. XXVIL ZOOLOGY. No. 173. 


CONTENTS. 
Page 


I. On the Gastric Giands of the Marsupialia. By Jamus 
JouNstone, Fisheries-Assistant, University College,, 
Liverpool. (Communicated by Prof. G. B. Howzs, 
Eero; Oeceinoae Ce late: L.) si... ccsseo nates: oneatee seamen 1 


II. On some Spiders from Chili and Peru collected by 
Dr. Plate of Berlin. By F. O. Prckarp-CamMeRipeE. 
(Communicated by Prof. G. B. Howes, F.RB.S., Sec.L.8.) 
Gear e Oa) eee Pa ees eel ude (a ean at vues see eee meee 15 


TIT. On the Zoology and Botany of the Altai Mountains. 
By Bod: Buwes, BORS:, F.U:S.0 2. ..see.: sioner 23 


IV. Some Observations on the Caudal Diplospondyly of 
Sharks. By W. G. Ripewoop, D.Se., F.LAS............. 46 


See Notice on last page of Wrapper. 


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IMPORTANT NOTICE. 


Oo—— 


Journal of Zoology. 


Volume XXVIII. commences with the present Part 178. 


The Journal (both Zoological and Botanical) is 
now issued in THREE PARTS PER ANNUM as 
follows :— 


Parr I., containing papers read from November to the middle 


of January, on April Ist. 


Parr II., containing papers read from the middle of January 
to the end of April, on July Ist. 


Parr III., containing papers read in May and June, on 


November Ist. 


A GENERAL INDEX to the first twenty Volumes of the 
Journal (Zoology) is now ready, and may be had on application, 
either in cloth or in sheets for binding. 


A new CATALOGUE of the LIBRARY may be had on applica- 
tion. Price to Fellows, 5s.; to the Public, 10s. 


Juty 1. Price 14s. 
THE JOURNAL 


THE LINNEAN SOCIETY. 


Vou. XXVII. ZOOLOGY. No. 174. 


CONTENTS. 
Page 
I. Report on the Marine and Freshwater Crustacea from 
Franz-Josef Land, collected by Mr. William 8. Bruce, 
of the Jackson-Harmsworth Expedition. By Tuomas 
Scott, F.L.S., Naturalist to the Fishery Board for 
Scotland) CElates 3-95)... . 06.4 era ee 60 


II. Recent Poritide, and the Position of the Family in the 
Madreporarian System. By Henry M. Burnarp, 
MI Mery ES ecehe kt sks cetleconece «Shae sob cake es pheeete 127 


IIT. Report on the Marine Mollusca obtained during the 
First Expedition of Prof. A. C. Haddon to the Torres 
Straits, in 1888-89. By James Cosmo Metviit, ~ 
M.A., I'.L.8., F.Z.S., and Ropert StanpeEn, Assistant-- 
Keeper, Manchester Museum. (Plates 10 &11.) ... 150 

NS 


IV. Note on the External Nares of the Cormorant. By 
NW eR vGRaer ee ACTS. rise ok. cose igen ee ee 207 


See Notice on last page of Wrapper. 


LONDON: 
SOLD AT THE SOCIETY’S APARTMENTS, BURLINGTON HOUSE, 

PICCADILLY, W., 

AND BY 
LONGMANS, GREEN, AND CO., 
AND 
WILLIAMS AND NORGATE. 

1899. 


LINNEAN SOCIETY OF LONDON. 


LIST OF THE OFFICERS AND COUNCIL. 
Elected 24th May, 1899. 


PRESIDENT. 
Albert C. L. G. Ginther, M.A., M.D., F.R.S8. 


VICE-PRESIDENTS. 


©. B. Clarke, M.A., F.R.S. A. D. Michael, F.Z.S., F.R. M.S. 
Trank Crisp, LL.B., B.A. G. M. Murray, F.R.S. 
TREASURER. 


Frank Crisp, LL.B., B.A. 


SECRETARIES. 


B. Daydon Jackson, Esq. | Prof. G. B. Howes, LL.D., F.R.S8. 
COUNCIL. 

C. B. Clarke, M.A., F.R.S. Prof. G. B. Howes, LL.D., F.R.S. 
Frank Crisp, LL.B., B.A. B. Daydon Jackson, Esq. 
Francis Darwin, M.B., F.R.S. A. D. Michael, F.Z.S., F.R.M.S. 
Prof. J. B. Farmer, M.A. H. W. Monckton, F.G.S. 
F. D. Godman, F.R.S. G. M. Murray, F.B.S. 
Henry Groves, Esq. A. B. Rendle, M.A., D.Sc. 


A.C. L. G. Giinther, M.A., M.D., F.R.S.| Howard Saunders, F'.Z.8. 
W. Percy Sladen, F.G.S. 


ASSISTANT SECRETARY. 
James Edmund Harting, F.Z.S. 


LIBRARIAN. CLERK. 
A. W. Kappel. A. R. Hammond. 


LIBRARY OOMMITTEE. 


This consists of nine Fellows (three of whom retire annually) and of the four 
officers ex officio, in all thirteen members. The former are elected annually 
by the Council in June, and serve till the succeeding Anniversary. The 
Committee meet at 4 P.M., at intervals during the Session. The Members for 
1898-99, in addition to the officers, are :— 


E. G. Baker, Esq. A. D. Michael, F.Z.S. 
O. B. Clarke, M.A., F.R.S. H. W. Monckton, F.G.S. 
Prof. J. B. Farmer, M.A. Howard Saunders, F.Z.S8. 


Prof. J. Reynolds Green,D.Sc.,F.R.8.} Roland Trimen, F.R.S8. 
W. B. Hemsley, F.R.S. 


Norr.—The Charter and Bye-Laws of the Society, as amended to 
the 19th March, 1891, may be had on application. 


‘NOLYONOW “M HOVUOH 


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‘6681 ‘WI0g [dy suipus reok 044 doy YUNODDY Ss yauASVAy, 


Journal of Zoology. > 


Volume XXVII. commenced with Part 173. 


The Journal (both Zoological and Botanical) is 
now issued in THREE PARTS PER ANNUM as 
follows :— 


Parr I., containing papers read from November to the middle 
of January, on April Ist. 


Parr II., containing papers read from the middle of January 
to the end of April, on July ist. 


Part III., containing papers read in May and June, on 


November lst. 


AS GENERAL FNDEX to the first twenty Volumes of the 


Journal (Zoology) may be had on application; either in cloth or 
in sheets for binding. 


A new CATALOGUE of the LIBRARY may be had on applica- 
tion. Price to Fellows, 5s.; to the Public, 10s. 


as 


S 0 &: °G e079 


Juxy l. Price 6s. 


THE JOURNAL 


THE LINNEAN SOCIETY. 


Vou. XXVIII. ZOOLOGY. No. 175. 


CONTENTS. 


Page 
I. On go-called ‘“ Quintocubitalism” in the Wing of 


Birds ; with special reference to the Columba, and 
Notes on Anatomy. By P. Cuaumers MircHet., 
NGAGE sei (elateste G43.) ikea) wields. se, 220 


II. Some Facts concerning the so-called ‘‘ Aquintocubit- 
alism’’ in the Bird’s Wing. By W. P. Pycrart, 
1538 PUSH ed Weuitetsyeg L212) Jer Rea eee 236 


III. On the Discovery and Development of Rhabdite- 
“cells” in Cephalodiscus dodecalophus, McIntosh. 
By F. J. Cox, University College, Liverpool. 
(Communicated by Prof. Howss, Sec. Linn. Soc.) 
leita Ea) ae es eee a wees oy) oo jean Har monies 256 


See Notice on last page of Wrapper. 


LONDON: 


SOLD AT THE SOCIETY'S APARTMENTS, BURLINGTON HOUSE, 
PICCADILLY, W.; 


AND BY 
LONGMANS, GREEN, AND CO., 


AND 


. WILLIAMS AND NORGATE. 
1899. 


LINNEAN SOCIETY OF LONDON. 


LIST OF THE OFFICERS AND COUNCIL. 
Elected 24th May, 1899. 


PRESIDENT. 
Albert C. L. G. Giinther, M.A., M.D., F.R.S. 


VICE-PRESIDENTS. 


©. B. Clarke, M.A., F.R.S. A. D. Michael, F.Z.S., E.R. M.S. 
Frank Crisp, LL.B., B.A. G. M. Murray, F.R.S. 
TREASURER. 


Frank Crisp, LL.B., B.A. 


SECRETARIES. 


B. Daydon Jackson, Esq. | Prof. G. B. Howes, LL.D., F.R.S8. 
COUNCIL. 

C. B. Clarke, M.A., F.R.S. Prof. G. B. Howes, LL.D., F.R.S8. 
Frank Crisp, LL.B., B.A. B. Daydon Jackson, Esq. 
Francis Darwin, M.B., F.R.S. A. D. Michael, F.Z.S., F.R.M.S. 
Prof. J. B. Farmer, M.A. H. W. Monckton, F.G.S. 
F. D. Godman, F.R.S. G. M. Murray, F.R.S. 
Henry Groves, Hsq. A. B. Rendle, M.A., D.Sc. 


A.C. L. G. Ginther, M.A., M.D., F.R.S.| Howard Saunders, F.Z.8S. 
W. Percy Sladen, F.G.S. 


ASSISTANT SECRETARY. 
James Edmund Harting, F.Z.S. 


LIBRARIAN. CLERK. 
A. W. Kappel. A. R. Hammond. 


LIBRARY COMMITTEE. 


This consists of nine Fellows (three of whom retire annually) and of the four 


officers ex officio, in all thirteen members. The former are elected annually 


by the Council in June, and serve till the succeeding Anniversary. The 
Committee meet at 4 p.m., at intervals during the Session. The Members for 


1898-99, in addition to the officers, are :— 


H. G. Baker, Esq. A. D. Michael, F.Z.8. 
O. B. Clarke, M.A., F.R.S. H. W. Monckton, F.G.S. 
Prof. J. B. Farmer, M.A. Howard Saunders, F.Z.S8. 


Prof. J. Reynolds Green,D.Sc.,F.R.S.} Roland Trimen, F.R.S. 
W. B. Hemsley, F.R.S. 


Norr.—The Charter and Bye-Laws of the Seciety, as amended to 


the 19th March, 1891, may be had on application. 


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‘6681 ‘uI0g [udy surpue seo 049 JOF JUNODOW 8,AaUASVAa, 


IMPORTANT NOTICHE. 


—_o—-—_ 
Journal of Zoology. 


Volume XXVIII. commenced with Part 173. 


The Journal (both Zoological and Botanical) is 
now issued in THREE PARTS PER ANNUM as 
follows :— 


Parr I., containing papers read from November to the middle 
of January, on April Ist. 


Part II., containing papers read from the middle of January 
to the end of April, on July Ist. 


Part III., containing papers read in May and June, on 


Wovember lst. 


A GENERAL INDEX to the first twenty Volumes of the 
Journal (Zoology) may be had on application, either in cloth or 
in sheets for binding. 


A new CATALOGUE of the LIBRARY may be had on applica- 
tion. Price to Fellows, 5s.; to the Public, 10s. 


iio 
t+ V 


NOVEMBER 1. Price As. 


THE JOURNAL 


‘OF 


THE LINNEAN SOCIETY. 


Vou. XXVII. ZOOLOGY. No. 176. 
CONTENTS. 
Page 


I. The Hdwardsia-stage of Lebrunia, and the Formation 
of the Gastro-celomic Cavity. By J. EH. Dunrpey, 
Assoc. Roy. Coll. Sci. (London), Curator of the 
Museum of the Institute of Jamaica. (Communi- 
cated by Prof. Howxs, Sec. Linn. Soc.) (Plates 
TUS preter she oats Aton at ot Gey A Nee aR em AR RSE 269 
IJ. The Position of Anomalurus as indicated by its 
Myology. By F. G. Parsons, F.R.C.S., F.L.S., 
Lecturer on Comparative Anatomy at St. Thomas’s 
Hospital and Hunterian Professor at the Royal 
College of Surgeonsyss 35 tes acres occ s oko eh cczenessscee 317 
III. On some Australasian Collembola. By the Rt. Hon. Sir 
Joun Luppock, Bart., M.P., F.R.S., LL.D., F.LS... 384 
IV. On the Characters of the Crustacean Genus Bathynella, 
Vejdovsky. By W. T. Cauman, B.Sc., Univ. Coll. 
Dundee. (Communicated by Prof. D’Arcy W. 
Tompson, C.B., F.L.8.) (Plate 20.) .................. 338 


See Notice on last page of Wrapper. 


LONDON: 
SOLD AT THE SOCIETY'S APARTMENTS, BURLINGTON HOUSE, 
PICCADILLY, W., 
AND BY 
LONGMANS, GREEN, AND CoO., 
AND 
WILLIAMS AND NORGATE. 


1899. 


LINNEAN SOCIETY OF LONDON. 


LIST OF THE OFFICERS AND COUNCIL. 
Elected 24th May, 1899. 


PRESIDENT. 
Albert C. L. G. Giinther, M.A., M.D., F.R.S. 


VICE-PRESIDENTS. 


C. B. Clarke, M.A., F.R.S. A. D. Michael, F.Z.S., F.R.M.S. 
rank Crisp, LL.B., B.A. G. R. M. Murray, F.R.S. 
TREASURER. 


Frank Crisp, LL.B., B.A. 


SECRETARIES. 


B. Daydon Jackson, Esq. | Prof. G. B. Howes, LL.D., F.R.S. 
COUNCIL. 

C. B. Clarke, M.A., F.R.S. Prof. G. B. Howes, LL.D., F.R.S. 
Frank Orisp, LL.B., B.A. B. Daydon Jackson, Esq. 
Francis Darwin, M.B., F.R.S. A. D. Michael, F.Z.S., F.R.MS. 
Prof. J. B. Farmer, M.A. H. W. Monckton, F.G.S. 
F. D. Godman, F-.R.S. G. R. M. Murray, F-.B.S. 
Henry Groves, Esq. A. B. Rendle, M.A., D.Sc. 


A.C. L. G. Ginther, M.A., M.D., F.R.S.| Howard Saunders, F.Z.S. 
W. Percy Sladen, F.G.S. 


ASSISTANT SECRETARY. 
James Edmund Harting, F.Z.8. 


LIBRARIAN. CLERK. 
A. W. Kappel. A. R. Hammond. 


LIBRARY COMMITTHE. as 


This consists of nine Fellows (three of whom retire annually) and of the four 
officers ex officio, in all thirteen members. The former are elected annually 
by the Council in June, and serve till the succeeding Anniversary. The 
Committee meet at 4 P.m., at intervals during the Session. The Members for 
1898-99, in addition to the officers, are :— 


HE. G. Baker, Esq. A. D. Michael, F.Z.S. 
O. B. Clarke, M.A., F.R.S. H. W. Monckton, F.G.S. 
Prof. J. B. Farmer, M.A. Howard Saunders, F.Z.S. 


Prof. J. Reynolds Green,D.Sc.,F.R.S.} Roland Trimen, F.R.S. 
W. B. Hemsley, F.R.S. 


Norr.—The Charter and Bye-Laws of the Society, as amended to 
the 19th March, 1891, may be had on application. 


LMONOW M HOVAOH 


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NOTICE. 


le} 


Journal of Zoology. 


The Journal (both Zoological and Botanical) is 
now issued in THREE PARTS PER ANNUM as 
follows :— 


Part I., containing papers read from November to the middle 
of January, on April Ist. 


Part II., containing papers read from the middle of January 
to the end of April, on July Ist. 


Part III., containing papers read in May and June, on 
November Ist. 


Volume SX XVII. commenced with Part 173. Owing to 
the amount of material accepted for publication it has been 
found necessary to duplicate Parts II. & III. The Volume will 
therefore consist of 5 parts, Nos. 173-177. 


A GENERAL INDEX to the first twenty Volumes of the 
Journal (Zoology) may be had on application, either in cloth or 
in sheets for binding. 


A new CATALOGUE of the LIBRARY may be had on applica- 
tion. Price to Fellows, 5s.; to the Public, 10s. 


DecemBeEr 30. Price }2s. 


THE JOURNAL 


OF 


THE LINNEAN SOCIEEY. 


Vou. XX VII. ZOOLOGY. No. 177. 
CONTENTS. 
Page 


Contributions to the Natural History of Lake Urmi, 
N.W. Persia, and its Neighbourhood. By Rozerr 
T. Ginrner, M.A., ¥.R.G.8., Fellow of Magdalen 
College, Oxford. (Communicated by the President.) 345 


(With Map and 9 Plates.) 


See Notice on last page of Wrapper. 


LONDON: 
SOLD AT THE SOCIETY'S APARTMENTS, BURLINGTON HOUSH, 
PICCADILLY, W., 
AND BY 
LONGMANS, GREEN, AND CO., 
AH D 
WILLIAMS AND NORGATE. __ 


1899. 


LINNEAN SOCIETY OF LONDON. 


LIST OF THE OFFICERS AND COUNCIL. 
Hlected 24th May, 1899. 


PRESIDENT. 
Albert C. L. G. Gimther, M.A., M.D., F.R.S. 


VICE-PRESIDENTS. 


©. B. Clarke, M.A., F.R.S. A. D. Michael, F.Z.8., F.R.M.S. 
Frank Crisp, LL.B., B.A. | G. R. M. Murray, F.RB.S. 
TREASURER. 


Frank Crisp, LL.B., B.A. 


SECRETARIES. 
B. Daydon Jackson, Esq. | Prof. G. B. Howes, LL.D., F.B.S. 
COUNCIL. 

©. B. Clarke, M.A., F.B.S. | Prof. G. B. Howes, LL.D., F.R.S. 
Frank Crisp, LL.B., B.A. | B. Daydon Jackson, Esq. 

Francis Darwin, M.B., F.B.S. | A. D. Michael, F.Z.S., F.R.M.S. 
Prof. J. B. Farmer, M.A. | H. W. Monckton, F.G.S. 

F. D. Godman, F.RB.S. | G. R. M. Murray, F.R.S. 

| 


Henry Groves, Hsq. A. B. Rendle, M.A., D.Se. 
A.C. L.G Ginther, M.A., M.D., F.R.S.| Howard Saunders, F.Z.S, 
W. Percy Sladen, I°.G.S. 


ASSISTANT SECRETARY. 
James Edmund Harting, F.Z.S8. 


LIBRARIAN. CLERK, 
A. W. Kappel. A. R. Hammond. 


LIBRARY COMMITTEE. 


This consists of nine Fellows (three of whom retire annually) and of the four 
oflicers ex officio, im all thirteen members. The former are elected annually 
by the Council in June, and serve till the succeeding Anniversary. The 
Committee meet at 4 p.m., at intervals during the Session. The Members for 
1898-99, in a:ldition to the officers, are :— 


lu. G. Baker, Esq. A. D. Michael, F.Z.S. 
©. B. Clarke, M.A., F.R.S8, H. W. Monckton, F.G.S. 
Prof. J. B. Farmer, M.A. Howard Saunders, F.Z.S. 


Prof, J. Reynolds Green,D:Sc.,F.R.S.} Roland Trimen, F.R.S. 
W’. B. Hemsley, F.R.S. 


Notn.—The Charter and Bye-Laws of the Society, as amended to 
the 19th March, 1891, may be had on application, 


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NOTICE. 


gues 
Journal of Zoology. 


Volume SAVII. commenced with Part 173. 


The Journal (both Zoological and Botanical) is 
mow issued in THREE PARTS PER ANNUM as 
follows :— 


Pant I., containing papers read from November to the middle 
ef January, on April Ist. 


Part ITI., containing papers read from the middle of January 
to the end of April, on July ist. 


Part III., containing papers read in May and June, oa 
Wovember ist. _ 


A GENERAL INDEX to the first twenty Volumes of the 
Journal (Zoology) may be had on application, either in eleth or 
in sheets for binding. 


A new CATALOGUE of the LIBRARY may be had on applica- 
tion. Price to Fellows, 5s.; to the Public, 10s. 


Apain 24. Price 12s. 


THE JOURNAL 694, 


OF 


THE LINNEAN SOCIETY. 


Vou. XXVIII. ZOOLOGY. No. 178. 


CONTENTS. 
Page 
I. On the Hyobranchial Skeleton and Larynx of the 
new Aglossal Toad, Hymenochirus Boettgeri. By 
W. G. Ripewoop, D.Sec., F.L.S., Lecturer on 
Biology at St. Mary’s Hospital Medical School, 
Pondone.. (Blate: Si ie ce. cies see ee ee 454, 
II. Note on the Carpus of the new Aglossal Toad, 
Hymenochirus Boettgerit. By W.G. RipEwoop, 
DSc Wels (Plate ol, fig. 5!) sa ae eee 460 
III. On the Eye-spot and Flagellum in Euglena viridis. 
By Harozp Wacer, F.L.8. (Plate 32.)........ 463 
IV. On some New Zealand Schizopoda. By Gro. M. 
Tomson, F.L.8. (Plates 33 & 34.)............ 482 
Y. On the Structure of Porites, with Preliminary Notes 
on the Soft Parts. By Henry M. Brrnarp, 
M.A. Cantab., F.L.S. (Plate 35.) ............ 487 
VI. The Air-bladder and its Connection with the Auditory 
Organ in Notopterus borneensis. By Prof. T. W. 
Brings, Se.D., F.L.8., Mason University College, 
Birmingham. (Plates 36 &37.) .............. 503 
VII. On the Presence of Nasal Secretory Sacs and a 
Naso-pharyngeal Communication in Teleostei, with 
especial reference to Cynoglossus semilevis, Gthr. 
By H. M. Kyuse, M.A. (Communicated by Prof. 
G. B. Howss, Sec. Linn. Soc.) (Plate 38.) .... 541 


Index, Titlepage, and Contents, 


See Notice on last page of Wrapper. 


LONDON: 
SOLD AT THE SOCIETY’S APARTMENTS, BURLINGTON HOUSE. , 
PICCADILLY, W., ; 
AND BY 
LONGMANS, GREEN, AND CO., 
AND \ 
WILLIAMS AND NORGATE. 


1900. 


LINNEAN SOCIETY OF LONDON. 


LIST OF THE OFFICERS AND COUNCIL. 
Elected 24th May, 1899. 


= PRESIDENT. 
Albert C. L. 4. Ginther, M.A., M.D., F.B.S. 


VICE-PRESIDENTS. 


©. B. Clarke, M.A., F.R.S. A. D. Michael, ¥.Z.S., F.R.M.S. 
Wrank Crisp, LL.B., B.A. G. R. M. Murray, F.B.S. 
TREASURER. 


Frank Orisp, LL.B., B.A. 


SECRETARIES. 


5. Daydon Jackson, Esq. | Prof. G. B. Howes, LL.D., F.R.S. 
COUNCIL. 

C. B. Clarke, M.A., F.R.S. Prof. G. B. Howes, LL.D., F.R.S. 
Frank Crisp, LL.B., B.A. B. Daydon Jackson, Esq. 

Francis Darwin, M.B., F.R.S. A. D. Michael, F.Z.S., F.R.M.S. 
Prof. J. B. Farmer, M.A. H. W. Monckton, F.G.S8. 

F. D. Godman, F.R.S, G. R. M. Murray, F.RB.S, 

Henry Groves, Esq. A. B. Rendle, M.A., D.Sc. 


A.C. L. G. Giinther, M.A., M.D., F.R.8.| Howard Saunders. E.Z8. « 
W. Percy Sladen, F.G.S. 


ASSISTANT SECRETARY. 
James Edmund Harting, F.Z.S. 


LIBRARIAN. CLERK. 
A. W. Kappel. A. R. Hammond. 


LIBRARY COMMITTEE. 


This consists of nine Fellows (three of whom retire annually) and of the four 
officers ew officio, in all thirteen members. The former are elected annually 
by the Council in June, and serve till the succeeding Anniversary. he 


Committee meet at 4 p.m., at intervals during the Session. The Members for 
1898-99, in addition to the officers, are :-— 


i. G. Baker, Hsq. A. D. Michael, F.Z.8. 
O. B. Clarke, M.A., F.R.S. H. W. Monckton, F.G.S. 
Prof. J. B. Farmer, M.A. Howard Saunders, F.Z.S. 


Prof. J. Reynolds Green,D.Sc.,F.R.S.| Roland Trimen, F.R.S. 
W. B. Hemsley, F.R.S. 


Notr.—The Charter and Bye-Laws of the Society, as amended to 
the 19th March, 1891, may be had on application. 


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i 


IMPORTANT NOTICE. 


m= OSS 
- Journal of Zoology. 


Volume XXVIII. commenced with Part 173. 


The Journal (both Zoological and Botanical) is 
now issued in THREE PARTS PER ANNUM as 
follows :— 


Parr I., containing papers read from November to tye middle 
of January, on April Ist. 


Parr II., containing papers read from the middle of January 
to the end of April, on July Ist. 


Parr III., containing papers read in May and June, on 
November Ist. 


A GENERAL INDEX to the first twenty Volumes of the 
Journal (Zoology) may be had on application, either in cloth or 
in sheets for binding. Price to Fellows, 15s, ; to the Public, 20s. 


A new CATALOGUE of the LIBRARY may be had on applica- 
tion. Price to Fellows, 5s.; to the Public, 10s 


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